The Pan-Pacific Entomologist

EDITORIAL BOARD

P. H. Arnaud, Jr., Editor

R. S. Lane, Associate Editor

D. F. Gross, Editorial Assistant

W. J. Pulawski, Treasurer J. A. Chemsak

R. M. Bohart J. A. Powell J. E. Hafernik, Jr.

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be addressed to Editor, Pacific Coast Entomological Society, California Academy

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Pacific Coast Entomological Society

OFFICERS FOR 1983

E. L. Smith, President W. J. Pulawski, Treasurer

H. I. Scudder, President-Elect V. F. Lee, Secretary

Statement of Ownership

Title of Publication: The Pan-Pacific Entomologist.

Location of Office of Publication, Business Office of Publisher and Owner: Pacific Coast Entomological

Society, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118.

Editor: P. H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San Francisco 94118

Managing Editor and Known Bondholders or other Security Holders: None.

This issue mailed Sept. 28, 1983

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, U.S.A.

Published by the

Pacific Coast Entomological Society

in cooperation with

The California Academy of Sciences

VOLUME FIFTY-NINE

1983

EDITORIAL BOARD

P. H. ARNAUD, JR., Editor

R. S. LANE, Associate Editor

D. F. GROSS, Editorial Assistant

R. M. BOHART

J. A. CHEMSAK

J. E. HAFERNIK, JR.

J. A. POWELL

W. J. PULAWSKI, Treasurer

A. S. MENKLE and E. E. GRISSELL

Co-editors of this volume

PUBLICATION COMMITTEE

1983

1984

1985

1986

K. W. Cooper

R. M. Bohart

R. S. Lane

H. V. Daly, Chairman

J. E. Hafemik, Jr.

J. G. Edwards

M. S. Wasbauer

R. W. Thorp

San Francisco, California

1983

CONTENTS FOR VOLUME 59

PREFACE._. 3

A BIOGRAPHY OF AN ENTOMOLOGIST. 4

BECHTEL, R. C.-See RUST, R. W. 246

BOHART, G. E. —R. M. Bohart—a brother’s perspective___ 4

BOHART, M. E. R.—R. M. Bohart—a wife’s perspective. 6

DAILEY, D. C., and C. M. SPRENGER—Gall-inducing cynipid wasps from

Quercus dunnii Kellogg (Hymenoptera)....._... 42

EIGHME, L. E Diodontusboharti, a new species from California’s north Coast

Range (Hymenoptera: Sphecidae)__ 50

EVANS, H. E.—The larva of Ammatomus icarioides (Turner) (Hymenoptera:

Sphecidae, Nyssoninae). 52

FERGUSON, G. R.—Two new species in the genus Philanthus and a key to

the politics group (Hymenoptera: Philanthidae)...._. 55

GRIGARICK, A. A., R. O. SCHUSTER, and D. R. NELSON-Heterotardi-

grada of Venezuela (Tardigrada). 64

GRISSELL, E. E.-See MENKE, A. S____ 3

GRISSELL, E. E.-R. M. Bohart: Recollections 1964-1973 . 16

GRISSELL, E. E. — Bohart tens, n. gen., with a review of Rhopalicus Foerster

and Dinotiscus Ghesquiere (Hymenoptera: Pteromalidae). 78

GRISSELL, E. E., and A. S. MENKE—R. M. Bohart—an overview... 22

GRISWOLD, T. L. —A new species of Bembix from Lower California (Hy¬

menoptera: Sphecidae).. 103

HORNING, D. S., JR., and R. O. SCHUSTER-Three new species of New

Zealand tardigrades (Tardigrada: Echiniscidae) .. 108

IRWIN, M. E.—The boharti species group of the genus Pherocera (Diptera:

Therevidae: Phycinae).. 113

KIMSEY, L. S.—The last grad-student years, 1973-1983 . 20

KIMSEY, L. S.—Review of the cuchroeine chrysidids (Hymenoptera: Chrysidi-

dae) ............ 140

MARSH, P. M . —Boharti ellus, a new genus of Doryctinae from South America

(Hymenoptera: Braconidae) .. 148

MATTHEWS, R. W. —Biology of a new Trvpoxylon that utilizes nests of Mi-

crostigmus in Costa Rica (Hymenoptera: Sphecidae).... 152

MENKE, A. S.-Memories of “Doc”- 1956 to 1967 . 10

MENKE, A. S.-See GRISSELL, E. E.... 22

MENKE, A. S. and E. E. GRISSELL-Preface...... 3

MENKE, A. S. and D. L. VINCENT—A review of the genus Polemistus in the

New World (Hymenoptera: Sphecidae). 163

MICHENER, C. D. —The classification of the Lithurginae (Hymenoptera: Mega-

chilidae).. 17 6

MILLER, D. R.—Key to North and Central American species of the mealybug

genus Hypogeococcus (Homoptera: Coccoidea: Pseudococcidae) with descrip¬

tions of four new species and redescription of the type species. 188

MOORE, C. G. —Habitat differences among container-breeding mosquitoes in

western Puerto Rico (Diptera: Culicidae).. 218

NELSON, D. R.-See GRIGARICK, A. A___ 64

PARKER, F. D. —A new Perdita from Utah’s San Rafael Desert (Hymenoptera:

Andrenidae). 229

POWELL, J. A. —Expanding geographical and ecological range of Platynota

stultana in California (Lepidoptera: Tortricidae)..... 233

PULAWSKI, W. J. —Identification and synonymies of two western Palearctic

Cerceris: maculata Radoszkowski and hathor n. sp. (Hymenoptera: Spheci¬

dae) _ 240

RUST, R. W., and R. C. BECHTEL—A fluorescing insect, Cysteodemus ar-

ruatus LeConte (Coleoptera: Meloidae). 246

SCHLINGER, E. I. —Some remembrances of the early U. C. Davis days, 1946—

1956

SCHLINGER, E. L—A new spider parasitoid, Ocnaea boharti, from Arizona

and New Mexico (Diptera: Acroceridae)___ 249

SCHUSTER, R. O. —R. M. Bohart’s impact on the U. C. Davis insect collec¬

tion ___ 21

SCHUSTER. R. O.-See GRIGARICK, A. A__ 64

SCHUSTER, R. O.-See HORNING, D. S., )R.. 108

SCHUSTER, R. O. —A new species of Macrobiotus from Tierra del Fuego

(Tardigrada: Macrobiotidae).. 254

SMITH, N. J.—New North American Pulverro Pate with a key to the species

(Hymenoptera: Sphecidae). 256

SNELLING, R. R.—Taxonomic and nomenclatorial studies on American polis-

tine wasps (Hymenoptera: Vespidae)_ 267

SPRENGER, C. M.-See DAILEY, D. C. .. 42

STANCE, L. A.—A synopsis of the genus Epanthidium Moure with the de¬

scription of a new species from northeastern Mexico (Hymenoptera: Megachil-

idae)_ 281

VINCENT, D. L.-See MENKE, A. S... 163

WASBAUER, M. S.—A redefinition of the Ageniella partita group with de¬

scriptions of two new species (Hymenoptera: Pompilidae). 298

INDEX TO NEW TAX A IN VOLUME 59 . 302

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 1-41

IN HONOR OF

RICHARD M. BOHART

ON THE OCCASION OF HIS 70TH BIRTHDAY

Dick Bohart has been a member of the Pacific Coast Entomological Society for

nearly half a century, and the Society takes great pleasure in dedicating this issue

to him on his 70th birthday, September 28, 1983.

VOLUME 59, NUMBERS 1-4

Preface

This special issue of the Pan Pacific Entomologist had its inception about two

years ago when we realized that Dick Bohart was approaching his 70th birthday.

In view of his outstanding career in entomology both as scientist and teacher, it

seemed appropriate to commemorate this event by dedicat ing an issue of the jour¬

nal to him, an issue filled with papers by his students, colleagues and friends. All

of us wish Dick well on his special natal anniversary and hope he will have many

more productive years with his beloved wasps.

The biography that begins this issue is not the usual type that accompanies

“Festschrifts." It is a multi-author concoction filled with anecdotes, plaudits, hu¬

mor and history. We have invited friends and relatives to write about their re¬

membrances of Dick, asking each one to cover a certain period in his life. A

summary of his impact in the scientific community, his qualities, accomplishments,

and a bibliography of Dick’s publications conclude the biography.

We have had help from a number of people in putting this issue together. Dave

Smith gave us insight into the world of copy editing. The following helped us with

biographical and bibliographical material: Lionel Stange, George “Ned" Bohart,

Roy Snelling, Frank Parker, Bob Bechtel, Patti Paige Adams, Margaret Bohart,

Bob Schuster, Bob Washino, Lynn and Bob Kimsey, Norm Smith and Ev Schling-

er. To all these people we extend our thanks.

Arnold S. Menke and E. Eric Grissell

Coeditors for this issue

Systematic Entomology Laboratory, USD A, ARS

% National Museum of Natural History

Washington, DC 20560

PAN-PACIFIC ENTOMOLOGIST

A Biography of an Entomologist

R. M. Bohart—A Brother’s Perspective

G. E. Bohart 1

Dick’s artistic talents are among my earliest recollections. I suppose he was 8

and I was 5 when I went from door to door in Washington, D.C. selling little

dishes he made from clay we dug from the street in front of our house. I recall

them as being handsomely decorated with clover leaf designs made from different

colors of clay. He retained this interest for many years. In his high school and

early college days, he made clay models of his own concepts of futuristic autos,

of animals in San Francisco’s Fleischacker Zoo, of my head, and of female figures

discovered in magazines. The many items were made from clays we dug out of

the Berkeley hills and, although they were never fired, some of them still remain

in his home as objets d’art.

His budding scientific interests were manifested early by experiments with quick

cooling of red hot clinkers taken from the furnace and dropped into the toilet. I

can still see our cat with her kittens swimming around in the flooded basement

that resulted from this experiment. Another endeavor involved anaesthetizing the

same cat with illuminating gas. Fortunately, she survived both ordeals.

Besides his talents as a sculptor and scientific experimenter, he was a candy

maker of no mean ability. On one occasion during his teen years he made a huge

batch of fudge on the “QT.” Suddenly, he spied mother coming home early. The

pan full of fudge went onto a pile of furniture stored behind a curtain in our

basement butterfly collection workshop. Since the candy was snatched off the

stove too soon, it never completely hardened and, weeks later, after the pan had

been accidentally jostled, mother spied a glacial tongue of fudge creeping out from

behind the curtain. The furniture was kind of sticky but mighty sweet and tasty.

Dick was indeed a “man for all seasons.” During his high school years, he and

I used to fire a football at each other’s heads at close range. This led to throwing

it for distance on a course we measured out on the street. Finally, it led to his

competing in the annual football team kick and throw contest. It must have been

embarrassing for the team and its coaches when Dick, as a sophomore in street

clothes, won the throwing event against the team’s star players. His name was

engraved on the silver trophy cup, but when he won the event the second year in

a row, someone dropped the cup and broke it. That was the end of this contest

at U.C. You can draw your own conclusions from this story.

Early on, Dick developed a keen appreciation of the female form. I believe it

was in his early teens that issues of “Paris Nights” and similar magazines began

turning up in various hiding places around the house. At first I was very shocked

and felt sure that he was destined for hell fire and damnation. However, before

very long, I, too, began to appreciate the attractions that lay between the maga-

1 Bee Biology and Systematics Laboratory, ARS, USDA, Utah State University, UMC 53, Logan,

Utah 84322.

VOLUME 59, NUMBERS 1-4

zine’s covers. His interest was not restricted to graphic representations. For ex¬

ample, I recall two young ladies up the block whose bedroom window faced ours.

Since they sometimes failed to pull down the shade, Dick began to make some

careful studies which resulted in his rather sexist nicknames for the girls, PPB,

for pretty pink bottom and PBB, for pretty big bottom. I cannot affirm or discredit

the accuracy of his designations since I could never get my hands on the field

glasses during critical observation periods. However, his subsequently demon¬

strated ability to discern minute distinguishing differences between insects leads

me to believe that he did not let his excitement interfere with the accuracy of his

observations.

Paleontology was another of Dick’s youthful interests. For example, when, as

a college freshman, he got a job as caretaker of a duck hunting club on Point

Reyes, he found dozens of Miocene marine fossils along a lonely stretch of beach.

He brought back numerous cetacean ear bones and several skulls of a dolphin

subsequently described as Echinorhynchodelphus pontereyensis by a professor in

the Paleontology Department. A monstrous whale skull imbedded in the beach

had to remain there. Rowing back to the clubhouse with his trophies, he spotted

some pine cones protruding from a sea cliff. Dr. Mason, the paleobotanist at U.C.,

was very excited with this find since it helped him develop a revised concept of

pine speciation and distribution in coastal California during the Pleistocene.

“Rock hounding” was a hobby we both carried on with our father who was a

research chemist, but more significantly to him, a frustrated would-be mineral¬

ogist. In 1926 we drove our old Hupmobile through Oregon and Washington

collecting agates from every gravel service station driveway encountered. We also

scoured such favorite rock hound haunts as Agate Beach and the Medford lava

beds where plenty of “good stuff’ could still be found. In later years, when Dad

put together a lapidary outfit, Dick, with his ready eye for symmetry, gave the

initial shape to most of our “gems.” I smoothed out the little ridges and flats, and

Dad, whose eyesight was not the best, did the polishing. I still have some polished

stones that Dick found in such far away places as our junior high school playground

in Berkeley and the dry bed of Putah Creek near Davis.

I should say something about Dick’s early experiences in entomology. Although

I claim to be the first of us to start collecting butterflies (at age 7), he soon “took

over” as big brother. We developed a great proficiency at picking “silverspots”

and swallowtails from butterfly bushes ( Buddleia ) with our hands without dam¬

aging the scale patterns. After moving to California, Dick vowed that we would

collect every kind of butterfly in the State. We had to sell a number of rare

“aberrations” to a wealthy amateur lepidopterist named Gunder before we could

finance trips to southern California to collect such rarities as the Sonora Blue and

the Pima Orangetip.

Dick’s final transition from butterfly collecting to a more scientific endeavor

took place while he was making an insect collection to satisfy requirements for

the U.C. field course in Entomology. He found a bee ( Andrena ) with an odd

triangular plate protruding from its abdomen. He pulled out a sack-like parasite

that soon produced a state of confusion in the Entomology Department. Dr. W.

B. Herms sent Dick to the Zoology Department where Dr. Sol F. Light pronounced

that it was not a helminth. Professor E. O. Essig recognized it as an insect and

suggested showing it to Dr. E. C. VanDyke, a noted coleopterist. He recognized

PAN-PACIFIC ENTOMOLOGIST

it at once as a female Sty lops (order Strepsiptera) and excited Dick’s interest in

the group to the point that he continued studying Strepsiptera and eventually

made them the subject of his Ph.D. thesis.

R. M. Bohart—A Wife’s Perspective

Margaret E. Russell Bohart 2

I have been asked to make a few comments about our early life together. I am

stunned to realize we have been going together fifty-one years. We’ve kept so busy

I guess we haven’t noticed the passing of time.

Dick and I met at Cal-Berkeley the fall of 1932, and I was soon following him

across the Berkeley hills and Antioch sand dunes as he pursued his interest of the

moment, butterflies and Strepsiptera. I have to admit that I was rather embarrassed

to be seen with this big guy swinging a butterfly net, but I outgrew that, and in

the end I have drawn my own snickers from young boys, as I, that gray-haired

lady under a pink flowered hat, swept up the wasps in an Argentine wash.

We were married Jan. 4, 1939 and settled in West Los Angeles where Dick was

teaching and doing research at UCLA. For me the fun came from the collecting

trips, and I was lucky enough to go on almost all of them. From Los Angeles we

had weekend camping trips to the Mojave and Borrego deserts, and sometimes

longer ones to the High Sierra, and the Grand Canyon of Arizona. After a couple

of years I tried my hand at collecting, as there was a limit to how much needl ework

I could do. I missed a lot of “bugs” in those days, and 1 never learned to pinch

the thorax. I have my own technique.

One of Dick’s research projects at UCLA was the sod webworm that was

destroying lawns. At one point we went out every night, all night, every hour on

the hour, to a private lawn to lie prone on the grass and, with the glow of a

flashlight, watch the worms snip off tender shoots and haul them back into their

homes. We also poured a solution on one square foot of sod, and then counted

the worms that rose to the surface. Great way to catch a cold.

In the early spring of 1942 our Los Angeles apartment burned. While it was

being redecorated we took a trip to Yosemite. We hauled his papers and typewriter

along. I’m the typist. One evening we were walking across a little bridge and a

mosquito got in Dick’s way. He examined it and then dashed it down, saying that

was one thing he was never going to work on. Fate interfered.

November 1942 Dick was drafted into the Army, but with some agile footwork

he got himself into the Navy as an Ensign, and by Dec. 5 we were at the Marine

base, Camp Lejeune, Jacksonville, North Carolina. He was to teach malaria and

mosquito control. A few months later he was transferred to Camp Perry, Wil¬

liamsburg, Virginia to do the same. Two thousand in a class, a new class every

five weeks. That’s where he got that voice heard a mile away. We didn’t have a

car so our collecting trips were small walks through the lovely woods. Of course

2 230 Shepherd’s Lane, Davis, California 95616.

VOLUME 59, NUMBERS 1-4

chiggers, sudden rain storms and humidity were something we had never had to

reckon with before. We enjoyed Williamsburg and all its history.

In the early spring of 1944, Dick was transferred to Washington, DC and

assigned to Naval Medical Research Unit #2 housed at the National Academy of

Sciences. Almost immediately he and his brother, Ned, and Ken Knight were

sent to Orlando and Tallahassee, Florida, where Dick studied mosquitoes under

Dr. Boyd. We stayed a month and then returned to Washington. Thus mosquito

taxonomy began in earnest and Dick was in his element. We spent our free time

at the Mellon Art Gallery and the Smithsonian. Couldn’t ask for anything more.

At this point he tried to make a taxonomist out of me, studying the bee genus

Nomadopsis, but it didn’t take. Ned, Ken Knight, Alan Stone, Karl Krombein

and other taxonomists were in DC as well as some of Dick’s cousins, so we enjoyed

the stay very much.

It was time to go overseas and in the fall of 1944 Dick was assigned to Guam

and then Okinawa, for malaria control, and I returned home to Fullerton, Calif.

Dick took with him all the equipment and photocopies of the literature necessary

to carry on the study of mosquitoes in those islands. Stan Bailey was working

with him, and they had good luck in cleaning up the mosquito problems of Guam.

They moved on to Okinawa a little too soon. The Marines hadn’t quite finished

taking it from the Japanese and Dick landed amid some shelling. His seabag came

up missing and he figured it was tossed overboard, because the outline of his

pistol was visible, and someone wanted the gun worse than he did. He lost all of

his equipment, literature and clothes, but fortunately nothing more serious oc¬

curred. Despite occasional pot shots from the Japanese, who were still holed up

in the hills, he seemed to really enjoy Okinawa, its beautiful beaches, fine swim¬

ming, and of course the insect collecting. Funny things happened, too. One of the

responsibilities was to eliminate mosquito larvae by covering any standing water

with xylene or a similar chemical. This assignment included outhouses. Yep,

someone dropped a lighted match and blew the place up, but luckily only his

dignity was bruised.

Dick could have been released from the Navy in the fall of 1945, but he agreed

to stay on a few more months to get his research written up, thus we returned to

Washington. We worked at the Academy every night. I did lots of typing of

literature, as Dick was collecting material for other projects. Photocopying was

still somebody’s dream. Somewhere in these years Dick wrote the “Mosquitoes

of the Philippines” (published in 1945), from specimens sent from the field.

The project came to an end, and Dick was released from the Navy in the spring

of 1946 with the rank of Lt. Commander. We returned to California where his

position at the University of California was waiting for him. He transferred from

the UCLA campus to the Davis campus, arriving there July 1946.

A new branch of life started that came to involve many students, particularly

graduate students with whom we came to have a close, personal contact. With

retirement I think we miss this the most. The research and the trips to the Hawaiian

and other Pacific Islands, Japan, British Isles, Western Europe, East Africa, Natal,

South Africa, Argentina, Brazil, Costa Rica, Venezuela, Mexico, Australia, Canada

and the United States, especially the beloved southwest, were a part of these years,

too. And the gleam in the eye for another trip has not disappeared.

Just so the modern girls won’t think I had no life of my own, I will tell them

PAN-PACIFIC ENTOMOLOGIST

Richard and Margaret Bohart at their home in Davis, July 1975. Photograph by Woj Pulawski.

I was a social worker, and had [and still has—edit.] an all absorbing hobby,

genealogy. We also entertained a great deal, played a lot of bridge, and raised

Pepe Le Moko, Annabel and Miss Bo, who would not like being left out. Those

three kittens were with us a total of thirty-one years.

I think the greatest thing has been that Dick was lucky enough to find an

occupation in this life that he has thoroughly enjoyed.

Some Remembrances of the Early U.C. Davis

Days, 1946-1956 3

Ev Schlinger 4

Although I didn’t know it, when I arrived at U.C. Davis in February 1946 as

a 17-year-old budding entomologist, I arrived a full four months before my future

major professor. Assistant Professor Richard M. Bohart. This was a time, near

the end of World War II, when U.C. Davis had about 100 students, only one of

whom was a woman. Needless to say, she was well escorted!

3 Thanks go to Bob Bechtel for supplying some of the memories recorded here.

4 Department of Entomology, University of California, Berkeley, California 94720.

VOLUME 59, NUMBERS 1-4

The first “post-war” Introduction to Entomology class (Ent. 1), a 5-unit semester

course, was to be taught in the spring of 1946 by Drs. W. Harry Lange, Jr., and

the late E. M. Stafford. There was a total of seven students in the class with two

of us majoring in Entomology and two professors to teach us. What a student-

teacher ratio.

In the spring of 1947, as I remember it, when “Doc” was asked to teach this

Ent. 1 course, he asked several of us “older” students to go along on a field trip

to the Sierras with his class to help collect insects to start to build an insect

teaching collection. I clearly remember that day when he warned all of us not to

drink the clear mountain water as it might be unsafe. It was a hot day, no large

animals were evident, and we just had one quick drink of the cool mountain

water—only to come down with dysentery the next day. I have never told Doc

about the dysentery, but I learned to take his words more seriously and eventually

became one of his first graduate students.

When I entered graduate school in the fall of 1950, there were six graduate

students, all of whom were under the late Dr. Stan F. Bailey, head of the de¬

partment. I well remember how Dr. Bailey insisted that each of us come to see

him at a specified time each Saturday morning to discuss our research with him.

He wouldn’t discuss research with us during the week, and needless to say, we

would much rather have been allowed to go collecting, or . . . Bailey insisted on

being responsible for all taxonomy grad students, but about six months after I

entered grad school Doc was permitted to have his first students. That was when

R. C. “Bob” Bechtel, A. D. “Biff” Telford, and I chose to work with him. In those

days, the graduate students had offices in the “quonset” building next to the

Entomology building. This was an army surplus, metal, pre-fab structure, which,

although freezing in the winter, kept us and our insects dry.

At that time, Doc was working on at least three separate projects; i.e., (1)

Taxonomy of Culicidae, (2) Taxonomy of Vespidae, and (3) the California Cherry

Tree’s insects and their control. He was also very much involved with working

up and collecting sphecid wasps. He took the three of us in stride even though

all of us had chosen thesis topics quite different than his; i.e., Bechtel had chosen

to work on Plastoceridae, Telford had chosen to study diapause phenomena in

marsh mosquitoes, and I had chosen the biology of spiders and their parasitoids.

Doc tried hard to turn Bechtel and me into “Hymenopterists,” and by allowing

us to help prepare and publish papers with him (Bechtel on California social

Vespidae and myself on California Oxybelus wasps), he taught us considerable

about taxonomic methodologies and Hymenoptera. Bechtel went on to study the

family Sapygidae, Telford eventually took on the chrysidid genus Parnopes, but

I finally settled on a taxonomic revision of the Acroceridae (Diptera). Jack Hall

and Bob Washino eventually joined Doc’s group as master’s candidates working

on bombyliid flies and mosquitoes, respectively.

The years 1950-1956 were exciting times for us, and other than hard day-to-

day laboratory revisionary work and field sampling, I think the events I remember

most were the special field collecting trips. Locally, in those days, Putah Creek

was close by and contained between its banks some most interesting insects. Doc

was able to join these trips frequently and even more often in the higher Putah

Canyon area, especially around Monticello near Pope Valley, with Samuel Spring

the favorite site. This latter locality, now covered by waters of the government’s

PAN-PACIFIC ENTOMOLOGIST

massive project called Lake Berryessa, became the type locality of a number of

his new species.

Other events remembered well were Doc and his competitive spirit on our

Departmental Softball Team which, by the way, we won most of the time, and

our “horseshoe” games after work just outside his office in the “old” one-story

Entomology building. Badminton was also one of Doc’s favorite pastimes and he

was instrumental in the establishment of the “Davis Badminton Club,” of which

he became president.

Those were wonderful years for all of us, and I am pleased to have been lucky

enough to have had the guidance and tutorship of one great guy and excellent

entomology professor, Dick Bohart.

Memories of “Doc”—1956 to 1967

Arnold S. Menke 5

My first exposure to “Doc” was on the entomology summer field course, Ent.

49, in 1956 at Tanbark Flat in the San Gabriel Mountains of southern California.

It was the second time he had been in charge, having first led the course in 1954.

Margaret accompanied him as did Bob Bechtel, the latter serving as his assistant.

Some of the students who took the course that summer were Lionel Stange, Mel

Sparks, Jerry Stage, Louie Shainberg, “Hal” Moffitt, Dick Bushing, “Hank” Mi-

chalk, Art Bartels, and Beverly Bartosh. I remember well the noxious rhagionid

biting flies at Tanbark Flat which made collecting around camp miserable at

times. Breakfast for Doc always seemed to be “Special K” and lunches often were

the infamous peanut butter and jam sandwiches. I recall that Doc had a large

disdain for milk claiming that it was only for kids. He used to tell us that it made

adults prone to colds and other problems. Instead Doc drank Pepsi or buttermilk,

the last being something of a contradiction. We could never understand how he

could drink that stuff.

Doc impressed all of us as a tireless collector, and he always put in a full day

of collecting. By his example and instruction we learned the many special tech¬

niques necessary to collect various kinds of insects. He was always stressing that

we should not just collect big showy beetles, wasps and butterflies, and he expected

everyone to catch small insects that required point mounting. During the evening

while we were pinning up our material, he would come around to see what we

had caught, and I recall his frequent question: “where are all the small Diptera?”

Doc alerted us to the tiny sphecids and other wasps that were found on the ground

and which required the “clap” technique to capture. He also drew our attention

to the wondrous nannofauna that frequented mat Euphorbia and other prostrate

plants that most of us would otherwise have ignored completely as potential

sources for insects. In his own way he was always stressing observation. I remem-

5 Systematic Entomology Laboratory, IIBIII, ARS, USDA, % U.S. National Museum of Natural

History, Washington, D.C. 20560.

VOLUME 59, NUMBERS 1-4

ber when he broke off a plant stem that had blooming dodder entwined around

it, showed it to some students, and asked if they knew what kind of plant it was.

Of course he was hoping to trick them into thinking that the dodder was part of

the plant stem, but at the same time he was pointing out that there was always

more to things than met the eye at first glance. He also took the opportunity to

emphasize that dodder blossoms attracted some interesting insects. Doc always

seemed to be able to collect more of anything than anyone else. I remember

catching a couple of rhipiphorids on an Enogonum plant, the first of these beetles

I had ever seen alive. Naturally I took my prizes over to Doc to show them off.

To my chagrin and amazement he produced a vial containing a dozen or more

rhipiphorids, telling me with a rather straight face that “they were revolving slowly

in a mass on a single Eriogonum blossom.” That was his sly style of humour.

Lionel Stange, my pseudo-brother and collecting chum, possessed an old 1947

Studebaker which he and I used during the course for weekend forays to collect

insects at distant favorite collecting haunts like Blythe on the Colorado River. I

think that Doc secretly enjoyed our initiative, but I know that we also caused him

some concern and problems. We always brought back stuff that no one else in

the class had seen and that engendered some jealousy among fellow students. Also

Lionel’s car was approaching a basket case. In fact it did die one night when

Lionel and I were driving down to Glendora. On a hairpin turn on that long

winding mountain road the right front axle snapped bringing us to a grinding halt

short of the dropoff. Doc was not too happy when we arrived back at Tanbark

Flat late that night courtesy of the Glendora police.

Doc had some very definite ideas about how insects should be mounted and

labeled. He always used #2 pins, and we marveled at his disdain for point punches.

He would cut his points from strips of paper with scissors. Doc insisted that insect

labels not exceed 12 mm in length, nor be more than 5 printed lines. During the

course Lionel Stange and I ran the label printing press instead of washing dishes.

My next encounter with Doc was at his invitation. In the Spring of 1957 when

I was finishing up my senior year at Berkeley and contemplating grad school, he

wrote to me asking if I would be interested in being his Research Assistant at

Davis to help him with teaching general entomology and systematic entomology.

To quote from his letter, “each of these jobs . . . [offers] . . . the finest possible

experience for an entomologist.” I had visited the Davis campus several times

and liked the country atmosphere and small student body (about 2000) far better

than the Berkeley fog and zoo, so I jumped at the opportunity. Thus Doc acquired

a water bug (belostomatid) lover. When I arrived at Davis in September, 1957,

he had just become Department Chairman.

Helping Doc with the labs in these courses proved to be a great experience, and

looking back on those days now, I realize that in the taxonomic course, especially,

he accomplished something that is missed in many such courses: he taught people

practical, everyday, taxonomic procedure, the real world of nomenclature and

how you deal with it, how descriptions and keys are written, how illustrations are

rendered, and how you put it all together in a publishable paper. Truly a rare

experience for those fortunate to have taken his taxonomy course. As he admitted

in his letter to me, “we run 112 [taxonomy] somewhat differently from Berkeley.

The student takes a special problem under close supervision of the lab instructor

and staff member.” Each student actually did a miniature (some not so small!)

PAN-PACIFIC ENTOMOLOGIST

research problem starting with a box of insects, usually representing a genus in a

family of Hymenoptera. The specimens had to be sorted to species and identified

(or declared to be new which was not infrequent), described, illustrated, a key to

them constructed, and the whole thing written up as a publishable paper.

Some students did in fact make valuable contributions and discoveries in Ent.

112 which they subsequently published, or which later blossomed into thesis

problems. For example, Paul Marsh discovered a new species of the sphecid genus

Prionyx which Doc later described and published as subatratus. Frank Parker

discovered excellent mouthpart characters that easily separated the very similar

females of two species of Prionyx. Frank subsequently published his findings which

essentially made Doc’s own recent paper on the genus obsolete. The chrysidid

genus Omalus was revised by Doc and Luciano Campos subsequent to the latter’s

discoveries while working on the genus in Ent. 112. As for myself, lab instructor,

I became enamored with the sphecid genus Ammophila which Dick James was

working with in the course. He discovered a new species which I eventually

described as formicoides, and I eventually revised the genus for my Ph.D. thesis.

Doc’s exam questions in taxonomy were famous. They always included at least

one thorny nomenclatorial question to solve, and he often had a number of obscure

terms to define, some of which bordered on craziness (hypodigm, cheironym,

natio, for example). Some of his questions were so subjective and broad in scope

that they were a bear to grade.

General Entomology lab involved identification of insect orders and common

families, and included a number of afternoon and weekend collecting trips. An

identified collection of a considerable number of orders and families was a pre¬

requisite to passing. Doc always took a shine to the ever present girls in his classes,

and I recall that he tended to be lenient toward them at grading time.

In 1959 I was Doc’s assistant on Ent. 49 which was held at the University of

California’s Santa Barbara campus located at Goleta. Doc also brought along Paul

Marsh who was “’selected” to run the label printing press. Possibly this was the

most popular summer field course ever because of the nearby beaches to which

most of us fled in the afternoons. Sunbathing, body surfing, and swimming made

for great tans, but we discovered early on why they drill for oil offshore. Removing

tar from feet after a trip to the beach became routine. We lived in a fraternity

house, the only time most of us, especially the women, had such an experience.

The men’s john was a single large room with a number of unenclosed toilets along

one wall. Nothing like communal elimination. Among the students taking the

course were Frank Parker, Gene Cherry, Patti Paige, Meredith Bruck, Wally

Steffan, Jim and Jack Bath, Ray Spore, Craig Campbell, and Bob Granados.

Collecting near the beaches was generally very good for wasps such as Ammophila,

and Doc discovered an Oxybelus nesting site adjacent to a lagoon behind the

beach. He assigned the task of watching Oxybelus nesting behavior to Paul Marsh.

I well remember how Paul spent many days sitting in the hot sun watching the

wasps’ activities while the rest of us hit the beaches for aquatic sports. Paul’s

observations were subsequently published in collaboration with Doc and that was

Paul’s first paper. One day Doc borrowed my VW beetle for a collecting trip.

Reports I got from those that accompanied him suggested that I was lucky that

the gear box was still intact after he returned.

Frank Parker and the girls had an interesting experience one night. There had

VOLUME 59, NUMBERS 1-4

been a “private party” in the girls’ room. After the other participants had gone

back to their respective sleeping quarters, a student, who shall remain nameless,

returned to the girls’ room around 2 AM thoroughly inebriated. The girls asked

him to leave and he subsequently fell down the stairs suffering a large, deep gash

on his forehead in the process. He was bleeding profusely and Patti sought Frank

Parker’s help. She found his room in the dark and woke him up. They got him

into Frank’s car, and Frank and the two girls drove to the campus infirmary. The

nurse on duty called the police after seeing the student’s condition. By chance the

police were looking for two guys and two women who had been causing some

disturbance in town. They took everyone to the police station for questioning.

Somehow Meredith and Patti, dressed in nightgowns, eventually convinced the

police to release them. They were taken back to the fraternity house in the wee

hours of the morning just in time to pretend to be getting up in preparation for

that day’s scheduled field trip. When Doc saw the fellow that morning with all

his stitches he was naturally concerned. Doc was told that the student “ran into

something in the dark” and that the wound was not serious. As far as I know,

Doc never knew the true story of that wild night.

The late 50’s and early 60’s saw Doc’s graduate student load mushrooming.

Paul Marsh was working on braconids, Helen Court was working on crabronine

sphecids, Frank Parker was working on astatine sphecids, Byron Chaniotis and

Fred litis were working on mosquitoes, Chet Moore was working on chrysidids,

Lionel Stange was working on ant lions, and I was doing a master’s thesis on a

genus of Belostomatidae. Obviously Doc didn’t insist that his students take on

thesis problems in the area of his first love, the predatory wasps.

Another aspect of his graduate student guidance that is noteworthy was the

rather limited amount of it. He tended to let his students pursue their thesis work

without much inspection of progress, etc., at least after initial discussions of

procedure. This meant that a student without strong inner drive (someone other

than a “born taxonomist”) would probably not survi ve long. This aspect of Doc’s

tutelage developed independence, discipline, and self initiative. The last is perhaps

exemplified by the fact that his students often published papers on various taxo¬

nomic projects “on the side” during their graduate tenure. They were able to do

this in part because Doc only expected his Research Assistants to teach laboratory

in his courses. Rarely did he ask his RA to do anything else. Thus the rest of your

time was free for other pursuits.

Doc commonly took on one of his students as junior author on some taxonomic

research problem. In my case I first worked with him on the sphecid genus

Palinodes, one of the spinoffs of Ent. 112. What an experience! (and not unique

to me): while looking at specimens under the microscope he verbally described

the creature, and I wrote it down to produce the description. That is not to imply

that I did not do some looking at material myself. In fact, in this and all subsequent

projects on which he and I collaborated, I was always impressed by the fact that

he would listen to and often agree with my ideas and viewpoints on various

matters. No one could accuse him of being dogmatic, stubborn maybe, but not

inflexible. An example of his continued growth as a scientist was his change in

attitude on subspecies. He was from the school that made every geographic color

form a different subspecies, but in later years he came to appreciate the clinal

nature common to many of these forms and his formal naming of them steadily

PAN-PACIFIC ENTOMOLOGIST

decreased. This change in philosophy was influenced in part by his students which

emphasizes his willingness to listen to new ideas. His de-emphasis of subgenera

in favor of species groups in later years is another example that comes to mind.

The research for and writing of the book “Sphecid Wasps of the World” by

Dick and me occupied a big chunk of both our lives—nearly 10 years in fact.

Perhaps it is appropriate here to review a little of its history. While I was finishing

up my Ph.D. thesis on Ammophila in 1965, Doc suggested that he and I collaborate

on a revision of the North American genera of Sphecidae for which he would

seek NSF support. Naturally his proposal appealed to me. After some thought I

suggested that from the standpoint of producing a viable classification it probably

would be better to expand our horizons. He agreed, and at first we set our sights

on a generic study of the Sphecidae of the New World. In the initial stages of our

NSF supported study we soon realized that only a worldwide revision would

produce a sound classification for the family, and that became our ultimate goal.

Our game plan was to divide the work between us, each having responsibility for

researching and writing up a number of the subfamilies. Of course we had help

from Doc’s students, past and present (Frank Parker, Dick Brumley, Lionel Stange,

Helen Court, Eric Grissell, among others), and colleagues (Wojciech Pulawski and

Jack van der Vecht in particular), but the main input was ours. Dividing up the

work between us may have resulted in some unevenness in treatment but it was

the only logical way to handle the immense job before us.

Initially Dick assumed that we could finish the project in about three years,

but expansion to a world study made this an impossibility. I don’t think either

of us (or anyone else that has tackled such a project) had any idea of the millstone

we were putting around our necks, nor the time it was going to take to remove

it. I worked on the project at Davis for three years until the NSF grant expired,

at which time I was forced to find employment elsewhere, namely with the Sys¬

tematic Entomology Laboratory in Washington, DC. My added responsibilities

slowed me down some, but Dick plowed forward with his usual determinedness.

I suspect that with my departure he probably worried some about keeping the

whole project going at a steady pace, especially with my propensity for getting

sidetracked on details. As I wrote up various sections I would forward rough drafts

to Dick and he would have them typed up. We kept in touch mostly by frequent

telephone calls, and fortunately I was able to visit Davis each year for periods of

two to four weeks. During these visits he and I accomplished a tremendous amount

of work. On one such trip we jointly worked up the Ampulicinae, and on another

we did most of the introductory chapters for the book. He made one trip to

Washington which gave us another opportunity to work together on the book.

By the spring of 1972 most of the taxonomic sections of the book were finished,

and in May of that year we showed parts of it to the editors of the University of

California Press which we hoped would publish it. They gave us tentative accep¬

tance and we set Sept. 1 as a deadline for ourselves to complete the whole book.

Dick was the instructor on Ent. 109 (formerly Ent. 49) that summer at Sagehen

Creek and he spent much of his time during the course going over completed

sections checking for consistency, errors, and making improvements rather than

collecting. He was otherwise kept plenty busy the rest of the summer finalizing

the subfamily Crabroninae, the last unfinished taxonomic section of the book. He

was also saddled with the job of arranging all the plates of figures and making

VOLUME 59, NUMBERS 1-4 15

Dick and Princess Silver Bow at the Bohart residence, Davis, April 29, 1973. Manuscript for the

Big Blue Book had just been completed! Photograph by Arnold Menke.

sure that figure numbers were entered in the ms. correctly—a horrendous task. I

busied myself with rendering drawings for my subfamily sections, checking species

checklists in the Crabroninae and Nyssoninae, writing up the catalog of generic

PAN-PACIFIC ENTOMOLOGIST

names and their synonyms, solving last minute nomenclatorial problems that

required ICZN petitions, surveying last minute morphological discoveries

subfamily by subfamily, and other odd jobs. Sept. 1 came and went, and by the

end of 1972 the book remained unfinished. A shortage of secretarial help at Davis

was partly responsible and this caused Dick to rent a typewriter and hire a typist

with personal funds in early 1973 in order to get the monster finished. We finally

got the completed manuscript to the Press around May 1, 1973. Publication was

then only three years away! Ahead were months of proof reading by Dick and

Margaret at his end, and by Karen and me 3000 miles away. Generally Karen

and I read the proofs first and then forwarded them to Dick for his turn at them.

Karen Menke also did the horrendous job of typing up all of the more than 14,000

3X5 cards for the taxonomic index, one name to a card. Dick took care of general

indexing, also a chore. All galleys had been proofed by mid 1974, and page proofs

were received in April 1975, reviewed and returned promptly. Later, while Dick

and Margaret were touring Africa in the fall of 1975 (those lucky dogs!) 1 had the

pleasure of reading and correcting the page proofs of the index! At last the mill¬

stone was gone, and our baby arrived between blue covers on April 27, 1976. It

has since become known in sphecidological circles as the Big Blue Book or simply

B 3.

The years spent working with Dick, first as a student and later as a colleague,

provide me with many happy memories. Dick is a remarkable guy and I especially

treasure the friendship that we share.

R. M. Bohart: Recollections 1964-1973

E. E. Grissell 6

My association with Dick Bohart began in the summer of 1964. As a soon-to-

be undergraduate at Davis, I had an invitation to visit the entomology department

and to “get acquainted.” Dick fortunately happened to be chairman at the time,

and he gave me both a warm welcome and a personal tour of the facilities. This

was my first exposure to one of Dick’s superior qualities . . . his love and enthu¬

siasm for insects of all kinds. It is all pervasive, and he has retained this attitude

for the 20 years I have known him. I’m certain it has been strongly felt by students

and colleagues alike from the earliest days of his career.

My second meeting with Dick occurred only a few months later when school

began. In retrospect the meeting was not quite as bad as it seemed at the time,

but it was slightly different than I expected. The enthusiasm was still there to be

certain, but Dick didn’t seem to remember who I was ... he couldn’t remember

my name. This bruised a rather young and fragile ego, but I soon learned a very

simple fact about Dick ... he can’t remember anyone’s name! For many years I

6 Systematic Entomology Laboratory, IIBIII, ARS, USDA, % U.S. National Museum of Natural

Elistory, Washington, D.C. 20560.

VOLUME 59, NUMBERS 1-4

have suspected that the first thing he does in the morning, upon arising, is to

memorize the name on his driver’s license so he might get at least one name

correct during the day. It has been secretly amusing, over the years, to see how

Dick manages to avoid the problem of introductions when the situation arises.

Most of his colleagues are understanding sorts, though, and they often begin a

conversation with their own name just to avoid possible confusion.

I was extremely fortunate, as a financially depauperate undergraduate, to receive

Dick’s early help in the form of a work-study position. Whether this opportunity

was decreed by destiny, divinity, or deviousness I know not. I only know that

help always came at the right time during my association with Dick and that the

experience of working for him was an education, not a job. In looking back, even

as a beginning student, my experiences with Dick have always been of a mutually

beneficial nature: each person gaining in some way from the association. Doc has

had this arrangement with many students, and most erf us have recognized the

benefits we’ve received. The fact is, that many of us have been dragged (albeit

screaming and kicking) through our first publication largely on Dick’s coat tails.

We simply learned by doing. Perhaps more importantly though, some of us have

learned the lesson exemplified by such generosity of time, knowledge, and ex¬

perience. In matters of research and publication, Doc always gave credit where

it was due, and usually gave a little extra for good measure.

During the mid 60's and early 70’s, Dick had a profusion of graduate students

working on taxonomic projects: Steve Buckett was working on noctuids, Mike

Gardner on millipeds, Charles Dailey on cynipids, Don Homing on chrysidids,

and I worked on chalcidoids. An “adopted” student, Dug Miller, worked on

mealybugs. Doc even had several students, David Levin, Dick Brumley, Edmundo

Rubio, and Joanne Slansky Wasbauer, who actually worked on sphecids. It is to

Dick’s credit that he gave each student the freedom to choose his or her own

project. He was equally as willing to suggest any number of potential projects as

well. Doc gave us the opportunity to work at our own pace which encouraged a

necessary amount of self-reliance and responsibility. Conversely, however, Doc’s

door was always open and he would stop his own work at a moment’s notice to

answer questions or give help if he could. Occasionally there would be a flurry of

pointed questions about “progress,” intended, I’m certain, to gently prod some

of the slower members of his entourage. Dick took an interest in each of his

students and tried to accommodate the individual idiosyncrasies which charac¬

terize most taxonomists.

For several years (1968-1971) I worked as Dick’s research and teaching assis¬

tant. This, again, was an education not only in taxonomy, but also in human

behavior. Under his guidance I learned to catalog, collect and identify Hymen-

optera, organize data, and write papers. At least I hope I learned. I also learned

how to translate what Dick “said” into what he actually “wanted.” This was

purely a matter of survival. In the beginning it was my reaction to immediately

do something if Dick asked me to. I soon learned, however, from his previous

assistant, to put off until tomorrow whatever today’s request might be. That way

Dick might have a few days to sort of “rearrange” his thoughts, as it were. And

it seemed as if he were constantly rearranging, much to the irritation of his

taxonomy students. Now that Dick is safely retired I can reveal the secret “code”

by which we assistants all lived. Doc was known to have his way of doing things,

PAN-PACIFIC ENTOMOLOGIST

and there seemed, to the uninitiated, to be a rather stubborn streak to his nature.

Those of us who lived by the code, however, knew that if Doc were “hit-up” with

a new idea during his “rearranging phase,” it often became incorporated into his

thoughts and could be prodded forth several days later as a worthwhile revelation.

It was his idea, of course, but that was actually a fair price to pay for getting things

done our way. Graduate students aren’t entirely stupid, you know!

The term paper in the undergraduate taxonomy course was a good example of

this phenomenon. It was universally acknowledged, even before a student took

the course, that the paper was an evil of unspeakable vileness. Some of us actually

enjoyed the paper, but most did not. As an assistant in the course for 3 years, I

was the intermediary between students and professor. This position was not the

least bit enviable, and the first year was, shall we say, a struggle (“It was hell,”

said a former assistant). While students at Berkeley were burning the library and

protesting every social evil, students in the taxonomy class at Davis were revolting

(to say the least) against writing descriptions and making distribution maps. Need¬

less to say, the assistant took the majority of the heat. By the second year, however,

by invoking “the code,” we managed to persuade Doc to alter the method of

presenting the term paper. This so unbalanced the students that before they could

organize themselves to riot, the course was over. Whereupon Doc and I, to il¬

lustrate one of our finer attributes, immediately took off for the deserts to do some

collecting. The ploy worked so well that we tried it again the third year, and as

far as I know, Doc changed the paper every year thereafter. I was fortunate to

learn the universal doctrine of teaching under Doc’s supervision: always leave the

students off-balance, it is best for everyone involved!

My fondest experiences with Dick are associated with the five-week summer

field course (then Ent. 49) and our numerous collecting trips together. In Ent. 49

Dick charged up a frenzy of entomomania among the undergraduates. This was

due in large part to genuine enthusiasm and a prodigious knowledge of insects.

Mention of this has been made in the previous paper by Arnold Menke, and I

can confirm the effects. Students were out day and night ravaging the countryside

for just “one more family.” One summer, we had a particularly knowledgeable

group of students, several of whom specialized in various micro-faunas. The

number of families in each student’s collection grew dramatically as battles raged

over where to find the smallest Diptera, Coleoptera, Lepidoptera, or Hymenoptera.

I can’t remember who had the biggest collection, but I do know who held the

record for “most countryside terrorized by a single entomologist”: Baldomero

Villegas! With two sweep nets going at once, bandoliers of cyanide vials over each

shoulder, and a gallon killing bottle under each arm, Baldo struck fear even in

the heart of Dick. Doc was forced to restrain the “great destroyer” while the other

students collected first. Then he would let Baldo loose, and there would be little

reason to return to that particular collecting site for several years. I know that

Dick secretely admired such enthusiasm in a student.

Shortly before I first took Ent. 49 in 1966, Dick had changed the course from

solely making an insect collection to including an ecological or biological project.

The project was designed to “encourage” students to look at living insects. Almost

anything would do, and if we didn’t have an idea, Doc would soon supply us with

one. Doc suggested that another student (Steve Clement) and I conduct obser¬

vations on the nesting habits of a little known wasp named Euparagia scutellaris.

VOLUME 59, NUMBERS 1-4

These observations formed the basis for my first publication (in coauthorship

with Clement). When asked if Dick would also like to be a coauthor, he char¬

acteristically declined on the pretext that he had had nothing to do with the paper.

Unlike many professors, Doc didn’t take credit for work he didn’t do.

A few years after taking the course, I had the privilege (at least it seemed so at

the time) of serving as the teaching assistant. I might say that having the respon¬

sibility for 14 students is not quite the same as being one of the students. Having

survived three years as intermediary' between Doc and students in the taxonomy

course had helped a bit, but I was not quite prepared for the food strikes, pranks,

and physical exhaustion which came with the summer students. Doc always took

it in stride, and in spite of suffering from a severe vitamin deficiency, could

outcollect and outlast any of us any day. However I always managed to outdo

Dick in one respect: it had something to do with automobiles. As I vaguely recall,

a single trip with Doc at the wheel taught one a certain respect for one’s life.

Whenever a trip was in the offing, 14 (of 14) students tried jamming themselves

into my station wagon. I still have heel marks in the back of my head to prove it.

Entomology 49 was a rewarding entomological and social experience for many

students ... a chance to “live, breathe, and eat bugs” as Doc would say. It was

undoubtedly the most concentrated dose of real-world entomology that a student

could receive.

Field trips with Doc were a special experience, shall we say, especially as they

required a one to one relationship. They also required a lot of driving, but I try

not to think much about that. Doc introduced me to the California desert regions,

early on. He took me on numerous trips to what appeared to be the most barren

places imaginable. I soon came to appreciate Doc’s ability to find insects in these

spots, however, and I must confess that the deserts are still my favorite domain.

At first it was Doc who had all the expertise in terms of how and where to collect,

but after a few years I think I showed him a thing or two about chalcidoids. With

a little practice, however, Doc was as proficient as I was, and he was again the

master collector. This, I think, is one of Dick’s secrets ... he is always willing to

learn new things about bugs. Collecting trips provided a chance for protracted

conversations, discussions of taxonomy, progress reports on the dissertation, com¬

parisons of collecting techniques, assorted braggings about the day’s catch, and

keeping up with departmental gossip. These trips were a superior educational

experience not to be duplicated in any classroom. Over the years, Dick has sup¬

plemented many a student’s classroom education with his numerous field trips.

Over the years, I’ve grown to appreciate Dick more and more as an entomologist

in the finest sense of the word. There is probably no aspect of entomology which,

if required, he could not master. He has worked upon a diversity of taxonomic

groups and in a diversity of fields from toxicology and economic entomology to

ethology, ecology, and taxonomy. I’ve also grown to realize that Doc always had

the interests of his students at heart, and in this respect something must addi¬

tionally be said about his wife of 45 years, Margaret. In the treatment of students

it is somewhat difficult to know where Doc leaves off and Margaret begins. In

many ways Margaret has been the unofficial overseer of student concerns, and I

know she has contributed much to their welfare. How many of us have enjoyed

the hospitality of the Bohart home over the years, both as students and long

afterwards? I was especially touched by the kindness of Dick and Margaret who

PAN-PACIFIC ENTOMOLOGIST

took me into their home for the last year of my graduate work. Such attitudes

toward students are the rule with Margaret and Dick and that makes them the

special people they are.

The Last Grad-Student Years, 1973-1983

Lynn Kimsey 7

I first became acquainted with “Doc” Bohart as an undergraduate in his Insect

Systematics class in 1973. By this time in his career he had earned a reputation

as a tough instructor and because of this the students in systematics, both graduate

and undergraduate, were terrified of him. Many GI’s from Viet Nam were begin¬

ning to come through the entomology program in this period; several told me

that even in the Army they had heard that Dr. R. M. Bohart ate undergraduates

for breakfast. Despite the students’ fears of Doc in systematics class, many of

them went into systematics or fields of research requiring this kind of background

because of his influence. Most students came away with an understanding of

systemaxics, species distinctions, and interspecies variation that they could have

gotten nowhere else.

After completing his systematics class I decided I would never go into System¬

atics as a field of study, and certainly never work on wasps. However, one year

later Doc proved to be quite persuasive; we co-authored a revision of the sphecid

genus Mellinus, and I began my career as a wasp systematist. I worked as a scientific

illustrator for Doc for about 6 years, drawing everything from mosquito larvae

to eumenid legs and sphecid genitalia. He was a demanding taskmaster, each

drawing had to be accurately and neatly done. The quality of the illustrations I

make today is largely due to his training.

In 1975 I entered graduate school and became his student in systematics. Al¬

though my thesis work was on orchid bees we began a series of cooperative papers

on the Chrysididae with the nebulous goal of straightening out the family. The

first was the marathon revision of the Western Hemisphere Hedychridium. Since

I was only being paid as a research assistant for 6 months he decided that we had

to complete this revision of 30+, mostly new species, during this period. We lived

and breathed Hedychridium for 6 months. That year I also was teaching assistant

for his summer field course at Sage Hen Creek. Every hour not spent collecting

(looking for specimens of Hedychridium ) at Sagehen was spent looking at, writing

about or drawing these wasps. Even now neither of us can look at Hedychridium

without flinching.

While I was a graduate student the number of systematics students began to

dwindle, primarily due to a growing lack of interest, even antagonism, toward

systematics in the scientific community. Baldo Villegas, Norm Smith, Joanne

Slansky (in absentia) and I were Doc’s graduate students in 1975. In 1978 Joanne

7 Department of Entomology, University of California, Davis, California 95616.

VOLUME 59, NUMBERS 1-4

completed her thesis, Baldo took a job with the California Department of Agri¬

culture, and Doc took on his last graduate student, Larry French, who is still

under his guidance. In January, 1979, Doc retired from the University of Cali¬

fornia and became Professor Emeritus. Norm Smith and I finished our theses in

1980 and 1979, respectively.

The years I’ve known Doc have been years of change for him. Although con¬

sidered stubborn by many, he is one of the most open-minded scientists about

his field of study I have ever met. For example, Doc, in spite of years of disdain

for certain new taxonomic approaches, finally adopted the use of cladograms in

his papers to demonstrate relationships between genera and higher categories. In

a lesser vein he now even uses a point punch to make points for mounting small

specimens. [This is one change that is difficult to believe! Editors.] As Doc neared

retirement he became more easy going and relaxed, and his students gradually

lost their fear of him although the awe remained. In short, he became a pussycat.

Some things never seem to change, however. Even at the last Sage Hen class he

taught (1979), his attitudes toward milk, buttermilk, peanut butter and jam sand¬

wiches, Special K, and EXTRA dry martinis (double, of course) remained un¬

changed. The same could be said for his infamous Sunday dinner at Sage Hen —

wine punch, concocted of lemon-lime soda and $2 a gallon red wine. This tradition

continued despite surreptitious additions of absolute ethanol to the punch by a

variety of students.

Since retiring, Doc still comes in to work practically every day or works at

home on Chrysididae and Sphecidae. We’re still pursuing the nebulous goal of

revising the family Chrysididae and have continued to do cooperative papers on

this group.

R. M. Boh art’s Impact on the U.C. Davis

Insect Collection

R. O. Schuster 8

In 1946 the Entomology Department made a commitment to establish a re¬

search collection by hiring A. T. McClay to curate what had been mainly a teaching

oriented collection. Mr. McClay brought to the department a fine personal col¬

lection of Coleoptera, and Dick contributed research material in the Diptera,

Hymenoptera, and Strepsiptera. Dick’s interests eventually polarized with the

aculeate wasps, and the collection has grown in this area to be among the most

comprehensive in the world.

In 1950 Dick became involved with Entomology 49 (now 109), a summer field

course stressing the biology, collecting techniques, and taxonomy of insects. Later,

he taught the course for both the U.C. Berkeley and Davis campuses. He provided

financial assistance for the development of the University Sage Hen Creek Field

8 Department of Entomology, University of California, Davis, California 95616.

PAN-PACIFIC ENTOMOLOGIST

Station near Truckee, California, and for many years the Entomology 109 course

has been conducted exclusively from Sage Hen. The insect fauna of the Sage Hen

area is extremely well known as a result of repeated and intensive collecting, and

the specimens collected by Dick, along with those of numerous students, are

deposited in the department’s collection. Throughout the years, specimens col¬

lected by Dick’s undergraduate and graduate students have contributed signifi¬

cantly to the development of the collection at Davis.

Field work in support of his research took him frequently, usually with co-

workers or students, to desert areas of the western United States. Expeditions to

Africa and South America were made in 1972 and 1975-1976, Venezuela in 1973,

and Australia in 1979. Margaret Bohart has been a frequent collecting companion

on these trips, and is a proficient collector. The resulting specimens have been

carefully taken, meticulously prepared, and of exceptional scientific value.

In addition to the specimens that have directly or indirectly resulted from Dick’s

research and teaching, he also has provided generous personal financial support

which has assisted in the collection’s becoming a major resource for systematic

entomology.

R. M. Bohart— an Overview

E. E. Grissell and A. S. Menke

During the past two years we have had occasion to contact many former stu¬

dents, colleagues, and friends of Dick Bohart. Through hours of conversations,

acres of correspondence, and pounds of manuscripts, we have come to learn that

there is an unsuspected uniformity of opinion about Dick among his associates.

We believe it is appropriate to present a short overview of the consensus which

has crystalized as we have put this “Festschrift” together.

One of the main points of agreement among Dick’s associates has been his

generosity toward students, friends, and colleagues. On the strictly practical side

this generosity has been expressed in the form of untold thousands of identifi¬

cations which Dick has given over many years to nearly anyone who has asked

for them. More than one curator has told us that they slip “extra” specimens into

a shipment which Dick has requested and that these additional specimens always

come back identified. The material is always identified in a quick, conscientious

manner whether it is a group he is currently working upon, or a group of some

past interest. The names are provided, in many cases, at the loss of personal

research time as well as a great amount of labor which often extends late into the

night at home. It is a labor of love, of course, and its breadth of coverage will

likely not be seen again in this day of specialization. He is among the last of those

“all knowledgeable” entomologists who can identify countless species in a wide

range of families.

Perhaps more important than the obvious public service that Dick has provided

over the past 50 years is the generosity of time which he has extended to students.

VOLUME 59, NUMBERS 1-4

Dick has always been available for questions, even at the expense of his own

work. Not only did he give us the feeling that our work was important (little did

we know!), but he exemplified what all of us should represent as taxonomists.

Perhaps Norm Smith, one of Dick’s later students, summed it up best when he

wrote to us:

“I think one thing that I respected most about Doc was that he was an excellent

example of what a University Professor should be. A young, impressionable

Ph.D. student could find no finer example to emulate. In academia, Doc

practiced what he preached and was able to set himself as a prime example

of what could be accomplished with dedication and perseverance in one’s

field of endeavor.”

Doc’s generosity to students has taken a variety of forms, from subsidizing field

trips at personal expense to helping with employment. But many students re¬

member him most as the motivating force behind their first publication. Many

of us have been junior authors with Dick, and it is safe to say, we think, that we

may not always have contributed our “fair” share toward the paper. On the other

hand, where other professors might insist on being a co-author on a student’s

dissertation, Dick would never consider it.

In co-authoring a paper, Dick was a considerable taskmaster, but at the same

time he was willing to listen to the other person’s point of view and to compromise.

Dick also read and reviewed a student’s paper with care and concern for the future

of that student. Roy Snelling wrote the following about the help that Dick gave

him as an early student:

“Always, he was remarkably patient and helpful to me, answering my naive

questions, identifying specimens, sending reprints, etc. He also had the no

doubt painful task of reading my first paper on Polistes systematics (I was

then a junior in high school). With admirable restraint and kindness, Dick

suggested I might profit from holding the paper a while until I had accu¬

mulated more data on certain of the forms; the paper has long since disap¬

peared into gracious oblivion.”

Dick is continuing his work, of course, with projects on Sphecidae and Chry-

sididae, but he is no longer taking graduate students. However, Dick’s former

students indirectly continue his work and have achieved some notoriety of their

own. Among these may be included the Chief of the USDA’s Systematic Ento¬

mology Laboratory and 3 Research Entomologists with the same organization;

the Chief of the USDA’s Bee Biology and Systematics Laboratory, Logan, Utah;

the Supervisory Research Entomologist of the Division of Vector-Borne Viral

Diseases, Ft. Collins, Colorado; the former Dean of the College of Agriculture,

Universidad de Zulia, Maricaibo, Venezuela; the former chairman of the De¬

partment of Entomology, University of California, Riverside; professors and as¬

sistant professors in various universities throughout the country, and several

entomologists in state agricultural departments.

Everyone we’ve contacted has commented upon Dick’s remarkably broad in¬

terest in insects ... an interest that has spanned more than 60 years. Although

recognized largely for his work with numerous groups of aculeate Hymenoptera,

Dick has also published papers on Thysanoptera, Lepidoptera, Diptera, Homop-

PAN-PACIFIC ENTOMOLOGIST

tera, Strepsiptera, and even Isopoda! His work has ranged from the toxicological

to the taxonomical. A glimpse through the appended bibliography will quickly

attest to Dick’s diversity as an entomologist. One notes several books or book

length treatises in several different groups of insects. Over the years his knowledge

and scientific stature in taxonomy have prompted many authors to name new

species and genera in his honor. He even has a family, Bohartillidae Kinzelbach

(Strepsiptera), named after him.

Perhaps less well known is the role that Dick has played in building the Uni¬

versity of California insect collection at Davis. This collection was in a fledgling

state in the late 40’s, and Dick, along with the help of A. T. McClay, its first

curator, and R. O. Schuster, its present curator, played a leading role in building

and developing the collection, often subsidizing its growth with personal funds.

Dick has always been a keen collector and a meticulous curator, and material

placed in the collection is of the highest quality. Today, the insect museum at

Davis is one of the outstanding university collections in the country.

As an aside from Dick’s career, few who have come into contact with him have

failed to note his rather acute sense of humor. A personal anecdote, concerning

long-known or “old-time” taxonomists, is a special pleasure coming from Dick.

We have also all fallen victim to the unexpected pun which could erupt at any

time or place. Better than trying to explain all of this we have been very fortunate

in obtaining a series of photos taken by Bob Kimsey during one of Dick’s taxonomy

classes in 1979. We think that Bob has done a remarkable job of capturing his

expressions and moods which most of us remember as students. The accompany¬

ing captions are doubly amusing when one considers that Dick wrote them himself.

Little did he know these photos would ever be made public, but how very well

he knows what his students are thinking!

We think anyone who knows Dick will agree that over his long and varied

career one of his finest assets has been Margaret, his wife. In fact we suspect that

many of Dick’s finer qualities have been encouraged and nurtured by Margaret.

Perhaps she has added just the right touch of “reason” to balance out Dick’s

somewhat over-zealous nature. Not to detract from Dick, but we can attest to

rather severe working conditions upon occasion; conditions which were mercifully

interrupted by coffee and cake, or a freshly baked pie upon the insistence of

Margaret. While Dick watched over our minds, Margaret often took care of our

nutritional needs and made certain Dick didn’t treat us too severely. She also

offered Dick’s friends, colleagues, students, and visitors the hospitality of her

home. We suspect that by taking exceptionally good care of Dick over these many

years, that Margaret has contributed more than her fair share to Dick’s accom¬

plishments. We know that she is a very capable collector in her own right. We

also know that Margaret was the precursor of the photocopying machine and that

for many years she “duplicated” all of Dick’s necessary research references on a

typewriter. Such duty goes a bit beyond normal marital laws, we feel. Perhaps

less well known is Margaret’s role as a guide during many of Dick’s collecting

trips. It is a well kept secret that Dick cannot tell left from right, especially while

driving. Margaret has been, on many occasions, the only means by which Dick

has been able to return home with any certainty. We all recognize that Dick would

not be the person he is without Margaret.

Of all that has been said of Dick Bohart, perhaps the simplist and most obvious

VOLUME 59, NUMBERS 1-4

“There is Little Bit of Ham in Every Professor—Some More than Others”

“You may as well resign yourselves, we are both stuck here for an hour”

“Pay close attention, or else!”

PAN-PACIFIC ENTOMOLOGIST

“This is really gruesome stuff’

VOLUME 59, NUMBERS 1-4 27

“I am absolutely serious about this!”

PAN-PACIFIC ENTOMOLOGIST

“My opinion of numerical taxonomy is hard to express in words”

VOLUME 59, NUMBERS 1-4

“I enjoyed the punch line too!”

PAN-PACIFIC ENTOMOLOGIST

conclusion is that he has an overwhelming and truly enthusiastic love of insects.

There is a realization among his friends that over the years, Dick’s enthusiasm

for insects, if anything, has grown stronger. We also know that this enthusiasm

has infected each of his students and colleagues in some manner and that we are

all the better for it. Not that many of us will duplicate Dick’s remarkably broad

knowledge, nor his zest for life ... for that is not taught. It is simply admired

and respected.

Bibliography of R. M. Bohart

The following list of papers by R. M. Bohart is complete through 1982. In an

effort to give correct citations, including mailing dates, we personally examined

nearly all of his publications. Some of his more obscure mosquito papers published

during WWII were found in the Medical Entomology Project at the U.S. National

Museum. Other papers were checked in the National Agriculture Library and the

library of the Smithsonian Institution. Lynn Kimsey checked a few obscure papers

for us, and also provided us with a copy of Dick’s list of publications. Papers not

seen by us are preceded by an asterisk. When known, the true date of publication,

that is, the mailing date, is cited at the end of each entry. Often this has meant

some sleuthing in journals. If a precise date could not be determined we have

simply indicated the month of issue in quotation marks at the end of the entry.

Circumstantial evidence suggests that some of Dick’s papers published in the

December issues of the Journal of Economic Entomology> may not have appeared

until early the following year.

We found several papers that Dick had overlooked when his list was made up.

Also, some papers were out of chronological sequence in his list. The result is

that the number we assign each paper in the following bibliography does not

necessarily coincide with those stamped on his reprints.

1936

1. A preliminary study of the genus Stylops in Calif. (Part I) (Strepsiptera,

Stylopidae). Pan-Pac. Ent. 12:9-18. March 20.

1937

2. A preliminary study of the genus Stylops in Calif. (Part II) (Strepsiptera,

Stylopidae). Pan-Pac. Ent. 13:49-57. “April.”

3. A new genus and species of Strepsiptera from Canada. Pan-Pac. Ent. 13:

101-105. “July.”

4. Synonymy of the genus Pseudoxenos Saunders (Strepsiptera, Xenidae) and

records of stylopized Hymenoptera from North Carolina. Psyche 44:132-

137. “December.”

1938

5. A synopsis of the genus Euparagia (Hymenoptera, Vespidae, Euparagiinae).

Pan-Pac. Ent. 14:136-139. August 19.

*6. Flower thrips in southern California on ornamentals. Floral West 4:4.

VOLUME 59, NUMBERS 1-4

1939

7. Taxonomy of the typical subgenus Odynerus in North America (Hymenop-

tera: Vesidae [sic]). Pan-Pac. Ent. 15:76-84. April 27.

8. Notes on Odynerus with a key to the North American subgenera and de¬

scription of a new subgenus (Hymenoptera, Vespidae). Pan-Pac. Ent. 15:

97-104. August 9.

9. A synopsis of the Odynerus boscii group in North America (Hymenoptera,

Vespidae). Bull. Brooklyn Ent. Soc. 34:245-251. November 13.

10. A new host plant and locality record for the Christmas berry thrips. J. Econ.

Ent. 32:880-881. “December.”

1940

11. A revision of the North American species of Pterocheilus and notes on related

genera (Hymenoptera, Vespidae). Ann. Ent. Soc. Amer. 33:162-208.

“March.” '

12. A preliminary study of the subgenus Leptochilus in North America (Hy¬

menoptera: Vespidae). Pan-Pac. Ent. 16:81-91. May 23.

13. Additional notes on Pterocheilus diversicolor Rohwer (Hymenoptera: Ves¬

pidae). Ann. Ent. Soc. Amer. 33:282. “June.”

14. Studies on the biology and control of sod webworms in California. J. Econ.

Ent. 33:886-890. “December.”

1941

15. Yellowjackets versus campers. Pan-Pac. Ent. 17:58. May 15.

16. A revision of the Strepsiptera with special reference to the species of North

America. Univ. Calif. Publ. Ent. 7:91-160. August 27.

1942

17. Life history of Diaspis boisduvalii and its control on Cattleya with calcium

cyanide. J. Econ. Ent. 35:365-368. “June.”

18. Plalynota stultana as a pest of field-grown carnations. J. Econ. Ent. 35:399-

403. “June.”

19. R. M. Bohart and A. Mallis. The control of pillbugs and sowbugs. J. Econ.

Ent. 35:654-658. “October.”

20. An analysis of the Odynerus congressus group of the subgenus Leptochilus

(Hymenoptera, Vespidae). Pan-Pac. Ent. 18:145-154. October 31.

1943

21. North American Stenodynerus of the anormis group (Vespidae: Hymenop¬

tera). Bull. Brooklyn Ent. Soc. 38:6-11. March 31.

22. A new generic name in Strepsiptera and description of a new species (Strep¬

siptera, Stylopidae). Bull. Brooklyn Ent. Soc. 38:12-13. March 31.

23. Calcium cyanide fumigation for the western thrips. J. Econ. Ent. 36:442-

444. “June.”

24. New species of Halictophagus with a key to the genus in North America

(Strepsiptera, Halictophagidae). Ann. Ent. Soc. Amer. 36:341-359. Octo¬

ber 4.

25. Bohart, G. E., P. C. Ting, R. M. Bohart, and C. K. Dorsey. Fundamentals

PAN-PACIFIC ENTOMOLOGIST

of field malariology and mosquito control—A reference and teaching sylla¬

bus. U.S. Naval Construction Training Center, Medical Dept., Camp Peary

[sic], Williamsburg, Va., 37 pp.

1944

26. Stenodynerus fundatus and related species in North America (Hymenoptera,

Vespidae). Pan-Pac. Ent. 20:69-75. April 29.

27. K. L. Knight, R. M. Bohart, and G. E. Bohart. Keys to the mosquitoes of

the Australasian Region. Natl. Research Council, Div. Med. Sci., Off. Med.

Inform., Washington, D.C., 71 pp. “July.”

28. R. M. Bohart and D. S. Famer. New culicine mosquitoes from the Philippine

Islands (Diptera, Culicidae). Proc. Biol. Soc. Wash. 57:69-73. October 31.

29. A. Stone and R. M. Bohart. Studies on mosquitoes from the Philippine

Islands and Australasia (Diptera: Culicidae). Proc. Ent. Soc. Wash. 46:205-

225. November 21.

30. D. S. Farner and R. M. Bohart. Three new species of Australasian Aedes.

Proc. Biol. Soc. Wash. 57:1 17-122. November 30.

1945

31. D. S. Farner and R. M. Bohart. A preliminary revision of the Scutellaris

group of the genus Aedes. U.S. Nav. Med. Bull. 44:37-53. “January.”

32. A synopsis of the Philippine mosquitoes. Bur. Med. and Surg., Navy Dept.,

Washington, D.C. Navmed 580:1-88. January.

33. A preliminary synopsis of the mosquitoes of Okinawa. U.S. Naval Med.

Res. Unit #2, Med. Sect. IsCom, Okinawa, 16 pp.

*34. Notes of disease-bearing mosquitoes in four specific areas: (a) Formosa; (b)

China coast from Amoy to Shanghai; (c) Ryukyu Islands (d) Bonin Islands.

Malaria and Epidemic Disease Control Bulletin #3, pp. 2-4.

35. New species and subspecies of Rygchium for North America (Hymenoptera:

Vespidae). Proc. Ent. Soc. Wash. 47:45-49. March 2.

36. H. S. Hurlbut and R. M. Bohart. Factors affecting the larvicidal action of

DDT on Culex quinquefasciatus. J. Econ. Ent. 38:725. “December.”

1946

37. A key to the Chinese culicine mosquitoes. Bur. Med. and Surg., Navy Dept.,

Washington, D.C. Navmed 961:1-23. “January.”

38. R. M. Bohart and R. L. Ingram. Four new species of mosquitoes from

Okinawa (Diptera: Culicidae). J. Wash. Acad. Sci. 36:46-52. February 15.

39. New species of mosquitoes from the Marianas and Okinawa (Diptera, Cu¬

licidae). Proc. Biol. Soc. Wash. 59:39-46. March 11.

40. Common caterpillars of the flower garden and their control. Southern Calif.

Homes and Gardens 3(6):7, 22.

41. Tsai-Yu Hsiao and R. M. Bohart. The mosquitoes of Japan and their

medical importance. Bur. Med. and Surg., Navy Dept., Washington, D.C.

Navmed 1095:1-44. September.

42. R. M. Bohart and R. L. Ingram. Mosquitoes of Okinawa and islands in the

Central Pacific. Bur. Med. and Surg., Navy Dept., Washington, D.C. Navmed

1055:1-110. October.

VOLUME 59, NUMBERS 1-4

43. A new species of Halictophagus parasitic on Cercopidae (Strepsiptera, Ha-

lictophagidae). Proc. Ent. Soc. Wash. 48:200-202. November 1.

44. B. V. Travis and R. M. Bohart. DDT to control maggots in latrines. J. Econ.

Ent, 39:740-742. “December.”

1947

45. Sod webworms and other lawn pests in California. Hilgardia 17:267-308.

“March.”

*46. Book review: Biologic des Abeilles, by Maurice Caullery, et al. Quart. Rev.

Biol. 22:222. “September.”

*47. Strepsiptera—An insect oddity. Ward’s Nat. Sci. Bull. 21:28-29. “Novem¬

ber.”

1948

48. Contributions toward a knowledge of the insect fauna of Lower California,

No. 9, Hymenoptera: Eumeninae. Proc. Calif. Acad. Sci. (4) 24:313-336.

“March.”

49. Organic phosphate insecticides: Mosquito larvae. Calif. Agric. 2(4):22-23.

“April.”

50. New North American Rygchium (Hymenoptera, Vespidae). Bull. Brooklyn

Ent. Soc. 43:80-87. July 1.

51. The genus Euparagia in North America (Hymenoptera: Vespidae). Pan-Pac.

Ent. 24:149-154. October 16.

52. The red Stenodynerus of Florida (Hymenoptera: Vespidae). Florida Ent. 31:

71-80. November 6.

53. A comparison of DDT and Dichlorodiphenyl Dichlorethane as larvicides

for Aedes mosquitoes. J. Econ. Ent. 41:834-835. “October.”

54. Differentiation of larvae and pupae of Aedes dorsalis and Aedes squamiger

(Diptera, Culicidae). Proc. Ent. Soc. Wash. 50:216-218. November 30.

1949

55. The subgenus Neoculex in America north of Mexico (Diptera, Culicidae).

Ann. Ent. Soc. Amer. 41:330-345. January 27 (incorrectly 1948).

56. Notes on North American Polistes with descriptions of new species and

subspecies (Hymenoptera, Vespidae). Pan-Pac. Ent. 25:97-103. July 29.

57. The genus Stenodynerus in southern Arizona (Hymenoptera, Vespidae).

Proc. Ent. Soc. Wash. 51:237-259. December 29.

1950

58. A synopsis of Maricopodynerus (Hymenoptera: Vespidae). Bull. Brooklyn

Ent. Soc. 45:17-25. February 18.

59. New species of solitary Vespidae from North America (Hymenoptera, Ves¬

pidae). Proc. Biol. Soc. Wash. 63:77-81. May 25.

60. R. M. Bohart and W. D. Murray. DDT resistance in Aedes nigromaculis

larvae. Proc. Papers 18th Ann. Conf. Calif. Mosq. Contr. Assoc., pp. 20-

22. “August.”

61. Observations on snow mosquitoes in California (Diptera: Culicidae). Pan-

Pac. Ent. 26:111-118. September 27.

PAN-PACIFIC ENTOMOLOGIST

*62. Turf pests. Northern Calif. Turf. Conf. Rept., pp. 28-31. “November.”

63. A new species of Orthopodomyia from California (Diptera, Culicidae). Ann.

Ent. Soc. Amer. 43:399-404. November 17.

64. North American additions to the genus Pterocheilus (Hymenoptera, Ves-

pidae). Proc. Biol. Soc. Wash. 63:195-200. December 29.

1951

65. Family Sapygidae, pp. 777-778. In: C. F. W. Muesebeck et al., Hymenoptera

of America north of Mexico. Synoptic catalog. U.S. Dept. Agric. Monograph

2. July 11.

66. Family Vespidae, pp. 875-907. In: C. F. W. Muesebeck et ah, Hymenoptera

of America north of Mexico. Synoptic catalog. U.S. Dept. Agric. Monograph

2. July 11.

67. The Myrmecolacidae of the Philippines (Strepsiptera). Wasmann J. Biol. 9:

83-103. May 25.

68. S. B. Freeborn and R. M. Bohart. The mosquitoes of California. Bull. Calif.

Insect Survey 1:25-78. June 8.

1952

69. The California species of mite-bearing Stenodynerus (Hymenoptera, Ves¬

pidae). Proc. Ent. Soc. Wash. 54:38-53. February 15.

1953

70. S. F. Bailey and R. M. Bohart. A mosquito survey and control program in

Guam. J. Econ. Ent. 45:947-952. February 9 (incorrectly 1952).

71. A new species of Culex and notes on other species of mosquitoes from

Okinawa (Diptera, Culicidae). Proc. Ent. Soc. Wash. 55:183-188. August

24.

72. R. M. Bohart, E. C. Mezger, and A. D. Telford. Observations on the seasonal

history of Aedes squamiger. Proc. Papers 21st Ann. Conf. Calif. Mosq. Contr.

Assoc., pp. 7-9. “November.”

73. Three new species of Stenodynerus from eastern United States (Hymenop¬

tera, Vespidae). Proc. Biol. Soc. Wash. 66:185-189. December 2.

1954

74. Cherry fruit sawfly. Calif. Agric. 8(3): 14. “March.”

75. Identification of first stage larvae of California Aedes (Diptera, Culicidae).

Ann. Ent. Soc. Amer. 47:355-366. August 17.

76. Some peculiarities in the distribution of mosquitoes in California. Proc.

Papers 22nd Ann. Conf. Calif. Mosq. Contr. Assoc., pp. 18-19. “July.”

77. S. F. Bailey, R. M. Bohart, and L. J. Booher. Mosquito control on the farm.

Calif. Agric. Exp. Sta. Circ, 439, 27 pp. “August.”

1955

78. Book review: Insects of Micronesia, vol. I, by J. Linsley Gressitt. Pan-Pac.

Ent. 31:172. December 13.

79. Strepsiptera, pp. 566-569. In: “A century of progress in the natural sciences,

1853-1953.” California Academy of Sciences, San Francisco. “November.”

VOLUME 59, NUMBERS 1-4

80. The role of taxonomy in relation to ecology and control. Proc. Papers 23rd

Ann. Conf. Calif. Mosq. Contr. Assoc, and 11th Ann. Meeting Amer. Mosq.

Contr. Assoc., pp. 97-98. “December.”

81. North American Leptochilus of the trinodus group (Hymenoptera, Vespi-

dae). Proc. Ent. Soc. Wash. 57:287-299. December 22.

1956

82. R. M. Bohart and A. H. Smith. Radioisotopes for evaluation of a fly nuisance

source. Calif. Vector Views 3:5, 7. “February.”

83. New species of mosquitoes from the southern Ryukyu Islands. Bull. Brook¬

lyn Ent. Soc. 51:29-34. May 11.

84. R. M. Bohart and E. I. Schlinger. Descriptions of two new western Oxybelus

(Hymenoptera: Sphecidae). Proc. Biol. Soc. Wash. 69:37-39. May 21.

85. A. H. Smith and R. M. Bohart. Evaluating fly nuisance source. Calif. Agric.

10(7): 11—12. “July.”

86. Identification and distribution of Aedes melanimon and Aedes dorsalis in

California. Proc. Papers 24th Ann. Conf. Calif. Mosq. Contr. Assoc., pp.

81-83. July 30.

87. E. C. Loomis, R. M. Bohart, and J. N. Belkin. Additions to the taxonomy

and distribution of California mosquitoes. Calif. Vector Views 3:37-45.

“August.”

88. R. M. Bohart and J. A. Powell. Observations on the nesting habits of Eu-

cerceris flavocincta Cresson (Hymenoptera: Sphecidae). Pan-Pac. Ent. 32:

143-144. August 14.

89. R. M. Bohart and E. I. Schlinger. New species of Oxybelus from western

North America (Hymenoptera: Sphecidae). Pan-Pac. Ent. 32:147-156. No¬

vember 7.

90. R. M. Bohart and E. I. Schlinger. An annotated synonymical list of North

American Oxybelus (Hymenoptera: Sphecidae). Pan-Pac. Ent. 32:157-165.

November 7.

1957

91. Insects of Micronesia: Culicidae. Insects Micronesia 12:1-85. January 21

(incorrectly 1956).

92. R. M. Bohart and R. C. Bechtel. The social wasps of California (Vespinae,

Polistinae, Polybiinae). Bull. Calif. Insect Survey 4:73-101. March 1.

*93. Five years of fly control on an agricultural university campus. Calif. Vector

Views 4:23, 25, 26. “April.”

94. R. M. Bohart and E. I. Schlinger. California wasps of the genus Oxybelus

(Hymenoptera: Sphecidae, Crabroninae). Bull. Calif. Insect Survey 4:103-

142. April 11.

95. Biological studies, mosquito control, and the entomologist. Proc. Papers

25th Ann. Conf. Calif. Mosq. Contr. Assoc., pp. 78-79. “July.”

96. R. M. Bohart and R. K. Washino. Differentiation of second and third stage

larvae of California Culex (Diptera: Culicidae). Ann. Ent. Soc. Amer. 50:

459-463. September 15.

1958

97. E. C. Loomis, R. M. Bohart, and J. N. Belkin. Insecticides for control of

PAN-PACIFIC ENTOMOLOGIST

flies on poultry ranches, dairies, homes, and food processing plants. Univ.

Calif. Agric. Ext. Serv. (unnumbered bulletin), 10 pp. “June” [also printed

in Calif. Vector Views 5:35-41 (June)].

98. A North American species of the genus Prosopigastra (Hymenoptera,

Sphecidae). Proc. Ent. Soc. Wash. 60:122-124. “June.”

99. R. M. Bohart and H. K. Court. New species of Lindenius from western

North America (Hymenoptera: Sphecidae). Pan-Pac. Ent. 34:161-167. July

22.

100. A new Priononyx and a key to the North American species (Hymenoptera:

Sphecidae). Bull. Brooklyn Ent. Soc. 53:90-93. October 15.

1959

101. New species of Aphilanthops from western North America (Hymenoptera,

Sphecidae). Ann. Ent. Soc. Amer. 52:105-108. January 20.

102. R. M. Bohart, D. H. Ecke, J. Fowler, and other collaborators. In: E. C.

Loomis, ed., A field guide to the common mosquitoes of California. Ent.

Committee, Calif. Mosq. Contr. Assoc., 27 pp. “February.”

103. A survey of the mosquitoes of the southern Ryukyus. Mosq. News 19:194-

197. “September.”

1960

104. R. M. Bohart and L. E. Campos. A review of the genus Omalus Panzer in

North America (Hymenoptera, Chrysididae). Ann. Ent. Soc. Amer. 53:235-

250. March 31.

105. Addendum to a review of the genus Omalus Panzer in North America

(Hymenoptera, Chrysididae). Ann. Ent. Soc. Amer. 53:435. May 31.

106. R. M. Bohart and P. M. Marsh. Observations on the habits of Oxybelus

sericenm Robertson. Pan-Pac. Ent. 36:115-118. August 9.

1961

107. R. M. Bohart and A. S. Menke. A review of the genus Palmodes in North

America (Hymenoptera: Sphecidae). Proc. Ent. Soc. Wash. 63:179-191.

November 22.

108. The art and practice of key-making in Entomology. Verhandl. XI Internat.

Kong. Ent., Wien, 1960. 1:17-19. December (incorrectly 1962).

1962

109. Gorytes (Gorytes) mcateei Krombein and Bohart, new species, pp. 15-16.

In: K. V. Krombein, Natural history of Plummers Island, Maryland, etc.

Proc. Biol. Soc. Wash. 75:1-17. March 30.

110. New species of black Tachysphex from North America (Hymenoptera,

Sphecidae). Proc. Biol. Soc. Wash. 75:33-39. March 30.

111. R. M. Bohart, D. H. Ecke, J. Fowler, and other collaborators. In: T. H.

Lauret, ed., A field guide to common mosquito-like gnats of California. Ent.

Committee, Calif. Mosq. Contr. Assoc., 19 pp.

112. R. M. Bohart and L. A. Stange. Lectotypes of Nearctic and Neotropical

Zethini in European and American museums (Hymenoptera: Vespidae).

Bull. Brooklyn Ent. Soc. 57:28-32. April 9.

VOLUME 59, NUMBERS 1-4

113. R. M. Bohart and G. E. Bohart. A revision of the Larropsis subgenus An-

cistromma Fox (Hymenoptera: Sphecidae). Proc. Ent. Soc. Wash. 64:21-

37. May 1.

114. The Tachytes pepticus group in North America (Hymenoptera: Sphecidae).

Pan-Pac. Ent. 38:117-129. June 29.

115. Book review: A catalogue and reclassification of the Indo-Australian lchneu-

monidae by Henry Townes. Bull. Ent. Soc. Amer. 8:93-94. “June.”

116. A new Strepsipteran parasitic on Coreidae (Strepsiptera: Halictophagidae

and Hemiptera: Coreidae). Proc. Ent. Soc. Wash. 64:91-94. August 27.

117. A review of the hexadentate species of Chrysis of America north of Mexico

(Hymenoptera, Chrysididae). Acta Hymenopterologica 1:361-375. Septem¬

ber 30.

118. Vector control research at the University of California, Davis. Proc. Papers

30th Ann. Conf. Calif. Mosq. Contr. Assoc., pp. 29-30. October 22.

1963

119. New species of Pseudomasaris, pp. 290-292, 298-299, 301. In: O. W. Rich¬

ards, The species of Pseudomasaris Ashmead (Hymenoptera: Masaridae).

Univ. Calif. Publ. Ent. 27:283-310. July 3.

120. R. M. Bohart and A. S. Menke. A reclassification of the Sphecinae with a

revision of the nearctic species of the tribes Sceliphronini and Sphecini

(Hymenoptera, Sphecidae). Univ. Calif. Publ. Ent. 30:91-182. October 15.

1964

121. New species of Chrysis in the lauta, propria and venusta groups from North

America (Hymenoptera: Chrysididae). Proc. Biol. Soc. Wash. 77:223-236.

October 30.

122. Mosquito and related arthropod control activities—University of California,

Davis. Proc. Papers 32nd Ann. Conf. Calif. Mosq. Contr. Assoc., pp. 29-

30. December 30.

1965

123. Synonymy of North American Odynerini described by Peter Cameron. Pan-

Pac. Ent. 31:107—113. June 11.

124. R. M. Bohart and L. A. Stange. A revision of the genus Zethus Fabricius

in the Western Hemisphere (Hymenoptera: Eumenidae). Univ. Calif. Publ.

Ent. 40:1-208. September 21.

125. R. M. Bohart and A. S. Menke. Stizus Latreille, [1802-1803] (Insecta, Hy¬

menoptera): proposed designation of a type-species under the plenary pow¬

ers. Z. N. (S.) 1710. Bull. Zool. Nomencl. 22:255-256. “November.”

126. R. M. Bohart and A. S. Menke. Diodontus Curtis, 1834 (Insecta, Hyme¬

noptera): Proposed designation of a type-species under the plenary powers.

Z. N. (S.) 1711. Bull. Zool. Nomencl. 22:257-258. “November.”

127. Progress in mosquito source reduction in California. Proc. Papers 33rd Ann.

Conf. Calif. Mosq. Contr. Assoc., pp. 30-31. “December.”

1966

128. New species of Ceratochrysis from North America (Hymenoptera: Chrysi¬

didae). J. Kansas Ent. Soc. 39:112-123. February 21.

PAN-PACIFIC ENTOMOLOGIST

129. The genus Neochrysis in America north of Mexico (Hymenoptera: Chrysid-

idae). Bull. Brooklyn Ent. Soc. 58:139-144. March 14 (incorrectly 1963).

130. Withdrawal of application for the designation of a type-species for Stizus

Latreille [1802-1803]. Z. N. (S.) 1710. Bull. Zool. Nomencl. 23:7-8. April

29.

131. A. S. Menke, R. M. Bohart, and J. van der Vecht. Podalonia Spinola, 1853:

Proposed suppression under plenary powers in favour of Podalonia Fernald,

1927, with Ammophila violaceipennis Lepeletier, 1845, as type species. Z.

N. (S.) 1735. Bull. Zool. Nomencl. 23:48-51. April 29.

132. New Stenodynerus from North America (Hymenoptera; Eumenidae). Proc.

Biol. Soc. Wash. 79:73-81. May 23.

133. F. D. Parker and R. M. Bohart. Host-parasite associations in some twig¬

nesting Hymenoptera from western North America. Pan-Pac. Ent. 42:91-

98. June 14.

134. R. M. Bohart and J. F. Holland. Nesting habits of Enchemicrum australe

Pate (Hymenoptera, Sphecidae, Crabroninae). Pan-Pac. Ent. 42:161. June

14.

135. A review of Aphilanthops and related genera (Hymenoptera: Sphecidae).

Proc. Ent. Soc. Wash. 68:158-167. June 29.

136. R. M. Bohart, C. S. Lin, and J. F. Holland. Bionomics of Oxybelus sparideus

at Lake Texoma, Oklahoma (Hymenoptera: Sphecidae: Crabroninae). Ann.

Ent. Soc. Amer. 59:818-820. July 15.

137. New species of cuckoo wasps bred from twigs in western North America

(Hymenoptera: Chrysididae). Proc. Biol. Soc. Wash. 79:131-134. August

15.

138. Strepsiptera. In: Encyclopedia Brittanica, vol. 21, 14th edit. W. Benton,

Chicago. “August” (reprinted in subsequent printings).

139. Book review: A catalogue and reclassification of the eastern Palearctic Ich-

neumonidae, by Henry Townes, Setsuya Momoi, and Marjorie Townes.

Bull. Ent. Soc. Amer. 12:358. “September.”

140. H. K. Court and R. M. Bohart. Systematic notes on crabronids with de¬

scription of a new species (Hymenoptera: Sphecidae). Pan-Pac. Ent. 42:329-

332. November 30.

141. G. E. Bohart and R. M. Bohart. A revision of the genus Larropsis Patton

(Hymenoptera: Sphecidae). Trans. Amer. Ent. Soc. 92:653-685. Decem¬

ber 16.

1967

142. New genera of Gorytini (Hymenoptera: Sphecidae: Nyssoninae). Pan-Pac.

Ent. 43:155-161. July 14.

143. R. M. Bohart and R. L. Brumley. Two new species of Hedychridium from

California (Hymenoptera: Chrysididae). Pan-Pac. Ent. 43:232-235. Octo¬

ber 30.

1968

144. New species of Nysson from southwestern United States (Hymenoptera:

Sphecidae). Pan-Pac. Ent. 43:315-325. February 28 (incorrectly 1967).

145. F. D. Parker and R. M. Bohart. Host-parasite associations in some twig-

VOLUME 59, NUMBERS 1-4

nesting Hymenoptera form western North America. Part II. Pan-Pac. Ent.

44:1-6. June 28.

146. New Hoplisoides from the United States (Hymenoptera; Sphecidae). Proc.

Ent. Soc. Wash. 70:287-292. October 4.

147. New Nyssoninae from North and South America (Hymenoptera: Spheci¬

dae). Pan-Pac. Ent. 44:228-236. October 21.

148. A synopsis of the American species of the genus Oryttus (Hymenoptera,

Sphecidae). Proc. Biol. Soc. Wash. 81:431-437. December 30.

1969

149. New species of Pseudoplisus from North and Central America. I. The P.

phaleratus group (Hymenoptera: Sphecidae). J. Kansas Ent. Soc. 41:494-

501. January 10 (incorrectly 1968).

150. R. M. Bohart and H. H. Laidlaw. Howard Lester McKenzie. Univ. Calif.,

In memoriam for 1969, pp. 102-104. “May.’"

151. R. M. Bohart and E. E. Grissell. New species of Psenini (Hymenoptera:

Sphecidae). Pan-Pac. Ent. 45:216-221. October 14.

152. New species of Pseudoplisus from North America, II (Hymenoptera: Spheci¬

dae). J. Kansas Ent. Soc. 42:392-405. October 27.

153. D. R. Miller, R. M. Bohart, and R. F. Wilkey. Howard Lester McKenzie,

Jr. 1910-1968. Pan-Pac. Ent. 45:245-259. December 8.

1970

154. R. E. Eastwood, J. R. Fowler, and other collaborators including R. M.

Bohart. In: J. L. Mallars and J. R. Fowler, eds.. Mosquito eating fishes in

California. Calif. Mosq. Contr. Assoc., Visalia, Calif., 27 pp. “June.”

155. New species, synonymy and lectotype designations in North American Bem-

bicini (Hymenoptera: Sphecidae). Pan-Pac. Ent. 46:201-207. August 10.

156. The genus Ochleroptera in New Guinea. Proc. Ent. Soc. Wash. 72:386-387.

September 25.

157. Systematics of mosquitoes. Proc. Papers 38th Ann. Conf. Calif. Mosq. Contr.

Assoc., p. 48. September 25.

1971

158. New species of Gorytini from western North America (Hymenoptera:

Sphecidae). Proc. Biol. Soc. Wash. 83:445-453. February 9.

159. R. M. Bohart and D. S. Horning, Jr. California bembicine sand wasps. Bull.

Calif. Insect Survey 13:1-49. December 30.

1972

160. R. M. Bohart and R. K. Washino. Changing perspectives in systematics of

mosquitoes in California. Proc. Papers 40th Ann. Conf. Calif. Mosq. Contr.

Assoc., pp. 126-127. August 23.

161. R. M. Bohart and E. E. Grissell. Nesting habits and larva of Pulverro mon-

ticola (Hymenoptera: Sphecidae). Pan-Pac. Ent. 48:145-149. September 28.

162. R. M. Bohart and R. O. Schuster. A host record for Fedtschenkia (Hyme¬

noptera: Sapygidae). Pan-Pac. Ent. 48:149. September 28.

PAN-PACIFIC ENTOMOLOGIST

163. New North American Philanthus (Hymenoptera: Sphecidae). Proc. Ent. Soc.

Wash. 74:397-403. December 30.

1974

164. A. S. Menke, R. M. Bohart, and O. W. Richards. Proposed emendation to

Nysson of the name Nysso Latreille, 1796 (Hymenoptera, Sphecidae). Z. N.

(S.) 2055. Bull. Zool. Nomencl. 30:217-218. June 28.

165. A. S. Menke, R. M. Bohart, and O. W. Richards. Euplilis Risso, 1826

(Hymenoptera, Sphecidae): Proposed suppression under the plenary powers

in favor of Rhopalum Stephens, 1829. Z. N. (S.) 2056. Bull. Zool. Nomencl.

39:219-220. June 28.

166. R. M. Bohart and A. S. Menke. Names of some North American vespoid

wasps (Hymenoptera: Vespidae and Eumenidae). J. Kansas Ent. Soc. 47:

459-469. October 30.

167. M. L. Siri and R. M. Bohart. A review of the genus Mellinus (Hymenoptera:

Sphecidae). Pan-Pac. Ent. 50:169-176. November 8.

168. A review of the genus Rhopalum in America north of Mexico (Hymenoptera:

Sphecidae). J. Georgia Ent. Soc. 9:252-260. November.

1975

169. Comment on the proposed suppression of Euplilus Risso, 1826 in favour

of Rhopalum Stephens, 1829. Z. N. (S.) 2056. Bull. Zool. Nomencl. 32:98.

“June.”

170. R. M. Bohart and E. E. Grissell. California wasps of the subfamily Philan-

thinae (Hymenoptera: Specidae). Bull. Calif. Insect Survey 19:1-92. Au¬

gust 1.

1976

171. C. J. Mitchell and R. M. Bohart. Mosquito control on the farm. University

of California, Div. Agric. Sci., Coop. Ext., Leaflet 2850, 14 pp. “January.”

172. R. M. Bohart and A. S. Menke. Sphecid wasps of the world. A generic

revision. University of California Press, Berkeley and Los Angeles, ix + 695

pp. April 27.

173. A review of the nearctic species of Crabro (Hymenoptera: Sphecidae). Trans.

Amer. Ent. Soc. 102:229-287. “July.”

174. R. M. Bohart and A. S. Menke. Reply to Dr. Krombein’s comment on our

proposal to suppress Euplilis Risso, 1826. Z. N. (S.) 2056. Bull. Zool. No¬

mencl. 33:68. “September.”

175. Comment on the application concerning Notozus Forster, 1853. Z. N. (S.)

2109. Bull. Zool. Nomencl. 33:72. “September.”

1977

176. R. M. Bohart and L. Stange. Liogorytes joergenseni (Brethes), a cicada killer

in Argentina (Sphecidae, Nyssoninae). Pan-Pac. Ent. 52:313. March 27 (in¬

correctly 1976).

177. R. M. Bohart and B. Villegas. Nesting behavior of Encopognathus ruftventris

Timberlake (Hymenoptera: Sphecidae). Pan-Pac. Ent. 52:331-334. March

27 (incorrectly 1976).

VOLUME 59, NUMBERS 1-4

1978

178. R. M. Bohart and N. J. Smith. A revision of Nearctic Ammoplanops (Hy-

menoptera, Sphecidae). J. Kansas Ent. Soc. 51:75-90. January 26.

179. R. M. Bohart and R. K. Washino. Mosquitoes of California. 3rd edit. Uni¬

versity of California, Div. Agric. Sci., Berkeley, Publication Number 4084,

viii 4-153 pp. March.

180. R. M. Bohart and M. E. Irwin. A study of stylopization in the bee genus

Dufourea. Pan-Pac. Ent. 54:98-102. September 5.

181. R. M. Bohart and L. S. Kimsey. A revision of the New World species of

Hedychridium (Hymenoptera, Chrysididae). Proc. Biol. Soc. Wash. 91:590-

635. December 6.

1979

182. Tachytes of South America (Hymenoptera, Sphecidae, Larrinae). Trans.

Amer. Ent. Soc. 104:435-505. February 20.

183. A. S. Menke and R. M. Bohart. Sphecid wasps of the world: Errors and

omissions (Hymenoptera: Sphecidae). Proc. Ent. Soc. Wash. 81:111-124.

February 28.

184. R. M. Bohart and J. D. McLaughlin. Evidence indicating Ammophila (Hy¬

menoptera: Sphecidae) as host of Spintharosoma (Hymenoptera: Chrysid¬

idae). Pan-Pac. Ent. 54:310. April 5 (incorrectly 1978).

185. R. M. Bohart and L. S. Kimsey. A key to the species of Ectemnius in America

north of Mexico with notes and description of a new species (Hymenoptera:

Sphecidae). Proc. Ent. Soc. Wash. 81:486-498. August 13.

1980

186. New genera and species of North American Chrysididae (Hymenoptera). J.

Kansas Ent. Soc. 53:132-136. January 2.

187. R. M. Bohart and L. S. Kimsey. A generic synopsis of the Chrysididae of

America north of Mexico. J. Kansas Ent. Soc. 53:137-148. January 2.

188. A review of the North American species of Dienoplus (Hymenoptera: Spheci¬

dae). Pan-Pac. Ent. 56:63-70. January 24.

189. The Middle American species of Stenodynerus (Hymenoptera, Eumenidae).

Polskie Pismo Ent. 50:71-108. March 30.

1981

190. L. S. Kimsey and R. M. Bohart. A synopsis of the chrysidid genera of

neotropical America (Chrysidoidea, Hymenoptera). Psyche 87:75-91. April

27 (incorrectly 1980).

1982

191. R. M. Bohart and L. S. Kimsey. A synopsis of the Chrysididae in America

north of Mexico. Mem. Amer. Ent. Inst. 33:1-266. March 8.

192. Leucodynerus, a new genus of solitary wasps from western North America

(Hymenoptera: Sphecidae [correctly Eumenidae]). J. Kansas Ent. Soc. 55:

442-446. July 20.

193. New species of Stictiella (Hymenoptera: Sphecidae). J. Kansas Ent. Soc. 55:

593-597. July 20.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 42-49

Gall-inducing Cynipid Wasps from Quercus dunnii Kellogg

(Hymenoptera)

D. Charles Dailey and Christine M. Sprenger

Sierra College, 5000 Rocklin Road, Rocklin, California 95677.

The cynipid fauna on the oak Quercus ( Protobalanus) dunnii Kellogg previously

has received little study partly because of its restricted ecological and geographical

distributions. Quercus dunnii occurs sporadically in the sagebrush-chaparral eco-

tone. Its range is a narrow horizontal band extending from southwest Arizona

and northern Baja California, Mexico, around the western edge of the Mojave

Desert (Tucker and Haskell, 1960), in San Benito County, California (Griffin and

Tucker, 1976), and north to the inner coast range west of Byron, Contra Costa

County, California (Tucker et al., 1982).

The only prior cynipid host record for Q. dunnii (Weld, 1957) listed Disholcaspis

chrysolepidis (Beutenmuller) and two undescribed stem galls. The present paper

reports an additional 8 unisexual and 1 bisexual generation gall-inducing cynipid

wasps including 4 newly described species ( Trichoteras burnetii, Loxaulus boharti,

Heteroecus fragilis and Heteroecus lyoni; all unisexual generations) and five new

host records. Type specimens placed in the U.S. National Museum and the Cal¬

ifornia Academy of Sciences are indicated with the abbreviations USNM and

CAS respectively. Those placed in the L. H. Weld collection are in the possession

of Robert J. Lyon, Los Angeles City College, Los Angeles, Calif.

Biogeography

Q, dunnii has a highly disjunct relict distribution, occasionally only single trees.

As is typical for island faunas each local oak population supports only a few

cynipid species, particularly in the northern end of the range. This scattered

distribution pattern provides a potential for rapid evolution and/or extinction of

new species and requires widespread collecting to determine the current total

fauna of this oak. The galls of numerous additional undescribed cynipids, probably

the alternate bisexual generations, have been seen or collected from Q, dunnii but

they will not be described until insects can be reared.

The 3 subgenera of American oaks have no gall-inducing cynipids in common,

but most gall-inducing cynipids occur on more than 1 oak species within a sub¬

genus. The cynipids on Lepidobalanus, or white oaks, and Erythrobalanus, or red

and black oaks, of the Southwest and the Pacific slope are generally distinctive,

with only 1 species known from both regions (R. J. Lyon, pers. comm.). But for

some reason a different pattern occurs in the Protobalanus oaks. Some of the

species listed in this paper on Q. dunnii occur in both the Pacific slope and the

Southwest. An even more widespread distribution of the gall-inducing cynipids

occurs on the more montane Quercus chrysolepis Leibmann. The cynipids on Q.

dunnii with the widest geographic distribution are those which also occur on other

Protobalanus oaks. The host patterns of these insects have not been generally

appreciated because Weld (1960) routinely listed Q. chrysolepis, Q. dunnii and

hybrids between these two species in Arizona all as Q. wilcoxii. In Weld (1957)

VOLUME 59, NUMBERS 1-4

Quercus palmeri is a synonym of Q. dunnii. The differences in the evolutionary

history of the three American oak subgenera and the atypical geographic pattern

of the Protobalanus cynipid faunas in these two regions are not yet understood.

Trichoteras burnetti Dailey and Sprenger, New Species

(Fig. 2)

Unisexual generation female holotype.— Head: pubescent, reticulate; from above

transverse; slightly narrower than thorax; cheeks widened behind eyes; occiput

not elevated above general head contour; malar space 0.4 eye height, not grooved;

antenna 13 segmented, filiform, segment 1 longer than 2, segment 3 equal to 4

or 1 plus 2,13 twice the length of 12. Thorax: mesoscutum polished, micropunctate

throughout, as broad as long, pubescent, profile normal; anterior parallel lines

absent; notauli complete, close together throughout and nearly meeting poste¬

riorly, twice as wide posteriorly as anteriorly; parapsidal lines and median groove

absent; scutellum pubescent, punctate; fovea indistinct, coriaceous, weakly lon¬

gitudinally ridged with slightly stronger median ridge; mesopleuron shiny, pu¬

bescent, punctate. Wings normal sized, hyaline, with fine pubescence, margin

ciliate, veins light brown, radial cell open, aerolet 0.1 first cubital cell length.

Claws toothed. Abdomen: propodeum smooth, polished, without median carina;

length less than head plus thorax, tergites II and III smooth, polished, pubescent

anteriorly; ventral spine stout but evenly tapering, 3 times as long as high or wide,

bristly, shorter than basal hind tarsal segment. Using maximum head width as

base, mesonotum length is 1.5, antenna 2.3, wing 5.5. Body length is 2.5 mm.

Color of head, thorax, legs, and abdomen light brown, antennae brown, darkest

distally.

Host. — Quercus dunnii.

Variation. — Size range of 28 specimens: 1.8-2.7 mm, average 2.1 mm.

Gall.— Yellow, highly pubescent aborted axillary bud or leaf petiole gall; poly-

thalamous with angular basal larval cells; diameter 1.0-3.0 cm (Fig. 2 and Weld,

1957, Fig. 163).

Biology. — Pupation occurs in September. Insects reared by Burnett emerged

during December and January and were observed to oviposit between axillary

bud scales but no galls developed.

Type material.— Holotype 2, 3 2 paratypes: CALIFORNIA, Riverside Co., ca.

12 km E Anza, reared from 5 galls coll, from a single plant (Dailey #1263).

Additional paratypes reared as follows: MEXICO, Baja California (Norte), El

Condor (12 galls and 29 2, Dailey #1168); CALIFORNIA, Riverside Co., 12 km

E Hemet (4 galls, 43 2), Dr. John Burnett. All material in Dailey collection except

holotype 2 and 2 para type 2 and galls in USNM; 2 paratype 2 and galls in CAS;

2 paratype 2 in Weld collection.

Distribution. — In addition to the above localities, T. burnetti was recorded by

Weld (1957) from galls collected in the San Jacinto Mountains, California on Q.

palmeri (=Q. dunnii).

Discussion. —Trichoteras burnetti is the only gall on Q. dunnii which is poly-

thalamous and covered with dense, thick pubescence and is the only bud or leaf-

petiole gall. It resembles Andricus lasius McCracken and Egbert which occurs on

other oak species. It is listed as species #2 in Weld’s manuscript key to Trichoteras

in the possession of Mr. Robert J. Lyon. Adult females may be distinguished from

PAN-PACIFIC ENTOMOLOGIST

related species by the shorter ventral spine which is only 3 times as long as wide

or high.

Etymology.— Named for Dr. John Burnett who first reared this species.

Trichoteras rotundula Weld: New Host Record

(Fig. 1)

This unisexual generation monothalamous, hollow, thin-walled, spherical leaf

gall with central larval cell supported by filaments is attached to lower veins,

occasionally in clusters (Fig. 1 and Weld, 1957, Fig. 91).

Host. —Quercus dunnii, Q. chrysolepis.

Biology. — Galls are fully grown by summer and females emerge in October and

November. The insect specimens range from 2.0 to 2.6 mm in body length.

Distribution.— Specimens emerged from galls collected near Hemet Lake, Riv¬

erside County, California by Dr. John Burnett. Similar galls were collected by us

at Sedona, Coconino County, Arizona and El Condor and Melling Ranch, Baja

California (Norte), Mexico.

Andricus chrysobalani Weld: New Host Record

This is a bisexual generation which emerges from cells in the inner margins of

the acorn cups and/or outer acorn hull during August. Insects emerged in mid-

August from galls collected east of Anza, Riverside County, California (Dailey

#1292). Similar galls were seen at Sedona, Coconino County, Arizona (Dailey

#1289).

Andricus reniformis McCracken and Egbert: New Host Record

Galls of Andricus reniformis have been collected at Sedona, Coconino County,

Arizona. These galls are similar to those of Disholcaspis truckeensis (Ashmead)

which Weld recorded on “ Quercus wilcoxii ” in Arizona. Weld’s listings of Q.

wilcoxii include Q. chrysolepis, Q. dunnii or hybrids between these two oaks.

Disholcaspis chrysolepidis (Beutenmuller)

Disholcaspis chrysolepidis is a unisexual generation which produces an elongate

stem gall, dark brown and convex on top and lighter in color and concave on

both sides (Weld, 1957, Fig. 52). It has been found 10 km east of Anza, Riverside

County, and 16 km northeast of Cajon Pass, San Bernardino County, California.

Disholcaspis truckeensis (Ashmead): New Host Record

Galls of Disholcaspis truckeensis were collected in the Haulaupi Mountains

south of Kingman, Mojave County and Sedona, Coconino County, Arizona and

from a single isolated tree 16 km northeast of Cajon Pass, San Bernardino County,

California. This gall is similar to those of the bisexual generation of Loxaulus

trizonalis Weld.

Hosts. — Quercus dunnii, Q. chrysolepis, and Q. vaccinifolia Kellogg.

The two species of Disholcaspis on Protobalanus oaks are distinctive from the

other members in this genus (Burnett, 1977). This is also true for the 5 species

of Andricus on Protobalanus oaks. All 7 species are sufficiently similar to each

VOLUME 59, NUMBERS 1-4

other that their life histories should be determined and their collective generic

assignment reassessed.

Loxaulus boharti Dailey and Sprenger, New Species

(Fig- 3)

Unisexual generation female holotype. — Head: coriaceous; from above massive;

broader than thorax; cheeks widened behind eyes; occiput rounded, only slightly

elevated above contour of head; malar space 0.3 eye height, grooved, inner-ocular

margins parallel; antenna 13 segmented, filiform, segment 1 longer than 2, segment

3 less than 4 or 1 plus 2, segment 13 twice the length of 12. Thorax: mesoscu-

tum strongly transversely coriaceous, bare, as broad as long; sides of pronotum

coriaceous; anterior parallel lines slightly less than 0.5 mesoscutum length; notauli

incomplete, extending from posterior 0.3 mesoscutum length with short shallow

depression anterior to that, wrinkled; parapsidal lines 0.4 mesoscutum length;

median groove absent; scutellum weakly reticulate, slightly longer than wide; fovea

indistinct, longitudinally wrinkled, separated by median wrinkle; mesopleuron

aciculate. Wings pubescent, margin ciliate, veins brown, radial cell closed, aerolet

0.25 cubital cell length, basal and first abcissa of radial cell clouded. Claws simple.

Abdomen: propodeum polished, slightly wrinkled, carinae straight, distinct, en¬

closing an area longer than wide, with distinct complete median carina; 2 tergites

visible in dorsal view, length less than head plus thorax, slightly longer than high,

smooth, polished, bare; ventral spine 3 times as long as high or wide. Using

maximum head width as base, mesonotum length is 1.3, antenna 2.2, wing 3.5.

Body length 1.8 mm. Color of head, thorax, legs, and abdomen yellow-brown;

antennal segments 1-4 yellow-brown, 5-13 brown and ribbed.

Host. —Quercus dunnii.

Gall. — Integral stem gall, larval cells 2 mm in diameter, develop in wood under

bark, stems slightly lumpy though lumps barely noticeable until insects emerge

(Fig. 3). Similar to gall of Loxaulus brunneus (Ashmead) on Quercus chrysolepis.

Variation. — Size range of 61 specimens 1.6-1.9 mm.

Biology. — Insects emerge in late March and early April from 2-year-old twigs

and only the basal half of prior year twigs. The oviposition site is unknown but

is probably in buds with bisexual generation emergence and oviposition in twigs

when one-half of the current year’s stem growth has occurred.

Type material. — Holotype 9 , 91 9 paratypes: CALIFORNIA, San Diego Co., 1

km W Jacumba, reared from galls collected by us from a single shrub. All material

in Dailey collection except holotype 9, 2 paratype 9 and 2 galls in USNM, 4

paratype 9 and 4 galls each in CAS and Weld collections.

Distribution. —Galls similar to the above were collected at El Condor, Baja

California (Norte), Mexico and 14 km east of Anza, Riverside Co., and 8 km west

of Byron, Contra Costa County, California on Quercus dunnii and from San Carlos

Canyon, Baja California (Norte), Mexico on Q. cedrosensis Muller. No insects

were reared.

Discussion.—Loxaulus boharti is the only integral stem gall known from Q.

dunnii. Adult females may be distinguished from L. brunneus (a similar gall-

inducer on Q. chrysolepis ) by the incomplete notauli and the ventral spine which

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-5. Cynipid galls. 1, Trichoteras rotundula. 2, T. burnetti. 3, Loxaulus boharti. 4, Heteroecus

fragilis. 5, H. lyoni.

is 3 times as long as wide or high. It keys to species 7 in Weld’s manuscript key

to Loxaulus in the possession of Robert Lyon.

Etymology .—This species is named for Dr. R. M. Bohart who has aided both

of us in our studies of cynipids.

Heteroecus fragilis Dailey and Sprenger, New Species

(Fig. 4)

Unisexual generation female holotype. — Head: coriaceous; from above trans¬

verse; slightly narrower than thorax; cheeks not widened behind eyes; occiput

flattened, elevated above the head contour; malar space 0.5 eye height, not grooved;

VOLUME 59, NUMBERS 1-4

antenna 14 segmented, filiform, segment 1 longer than 2, segment 3 longer than

4 or 1 plus 2, segment 14 one and one-third times the length of 13. Thorax:

mesoscutum coriaceous throughout; as broad as long, sparsely pubescent; anterior

parallel lines distinct, slightly longer than 0.5 mesoscutum length, weak transverse

rugosity between lines; notauli incomplete, extending from posterior 0.6 meso¬

scutum length, sharp edged, pitted; parapsidal lines distinct, from posterior 0.5

mesoscutum length; median groove weak, 0.2 mesoscutum length; scutellum ir¬

regularly rugose, sparsely pubescent; fovea wrinkled, polished, bare, separated by

a ridge; mesopleuron aciculate, bare. Wings hyaline, setae not projecting beyong

margin, veins yellow, radial cell open, aerolet 0.3 length of first cubital cell. Claws

simple. Abdomen: propodeum with weak median carina; length greater than head

plus thorax, tergum II smooth, polished, bare dorsally, slightly pubescent ante-

riolaterally, micropunctate on posterior margin; ventral spine 10 times as long as

high. Using maximum head width as base, mesonotum length is 1.3, antenna

2.3, wing 4.0. Body length is 2.7 mm. Color of head, thorax, legs, and abdomen

dark red-brown, first six antennal segments yellow-brown, antennal segments 7-

14 brown.

Host. — Quercus dunnii.

Gall. — Detachable, monothalamous stem gall with a thick wall formed from

many small, fragile, sponge-like chambers. Yellow, spherical, 5-8 mm in diameter

(Fig. 4).

Biology. — Insects had emerged from most galls collected 29 March 1972, how¬

ever the collection did yield one adult insect. Oviposition site of the unisexual

generation is unknown. Gall position indicates the eggs for the unisexual gener¬

ation are laid in buds in the spring or early summer.

Type material. — Holotype 9: MEXICO, Baja California {Norte), El Condor,

reared from gall on Quercus dunnii (Dailey #1239). Paratype galls: ARIZONA:

3, Mojave Co., Haulaupi Mountains. CALIFORNIA: 1, San Luis Obispo Co.,

Paso Robles. MEXICO: Baja California (Norte): 3, Mike’s Sky Ranch; 48, El

Condor; 4, Melling Ranch. All specimens in Dailey collection except holotype 9

and 4 paratype galls in USNM, 4 paratype galls in CAS, and 2 paratype galls in

Weld collection.

Discussion.— Heteroecus fragilis is the only gall on Q. dunnii which is mono¬

thalamous, spherical, and has thick, spongy walls (Fig. 4). Trichoteras burnetti

(Fig. 2) is spherical but is polythalamous with thick pubescence and T. rotundula

(Fig. 1) is spherical but is hollow, with paper-thin walls. Disholcaspis truckeensis

is spherical but is polythalamous.

Etymology. — This species is named for the fragile gall texture.

Heteroecus lyoni Dailey and Sprenger, New Species

(Fig. 5)

Unisexual generation female holotype.— Head: coriaceous; from above trans¬

verse; narrower than thorax; cheeks not widened behind eyes; occiput flattened,

elevated above contour of head; malar space 0.5 eye height, not grooved; antenna

filiform, 14 segmented, segment 1 longer than 2, segment 3 longer than 4 or 1

plus 2, segment 14 one and one-half times the length of 13. Thorax: mesoscutum

coriaceous, broader than long, sparsely pubescent; anterior parallel lines distinct,

more than 0.5 mesoscutum length; notauli incomplete, from posterior 0.7 meso-

PAN-PACIFIC ENTOMOLOGIST

scutum length, smooth, polished; parapsidal lines distinct, extending from pos¬

terior 0.5 mesoscutum length; median groove indistinct, 0.1 mesoscutum length;

scutellum rugose, sparsely pubescent; fovea smooth, polished, bare, separated by

a ridge; mesopleuron aciculate. Wings hyaline, setae not projecting beyond margin,

veins yellow-brown, radial cell closed, aerolet 0.2 first cubital cell length. Claws

simple. Abdomen: propodeum reticulate, pubescent, without median carina; length

greater than head plus thorax; tergum II smooth, polished, bare, very short pu¬

bescence on ventral-posterior edge of tergum II; ventral spine long and thin, 10

times as long as high or wide, evenly tapering, sparsely pubescent. Using maximum

head width as base, mesonotum length is 1.4, antenna 2.5, wing 5.4. Body length

is 2.5 mm. Color of head, thorax, and abdomen is red-brown, legs and antennae

are yellow.

Host. — Quercus dunnii.

Gall.— Detachable, monothalamous, hard, brown stem gall about as wide as

tall with flaring sides and flat top 5-11 mm in diameter with an elevated central

spot (Fig. 5, see also Weld, 1957, Fig. 164).

Variation. — Body length range of 27 specimens is 2.3 to 2.7 mm, average 2.6

mm. Both insects from Paso Robles have lighter colored wing veins. The color

range for specimens from 8 miles east of Hemet is from amber to dark brown.

Mesoscutum texture occasionally approaches weak transverse rugosity. Galls from

Paso Robles and Clear Creek, San Benito County, California and some from El

Condor, Baja California Norte, Mexico have nearly straight ribbed sides and a

depressed central top. Those developing from axillary buds typically have a groove

on the side adjacent to the stem.

Biology. — Mature galls were collected in November and insects emerged indoors

in the rearing containers in late winter or early spring. The oviposition site and

alternate generation are unknown.

Type material. — Holotype 2, 12 2, 3 gall paratypes: MEXICO, Baja California

(Norte), El Condor, C. Dailey. Additional paratypes as follows: MEXICO, Baja

California (Norte), Mike’s Sky Ranch, CALIFORNIA, 5 2, 28 galls, Riverside

Co., 10 km E Anza, C. Dailey; 2 2, 68 galls, San Luis Obispo Co., Clear Creek,

J. Tucker. All specimens in Dailey collection except holotype 2, 4 paratype 2 and

galls each in USNM, 4 paratype 2 and galls each in CAS and Weld collections.

Discussion.—Heteroecus lyoni is the only gall on Q, dunnii with the peculiar

flaring sides and flat top as shown in Fig. 5. Disholcaspis chrysolepidis might

rarely be confused with H. lyoni, but the former is mushroom-shaped and usually

occurs clumped together in rows.

Etymologer — This species is named in honor of Mr. Robert J. Lyon who has

worked out many cynipid life cycles including the first Heteroecus life history. He

has provided much valuable assistance and encouragement in these studies.

Heteroecus pacificus Kinsey: New Host Record

Galls of this species were seen at Sedona, Coconino County, Arizona (Dailey

#1289). The known hosts now include Quercus dunnii, Q. chrysolepis, Q. tomen-

tella Engelmann, and Q. vaccinifolia Kellogg.

Acknowledgements

The authors are grateful to Dr. Richard M. Bohart, whose efforts as an instructor

made this paper possible, to Dr. John M. Tucker for the Clear Creek, California

VOLUME 59, NUMBERS 1-4

specimens, Dr. John Burnett for use of his reared specimens and Robert J. Lyon

for use of the Weld collections and manuscript keys, and M. Lynn Kimsey for

illustrations.

Literature Cited

Burnett, John A. 1977. Biosystematics of the new, Oak-Gallwasp genus Weldia of Western North

America. Ph.D. Thesis, University of California, Riverside, California, 257 pp.

Griffin, James R., and John M. Tucker. 1976. Range extension for Quercus dunnii in central Cali¬

fornia. Madrono 23:295.

Tucker, John M., and Horace S. Haskell. 1960. Quercus dunnii in Arizona. Brittonia 12:196-219.

Tucker, J. M., G. H. Holstein, Heather Stout, and Neil Havlik. 1982. Another range extension for

Quercus dunnii in Central California. Aliso 10:321-328.

Weld, L. H. 1957. Cynipid galls of the Pacific slope. Privately printed. Ann Arbor, Michigan, 64

pp.

-. 1960. Cynipid galls of the Southwest. Privately printed. Ann Arbor, Michigan, 54 pp.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 50-51

Diodontus bo hard , a New Species from California’s North

Coast Range (Hymenoptera: Sphecidae)

Lloyd E. Eighme

Pacific Union College, Angwin, California 94508.

Collecting trips to the more remote portions of California’s Siskiyou Mountains,

Marble Mountains, Salmon Trinity Mountains, and Yolla Bolly Mountains since

1965 have added more than 30,000 specimens to the insect collection at Pacific

Union College, Angwin, Napa County, California. Most of the collecting expe¬

ditions involved small groups of students who, with the author, spent about ten

days at a time backpacking into the higher elevations in search of insects. The

North Coast Ranges of California have a unique flora that has been described as

rich in narrowly endemic species which often appear to be relicts restricted to a

specific ecosystem (Whittaker, 1961). We assumed that in an area so vegetatively

and geologically distinctive, we would also find a characteristic insect fauna. More

than 15 years of collecting in these mountains has resulted in a number of new

species and new distributional records. We suspect that there are more, but large

portions of the collection have not yet been determined to species and we are still

looking for systematists who would be willing to determine material for us.

Determination of wasp specimens from our collections has been done mostly

by Dr. Richard Bohart. My first contact with him was in connection with a new

species of Pulverro that I discovered in the Salmon Trinity Mountains in 1966

(Eighme, 1968). I was a bit hesitant to attempt my first description and publication

of a new species, but Dr. Bohart took care of that with his good natured assurance

and assistance. My immediate reaction was one of confidence with such authority

as he telling me I had a good species. Pulverro monticola Eighme has since been

collected over a wide area of northern California mountains.

It was Richard Bohart who encouraged me to work on the genus Diodontus. I

am grateful to him for introducing me to this fascinating group of wasps and I

take pleasure in naming this distinctive new species in his honor.

Diodontus boharti Eighme, New Species

Holotype male.— Black; mandibles (central portion), palps, apical Vi ofpronotal

lobe, spot on anterior portion of tegula, front side of all tibiae golden yellow; basal

tarsomere of foreleg dusky yellow; wings dusky, iridescent, wing veins and stigma

dark brown; labrum shallowly notched, clypeal teeth prominent; flagellomeres

V-X with apical margins arcuate, flagellomeres VTII-X with oval, smooth, shiny

tyli, flagellomere XI with small basal tylus and apical smooth shiny spot; frons

and vertex with large punctures separated by less than puncture diameter, mi¬

crosculpture finely linear, upper Vi of frontal line a weak carina; pronotal collar

with humeral angles sharp but not produced upward; scutum with dense punctures

slightly smaller than those on the head, no microsculpture, admedian lines prom¬

inent, raised and broader than the notauli which are also prominent, parapsidal

lines obscure; scutellum with faint median line; propodeum coarsely reticulate

VOLUME 59, NUMBERS 1-4

with blunt lateral spine posterad from the spiracular flange; abdomen with fine

setigerous punctures connected by fine reticulated microsculpture, tergum VI with

spinose tubercules on posterior margin bearing 3 spines, sternal brushes weak;

basal tarsomeres straight; omaulal ridge strong, sharp, joining postspiracular Ca¬

rina by a lateral extension at right angle.

Allotype female. — Black; mandibles (central portion), spot on anterior portion

of tegula golden yellow; palps brown; tibiae orange-brown with darker spot on

posterior side; wings dusky, iridescent, wing veins and stigma black; clypeal teeth

prominent, lateral teeth twice as long as medial one; frontal spine prominent but

blunt, orbital foveae narrow, extending from mid-orbit to vertex; frons and vertex

with widely scattered large punctures, microsculpture fine reticulation; pronotal

collar with coarse longitudinal striations at crest, humeral angles slightly winged;

scutum with widely scattered punctures of sam e size as those on vertex plus smaller

punctures densely concentrated on anterior portion, admedian lines distinct, no-

tauli faint, parapsidal lines long, distinct; scutellum punctured like posterior scu¬

tum, propodeum coarsely reticulate with blunt lateral spine; abdomen finely punc¬

tured with reticular microsculpture; omaulal carina similar to male but not as

prominent; pygidium flat with lateral border of coarse white setae.

Type material— Holotype 6 and allotype 2: CALIFORNIA, Siskiyou Co., Bear

Basin, 7000', August 9, 1967, Lloyd Eighme, deposited at California Academy

of Sciences, San Francisco. There are 21 paratypes as follows (all from California,

all deposited at Pacific Union College, except as noted): Del Norte Co., Stevens

Camp; Glenn Co., Plaskett Meadows; Siskiyou Co., Rattlesnake Meadow; Trinity

Co., Deadfall Lakes, Mirror Lake, Red Mt. Meadows, Swift Creek (University of

California, Davis), Ward Lake.

Discussion. — The coarse sculpturing of the frons and vertex is distinctive in

this species. D. boharti resembles D. vallicolae in some ways but differs markedly

in that it lacks the modified scutal margin seen in D. vallicolae, and the male has

spinose tubercles on tergum VI which relate it to a different species group. Col¬

oration may vary from the holotype in that darker specimens have no yellow on

the pronotal lobe or tegula. Varying amounts of yellow pigmentation is evident

in most species of this genus, so that structural characteristics have been selected

as much as possible to separate the species. I originally recognized this new species

from a few specimens I collected in Bear Basin, Siskiyou County in 1967. One

of my students, Terry Griswold, collected 23 specimens in the Swift Creek area

of Trinity County in 1972, and since then other specimens have been taken at

various sites in the North Coast Range.

Literature Cited

Eighme, Lloyd E. 1968. A new species of Pulverro from California. Pan-Pac. Ent. 44:261-264.

Whittaker, R. H. 1961. Vegetation history of the Pacific Coast States and the “central” significance

of the Klamath Region. Madrono 16(l):5-23.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 52-54

The Larva of Ammatomus icarioides (Turner)

(Hymenoptera: Sphecidae, Nyssoninae)

Howard E. Evans

Department of Zoology and Entomology, Colorado State University, Fort Col¬

lins, Colorado 80523.

There are few groups of insects more rewarding to ethologically inclined biol¬

ogists than digger wasps. They were favorites of Fabre, and since his time they

have been the subject of innumerable books and scientific papers. Yet for many

years Sphecidae were taxonomically a forbidding group, in which accurate species

identifications were hard to come by and the higher classification fraught with

difficulties. Problems still exist, and perhaps always will, but the picture has

changed dramatically for the better, thanks in large part to Richard Bohart and

the many students he has trained. The rewards of this research and teaching will

be reaped for many years in terms of increased depth of knowledge of these most

elegant of insects.

The present paper purports to fill only a tiny gap in our understanding of sphecid

phylogeny. In fact it poses a puzzle, for the larvae of Ammatomus have several

unusual features which tend to contradict evidence from ethology and from adult

structure. Nesting data (Hook, 1981) suggest that Ammatomus icarioides is a fairly

typical member of the tribe Gorytini, while adult morphology suggests that it

belongs to a complex of gorytine genera (including Sphecius, Kohlia, and others)

having a rather robust thorax, approaching the Stizini (Bohart and Menke, 1976).

This study is based on seven larvae from Brisbane, Queensland, Australia,

taken by Allan Hook in his recent biological study. Several cocoons from this

study were also available. Voucher specimens have been placed in the Colorado

State University Collections. Associated adults were identified by comparison

with the type specimen in the British Museum (Natural History).

Description of Mature Larva

(Figs. 1-7)

Body length 16 mm; maximum width 4.2 mm (Fig. 4). Integument smooth and

largely devoid of setae and spines, although under high magnification scattered

minute setae and surface granules can be detected. Each body segment divided

dorsally into two annulets by a transverse crease, the most posterior annulet with

a pair of weakly defined transverse welts; pleural lobes well developed; anus

terminal. Ten pairs of equally developed spiracles present; atria lined with hex¬

agonal cellules; opening into subatrium armed with a circlet of spines. Maximum

width of head 1.04 mm; length to apex of clypeus 1.08 mm, to apex oflabrum

1.3 mm. Front of head with five somewhat nipple-shaped callosities, three of

them in a series just below and mesad of antennae, two somewhat above antennae

(these callosities are slightly brownish in color in two specimens, unpigmented in

the other five). In lateral view (Fig. 1) only two of these callosities are visible.

Coronal suture distinct; parietal bands indicated by short, unpigmented streaks.

VOLUME 59, NUMBERS 1-4

Figs. 1-7. Ammatomus icarioides. 1-6, mature larva; 7, cocoon. 1, head, lateral view. 2, antenna.

3, mandible. 4, body, lateral view. 5, labium and maxilla, oral aspect. 6, labrum (left half) and

epipharynx (right half). 7, diagrammatic cross section of cocoon pore.

Antennal orbits small, circular, containing a circular brown area in the center;

this brown disc is slightly elevated above the membrane of the orbit and bears

three small sensilla (Fig. 2). Head with scattered punctures, some of which bear

minute setae no more than 3 times as long as width of puncture; clypeus with 24

punctures in lateral groups of 12 each.

PAN-PACIFIC ENTOMOLOGIST

Labrum 0.43 mm wide, apical margin straight, with a row of 22 small sensory

cones; surface strongly convex, with 20 punctures on each side, most of them

bearing short setae; apical margin also with several setae; surface papillose me-

dioapically (Fig. 6). Epipharynx strongly spinulose, sensory areas each with 7

pores. Mandibles 2.1 times as long as their greatest width, weakly tridentate, each

with 5 punctures toward the base laterally, punctures bearing minute setae (Fig.

3). Maxillae with small setae laterally, extensively papillose but papillae grading

into small spinules on mesal margin; palpi and galeae prominent, palpi slightly

longer and stouter than galeae (Fig. 5). Labial palpi equal in length to galeae, but

somewhat stouter; spinnerets paired, not exceeding palpi; labium with several

prominent apical setae.

Discussion

Although the nesting biology of Ammatomus icarioides appears to differ in no

important ways from that of Gorytes and related genera, the larva will not key

readily to Gorytini in either the artificial key of Evans (1959) or the tables of

Evans and Lin (1956). This is because of the absence of a distinct antennal papilla

and the five protuberances on the front of the head. These protuberances are not

as distinct as in Alyssonini and Nyssonini, but they are better developed than the

“two pairs of vertical depressions” and the resulting interspaces described for

Goiytes. The unusual antennae are difficult to explain. They appear to represent

a stage between the development of a true papilla and the presumably ancestral

condition, in which the sensilla merely arise from the membrane of the orbits.

The cocoons provide no assistance in solving this puzzle. As noted by Hook

(1981), there is a single pore with a raised rim at one end. This is actually at the

posterior end, opposite the cap, and resembles the pore in the cocoon of Gorytes,

as figured by Evans (1966). This pore penetrates both the outer and inner layers

of the cocoon and has a thickened rim (Fig. 7). It may represent a precursor of

the more elaborate pores of Sphecius and of Bembicini, which are distributed

around the widest part of the cocoon. In any case, there seem to be no important

differences between the cocoons of Ammatomus and those of Gorytes.

We are left with the conclusions that the larvae of Ammatomus have probably

been subjected to somewhat different selection pressures than those of other Gor¬

ytini and that the genus perhaps stands somewhat farther apart phylogenetically

from Gorytes and from Sphecius than usually recognized.

Literature Cited

Bohart, R. M., and A. S. Menke. 1976. Sphecid wasps of the World. A generic revision. Univ. Calif.

Press, Berkeley, 695 pp.

Evans, H. E. 1959. Studies on the larvae of digger wasps (Hymenoptera, Sphecidae). Part V: Con¬

clusion. Trans. Amer. Ent. Soc. 85:137-191.

-. 1966. The Comparative Ethology and Evolution of the Sand Wasps. Harvard Univ. Press,

Cambridge, Mass., 526 pp.

Evans, H. E., and C. S. Lin. 1956. Studies on the larvae of digger wasps (Hymenoptera, Sphecidae).

Part II: Nyssoninae. Trans. Amer. Ent. Soc. 82:35-66.

Hook, A. 1981. Nesting biology of Tanyoprymnus moneduloides and Ammatomus icarioides. Ann.

Ent. Soc. Amer. 74:409-411.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 55-63

Two New Species in the Genus Philanthus and a Key to the

politus Group (Hymenoptera: Philanthidae ) 1

George R. Ferguson

Systematic Entomology Laboratory, Department of Entomology, Oregon State

University, Corvallis, Oregon 97331.

Bohart and Grissell (1975) arranged the North American Philanthus into seven

species groups and provided a key for the identification of the species. An un¬

described species belonging to their P. politus group was contained in a collection

of philanthine wasps received for identification in 1980 from F. D. Parker (USDA,

Bee Biology and Systematics Laboratory, Utah State University, Logan). Since

this new species was difficult to place in their key, a study of the P. politus group

was undertaken with the objective of preparing a key to accommodate the new

species. A consequence of this study was the finding that the holotype of Philanthus

serrulatae Dunning is a male of the species previously known as Philanthus sioux-

ensis Mickel, a member of the P. politus group. The species in the P. pacificus

group to which Bohart and Grissell (1975) had applied the name P. serrulatae,

nec Dunning, is thus left without a name.

In Part I of this paper P. siouxensis is synonymized under P. serrulatae, and

Philanthus boharti new species is described in the pacificus group to provide a

name for the species previously known as P. serrulatae, nec Dunning. In Part II

Philanthus parkeri new species is described in the politus group; the politus group

is redefined to include Philanthus ventilabris Fabricius; and a key to the politus

group is presented. Abbreviations used for institutional repositories are explained

in Acknowledgments.

Part I. The Identity of Philanthus serrulatae Dunning and

Description of a New Species in the Philanthus pacificus Group

Strandtmann (1946) treated several taxa in the genus Philanthus as subspecies

of Philanthus politus Say. Bohart and Grissell (1975) synonymized Philanthus

texanus Banks under P. politus and elevated the remainder of Strandtmann’s

subspecies to species. They placed P. politus, P. psyche Dunning, P. siouxensis,

and P. tarsatus H. Smith in their politus group; and they placed P. barbiger Mickel,

P. pacificus Cresson, P. pulcher Dalla Torre, and P. serrulatae in their pacificus

group. The politus group has a metapleural lamella (= metanotal lamina of Strandt¬

mann), whereas this structure is lacking in the pacificus group. This placement of

P. serrulatae was in accord with that of Strandtmann (1946 in key, couplet 27)

who stated “No metanotal lamina . . .” in characterizing the species.

Four of the specimens to which Strandtmann (1946) referred in his discussion

of P. serrulatae were studied (OSU). Three of these (10 mi. e. Santa Fe, New

Mexico; Lusk, Wyoming; Clear Creek, Colorado) have a metapleural lamella and

are males of Philanthus siouxensis. The male from Clear Creek, Colorado, bears

1 Oregon Agricultural Experiment Station Technical Paper No. 6589.

PAN-PACIFIC ENTOMOLOGIST

a label “Compared with type P. serrulatae D.” The fourth specimen (Imperial

County, California) is a male without a metapleural lamella and is conspecific

with P. serrulatae, nec Dunning, as interpreted by Bohart and Grissell (1975). An

additional five males, from Imperial County and Los Angeles, California [USNM],

that Strandtmann (1946) had referred to P. serrulatae were also examined and

found to be conspecific with the P. serrulatae, nec Dunning, of Bohart and Grissell

(1975). It was apparent that Strandtmann had two species before him. The male

holotype of P. serrulatae was studied and found to be a male of P. siouxensis and

conspecific with the above males from New Mexico, Wyoming and Colorado.

Philanthus serrulatae Dunning

Philanthus serrulatae Dunning, 1898:154. Male holotype, Denver, Colorado;

ANSP.

Philanthus siouxensis Mickel, 1916:406. Female holotype, Harrison, Nebraska;

NEB. Bohart and Grissell, 1975:6,7 (in key); Bohart and Menke, 1976:566;

Krombein, 1979:1725. New Synonymy.

Philanthus politus serrulatae, Strandtmann, 1946:83 (in part).

Philanthus politus siouxensis, Strandtmann, 1946:91.

Specimens examined. — 71 S, 82 9, including the male holotype of P. serrulatae

and two female paratypes of P. siouxensis.

Distribution. — From South Dakota and Wyoming south through the Rocky

Mountains and high plains to Arizona, New Mexico and Texas. In Mexico it

occurs in the states of Chihuahua, Coahuila, D urango, San Luis Potosi, Zacatecas,

Aguascalientes and Mexico. From Colorado northward dates of capture are mostly

in July, whereas it has been collected from April to October in the southern part

of its range.

Discussion.— The above synonymy leaves the P. serrulatae, nec Dunning, of

Bohart and Grissell (1975) without a name. It is a member of the pacificus group,

as defined by Bohart and Grissell (1975), and is described herewith.

Philanthus boharti Ferguson, New Species

Philanthus politus serrulatae, Strandtmann, 1946:83 (in part).

Philanthus serrulatae, Bohart and Grissell, 1975:17; Bohart and Menke, 1976:

566; Krombein, 1979:1725.

Bohart and Grissell (1975) keyed, figured and discussed this species under the

name P. serrulatae (nec Dunning) and both sexes run to P. serrulatae in their key.

Diagnosis. — Both Sexes: Length 7-9 mm; inner eye margins parallel below the

emargination; maximum head width 1.5 times width of face at apex of eye emar-

gination; interocellar distance greater than ocellocular distance; metapleural la¬

mella absent; forewing costa whitish, at least at base; punctures on terga II—III

about same size as scutal punctures; pale transverse band on terga IV-V biemargin-

ate anteriorly; pale markings whitish with few tinges of yellow. Male: clypeal

brushes whitish, separated medially by half the length of a brush; malar space

about equal to width of pedicel; upper half of face below midocellus with punctures

separated by 1 or 2 puncture diameters and microridges weak or obscure; vertex

width at midocellus greater than half the interocular distance at widest part of

clypeus; femora usually with a red tinge at juncture of black basal portion and

VOLUME 59, NUMBERS 1-4

white apical portion. Female: malar space less than half the width of pedicel; area

laterad of midocellus sparsely punctured; femora usually black, red and white, or

all red.

7>/?es.-Holotype <3: CALIFORNIA, San Diego Co., Borrego Springs, April 26,

1964 (G. & A. Ferguson, CAS). Paratypes: 100 <3, 87 2 from CALIFORNIA, San

Diego Co., as follows: 3 <3, 4 2, same data as holotype; 2 <3, 3 2, same as holotype

except April 27, April 28, April 29; 1 <3, 1 2, Borrego Springs, III-30-60 (M.

Wasbauer); 1 2, Borrego Springs, III-31-73 (C. Goodpasture); 3 <3, Borrego Valley,

sweeping alfalfa, III-26-59 (A. A. Grigarick); 59 <3, 43 2, Borrego Valley, IV-11-

62, IV-6-64, V-23-64, IV-11-69, IV-9-70, IV-2-73 (R. M. Bohart, C. Goodpasture,

E. E. Grissell, J. E. Slansky, M. Wasbauer); 15 6, 9 2, Borrego Valley, dunes, IV-

18-57, IV-20-57 (R. C. Bechtel, R. M. Bohart, J. C. Hall, H. E. Moffitt, E. I.

Schlinger); 8 <3, 9 2, Borrego Valley, Coyote Creek, III-26-59, IV-5-63 (R. M.

Bohart, A. A. Grigarick, M. E. Irwin, F. D. Parker); 1 <3, Borrego State Park, III-

28-77 (J. Slansky, M. Wasbauer); 1 <3, Borrego State Park, IV-17/20-69, Acacia

greggi (R. R. Pinger); 6 2, Borrego State Park, IV-17/20-69 (M. S. & J. S. Was¬

bauer); 7 6, 11 2, Borego [sic], IV-1-53, IV-25-54, IV-27-54, IV-29-54, IV-24-55,

IV-26-55, IV-27-55 (P. D. Hurd, M. Wasbauer). [AZS, UCB, UCD, CAS, CD A,

UID, LAM, OSU, USNM, USU].

Other specimens examined. — 131 <3, 121 2 in addition to the type series. [AZS,

UCB, UCD, CAS, CDA, UID, LAM, OSU, USNM, USU].

Distribution. Mostly in Lower Sonoran deserts from San Diego and Imperial

Counties north to Mono County, California; Clark County, Nevada; Washington

County, Utah; and Mohave and Yuma Counties, Arizona. In Mexico, it occurs

in northwestern Sonora (El Golfo; Sonoita; 39 mi. n. Puerto Penasco); Baja Cal¬

ifornia Norte (Bahia de los Angeles; Rosarito; Mexicali; San Felipe); and in Baja

California Sur (Guerrero Negro) near the border with Baja California Norte.

Flight period. Most dates of capture are from March to May with a peak in

April. A partial second brood is indicated by several October captures as follows:

California: 2 2, San Bernardino Co., 25 mi. s. Ivanpah, X-12/13-58 [UCB]; 1 2

Imperial Co., 2 mi. s. Palo Verde, X-18-59 [UCB]; 1 <3, San Diego Co., 13 mi. e.

Borrego Springs, X-7-67 [AZS]. It may also be bivoltine in the Baja California

peninsula as indicated by a male captured on 11-24-74 (Guerrero Negro) [USU]

and a female taken on VIII-5-73 (22 mi. s. Rosarito) [LAM]. A male and 2 females

were collected at black light near El Golfo, Sonora, Mexico, IV-10-73 [CDA].

Etymology. —I am pleased to dedicate this species to Dr. Richard M. Bohart

in recognition of his contributions to our knowledge of aculeate Hymenoptera,

and especially for his helpful advice and encouragement in behalf of my current

studies. Upon calling the P. serrulatae synonymy to his attention he suggested

that I proceed to publish it.

Discussion. Among the small, finely punctured species of Philanthus with an

anteriorly biemarginate pale band on terga IV-V, P. boharti closely resembles P.

psyche and P. serrulatae in the politus group, and P. pacificus (= P. arizonae

Dunning) and P. pulcher in the pacificus group. Since the metapleural lamella is

sometimes quite small in P. psyche and P. serrulatae , the following comments

may be helpful in separating the species. The metapleural lamella is almost always

pigmented in P. psyche, and in these cases recognition of the structure is relatively

easy.

PAN-PACIFIC ENTOMOLOGIST

The distinct malar space separates males of boharti from pacificus and psyche,

but not from pulcher and serrulatae. In pulcher the forewing costa is reddish to

brown to the base, and the interocellar distance is no greater than the ocellocular

distance. In serrulatae the ocellocular distance is greater than the diameter of the

midocellus, whereas in boharti the ocellocular distance and the midocellus di¬

ameter are subequal. In females the interocellar distance and ocellocular distance

are subequal in pulcher and serrulatae, whereas the interocellar distance is always

distinctly greater than the ocellocular distance in boharti, pacificus, and psyche.

The metapleural lamella separates psyche from the other two species.

Bohart and Grissell (1975) discussed the problem of separating females of

boharti from red-legged females of pacificus. The problem is further complicated

by the presence of females of boharti without reddish markings on the femora

(Old Woman Springs, San Bernadino County, California, IV-17-62; UCB). In

boharti the tergal punctures are somewhat sparser and more irregularly spaced

than in pacificus (see Bohart and Grissell, 1975, Figs. 25-28). Head measurements

show that the eyes of pacificus are somewhat more swollen than in boharti. The

ratio between least vertex width and width of face at the apices of the eye emar-

ginations is the same for both species, but the maximum head width is 1.6 times

the facial width in pacificus and only 1.5 times the facial width in boharti. There

is virtually no overlap in the measurements (n = 10). In addition, the two species

are largely allopatric with boharti being a Lower Sonoran species and pacificus an

Upper Sonoran and Transition zone form.

Part II. The Philanthus politus Group

In this section Philanthus parkeri new species, belonging to the politus group,

is described; the politus group is redefined to include Philanthus ventilabris; and

a key to the politus group is presented.

Philanthus parkeri Ferguson, New Species

(Figs. 1, 2)

Male. — Length 5-7 mm, forewings 4-6 mm; malar space at least 3 A as long as

width of pedicel; clypeal brushes whitish, separated medially by half the length

of a brush; antennal sockets separated from eye margins by about 2 socket di¬

ameters and from each other by about 3 socket diameters; maximum width of

head 1.5 times width of face at apex of eye emargination and 2.7 times least width

of vertex; face at apex of eye emargination 1.8 times least width of vertex; ocel¬

locular distance about equal to diameter of lateral ocellus and distinctly less than

diameter of midocellus; flagellomere I about as long as combined length of scape

and pedicel; interantennal groove weak to absent; upper face densely punctured

with numerous microridges, sculpture continuing into area between midocellus

and eye margin; ocellar triangle with few large, setigerous punctures of about the

same size as those behind the ocelli; pubescence of lower face and clypeus bristly,

recumbent to sub-recumbent; dense, setigerous punctures of lower face continuing

onto adjacent margins of clypeus, remainder of clypeus very sparsely punctate;

pronotal ridge thick, rounded; scutal punctures dense antero- and posteromedially,

separated and irregular elsewhere, pit-like; scutellum sparsely punctate, groove

obscure or absent; metapleural lamella prominent, pigmented; subalar carina

variably pigmented, with prominent translucent lamellae laterally; propodeal en-

VOLUME 59, NUMBERS 1-4

1 2

Figs. 1, 2. Philanthus parkeri, dorsal view of head. 1, male. 2, female. Actual head width 1.9 mm

for male and 2.2 mm for female.

closure with large patch of punctures on each side, furrow more densely and

irregularly sculptured; many punctures on tergum II smooth-rimmed and larger

than scutal punctures; sterna III to V with coarse punctures mostly separated by

less than 1 puncture diameter; lateral pubescence on sterna IV and V whitish, not

dense; without patches of dense pubescence on sterna VI or VII.

Female.— Length 6-8 mm, forewings 5-7 mm; malar space to % pedicel

width; antennal sockets separated from eye margin by about 1.5 socket diameters

and from each other by about 2.5 socket diameters; maximum width of head 1.6

times width of face at apex of eye emargination and 2.6 times least width of vertex;

ocellocular distance 1.1 times diameter of lateral ocellus and 0.9 times diameter

of midocellus; interocellar distance 1.2 times diameter of midocellus; flagellomere

I longer than scape; sculpture as in male.

Coloration. —Vale markings mostly bright yellow with occasional whitish

blotches; males almost entirely yellow in dorsal view, with black band across the

ocellar area often present, three narrow black stripes usually indicated on scutum;

median black stripe on popodeum usually present; prosternum, mesosternum

and metapleural area black; legs mostly yellow except femora basally, trochanters

and large spot on coxae black; tarsi yellow to reddish; wings clear hyaline, veins

fulvous to reddish; narrow anterior black bands on apical terga; sterna II to V

with yellow bands or large spots. Female colored much like male except that black

area of vertex extends laterally down inner orbits to antennal sockets, scutum

with 3 broad longitudinal black stripes, and more extensively black on propodeum.

Tergum VI in female and terga VI and VII in male extensively yellow. Terga IV

and V not biemarginate anteriorly.

Types. — Holotype 6: UTAH, Emery Co., 2 mi. n. Goblin Valley State Preserve,

5000 ft., Aug. 25, 1980 (A. S. Menke, F. D. Parker, Kurt A. Menke, USNM).

Paratypes: 26 6, 67 5, all from Emery Co., UTAH, as follows: 5 6, 3 2, same data

as holotype; 8 6, 63 2, Goblin Valley, in sand dunes, IX-16-79, IX-16-80 (T.

Griswold, F. D. Parker, D. Veirs); 13 6, 1 2, San Rafael Desert, Little Gilson

Butte, 5000-5100 ft., VIII-19, 24-27-1980 (A. S. Menke, F. D. Parker, Kurt A.

Menke) [UCD, CAS, UID, OSU, USNM, USU].

Other specimens. —21 6, 6 2, as follows: ARIZONA: Coconino Co.: 15 mi. n.

PAN-PACIFIC ENTOMOLOGIST

The Gap, IX-21-66, 1 2 (R. Rust, P. Torchio, G. Wood, N. Yousef, USU). IDAHO:

Butte Co.: 7 mi. e. Howe, IX-6-67, 12 <5 (W. F. Barr, UID). Lincoln Co.: 5 mi.

e. Dietrich, IX-2-65, 1 <3, 1 2 (R. L. Westcott, UID); 7 mi. e. Dietrich, VIII-8-69,

3 6 (W. F. Barr, UID). Owyhee Co.: Sand Dune Lake, IX-9-63, 1 3 (W. F. Barr

and G. B. Hewitt, UID). NEBRASKA: Sioux Co.: Glen Canyon, VIII-25-59, 2

2 (W. E. LaBerge and O. W. Isacson, NEB). NEW MEXICO: Torrance Co.: Town

of Gran Quivira, 6500 ft., VIII-20-67, 1 S (H. B. Leech, CAS). Duran, VII-13-

59, 3 5 (E. G. Linsley, UCB). UTAH: Dixie State Park, VI-13-61, 1 6 (G. E.

Bohart, OSU). Millard Co.: Flowed, VIH-20-61, 1 2 (G. E. Bohart, OSU). Juab

Co.: 12 mi. s. Eureka, VII-18-58, 1 6 (J. W. MacSwain, UCB). Washington Co.:

1 mi. s. Toquerville, IX-5-79, 4 6, 1 2 (J. C. and E. M. Hall, UCR).

Variation.— Most of the major collections in the western United States and

several national collections have been searched for additional specimens. Thirty-

three specimens were found which I am unable to separate from the type series

on morphological grounds, but they differ in coloration. The specimens from

outside the type locality are less maculated than the type series. Pale markings

are mostly whitish with some pale yellowish blotches. The four pale spots on the

scutum tend to form longitudinal stripes, narrow to moderate in width. The

mesopleuron and propodeum are pale spotted but not extensively so. The pale

bands on terga III to V are biemarginate anteriorly. In the female the pale band

on tergum I is interrupted medially, and the band on tergum II tends to enclose

black spots laterally. Terga VI and VII are immaculate in the male.

A high degree of intraspecific color variation is well known in Philanthus and

related genera from western North America, and several other species of Philan¬

thus collected in Emery County, Utah, are among the more highly maculated

specimens of the respective species. I therefore conclude that the type series and

the other specimens are color variants of the same species.

Distribution .—From southern Idaho to northern Arizona west of the Rocky

Mountains; and western Nebraska and central New Mexico east of the Rocky

Mountains.

Flight period .—The type series was collected between August 19 and September

16 with August collections composed mostly of males and September collections

composed mostly of females. The specimens not included in the type series were

collected between August 8 and September 21, with the exception of one male

collected in June (Utah) and three males collected in July (New Mexico). The

species is apparently univoltine.

Etymology :.—I am pleased to name this species for Dr. Frank D. Parker who

sent me the first specimens collected with the suggestion that they might represent

a new species.

Discussion .—The politus group and the monotypic ventilabris group of Bohart

and Grissell (1975) have a metapleural lamella which is absent in other North

American species groups. Elsewhere in the Philanthidae the metapleural lamella

occurs only in the genera Ctypeadon and Listropygia of the subfamily Aphilan-

thopinae (Bohart and Menke, 1976). The politus and ventilabris groups also have

short, pale clypeal brushes in the males, and the inner margins of the lower eye

lobes are parallel in both sexes.

In P. ventilabris the transversely grooved pronotal collar and contiguous coarse

tergal punctures of both sexes, and the enlarged, flattened apical flagellomere of

VOLUME 59, NUMBERS 1-4

the male are autapomorphies, which I do not believe are sufficient in themselves

to justify a group separate from the other species having a metapleural lamella.

Philanthus albopilosus Cresson was included in the politus group by Bohart and

Grissell (1975), as it is here, but it too has certain unique characters. The thin

head and wide vertex combined with an unusually wide interocellar distance are

autapomorphies which serve to isolate the species from all others in the genus.

The group relationship is based on the same shared characters mentioned above.

In the key T have placed P. ventilabris and P. albopilosus in separate monotypic

subgroups to indicate their more isolated positions.

The politus subgroup forms a compact, homogeneous group of five closely allied

species. Of these, P. politus, P. serrulatae (= P. siouxensis), and P. tarsatus are

separated from each other primarily on the basis of body sculpture and color

characters, since the ocellocular proportions and head, face and vertex ratios are

very similar in the three species.

P. psyche and P. parkeri n. sp. have a very short ocellocular distance, but they

have arrived at this condition by two different routes. In males of psyche the

maximum width of the head is 1.7 times the width of the face (measured at the

apices of the eye emarginations), whereas this ratio is 1.5 in the other four species

of the subgroup. The width of the face is 1.4 times the least vertex width in

psyche— as it is in politus and serrulatae. This indicates that the higher head/face

ratio in psyche is due to larger eyes relative to the other two species. The reduced

malar space and reduced ocellocular distance in psyche have apparently resulted

from this enlargement of the eyes.

In males of parkeri the head/face ratio is 1.5, as in politus, serrulatae, and

tarsatus ; but the face/vertex ratio has increased to 1.8 compared to 1.4 in politus,

serrulatae and psyche, and 1.5 in tarsatus. The vertex has narrowed in parkeri

with respect to the face, but the eyes have not perceptibly enlarged. The malar

space remains, and both the interocellar and ocellocular distances have been

reduced.

P. parkeri is presumed to be more closely related to P. tarsatus than to any

other species in the subgroup because of the coarse punctures on the sterna of the

two species. The denser sculpture of the upper face and ocellar area of the two

species is shared with P. politus but not with P. serrulatae or P. psyche . In addition,

the face/vertex ratio of P. tarsatus is 1.5, indicating a trend toward a narrowing

of the vertex which reaches its culmination in the very narrow vertex of P. parkeri.

The narrow vertex and compact ocellocular area of P. parkeri are unique within

the politus group and make it difficult to place in Bohart and Grissell’s (1975)

key. The following key is modified and adapted from their key to accommodate

P. parkeri.

Key to Species of the Philanthus politus Group

1. Pronotal collar thick, shallowly, transversely grooved dorsally; intero¬

cellar distance less than ocellocular distance ( ventilabris subgroup) . . .

. ventilabris Fabricius

- Pronotal collar rounded, convex; interocellar distance equal to or greater

than ocellocular distance. 2

2. Head thin in side view, not swollen behind the eyes; interocellar distance

greater than distance from lateral ocellus to posterior margin of head,

PAN-PACIFIC ENTOMOLOGIST

and greater than two lateral ocellus diameters (albopilosus subgroup)

. albopilosus Cresson

- Head swollen behind the eyes; interocellar distance about equal to or

less than distance from lateral ocellus to posterior margin of head, and

not greater than two lateral ocellus diameters {politus subgroup). 3

3. Males. 4

- Females. 8

4. Ocellocular distance about equal to lateral ocellus diameter and slightly

less than midocellus diameter . 5

- Ocellocular distance distinctly greater than lateral ocellus diameter and

at least slightly greater than midocellus diameter . 6

5. Malar space less than half the width of antennal pedicel; punctures of

tergum II uniformly distributed and about same size as scutal punctures

. psyche Dunning

- Malar space at least % the width of antennal pedicel; punctures of tergum

II somewhat unevenly distributed, of uneven size, with numerous punc¬

tures smooth-rimmed and larger than scutal punctures .

. parked Ferguson, new species

6. Upper face and area laterad of midocellus sparsely punctate with many

large polished areas . serrulatae Dunning

- Upper face densely punctured, interspersed with many fine ridges, sculp¬

ture extending into area laterad of midocellus . 7

7. Hypoepimeron with dense, contiguous punctures and many microridges;

pale band of tergum I broadly contiguous. tarsatus H. Smith

- Hypoepimeron polished at least on anterior half, with scattered fine

punctures; pale band of tergum I interrupted. politus Say

8. Ocellocular distance about equal to diameter of midocellus; forewing

costa whitish on basal half; conspicuous, recumbent, bristly pubescence

on lower face between upper clypeal lobe and eye margin . 9

- Ocellocular distance distinctly greater than diameter of midocellus; fore¬

wing costa reddish or brown to base; pubescence of lower face fine, not

recumbent .. 10

9. Interocellar distance about twice the diameter of the lateral ocellus;

sterna III-V sparsely, shallowly punctate; width of face at apices of eye

emarginations 1.4 to 1.5 times least width of vertex . psyche Dunning

- Interocellar distance much less than twice the diameter of lateral ocellus;

sterna III-V coarsely, rather densely punctate; width of face at apices of

eye emarginations 1.7 to 1.8 times least width of vertex .

. parked Ferguson, new species

10. Hindtarsi black or heavily infuscated; punctures of mesopleuron, es¬

pecially hypoepimeron, dense, contiguous, with numerous microridges

. tarsatus H. Smith

- Hindtarsi yellowish to reddish; punctures of mesopleuron well separated

by thick, polished ridges and anterior part of hypoepimeron shiny with

sparse punctures . 11

11. Hindfemora red, or black and red; scutal punctures sparse and irregular

with numerous large polished areas . serrulatae Dunning

- Hindfemora black and yellow; scutal punctures dense and evenly spaced

at least over anterior Vs of scutum . politus Say

VOLUME 59, NUMBERS 1-4

Acknowledgments

I am indebted to the following individuals and institutions for the loan of

specimens used in this study (abbreviations as used in the text are in brackets):

D. Otte, Academy of Natural Sciences of Philadelphia [ANSP]; F. F. Hasbrouk,

Arizona State University, Tempe [AZS]; J. A. Powell, University of California,

Berkeley [UCB]; R. M. Bohart, R. O. Schuster, University of California, Davis

[UCD]; S. I. Frommer, University of California, Riverside [UCR]; W. J. Pulawski,

California Academy of Sciences, San Francisco [CAS]; M. S. Wasbauer, California

Department of Food and Agriculture, Sacramento [CDA]; W. F. Barr, University

of Idaho, Moscow [UID]; R. R. Snelling, Los Angeles County Museum of Natural

History, Los Angeles [LAM]; B. C. Ratcliffe, University of Nebraska State Mu¬

seum, Lincoln [NEB]; J. D. Lattin, Oregon State University, Corvallis [OSU]; A.

S. Menke, Systematic Entomology Laboratory, USDA, % United States National

Museum of Natural History, Washington, D.C. [USNM]; F. D. Parker, Utah State

University, Logan [USU].

I thank Bonnie T. Hall, Systematic Entomology Laboratory, Oregon State Uni¬

versity, for preparation of the illustrations.

Literature Cited

Bohart, R. M., and E. E. Grissell. 1975. California wasps of the subfamily Philanthinae. Bull. Calif.

Insect Survey 19:1-92.

Bohart, R. M., and A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. University

of California Press, Berkeley and Los Angeles, ix + 695 pp.

Dunning, S. N. 1898. Notes on Philanthus. Can. Ent. 30:152-155.

Krombein, K. V. 1979. Sphecoidea, pp. 1573-1740. In K. V, Krombein, Paul D. Hurd, Jr., David

R. Smith, and B. D. Burks, Catalog of Hymenoptera in America north of Mexico, v. 2:1199—

2209. Smithsonian Inst. Press, Washington, D.C.

Mickel, C. E. 1916. New species of Hymenoptera of the superfamily Sphecoidea. Trans. Amer. Ent.

Soc. 42:399-434.

Strandtmann, R. W. 1946. A review of the North American species of Philanthus north of Mexico.

Ohio State Univ. Press, Columbus, 126 pp.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 64-77

Heterotardigrada of Venezuela (Tardigrada)

A. A. Grigarick, R. O. Schuster, and D. R. Nelson

(AAG, ROS) Department of Entomology, University of California, Davis, Cal¬

ifornia 95616; (DRN) Department of Biological Sciences, East Tennessee State

University, Johnson City, Tennessee 37614.

About 100 species of tardigrades are known from South America from the

countries of Argentina, Bolivia, Brazil, Chile, Columbia, and Paraguay (Ramaz -

zotti, 1972). In 1979 tardigrades were collected primarily in mosses and lichens

from 10 sites in the northwestern section of Venezuela. The area collected rep¬

resents only one-third of the country but over 20 species were identified in the

class Eutardigrada and 14 in the Heterotardigrada. Only Brazil shows a larger

species representation in the literature.

The Heterotardigrada found in Venezuela includes species in Oreella, Mopse-

chiniscus, Bryodelphax, Echiniscus, and Pseudechiniscus. Of the 14 species, five

are new—one species each of Oreella and Bryodelphax, and three species of

Echiniscus in the arctomys complex. Holotypes are deposited at the University

of California, Davis, and voucher specimens at the University of Zulia, Maracaibo.

Materials and Methods

Mosses, lichens, and liverworts were removed from various plant and mineral

substrates, placed in paper bags, and air dried—at times with artificial heat. Spec¬

imens were processed in the United States and prepared for light microscopy and

scanning electron microscopy as indicated in Horning, Schuster and Grigarick

(1978). One or more species of Heterotardigrada were collected from the following

sites:

El Tucuco, Zulia. VI-24-79. Tropical forest in foothills of Sierra De Perija, bor¬

dering a grassland biome, alt. 700 m, precipitation 2000 mm.

La Carbonera, Merida. VI-30-79. Subtropical cloud forest in Sierra Nevada range

of Andes, alt. 2000 m, precipitation 1800 mm.

Merida, Merida. VII-3-79. Sierra Nevada range of Andes, alt. 1627 m, precipi¬

tation 1800 mm.

La Mucuy, Merida. VII-4-79. Sierra De St. Domingo range of Andes, alt. 1800

m, precipitation 1800 mm.

Barrancas, Barinas. VII-6-79. Foothills of Sierra De St. Domingo range and edge

of Llanos Planes, alt. 200 m, precipitation 1280 mm.

Yacambu, Lara. VII-9-79. Subtropical rainforest, alt. 1500 m, precipitation 1300

mm.

Rancho Grande, Aragua. VII-14-79. Tropical rainforest, alt. 1130 m, precipitation

1720 mm.

Tardigrades in the class Heterogardigrada are recognized by the presence of a

lateral clava and cirrus between the head and scapular segments. Both families,

Echiniscidae and Oreellidae, of the Order Echiniscoidea occur in Venezuela.

VOLUME 59, NUMBERS 1-4

The Echiniscidae typically have hve transverse segments, designated by letters

A through E, also referred to as the head (A), scapular (B) and terminal (E)

segments. Median plates are numbered from anterior to posterior. Lateral spines

occur at the post-lateral angles of the transverse segments and bear the same letter.

Dorsal spines are identified by the segment letter and superscript “d.” Pseud-

echiniscus has an additional segment (pseudosegmental plate) between segments

D and E. The following keys will distinguish the taxa found in this study.

Key to Families and Genera

1. Dorsum without distinct plates .Oreellidae— Oreella

- Dorsum with distinct plates .Echiniscidae —2

2. Head without buccal cirri . Mopsechiniscus

- Head with buccal cirri . 3

3. Pseudosegmental plate present. Pseudechiniscus

- Pseudosegmental plate absent . 4

4. Median plates 1 and 2 transversely divided . Bryodelphax

- Median plates 1 and 2 not divided . Echiniscus

Key to Pseudechiniscus

1. Posterior margin of pseudosegmental plate with two spines or lobes;

granulation large (0.7-1.25 pm)’, cuticle without pores .

. P. novaezeelandiae (Richters)

- Posterior margin of pseudosegmental plate without spines or lobes; gran¬

ulation small (0.5-0.8 pm); cuticle with pores . .P. suillus facettalis Peterson

Key to Echiniscus

1. Lateral spines present only on plate A (cirrus A). 2

- Lateral spines present on plate A and one or more other plates. 6

2. Cuticle not green . 3

- Cuticle green . 5

3. Cuticle of dorsal plates consists of pores (light spots) and granules (dark

spots); spines smooth . 4

- Cuticle of dorsal plates consists of pores only; E spines barbed dorsally

and sometimes ventrally . E. angolensis da Cunha and Ribeiro

4. Some dorsal or lateral spines as long as spine E; legs II and III without

minute spine . E. virginicus Riggin

- Dorsal and/or lateral spines shorter than spine E; legs II and III with

minute spines. E. perannatus Murray

5. Cuticle of dorsal plates with pores . E. viridissimus Peterh

- Cuticle of dorsal plates without pores. E. viridus Murray

6. Cuticle of dorsal plates with pores (light spots only). 7

- Cuticle of dorsal plates with pores (light spots) and granules (dark spots)

.. 8

7. Scapular and terminal plates with longitudinal and transverse bands lack¬

ing pores . E. kofordi Schuster and Grigarick

- Scapular and terminal plates without such bands.

. E. mosaicus Grigarick, Schuster, and Nelson, n. sp.

8. Cuticle of dorsal plates with regularly distributed pores; anterior margins

PAN-PACIFIC ENTOMOLOGIST

of plates C and D with pores.

. E. aliquantillns Grigarick, Schuster, and Nelson, n. sp.

- Cuticle of dorsal plates with irregularly distributed pores; anterior margins

of plates C and D without pores.

. E. marginoporus Grigarick, Schuster, and Nelson, n. sp.

Genus Oreella Murray, 1910

The genus Oreella is in the family Oreellidae. It is distinguished from the other

genera in the family, which are aquatic, by its indistinctly eight-segmented body

lacking dorsal plates and by the presence of a short median caudal projection.

According to the diagnostic characters of the genus (Ramazzotti, 1972:136), the

claws are not spurred; however, the inner claws of the New Zealand specimens

(Horning et al, 1978) of Oreella minor Ramazzotti do have spurs, as does the

single specimen of Oreella from Venezuela.

Oreella breviclava Grigarick, Schuster, and Nelson, New Species

(Figs. 1, 2)

Holotype female.— Length excluding legs IV, 174 gm; including legs IV, 191

gm: width, 81 gm. No eyes. Cuticle colorless, with very fine granulation over

entire surface; small hemisphericle tubercles visible along margins of cuticle,

present on dorsal and lateral cuticle; short, median, conical, caudal papilla between

legs IV. Head with acuminate buccal cirri; internal cirrus 10 gm long; external

cirrus, 14 gm long; buccal papilla 5.7 ^m long by 2 ^.m wide. Cirrus A, 19 gm

long; clava short, 5 jum long by 2 gm wide. Leg I apparently without spine; papilla

on leg IV 2 gm long by 1 gm wide; without dentate fringe on legs IV. Strong spurs

on internal claws of all legs; claws 10 gm long.

Etymology.— Latin: brevi, short; clava, for the short clava.

Discussion.—A single specimen was collected from one locality in Venezuela.

It is very similar to Oreella minor but differs in the following respects: relative

lengths of the internal and external cirri, the presence of buccal papillae, claw

length, the absence of a spine on leg I, and the length of the clava. The internal

cirrus is half the length of the external in O. minor, but greater than half the length

(%) in the Venezuelan specimen. In his illustration of O. minor from Chile,

Ramazzotti (1964) omitted the buccal papillae because he was unable to detect

them on the eight specimens he studied; our specimen has distinct buccal papillae.

The claws of O. minor are smaller (3-4 gm) than those of our specimen (10 gm).

An obvious spine is present on leg I of O. minor. ; no such spine is apparent on

our specimen. Ramazzotti (1972) suggested that specimens with long, more or

less curved clavae (11-18 jtim) are probably males of O. minor, while those with

short straight clavae (3.5-5 gm) are females, corresponding to the hypothesis for

Halechiniscus proposed by Richters (1909) and Schulz (1955). The specimen from

Venezuela is a female based on the structure of the gonopore, and therefore may

actually be O. minor if Ramazzotti’s hypothesis is correct.

Another species, Oreella vilucensis Rahm, was described from Chile. This species

is characterized by the presence of two lateral conical spines and the absence of

clavae, which, if correct, would be unique in the Echiniscoidea.

A third species, Oreella mollis Murray, is a larger species that has a long thin

VOLUME 59, NUMBERS 1-4

clava, very long cirri A, no fine granulation (although it may have been over¬

looked), marked body segmentation, and internal claws without spurs.

Distribution. — Holotype, La Carbonera, 1 specimen.

Genus Mopsechiniscus du Bois-Reymond Marcus, 1944

The genus Mopsechiniscus in the family Echiniscidae is characterized by the

absence of the buccal cirri and the presence of two lateral oval cephalic structures

and a small median process dorsal to the buccal aperture. Two species have been

reported from South America: Mopsechiniscus imberbis (Richters) from Chile and

Brazil; Mopsechiniscus granulosus Mihelcic from the Andes in Argentina.

Mopsechiniscus imberbis (Richters), 1907

(Figs. 3-5)

Our specimens illustrate a wide range of variability in the presence and lengths

of the spines and filaments. Ramazzotti (1972) reported variations found in three

populations of M. imberbis, from South Georgia, Brazil and Chile. The South

Georgia specimens have lateral C spines and D filaments; the Brazilian, with C

spines only (sometimes missing); and the Chilean, with lateral C and D filaments

and sometimes small B spines. Our specimens have a combination of no spines;

lateral spines at D only or C and D; short lateral spines at C and D and a strong

spine at the posterior margin of the pseudosegmental plate; or strong or short

spines only on the pseudosegmental plate. A single larva has dorsal spines at the

posterior margins of median plates 1 and 2 and the pseudosegmental plate. In

general, the number and length of the spines decrease with an increase in the body

length of the animal. The opposite situation occurs in other armored tardigrades

such as Echiniscus and Pseudechiniscus. Our specimens also have spurs at the

base of the inner claws on all legs, but they are much less developed on legs I-

III. The cuticle is ornamented with irregular-sized raised dark granules.

The cuticle of Mopsechiniscus granulosus is ornamented, according to Mihelcic

(1967), inside the cuticle itself, not on the surface. There are short B spines, long

lateral C and D filaments, and the caudal margin of the pseudosegmental plate

has two lobes bearing a distal spine.

Distribution. — La Carbonera, 5 specimens; Merida, 1.

Genus Pseudechiniscus Thulin, 1911

The genus Pseudechiniscus in the family Echiniscidae is characterized by the

presence of a transverse pseudosegmental plate between the second paired plates

(D) and the terminal plate. Six species have been reported from South America

from Chile, Bolivia, Argentina, Brazil, and Columbia. The two species collected

in Venezuela were recovered from samples at four of the seven areas heterotar-

digrades were found. Most of the collections were made in the higher mountainous

areas of the Andes.

Pseudechiniscus novaezeelandiae (Richters), 1908

(Figs. 6-8)

Our specimens agree with the description in Ramazzotti’s monograph (1972),

and with specimens from New Zealand in our collection. The cuticular granulation

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-11. 1, 2. Oreella breviclava. 1, dorsal aspect; 2, detail of legs and claws IV. 3-5. Mops-

echiniscus imberbis. 3, dorsal aspect; 4, cuticle of scapular plate; 5, exterior and interior claws leg IV.

6-8. Pseudechiniscus novaezeelandiae. 6, dorsal aspect; 7, cuticle of scapular plate; 8, exterior and

interior claws leg IV. 9-11. Pseudechiniscus suillus facettalis. 9, dorsal aspect; 10, cuticle of scapular

plate; 11, exterior and interior claws leg IV.

is coarser than on Pseudechiniscus s. facettalis. The granules are larger on the

interior margin of the scapular plate, median plates 1 and 2 and the anterior halves

of the paired plates. Two spines, variable in size, are usually, but not always,

present on the posterior margin of the pseudosegmental plate. A thin basal spur

is present on the inner claw of all legs.

Discussion.—Pseudechiniscus n. novaezeelandiae has been previously reported

VOLUME 59, NUMBERS 1-4

in South America from Brazil, Columbia and Argentina. It is difficult to distinguish

from the subspecies P. n. marinae Barros and P. pseudoconifer Ramazzotti and

according to Ramazzotti (1972), the specimens from Brazil identified as P. n.

marinae by Barros were probably P. pseudoconifer. The cuticular granule size and

absence of cuticular pores are useful in separating P. n. novaezeelandiae from P.

s. facettalis.

Distribution .—La Mucuy, 26 specimens; Merida, 13; Barrancas, 1; La Carbo-

nera, 5.

Pseudechiniscus suillus facettalis Petersen, 1951

(Figs. 9-11)

Our. specimens agree with the description in Ramazzotti’s monograph (1972).

The terminal plate is faceted, and there is a tiny spur at the base of the inner

claws on all legs. The cuticle has a fine granulation with ridged areas of thickened

cuticle; a few pores are scattered irregularly in the cuticle.

Discussion .—The suillus group includes Pseudechiniscus suillus facettalis Pe¬

tersen and Pseudechiniscus suillus franciscae Barros. The nominate subspecies is

known from five South American countries. The subspecies P. s. facettalis has

been reported from Tierra del Fuego (Ramazzotti, 1972). Originally described

from Brazil, P. s. franciscae differs from our specimens by the presence of smooth

cuticle between the plates and the absence of spurs on all claws. A similar species,

Pseudechiniscus juanitae Barros, also described from Brazil, has “large” granules,

especially on the pseudosegmental plate and the terminal plate (Barros, 1939).

Distribution. — La Mucuy, 31 specimens; Merida, 39; Barrancas, 1; La Carbo-

nera, 6.

Genus Bryodelphax Thulin, 1928

The genus Bryodelphax is in the family Echiniscidae. Bryodelphax is distin¬

guished from other genera in the family by the transverse division of the first and

second median plates with the third median plate undivided and highly developed;

in addition, the terminal plate lacks the two notches along the posterior margin.

Bryodelphax crossotus Grigarick, Schuster, and Nelson, New Species

(Figs. 12-14)

Holotype female. —Length excluding legs IV, 135 pm. Cuticle colorless, with

granulation and pores on all plates, forming indistinct ridges on scapular and

terminal plates. Median plates 1 and 2 divided, median plate 3 undivided, tri¬

angular in shape. Head with acuminate buccal cirri internal cirrus 5 pm long,

external cirrus, 10 pm long; buccal papilla 2 jum long by 1.9 pm wide. Cirrus A,

30 pm long; clava, 2.5 pm long by 1.9 pm wide. No apparent spine on leg I; papilla

on leg IV, 4.8 pm long by 2.8 pm wide; non-dentate fringe on leg IV, thickened

irregular, almost scalloped edge, granulate above. Claws 8 pm long; inner claws

with spurs at base on all legs; all legs microgranulate.

Etymology. — Greek: krossotus, fringed, for the non-dentate fringe on legs IV.

Discussion. —Bryodelphax ortholineatus Bartos, B. parvulus Thulin and B. do-

minicanus Schuster & Toftner are species from South or Central America that

have the dorsal collar of leg IV with a smooth distal margin. This margin is

somewhat irregular on B. crossotus but not dentate. The cuticles of ortholineatus

PAN-PACIFIC ENTOMOLOGIST

Figs. 12-25. 12-14. Bryodelphax crossotus. 12, dorsal aspect; 13, cuticle of scapular plate; 14,

claws of leg IV. 15-17. Echiniscus viridis. 15, dorsal aspect; 16, cuticle of scapular plate; 17, claws leg

IV. 18-21. Echiniscus viridissimus. 18, dorsal aspect; 19, cuticle of scapular plate; 20, dorsal denticles

leg IV; 21, claws leg IV. 22-25. Echiniscus augolensis. 22, dorsal aspect; 23, cuticle of scapular plate;

24, claws leg IV; 25, dorsal denticles leg IV.

and parvulus and the shape of the median plates are different from crossotus. This

species is similar to dominicanus but the cuticle does not have the large depressions

(fossae) of dominicanus.

Distribution. — Holotype, Rancho Grande; paratypes: El Tucuco, 13 specimens;

Rancho Grande, 1.

Genus Echiniscus C. A. S. Schultze, 1840

The genus Echiniscus is in the family Echiniscidae Thulin, 1928. Echiniscus is

distinguished from the other genera by undivided median plates 1, 2, and 3, and

VOLUME 59, NUMBERS 1-4

by the presence of notches along the posterior margin of the terminal plate. In

Ramazzotti’s key (1972), species of Echiniscus are grouped according to Marcus:

the groups arctomys, spinulosus, scrofa, blumi, trisetosus, and biunguis. The arc-

tomys group consists of the species of Echiniscus that have no lateral appendages

other than cirrus A, although dorsal appendages may be present. The other groups

are distinguished by the number and type (spine, filament) of lateral appendages.

Nine species of Echiniscus were collected in Venezuela, three of which are new

species in the arctomys group.

Echiniscus viridis Murray, 1910

(Figs. 15-17)

Our specimen fits the description in Ramazzotti’s monograph (1972). Cirrus A

is 38 jum long. The cuticle is green and ornamented with clusters of small dark

green granules which form slight tubercles which appear as dark spots. A minute

basal spur is present on the inner claws of all legs. The dentate fringe has short

uneven spines with very irregular spacing in between the spines; the arrangement

of these dorsal leg spines is very distinctive and unique in the species complex.

Discussion.— Echiniscus viridis, known from Brazil, is one of four described

species in the viridis complex. The complex consists of green Echiniscus with no

lateral or dorsal appendages other than cirrus A. Echiniscus perviridis Ramazzotti

has a pattern of raised granules similar to that of E. viridis ; however cirri A are

up to 150-170 ym long. Echiniscus rufoviridis Bois-Reymond Marcus, described

from Brazil, is green caudally only, with short cirri A (30-40 /urn). The cuticular

pattern of Echiniscus viridissimus Peterfi consists of both pores and dark raised

granules. The cuticle of E. viridis is pigmented in areas indicated by denser stip¬

pling (Fig. 15) and the dorsal plates, except for the head, show a rather uniform

pattern of discrete granules.

Distribution. — Rancho Grande, 1 specimen.

Echiniscus viridissimus Peterfi, 1956

(Figs. 18-21)

Our specimens fit the description of Ramazzotti’s monograph and agree with

determined specimens in the collections at ETSU and UCD. Cirrus A is 75 /urn

long. The cuticle is green and ornamented with a very fine regular granulation.

The scapular and terminal plates have pores, which appear as light spots. The

paired plates have an anterior zone of dark granules which appear separated from

the posterior zone by a transverse stripe.

Discussion.— The dorsal plating of this species is greenish in the same areas as

E. viridis. Whereas the cuticular pattern of the dorsal plates of E. viridis consists

of discrete granules, that of E. viridissimus is predominantly of pores (light spots)

except for a minutely granular head and granules on the anterior of plate pairs C

and D and on the area between plates D and the terminal plate.

Distribution. — Rancho Grande, 11 specimens.

Echiniscus angolensis da Cunha and Ribeiro, 1964

(Figs. 22-25)

Body length excluding legs IV, 225 /urn. Cuticle with pores only, slightly larger

on median plates. Cephalic plate with rostral zone of pores. Cirrus A, 29 /urn long.

PAN-PACIFIC ENTOMOLOGIST

Short C, D spines (5-6 ^urn); robust D d spine, 17 pm, may have rough or spiny

appearance; robust, rough (spiny) E spine, 16 pm. Terminal plate with distinct

notches. Dentate fringe on legs IV with 16-17 slender, sharp, short teeth. Very

slender spur on internal claws of all legs.

Discussion.— The distinction between Echiniscus angolensis from West Africa

and Echiniscus crassispinosus fasciatus Marcus from Brazil and East Africa is

difficult. Based on the illustrations in Ramazzotti’s monograph (1972), the Ven¬

ezuelan specimens more closely resemble E. angolensis, although there are some

discrepancies not yet resolved. The description of E. angolensis does not mention

the presence of the C spine, yet it is clearly illustrated in their drawing; the C d

spine which is mentioned is very small or missing on the illustration.

We have not seen the type of either species and have elected to use the name

E. angolensis at this time.

Distribution. — Merida, 4 specimens.

Echiniscus virginicus Riggin, 1962

(Figs. 26-29)

Cuticle with characteristic ornamentation of pores and granules. Area of median

plate 3 also ornamented. Posterior margins of paired plates minutely scalloped.

The dorsal and lateral spines are variable, both in occurrence and development.

Lateral spine B is sometimes absent on one or both sides; C, D, E are present.

Dorsal spines C d and D d are of variable length and rarely absent; the spines may

be the same length or either may be considerably shorter than the other. Spur

present at base of internal claws of all legs. Leg I with spine and leg IV with large

papilla. Dentate fringe with 8-10 teeth.

Discussion .—The Venezuelan specimens agree with Riggin’s holotype deposited

in the Smithsonian Institution and with other specimens from Florida and Ten¬

nessee. The cuticular ornamentation differs from that of Echiniscus quadrispinosus

brachyspinosus Bartos, which Ramazzotti considered the same as E. virginicus.

Distribution. — La Carbonera, 9 specimens; La Mucuy, 15; Merida, 19; Barran¬

cas, 4; El Tucuco, 1; Yacambu, 5; Rancho Grande, 2. Echiniscus virginicus was

the most widely distributed heterotardigrade and was only surpassed by Pseud-

echiniscus suillus facettalis in numbers collected.

Echiniscus perarmatus Murray, 1907

(Figs. 30-33)

The Venezuelan specimens agree with the description in Ramazzotti’s mono¬

graph (1972). The cuticle is ornamented with a very line uniform granulation,

larger regularly distributed granules, and a few light pores. Each pore appears

bordered by four of the larger granules. Although pores are not mentioned in the

original description, they are present as surface structures. Short C, D teeth are

present and occasionally very short D d spines. The E spines are curved and as

long or longer than cirrus A. A distinguishing character is the presence of a short

thorn-like papilla on legs I—III; legs II and III should be observed carefully.

Discussion .—The species was described from South Africa and another report

from Hawaii is uncertain. Dr. Deidre Christenberry has collected E. perarmatus

from the Southeastern United States. Her specimens, deposited at the Smithsonian

Institution, agree closely with ours.

VOLUME 59, NUMBERS 1-4

Figs. 26-41. 26-29. Echiniscus virginicus. 26, dorsal aspect; 27, cuticle of scapular plate; 28, dorsal

denticles of leg IV; 29, internal and external claws leg IV. 30-33. Echiniscus perarmatus. 30, dorsal

aspect; 31, cuticle of scapular plate; 32, dorsal denticles of leg IV; 33, internal and external claws leg

IV. 34-37. Echiniscus aliquantillus. 34, dorsal aspect; 35, cuticle of scapular plate; 36, internal and

external claws leg IV; 37, dorsal denticles leg IV. 38-41. Echiniscus kofordi. 38, dorsal aspect; 39,

cuticle of scapular plate; 40, dorsal denticles of leg IV; 41, internal and external claws leg IV.

Distribution. — Yacambu, 10 specimens.

Echiniscus aliquantillus Grigarick, Schuster, and Nelson, New Species

(Figs. 34-37)

Holotypefemale .—Length excluding legs IV, 185 jum. Cuticle colorless; all plates

with very fine regular granulation and random distribution of small but distinct

pores. Cephalic plate with rostral zone of pores and granules and posterior band

PAN-PACIFIC ENTOMOLOGIST

of very fine granulation and very small pores. Pattern of granulation and pores

similar on median plates 1 and 2; median plate 3 absent or not well-defined but

the area between the paired plates and the terminal plate has the same cuticular

pattern as the other median plates. Paired plates have clear zone separating the

anterior (with few pores) from the posterior part. Granulation and pores extend

onto legs. Posterior margin of median plate 2 and paired plates D is scalloped.

Head with acuminate buccal cirri; internal cirrus 14 pm long; external cirrus, 16

pm long; buccal papilla, 7 pm long by 3 jum wide. Cirrus A, 34 pm long; clava,

7.6 pm long by 2.4 pm wide, longer and thinner than buccal papilla. Slender spine

on leg I; papilla on leg IV 5 pm long by 2 p m wide. Dentate fringe on leg IV with

10-12 short sharp spines; area above fringe with pores and granules. Spurs on

internal claws of all legs, very distinct and well-developed on legs IV; claws 12

pm.

Etymology.— Latin: aliquantillus, very small; named for the numerous small

pores.

Discussion. — Several species in the arctomys group are very similar but can be

distinguished by their cuticular patterns. Previously named species differ from

the Venezuelan specimens in the following ways. Echiniscus arctomys Ehrenberg

and Echiniscus wendti Richters, both reported from South America, have gran¬

ulations but no pores; both have longer cirri A, especially E. wendti which has a

cirrus A length up to 55-70% of the body length. Echiniscus phocae Bois-Reymond

Marcus, described from Brazil, has a characteristic ornamentation consisting of

circular groups of small granules. Mihelcic (1967) described several similar species

from Argentina, including Echiniscus bellus, insuetus, roseus, and speciosus\ these

species are practically indistinguishable on the basis of their descriptions, and

slides identified and labeled by Mihelcic do not correspond to each other or the

descriptions. All have cirrus A much longer than in the Venezuelan specimens.

The cuticular pattern of Echiniscus kerguelensis Richters, reported from Aus¬

tralia and SW Africa at 1600-2000 m and other localities, most clearly resembles

that of the Venezuelan specimens. The description in Ramazzotti (1972), however,

is unclear and may represent a combination of two or more species. For example,

the ornamentation can appear granulated or porous; the two paired plates have,

or do not have, a smooth unornamented transverse stripe; lateral cirrus A is 50-

80 jum; claws, 18 pm long. The Venezuelan specimens have very fine granulation

and pores, an unomamented stripe on the paired plates, shorter cirrus A, and

shorter claws.

Distribution. — Holotype, La Carbonera; paratypes: La Carbonera, 36 speci¬

mens; Merida, 4.

Echiniscus kofordi Schuster and Grigarick, 1966

(Figs. 38-41)

This species has only lateral cirrus A. The dorsal plates are granular with

distinctive patterns (fine stipple on Fig. 38) on the head, scapular and terminal

plates, and lateral margins of C and D.

Discussion .—This species is related to E. tesselatus J. Murray but differs by

having a shorter spine A and by the divisions of the scapular and terminal plates.

The single specimen compared favorably with the type from the Galapagos Is¬

lands.

VOLUME 59, NUMBERS 1-4

Figs. 42-51. 42-46. Echiniscus marginoporus. 42, dorsal aspect; 43, cuticle of scapular plate; 44,

cuticle of anterior margins of plates C and D; 45, dorsal denticles of leg IV; 46, internal and external

claws leg IV. 47-51. Echiniscus mosaicus. 47, dorsal aspect; 48, 49, variation in pores of cuticle of

scapular plate; 50, dorsal denticles of leg IV; 51, internal and external claws leg IV.

Distribution. — El Tucuco, 1 specimen.

Echiniscus marginoporus Grigarick, Schuster, and Nelson, New Species

(Figs. 42-46)

Holotypefemale .—Length excluding legs IV, 200 pm. Cuticle colorless; cuticular

pattern consists of granulation and pores. Cephalic plate consists of a rostral zone

with regular granulation and a few small pores, separated by a clear zone from a

posterior band of granules and pores. Scapular plate with regular granulation and

marginal pores (Fig. 43), especially on posterior part, and irregular median row

of pores. Anterior portion of plates C and D with irregular shaped granules only

(no pores) and posterior portion with granules and marginal pores, especially

along posterior margin. Paired plates entirely granulated, no unornamented stripe

present. Median plates 1 and 2 with regular granules and pores; median 3 absent

but area between D and E plates with granules. Terminal plates with granules and

scattered small pores. Head with internal acuminate buccal cirrus 12 pny external

cirrus, 18 pnv, buccal papilla 6.6 pm long by 2.8 ^m wide. Spine on leg I a small

thorn; papilla on leg IV present. Dentate fringe on leg IV with 7-8 well-defined

large sharp teeth; area above fringe granulated. Dorsal surface of all legs granulate;

internal claws of all legs with spurs, internal claws, 16 pm\ external claws, 14 pm.

Etymology.— Latin: marginoporus was named for the marginal pores on the

plates.

Discussion .—The Venezuelan specimens are in the arctomys group, but the

cuticular pattern differs from all other described species. See also the discussion

of Echiniscus aliquantillus n. sp.

PAN-PACIFIC ENTOMOLOGIST

Distribution. — Holotype, La Mucuy; paratypes: La Mucuy, 2 specimens; Rancho

Grande, 1; Barrancas, 2.

Echiniscus mosaicus Grigarick, Schuster, and Nelson, New Species

(Figs. 47-51)

Holotype female .—Length excluding legs IV, 190 p m; including legs IV, 218

jum. Cuticle colorless. Cephalic plate with small pores and granules on rostral zone

of head. Scapular plate with large pores 1-2.8 pm, and large black granules 2-3

pm; 4-6 pores surround granule, larger on posterior margin (Figs. 48-49). Paired

plates with clear (unornamented) transverse stripe that separates smaller anterior

part from posterior portion of plate. Median plates 1 and 2 with pattern similar

to posterior margin of scapular plate. Median 3 small but area has pattern similar

to other median plates. Terminal plate with pronounced pores. Tnterpore area

triangular or hexagonal. Cuticular pattern forms a mosaic appearance; hexagonal

pattern is more apparent in some areas where pore size is smaller. Interpore areas

are joined, forming a reticulate pattern. Slight lateral hemispherical pouch may

be seen on some specimens between legs II and III and between legs III and IV.

Head with acuminate buccal cirri, internal cirrus, 15 pm long; external cirrus, 19

pm long; buccal papilla prominent, 9.5 long by 5.7 pm wide, larger than clava.

Cirrus A, 33 pm; clava, 7.6 pm long by 3.8 pm wide. Spine on leg I a short thorn;

papilla on leg IV 3.8 pm long by 2 ^m wide. Dentate fringe on leg IV with 10-

12 sharp spines of irregular size. Spurs on internal claws of all legs; internal claws,

14 pm; external claws, 13 ^m.

Etymology.— Latin: mosaicus, mosaic; named for the mosaic cuticular pattern.

Discussion .—The cuticular pattern of E. mosaicus somewhat resembles that of

Echiniscus elegans Richters and Echiniscus reticulatus Murray, and is reminiscent

of the gameboard for Chinese checkers. The smaller sized cuticular pattern and

transverse clear stripes on the paired plates of E. mosaicus readily distinguish it

from these species.

Distribution.— Holotype, La Carbonera; paratype, Merida, 1 specimen.

Acknowledgments

We wish to thank Dr. Richard Bohart for partially financing and initiating the

arrangements that led to the collections of tardigrades reported in this paper. The

arrangements were made with Dr. Edmundo Rubio, the Dean of the Faculty of

Agronomy of the University of Zulia and formerly a student of Dr. Bohart.

Financing and assistance within Venezuela were provided by the University of

Zulia. Mr. Guillermo Alvarado Duran and many of the University staff were of

great help during the collecting trips. Facilities were also provided by the Uni¬

versity of Los Andes and the University of Central Venezuela at Maracay. Mr.

Robert Brooks and Mr. John McLaughlin (UCD) assisted with the tardigrade

collections and Mr. Chris Williams (ETSU) helped with the sorting.

Literature Cited

Barros, R. de. 1939. Pseudechiniscus juanitae, nova especie de Tardigrado. Bol. Biol. S. Paulo (N.S.)

3:52-54.

Homing, D. S. Jr., R. O. Schuster, and A. A. Grigarick. 1978. Tardigrada of New Zealand. New

Zealand J. Zool. 5:185-280.

VOLUME 59, NUMBERS 1-4

Mihelcic, F. 1967. Ein Beitrag zur Kenntnis de Tardigrada Argentiniens. Veroff. Zool. Bot. Ges.,

Wien 107:43-56.

Ramazzotti, G. 1964. Tardigradi del Cile, III con deserizione delle nuove specie Oreella minor e

Pseudechiniscus lateromamillatus. Atti Soc. Ital. Sci. Nat. 103:347-355.

-. 1972. II Phylum Tardigrada. Mem. 1st. Ital. Idrobiol. 28:1-732.

Richters, F. 1909. Tardigraden-studien. Ber. Senekend. Naturf. Ges. 40:28-49.

Schulz, E. 1955. Studien an marinen Tardigraden. Kieler Meeresf. 11:74-79.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 78-102

Boharticus, N. Gen., with a Review of Rhopalicus Foerster and

Dinotiscus Ghesquiere (Hymenoptera: Pteromalidae)

E. E. Grissell

Systematic Entomology Laboratory, IIBIII, ARS, USD A, % U.S. National Mu¬

seum of Natural History, Washington, D.C. 20560.

This paper represents a preliminary step toward integrating knowledge of the

Nearctic and Palearctic faunas of Pteromalidae. It presents a review of the known

species of Rhopalicus Foerster and Dinotiscus Ghesquiere, in which a new Nearctic

species of Rhopalicus is described ( zolae ), the Palearctic species Dinotiscus apon ius

is reported for the first time from the Nearctic, two Nearctic species of Dinotiscus

(acutus, polygraphi) are synonymized under a Palearctic name (D. eupterus), the

Nearctic species D. burkei is synonymized with D. dendroctoni, and keys to the

11 known species of both genera are given. All species are associated with wood-

infesting beetles. Additionally, the new genus Boharticus is described for 4 new

species associated with Cecidomyiidae on Juniperus.

The findings of this paper are based upon a comparison of type-specimens and

identified material from both the British Museum (Natural History) (hereafter

BMNH) and the United States National Museum (USNM). Additional material

was borrowed from the Florida State Collection of Arthropods (FSCA), Gaines¬

ville, Florida. Host records are mainly from Burks (1979) and Graham (1969),

but nomenclatural changes for these records have been made based upon newly

published references (Wood, 1982; O’Brien and Wibmer, 1982). In the case of

generic synonymy, I have relied upon the taxonomic conclusions reached by

Graham (1969) who, for many years, has studied the types of European Pter¬

omalidae. His research furnishes a base for much future pteromalid work.

Many years ago, when I began my chalcidoid studies, I desperately needed a

starting point such as that provided by Graham (1969). By taking up a group

wherein scarcely anyone was working, I assured myself of one thing ... no help

whatsoever! At least it seemed that way at the time. But upon the deliberations

of numerous passing years, I now realize that I really did have one resource of

help and a steady one at that. It was the source of direction, of purpose, and of

inspiration that came from Dick Bohart, an entomologist’s entomologist. Without

“Doc” there somewhere, always willing to help, I should surely have stumbled

once too often. Dick and his wife, Margaret, were generous to me in particular,

and to their students in general. I take great satisfaction in dedicating this work

to both of them.

Pteromalid Classification

Before presenting the descriptive aspects of this paper, a few general comments

should be made about the state of our knowledge of pteromalid taxonomy. Current

world figures place the number of valid pteromalid species at 3111 and the number

of genera at 611 (J. S. Noyes, pers. comm.). In the Nearctic there are nearly 400

VOLUME 59, NUMBERS 1-4

species in 129 genera (Burks, 1979). These figures are misleadingly low if we

consider the potential number of undescribed taxa and the fact that the group has

been poorly collected and studied.

In the Nearctic there has been little revisionary work which would serve as a

basis for the study of pteromalid taxonomy. In the Palearctic, however, Graham

(1969) published a monograph of the Pteromalidae of northwestern Europe in

which he provided keys to all European genera. Comparison of Graham’s work

with the most recent Nearctic catalog (Burks, 1979) shows that over two-thirds

(ca. 80 of 110) of the genera treated by Graham are Holarctic. Therefore Graham’s

work could serve an important function in first helping us to understand the

Nearctic fauna, and more importantly, in providing a starting point for the in¬

tegration of the Nearctic and Palearctic faunas.

As important as Graham’s work is, however, its indiscriminate use may cause

problems. The description of the new genus Boharticus has highlighted some of

these problems and I point them out here for precautionary reasons. To begin

with, relationships of genera and diagnostic characters for them are not discussed

in Graham’s text. Although genera are apparently arranged in a phylogenetic

system within the text, there is little or no discussion of relationships between

them. To define genera it is therefore necessary to search through the keys for

useful characters. However, when one does this for well-known genera with large

numbers of species, such as Chlorocytus Graham, Pteromalus Swederus, and

Mesopolobus Westwood (ca. 120 species total), it is apparent that they are not

clearly defined. For example, these 3 genera run to 24 different couplets in the

key. The conclusion one draws from this is that species grouped in each of these

genera cannot be defined by shared, unique characters (i.e., synapomorphies). A

more serious problem is that a large proportion of genera (54 of 128 Palearctic

genera) are represented by only 1 species each and that these genera are often

distinguished by a character (or combination of characters) which would be only

of specific value in one of the larger three genera listed above. For example,

characters associated with pronotal collar, clypeus, fiagellomere ratios, color, and

abdominal shape may be used to distinguish groups of species within genera (via

numerous couplets in the key) as well as to define genera with single species.

The 4 species which I place in Boharticus, new genus, serve to illustrate the

rather poor definitions and lack of understanding we have within the Pteromalidae.

Running these species individually through Graham’s key results in one being

placed in Dinotiscus Ghesquiere, two being placed in Rhopaliens Foerster, and

one being placed in Trychnosoma Graham. For reasons I will discuss under the

description of Boharticus, 1 believe the species of this genus represent a natural

grouping based upon thoracic and host-related characters. However, the act of

defining Boharticus causes the distinctions between Dinotiscus and Rhopaliens to

become unclear. That is, the latter two genera now are defined by characters which

appear to be more appropriate for separating species. At present, I can do little

to resolve the generic problems between Rhopalicus and Dinotiscus. Although I

have seen all the species of these genera and I have provided a modification of

Graham’s key which will separate them, their true status will require a major

taxonomic revision of the entire family. Such a revision will require analysis of

new sets of characters (such as the thoracic characters used to define Boharticus)

as well as reexamination of previously used ones.

PAN-PACIFIC ENTOMOLOGIST

Boharticus Grissell, New Genus

Type-species. — Boharticus richardi Grissell, new species.

Occiput without carina; clypeus entire; lower face not protuberant at level of

toruli; antenna with 2 anelli, 6 flagellomeres, and 3 segmented club; torulus 2 to

3 diameters above level of ventral margins of eyes; both mandibles with 3 den¬

ticles, uppermost denticle truncate. Almost entire thorax similarly and heavily

reticulate including postalar plate (Fig. 12), upper epimeron, dorsellum, and pro¬

podeum; scutum with notauli on anterior half; frenum weakly indicated by finer

sculpture on area anterior to it; dorsellum well developed (i.e., not narrowed or

reduced) and protruding at least as far as apex of scutellum, not separated from

scutellum by costate groove; propodeum with or without complete median carina,

without plicae or costula; nucha represented by narrow, lunate strip which may

be triangularly enlarged medially in some species (e.g., Fig. 11); supracoxal flange

not joined to nucha (but may be joined to inner one-third of petiolar foramen,

Figs. 1, 3); hindcoxa with or without setae dorsally; forefemur not enlarged or

emarginate ventrally; hind tibia with one apical spur; forewing with maculae,

postmarginal and marginal veins longer than stigmal, postmarginal equal to or

shorter than marginal, stigma not enlarged. Gaster lanceolate, petiole transverse

and scarcely visible, terga entire on posterior margins, bristles of cerci equal in

length, ovipositor not exserted, tip of hypopygium not (or barely) reaching middle

of gaster.

Etymology?.— Named in honor of Richard M. Bohart on the occasion of his

70th birthday. The gender is masculine.

Discussion.— The genus Boharticus is recognized as monophyletic based upon

the following synapomorphies:

1) All species share a structurally unique thorax as follows:

a) On the propodeum in dorsal view, the area between the nucha and the

supracoxal flange is not connected by a carina (thus the propodeum

appears to turn under at this point). In B. margaretae this connecting

carina is absent (Fig. 2). In B. richardi, nanellus, and apilosus, however,

the carina curves around the inner margin of the coxal cavity and joins

the propodeal foramen in the inner one-third (Figs. 1, 3; compare this

with Fig. 6 for Rhopalicus and Dinotiscus)',

b) The thorax is heavily and similarly reticulate on nearly every surface,

but especially on rhe upper epimeron, the dorsellum, the postalar plate

(Fig. 12), and the propodeum.

2) All species are associated biologically with gall-formers on the plant genus

Juniperus. The gall-formers are known to be Cecidomyiidae for 3 species

of Boharticus. The fourth species is known only from juniper “berries,” but

this is probably an error in identification because the galls of several ceci-

domyiids (e.g., Walshomyia ) resemble fruiting structures.

If the members of Boharticus are placed within the context of Graham (1969),

3 of the 4 species would key to couplet 205 which includes Dinotiscus and Rho¬

palicus, and the fourth would key to Trychnosoma Graham (couplet 110; for

discussion see B. apilosus below). Boharticus differs from both Rhopalicus and

Dinotiscus as follows:

VOLUME 59, NUMBERS 1-4

Boharticus

Dinotiscus

Figs. 1-6. Ventral view of propodeum and posterior apex of thorax (cx 3 = hindcoxal foramen;

n ~ nucha; p = propodeum; pf= propodeal foramen; sf= supracoxal flange).

1) All species of these latter 2 genera (as well as other related Pteromalinae

genera) share the following thoracic characters:

a) In dorsal view, the supracoxal flange is connected to the nucha by a carina

(ventrally the carina may be seen to join the nucha in the outer one-third,

Fig. 6);

b) The thorax is not everywhere heavily or evenly reticulate; at least the

postalar plate (Figs. 8, 10) is vertically carinate (may be smooth on outer

margin or with vertically elongate reticulations on inner margin), the

upper epimeron and/or dorsellum may be smooth, and the propodeum

is less heavily sculptured than the apex of the scutellum.

2) All species of these genera are parasitic upon wood-infesting beetles of the

families Scolytidae and Curculionidae.

The propodeal character involving the supracoxal flange is not the easiest of

characters to interpret, and I offer these additional notes as a guideline. Initially,

in trying to assess the structure of the flange, it was necessary to remove the

abdomen of each specimen for a better view. However, the flange may be seen

in dorsal view, without removing the abdomen, by tilting the specimen forward.

In genera such as Dinotiscus and Rhopalicus the flange is easily seen from this

position (Figs. 7, 9) partially because it is well developed and partially because it

is relatively near to the posterior margin of the propodeum (Fig. 6). In Boharticus,

the flange is either absent (Fig. 2) or weakly developed and positioned toward the

anterior of the propodeal foramen (Figs. 1, 3). In ventral view, and with the

Figs. 7-12. Scanning electron micrographs. 7, 8, Rhopalicus pulchripennis. 9, 10, Dinotiscus thom-

soni. 11, 12, Boharticus richardi. (Left, propodeum; right, postalar plate.)

abdomen removed, the supracoxal flange of Rhopalicus and Dinotiscus is contin¬

uous with the nucha and there is virtually no sclerite between the propodeal

foramen and the hindcoxal foramen (Fig. 6). In Boharticus, however, the supra¬

coxal flange (if present) does not join the nucha, but instead joins the propodeal

foramen and there is a sclerotized area between the propodeal and hindcoxal

foramina (Figs. 1-3).

It is not known how useful this character will be throughout the Pteromalinae.

However, in examining many species of Mesopolohus Westwood, which share the

narrowed nucha of Boharticus, Dinotiscus, and Rhopalicus, it was found that the

character was stable for the genus, although it was expressed in a manner inter¬

mediate (Fig. 5) to Boharticus (Fig. 1-3) and Rhopalicus-Dinotiscus (Fig. 6). In

VOLUME 59, NUMBERS 1-4

Mesopolobus there is a sclerotized area between the foramina (as in Boharticus)

but the supracoxal flange joins the nucha (as in Rhopalicus-Dinotiscus). In Pter-

omalus, a genus separated from the above mentioned genera by virtue of an

elongate, sculptured nucha, the supracoxal flange is also consistently well devel¬

oped, elongate, and joins the nucha as in Mesopolobus (cf. Figs. 4 and 5). This

character is constant for the species of Pteromalus I have examined.

For practical purposes the following couplets are proposed to replace those

numbered 205 to 206 in Graham (1969:387). This replacement is based upon all

known species of Boharticus, Dinotiscus, and Rhopalicus.

205 (53, 124) Hindtibia with 2 apical spurs (Palearctic) . . Acrocormus Foerster

Hindtibia with 1 apical spur .206

206 (205) Nearly entire thorax similarly and heavily reticulate (note

especially postalar plate (Fig. 12), upper epimeron, dorsellum,

and propodeum); supracoxal flange not joined directly to nu¬

cha (Figs. 1-3; may be joined to lower third of propodeal

foramen), in dorsal view flange not easily seen.

. Boharticus Grissell

- Thorax not similarly reticulate throughout, postalar plate ver¬

tically carinate (Figs. 8, 10; may appear elongately reticulate

towards inner margin and smooth along outer), epimeron may

be smooth above, propodeum alutaceous medially or at least

less heavily sculptured than apex of scutellum; supracoxal

flange joined to nucha and visible in dorsal view (Figs. 7, 9)

.207

207 (206) Pronotum carinate; propodeal nucha medially a semicircular,

narrowed carina (Fig. 9) . Dinotiscus Ghesquiere

- Pronotum without carina; propodeal nucha medially expand¬

ed as triangular area (Fig. 7). Rhopalicus Foerster

Key to Female Boharticus

1. Basal cell of forewing essentially bare (with 0 to 10 setae, but if ques¬

tionable then wing spot parallels marginal vein and parastigma (Fig. 16));

pronotal collar carinate . 2

- Basal cell of forewing fairly evenly setose (usually over 10 setae, but if

questionable, as in some specimens less than 1 mm, then wing spot

confined to stigmal area. Fig. 14); pronotum without carina. 3

2. Flindcoxa with basal setae; forewing with maculae expanded at both

stigmal and parastigmal areas (Fig. 13) . margaretae Grissell, n. sp.

- Hindcoxa without basal setae; fore wing with macula expanded only at

stigmal area (Fig. 16) . apilosus Grissell, n. sp.

3. Forewing macula surrounding stigmal vein and paralleling marginal vein

(Fig. 15); abdomen about 3X as long as wide; nucha expanded medially

as carina which reaches (or nearly) dorsum of propodeum (Fig. 11) ...

. richardi Grissell, n. sp.

- Forewing macula restricted to stigmal area (Fig. 14); abdomen less than

1.5X as long as wide; nucha parallel-sided medially, not extending dorsally

as carina . nanellus Grissell, n. sp.

PAN-PACIFIC ENTOMOLOGIST

Boharticus richardi Grissell, New Species

(Figs. 3, 11, 12, 15, 19)

Holotype female.— Body length 5.8 mm. Head and thorax (excluding legs be¬

yond coxae) metallic green; abdomen metallic red with green reflections; femora

and tibiae smoky orange; scape, tegula, and tarsomeres 1-4 straw-yellow; antenna

and tarsomere 5 brown, wing veins brown becoming slightly lighter towards

postmarginal. Head, thorax, and abdomen (except posterior margin of terga nearly

smooth) with reticulate sculpture. Face as shown in Fig. 19; toruli 2X own diameter

above ventral margin of eyes (5:10); intermalar distance 2. IX malar distance (32:

15); eye 2.3X length of malar distance (34:15); ratio of lateral ocellus diameter:

ocellocular distance : postocellar distance as 5:10:14; ratio scape : pedicel: anel-

li: flagellomeres 1-6 : club as 50:14:3:3:17:16:16:14:13:13:27, flagellomeres start¬

ing 1.6X as long as wide (FI = 16:10) and becoming as long as wide (F6 = 13:

13). Pronotal collar not margined anteriorly, dropping almost perpendicularly to

neck; nucha angled medially forming an acute triangle which narrows and con¬

tinues to dorsal margin of propodeum (Fig. 11); basal fovea about size of spiracle

and midway between spiracle and median carina, apical fovea about as large as

basal one, 10-15 setae present posterior to spiracle, callus evenly setose; epimeral

scrobe nearly straight, situated at midpoint of epimeron and occupying about V 7

of length; wing veins with ratio of submarginal: marginal: postmarginal: stigmal

as 48:16:16:10; basal cell with at least 25 setae, setae present on cubital and basal

veins, costal cell with complete anterior setal row beneath and several rows beneath

on distal Vi; maculations as shown in Fig. 15; hindcoxa with setae basally. Ab¬

domen keel-shaped (i.e., somewhat laterally compressed but dorsally flat), with

heavy, reticulate sculpture, with ratio of Tl-7 : ovipositor sheaths as 15:10:8:8:

12:17:15:3; ratio length : width as 43:14; ratio abdomen : thorax as 43:23.

Male. — Unknown.

Type material.— Holotype 2, USNM #100592; CALIFORNIA, Fresno Co.,

Coalinga, 21 March 1960, H. L. Wilson, ex Juniper galls. Paratypes: 7 2, same

data as holotype; 1 2, same data except 20 March 1959, ex “Juniper berries;” 3

2, CALIFORNIA, Los Angeles Co., Vasquez Rocks, 25 January 1963, ex pine-

cone gall on Juniperus californica; 2 2, CALIFORNIA, Stanislaus Co., Del Puerto

Canyon, E. E. Grissell, R. F. Denno, em. 15 March 1971 ex cecidomyiid cone

gall on Juniperus californica collected 6 March 1971. All specimens in USNM

except 2 2 in British Museum (Natural History). (The “pine-cone” gall is made

by a species of Cecidomyiidae in the genus Walshomyia according to R. J. Gagne,

personal communication. The juniper “berry” record is probably a mistake, as

some Walshomyia galls have the appearance of a fruiting structure.)

Etymology’.— Named for Richard M. Bohart, mentor, colleague, and friend.

Variation.— Females of B. richardi range in length from 4.9 to 6.2 mm. Color

of the head and thorax is metallic green in most specimens (15) but 1 specimen

is metallic blue. The abdomen is metallic red in 13 specimens and metallic green

tinged red in 2. The maculation pattern of the forewing is consistent, but the

intensity varies from very dark brown (with distinct edges) to a washed out brown

(with indistinct edges, i.e., the pattern fades into the surrounding clear areas). The

number of setae of the basal cell is variable, but the cell is fairly evenly covered

with 20 or more setae. All the specimens have the nucha extended dorsally as a

keel or carina, but in 2 specimens this carina fades toward the dorsal propodeal

PAN-PACIFIC ENTOMOLOGIST

margin and does not reach it. The specimens from Del Puerto Canyon have the

nucha less pronounced than other specimens, but the carina continues to the

propodeal margin. The basal fovae very from a simple depression about the size

of the spiracle to a depression several times as large with 2 or 3 carinae present.

The epimeral scrobe may be short as in the holotype or it may occupy half of the

epimeral length.

Discussion.— This is the most striking member of the genus. It is the largest

species (to over 6 mm), and the abdomen is generally metallic red which contrasts

distinctly with the green thorax. The abdomen is considerably elongate (3X longer

than wide, 2X longer than the thorax), keel-shaped, and as heavily sculptured as

the thorax. Although richardi shares the setose basal cell with nanellus, the two

do not appear particularly related. They differ in a number of characters as follows:

in richardi the eye height is less than 2.5X the malar distance (nearly 3.5X in

nanellus ), FI is over 1.5X longer than wide and the flagellomeres become quadrate

at F6 (FI is quadrate in nanellus, and flagellomeres become wider than long by

F6), the fore and midcoxae are green (yellow in nanellus), the abdomen is keel¬

shaped and flat dorsally with heavy, reticulate sculpture (dorso-ventrally sunken

in nanellus), and T1 and 7 are subequal (T1 is 2.5X as long as T7 in nanellus).

Additional characters may be found in the key.

Boharticus margaretae Grissell, New Species

(Figs. 2, 13, 17)

Holotypefemale. — Body length 3.0 mm. Head and thorax (excluding legs beyond

coxae) metallic green; abdomen blackish purple; femora, tibiae, and tegula, smoky

orange; scape and tarsi straw-yellow; antenna and wing veins brown, fore wing

with 2 brown spots. Head, thorax, and abdomen (except posterior margins of

terga smooth) covered with reticulate sculpture. Face as shown in Fig. 17; toruli

2.7X own diameter above ventral margin of eyes (6:16); intermalar distance 2.3X

malar distance (21:9); eye 2.9X length of malar distance (26:9); ratio of lateral

ocellus diameter: ocellocular distance: postocellar distance as 6:12:21; ratio scape:

pedicel: anelli: flagellomeres 1-6 : club as 40:14:2:2:10:9:9:9:9:8:17, flagello¬

meres starting slightly longer than wide (FI = 9:7) and becoming slightly wider

than long (F6 = 9:8). Pronotal collar margined anteriorly, dropping almost per¬

pendicularly to the neck, collar laterally rounded off; nucha evenly curved and

parallel-sided medially, disappearing at lateral edges, basal foveae essentially ab¬

sent, apical fovea well developed, 3 or 4 long setae present posterior to inner

margin of spiracular sulcus, callus with about 5 setae; epimeral scrobe a long,

crescentric groove reaching from near posterior margin to anterior margin; wing

veins with ratio of submarginal: marginal: postmarginal: stigmal as 45:21:19:14;

basally, cubital and basal veins without setae, wing essentially bare from speculum

to base (only a few setae present near intersection of cubital and basal veins, and

costal cell with anterior row beneath); maculations as shown in Fig. 13; hindcoxa

with setae basally. Abdomen keel-shaped, dorsally flattened, with reticulate sculp¬

ture, with ratio T1-7 : ovipositor sheaths as 12:12:14:13:10:8:7:6; ratio length:

width as 40:18; ratio abdomen: thorax as 40:26.

Male— Unknown.

Type material. — Holotype female, USNM #100593; COLORADO, El Paso

Co., Garden of the Gods, 20 July 1915, J. H. Pollock, [ex “cecidomyiid larvae

VOLUME 59, NUMBERS 1-4

in old buds”], Juniperus monosperma, “Hopkins U. S. 12946a.” Paratypes: 8 2,

same data as holotype; 1 2, ARIZONA, Mohave Co., 8 miles SW Peach Springs,

4500', 7 September 1964, C. W. O’Brien, ex Juniperus. All specimens in USNM,

except 1 2 in British Museum (Natural History). (Note: The bracketed material

above was taken from Hopkins cards on file at the USNM; according to these

cards, the gall material was collected on 1 July 1915 and parasites emerged on

20 July.)

Etymology*.— This attractive little species is named for Margaret Bohart who,

for many years, has shown much generosity and hospitality to Dick’s students.

Variation. —Females vary from 2.3 to 2.7 mm. The thorax varies from metallic

green to metallic blue. The wing maculations are equally intense on all of the

specimens. On several specimens the basal foveae are indicated by a slight depres¬

sion bordered on the inner side by a short carina; the pit is equal in size or smaller

than the spiracle.

Discussion.— This species is similar to apilosus in having the basal cell of the

forewing nearly asetose and in the carinate pronotum. There are rather striking

differences between the species especially in forewing maculations and hindcoxal

setae (as stated in the key). The flagellomeres of margaretae range from barely

longer than wide (FI) to wider than long (F6), whereas in apilosus, FI is nearly

2X as long as wide and F6 is longer than wide. Also in margaretae FI is cylindrical

but in apilosus it is constricted basally (Fig. 20b). Additionally, the abdomen of

margaretae is rigid with dull reticulate sculpture somewhat as on the thorax, but

in apilosus the abdomen is distorted and sunken with shiny, alutaceous sculpture.

Boharticus nanellus Grissell, New Species

(Figs. 1, 14, 18)

Holotype female.— Body length 1.7 mm. Head, thorax, and upper % of hindcoxa

metallic green; yellow are: scape, fore and midcoxae (and apex of hindcoxa), legs

(except tarsomere 5 dark brown), and abdomen (which is smoky dorsally); wing

veins and tegula straw colored; flagellum brown. Head and thorax evenly covered

with reticulate sculpture; abdomen evenly covered with reticulations but not as

raised as on thorax (also difficult to see with reflected light because of yellow

background). Face as shown in Fig. 18; toruli nearly 3X own diameter above

ventral margin of eyes (4:11); intermalar distance 2.4X malar distance (22:9); eye

2.4X length of malar distance (22:9); ratio lateral ocellus diameter: ocellocular

distance : postocellar distance as 5:10:15; ratio scape : pedicel: anelli: flagello¬

meres 1-6: club as 23:8:1:1:5:5:4:4:5:5:12, flagellomeres starting IX as long as

wide (FI = 5:5) and becoming wider than long (F6 = 6:5). Pronotal collar in dorsal

view dropping perpendicularly to the neck but without carina; nucha a narrow

carina which is scarcely widened medially, basal and apical fovae about the size

of spiracle (not very well developed); 2 or 3 long setae present posterior to spiracle,

callus with about 5 or 6 setae; epimeral scrobe poorly defined, situated at midpoint

of epimeron and occupying about V 7 the length; wing veins with ratio of sub¬

marginal: marginal: postmarginal: stigmal as 55:30:28:20; wing heavily setose

with areas bare as follows: speculum, area beneath cubital vein, basal half of costal

cell (except row of setae along lower foremargin); maculations as shown in Fig.

14, weakly expressed and diffuse; hindcoxa with several setae basally. Abdomen

PAN-PACIFIC ENTOMOLOGIST

dorsoventrally sunken, with ratio Tl-7 : ovipositor sheaths as 16:12:10:10:11:14:

6:1; ratio length : width as 40:32; ratio abdomen : thorax as 40:33.

Allotype male. — Body length 1.5 mm. Color as for female except abdomen

metallic reddish. Similar to female (including ratios) except as follows: flagello-

meres with recurved setae which are about as long as segment; wing maculation

restricted to weak stain posterior to stigma; ratio Tl-6 as 14:5:4:4:3:2.

Type material. — Holotype female, USNM #100594; OHIO, Wayne Co., Woos¬

ter, 25 June 1961, J. E. Appleby, ex tip midge on Juniperus horizontalis. Allotype

6, 1 2, 2 6 paratypes same data as holotype; 1 $, 1 8 paratype same data as holotype

except ex Juniperus virginiana ; 63 2, 66 8 paratypes, FLORIDA, Alachua Co.,

Gainesville (Doyle Conner Building), 15-26-XI-1973, E. E. Grissell, sweeping

Juniperus salicicola (Small) Bailey. Holotype $ and paratypes from Ohio in USNM;

paratypes from Florida in FSCA, USNM, BMNH, and the Canadian National

Collection, Ottawa.

Etymology.— From the diminuative (- ellus ) of nanus—a dwarf, in reference to

the small size of this species.

Variation. — Males and females from the Ohio population are generally larger

in size than the Florida population. Males from Ohio range in length from 1.3 to

1.5 mm and females are about 1.6 mm long. Females from Florida, however,

range in size from 0.8 to 1.5 mm and males from 0.7 to 1.0 mm. The wing

maculation varies considerably even within the 3 females from Ohio. In the

holotype and 1 female from J. virginiana the forewing spot is weak and occupies

the anterior half to third of the wing; in 1 female from J. horizontalis the spot

nearly reaches the hindmargin of the wing (but with a hyaline break at one-third

the distance from the posterior margin). Larger females from Florida (ca. 1.5 mm)

have the wing spot as for the holotype, but small females (ca. 0.8 mm) have the

spot restricted to a weak stain posterior to the stigma. In males the spot resembles

that of small females or is absent. In general, as the specimens become smaller

(either sex), the yellowish coloring of the abdomen and hindcoxae becomes darker.

The mid and forecoxa, legs, and scape remain yellow even in the smallest male.

In small specimens, the number of setae in the basal cell is reduced to about 10,

but the setae are fairly evenly spaced. In both males and females, the epimeral

scrobe may be absent regardless of the size.

Discussion.— This is the smallest species of the genus, being less than 2 mm,

and the only species for which the male is known. For a discussion see B. richardi.

Boharticus apilosus Grissell, New Species

(Figs. 16, 20)

Holotype female.— Body length 3.2 mm. Head and thorax metallic green; scape

and abdomen blackish green; legs yellowish orange except femora with greenish

reflections, tibiae smoky, foretarsi and mid and hind tarsomeres all darkish brown;

antennae, wing veins, and wing maculation brown. Head and thorax evenly cov¬

ered with reticulate sculpture; abdomen alutaceous. Face as shown in Fig. 20a;

toruli 2X own diameter above ventral margin of eyes (3:6); intermalar distance

2. IX malar distance (19:9); eye 2.6X length of malar distance (42:16); ratio of

lateral ocellus diameter: ocellocular distance : postocellar distance as 3:6:9; ratio

scape : pedicel: anelli: flagellomeres 1-6 : club as 36:10:2:2:11:13:12:12:12:10:20;

flagellomeres starting ca. 2X longer than wide (FI = 11:6) and remaining longer

VOLUME 59, NUMBERS 1-4

than wide (F6 = 10:8); FI constricted at base (Fig. 20b). Pronotal collar dropping

perpendicularly to the neck, with carina along anterior margin; nucha a narrow

carina which is slightly produced medially to form frail carina which reaches half

way to anterior margin of propodeum, basal and apical foveae about size of spiracle

(not well developed); ca. 6 long setae present posterior to spiracle, callus with

about 10 setae; epimeral scrobe a long, crescentric line reaching from posterior

margin to anterior; wing veins with ratio of submarginal: marginal: postmargi nal:

stigmal as 45:30:25:18; basal cell with 6 to 8 setae, speculum and area below

cubital vein bare, costal cell with row of setae along anterior margin on underside

and 2 sparse rows in distal Vi on underside; maculations as shown in Fig. 16;

hindcoxa without setae basally (2 or 3 setae distally). Abdomen keel-shaped,

dorsally sunken due to lightly sclerotized terga, with ratio Tl-7 : ovipositor sheaths

as 18:6:19:9:16:15:9:6; ratio length : width as 49:15; ratio abdomen: thorax as

49:25.

Male. — Unknown.

Type material. — Holotype 9, USNM #100595; ARIZONA, Santa Cruz Co.,

Patagonia, 3 October 1946, Nog. #65320, ex “Juniper berries .” Paratypes: 5 9,

same data as holotype. All specimens in USNM.

Etymology. — From pilosus —hairy and a— without, in reference to the lack of

setae on the base of the hindcoxa.

Variation .—The number of setae in the basal cell ranges from 4 to 8. The

abdomen of the type is distorted so that T2 and T4 appear shorter than for other

specimens. From the specimens at hand, T2-6 may be subequal, or T2 and T4

may be equal and shorter than T3, T5, and T6. The abdomen of this species is

weakly sclerotized and tends to shrivel.

Discussion. — Boharticus apilosus is the only species of the genus with the first

flagellomere constricted basally (Fig. 20b), and also the only one with no setae

on the base of the hindcoxa. For additional characters see B. margaretae.

Because of the lack of setae on the hindcoxa, this species would key to the genus

Trychnosoma Graham in Graham’s key (1969) to the European Pteromalidae. It

differs from that genus, however by the following: 3 denticles in both mandibles

(4 on right mandible of Trychnosoma), clypeus reticulate (radiating striae in Trych¬

nosoma), forewing with maculations (hyaline in Trychnosoma ), and the supracoxal

flange not joining the nucha (joining in Trychnosoma).

Rhopalicus Foerster

Rhopalicus Foerster, 1856:66, 70.

Type-species: Cleonymus maculifer Foerster, 1840. Monotypic.

This genus is known from 4 species as follows: tute/a (Walker) is Holarctic,

pulchripennis (Crawford) is primarily confined to the northern Nearctic, and brev-

icornis Thomson and guttatus (Ratzeburg) are Palearctic. A fifth species is herein

described as new and is found throughout the southeastern Nearctic. Rhopalicus

may be defined as follows: clypeus emarginate, with 2 well-defined lobes (Figs.

21a, 22a); venter of torulus less than own diameter above line connecting ventral

margins of eyes or slightly below; pronotum without carina; thorax not similarly

reticulate throughout, at least postalar plate vertically carinate (may appear elon-

gately reticulate towards inner margin and smooth along outer), epimeron may

PAN-PACIFIC ENTOMOLOGIST

be smooth above, propodeum nearly shiny or alutaceous medially (but as reticulate

as scutellum in zolae and some pulchripennis)\ nucha medially expanded as tri¬

angular area which narrows to median carina and meets (or nearly) anterior margin

of propodeum (Fig. 7); supracoxal flange joined to nucha (as in Fig. 6); forewing

with or without maculations; postmarginal and marginal veins longer than stigmal,

and postmarginal at least slightly longer than marginal, stigma not enlarged in

female, but may be elongate as in Fig. 24c (in male slightly enlarged in tutela,

Fig. 24a).

My concepts of the species discussed below are based upon Graham’s work

(1969: extant types examined), my identification of Palearctic specimens based

upon his key, the types of the Nearctic species, and the lectotype of tutela (Walker)

(BMNH).

Key to Female Rhopalicus

1. Lower surface of forewing costal cell with 2 or more rows of setae in

proximal half; upper epimeron smooth; macula of wing (rarely absent)

associated with stigma (Holarctic). tutela (Walker)

- Lower surface of forewing costal cell with 1 row of setae in proximal half;

upper epimeron similarly reticulate to lower portion; macula of wing

absent or associated with marginal vein . 2

2. Scape length at most 3 A distance between top of torulus and venter of

midocellus (Fig. 22a); basal vein without setae (Nearctic) .

. zolae Grissell, n. sp.

- Scape length subequal to distance between top of torulus and venter of

midocellus (Fig. 21a); basal vein with setae . 3

3. Forewing hyaline; flagellomeres 1-6 together shorter than distance be¬

tween eyes (dorsal view) (Palearctic). quadratus (Ratzeburg)

- Forewing with some form of spot; flagellomeres 1-6 together longer than

distance between eyes. 4

4. Forewing with infumate spot confined to basal half of marginal vein;

dorsellum separated from scutellum by pitted groove (Palearctic) .

. guttatus (Ratzeburg)

- Forewing with infumate spot beneath entire length of marginal vein;

dorsellum continuous with scutellum (at most a weak line delimits the

suture) (Nearctic). pulchripennis (Crawford)

Rhopalicus zolae Grissell, New Species

(Fig. 22)

Holotypefemale.— Body length 2.7 mm. Head and thorax metallic blue-green;

legs, apices of fore and midcoxae, and scape concolorus golden-yellow; flagellum,

tegula, and abdomen dark brown; base of fore and midcoxae and entire hindcoxa

metallic blue to violet; submarginal and stigmal veins brownish white, marginal,

postmarginal, and stigma pale brown. Head and thorax nearly evenly covered

with reticulate sculpture (except postalar plate vertically carinate); abdomen weak¬

ly sclerotized and shriveled, polished to faintly alutaceous. Face as shown in Fig.

22a; torulus slightly below line connecting ventral margin of eyes; intermalar

distance nearly 2X malar distance (17:9); eye 2.3X length of malar distance (21:

9); ratio of lateral ocellus diameter: ocellocular distance : postocellar distance as

VOLUME 59, NUMBERS 1-4

21 pulchripennis 22 zolae

Figs. 21-24. Rhopalicus spp. 21, 22, a = face, frontal view; b = antennae, side view (to same scale

as face). 23, 24, stigma and stigmal vein of forewing. 24, a = male; b, c = female showing variation.

6:12:22; ratio scape : pedicel: anelli: flagellomeres 1-6: club as 26:9:1:2:9:8:7:7:

6:6:10, flagellomeres starting IX as long as wide (FI = 9:9) and becoming 0.6X

as long as wide (F6 = 6:10); Fl-6 0.9X distance between eyes dorsally (44:50);

scape 0.72X as long as distance between top of toruli and venter of midocellus

(26:36) and 0.65X eye height (26:40). Thorax dorso-ventrally flattened, scutellum

and scutum in same plane, propodeum with complete median carina; wing veins

with ratio of submarginal: marginal: postmarginal: stigmal as 43:16:21:15; basal

cell, basal vein, and cubital veins without setae, costal cell with complete anterior

row below, and distal half with 1 additional row; weakly defined pale brown spot

PAN-PACIFIC ENTOMOLOGIST

present below marginal vein. Abdomen with first tergum 0.14X abdomen length

(7:35), abdomen 1.5X longer than wide (35:23) and 1.5X longer than thorax (35:

23) .

Allotype male. — Length 2.3 mm. Color and sculpture as for female, except pale

yellow band present on distal half of tergum I and all of tergum II. Torulus slightly

above a line connecting ventral margins of eyes; malar and eye ratios as for female;

ratio scape : pedicel: anelli: F1-F6 : club as 30:10:1:2:10:9:8:8:7:7:17; flagello-

meres starting 1.4X as long as wide; Fl-6 1.13X least interocular distance (27:

24) ; scape 0.84X as long as distance between top of toruli and venter of midocellus

(26:31) and 0.70X eye height (26:37). Thorax and propodeum as for female; wing

veins with same ratio as female, basal cell with several setae distally and basal

vein with row of setae, costal cell as for female except distal half with 3 rows of

setae instead of 2. Abdomen with first tergum 0.40X abdomen length (20:50),

abdomen 1.7X longer than wide (50:24) and 0.96X as long as thorax (50:52).

Type material.— Holotype 2 USNM #100695; TEXAS, “Southeast Texas,” Mar.

1962 to Oct. 1963, R. C. Thatcher, ex loblolly pine with Dendroctonus frontalis

Zimmermann. Allotype <3, 8 2 paratypes and 3 6 paratypes with same data; 6

paratypes as follows: 1 2 , MISSISSIPPI, Amite Co., 7 May 1965, N. A. Overgaard,

ex pine infested with southern pine beetle; 1 2 , TEXAS, Hardin Co,, June 1960,

ex pine infested with Ips, Hopkins No. 38912; 1 2 , GEORGIA, Dekalb Co., Stone

Mountain, 26 Oct. 1947, P. W. Fattig; 1 2 , GEORGIA, Clark Co., Oct. 1965, R.

T. Franklin, ex Ips infested log of Pinus echinata; 2 2 , NORTH CAROLINA,

Durham Co., Durham, 30 May 1942, W. Haliburton, ex Ips avulsus LeConte.

All specimens in USNM.

Etymology?.— Named for my mother who unselfishly supported a young lad’s

interest in bugs.

Variation.— Females of zolae range in length from 2.5 to 3.2 mm. The intensity

and size of the forewing maculation varies from an extremely faint stain, confined

to the apical fourth of the wing, to a distinct dark spot on the apical fourth which

fades gradually and posteriorly to the apical third. The abdomen of all specimens

is shriveled, but T1 is about one-sixth the total length and is slightly emarginate

posteromedially. In females, the scape ranges from 0.68 to 0.75 times the distance

from the top of the toruli to the venter of the midocellus (n = 10; x = 0.72 ±

0.03); in males the range is 0.84 to 0.87 times the distance (n = 4; x = 0.86 ±

0.01). In females the length to width ratio of FI varies considerably and ranges

from 0.93 to 1.5 times as long as wide (n = 10; x= 1.2 ± 0.20). F6 ranges from

0.50 to 0.67 times as long as wide {n = 10; x = 0.62 ± 0.06). The few males

available for study have FI and F6 relatively longer than females with FI having

a range of 1.4 to 1.7 times longer than wide ( n = 4; x = 1.5 ± 0.14) and F6 0.75

to 0.88 times as long as wide (n = 3; x = 0.83 ± 0.07).

Distribution.— This species ranges throughout the southeastern United States,

from North Carolina south to Georgia and east to Texas.

Hosts. —Rhopalicus zolae is associated with pine log rearings usually containing

Ips and Dendroctonus frontalis. One pair of specimens bears the label “ex Ips

avulusf but I doubt that the association was actually proven.

Discussion.— Among Nearctic Rhopalicus, zolae and pulchripennis are super¬

ficially similar in appearance. The first species is apparently confined to the south¬

eastern United States, whereas the latter is largely northern in range. The species

VOLUME 59, NUMBERS 1-4

do overlap, however, and I have seen a few specimens of pulchripennis from Texas

and Mississippi. I know of at least two instances where zolae has been misidentified

as pulchripennis, and I am certain that some published records for pulchripennis

actually refer to zolae. This is especially true of papers treating the southern pine

beetle, Dendroct onus frontalis (e.g., Overgaard, 1968; Moser et al., 1971; Berisford,

undated). Specimens (in USNM) reared by Overgaard in Mississippi are zolae,

but they may have been reported as pulchripennis in the paper cited above.

The two species are fairly easily distinguished as follows: zolae has the length

of the scape at most three-fourths that of the distance to the venter of the midocel¬

lus (x = 0.72 ± 0.03; range 0.68 to 0.75; n = 10) (Fig. 22a), whereas pulchripennis

has the scape essentially as long as the distance to the midocellus (x = 1.1 ± 0.08;

range 0.93 to 1.2; n — 10) (Fig. 21a). This difference is actually a result of the

placement of the toruli of zolae at a point slightly below a line connecting the

ventral margins of the eyes, whereas in pulchripennis the toruli are distinctly

above such a line. In zolae the flagellomeres are generally wider than long (Fig.

22b) and in pulchripennis they are generally longer than wide (Fig. 21b), but

measurements tend to overlap on small specimens (ca. 3.0 mm) of pulchripennis,

thus making this character not reliable in every case. In addition, zolae has no

setae on the basal vein of the forewing, whereas pulchripennis has a row of setae

present.

Both zolae and the Palearctic species quadratus share the shortened scape and

flagellomeres just mentioned as well as having a dorsoventrally flattened scutum

and scutellum (i.e., both essentially in the same plane). In the Palearctic the degree

of convexity of the scutellum can be used to separate species (as, for example,

quadratus from guttatus) but in the Nearctic pulchripennis varies enough to make

the use of this character difficult within the framework of a Holarctic key.

Rhopalicus quadratus (Ratzeburg)

(Fig. 23)

Pteromalus quadratus Ratzeburg, 1844b:203, 2.

Pteromalus neostadiensis Ratzeburg, 1844b:204, 2.

Rhopalicus brevicornis Thomson, 1878:43, 2.

Discussion.—Rhopalicus quadratus is the only species with a hyaline forewing.

The shortened flagellum (FI -6 shorter than distance between eyes) which separates

quadratus from guttatus and pulchripennis is shared with zolae, but the two are

readily separated by the characters given in the key.

Material examined.— I have seen 37 specimens of this species from Europe

(BMNH, USNM).

Distribution. —Northwestern Europe.

Hosts. — Hedqvist (1963:80-81) gave a detailed list of the hosts for this species.

All known hosts are Scolytidae of the genera Tomicus, Carphoborus, Hylurgops,

Ips, Orthotomicus, Phloeosinus, and Pityogenes.

Rhopalicus guttatus (Ratzeburg)

Ichneumon (. Pteromalus) guttatus Ratzeburg, 1844a:29 (pi. 8, fig. 5), 2.

Discussion. — This species and pulchripennis share the lengthened flagellum (FI-

PAN-PACIFIC ENTOMOLOGIST

6 longer than distance between the eyes) and the convex scutellum. I can find no

differences between them except those noted in the key.

Material examined.— Thirteen specimens from Europe determined by Boucek

(BMNH) and one determined as a homotype by Ruschka (USNM).

Distribution. — Reported from Britain, Germany, Sweden, and Czechoslovakia

(Graham 1969:415), and known additionally from Portugal, Yugoslavia, and

Turkey (based upon specimens in BMNH).

Hosts. — Hedqvist (1963:82-83) reported that guttatus attacked several species

of Pissodes (Curculionidae), but he discounted a record for Ips (Scolytidae).

Rhopalicus pulchripennis (Crawford)

(Figs. 7, 8, 21)

Spintherus pulchripennis Crawford, 1912:168-169, 9 . Holotype 9 , USNM [ex¬

amined].

Rhopalicus americanus Girault, 1916:296-297, 9. Holotype 9, USNM [examined].

Discussion.— Gahan (1924:16) synonymized americanus with pulchripennis. I

agree with this conclusion based upon an examination of the type material. In

the Nearctic, pulchripennis is the only species to combine both a spot beneath

the marginal vein and the presence of setae on the basal vein. For additional

remarks see zolae.

Material examined.—In. addition to the types of pulchripennis (7 9 ) and ameri¬

canus (1 9 ), I have examined 123 specimens from North America.

Distribution.— Throughout southern Canada and the northern United States

with infrequent (but confirmed) records in Texas, Mississippi, and Virginia.

Hosts.— Recorded from numerous Coleoptera as follows: Scolytidae: Dendroc-

tonus brevicomis LeConte, D. murrayanae Hopkins, D. ponderosae Hopkins,

Pseudohylesine nebulosus (LeConte), Scolyte unispinosus LeConte; Curculioni¬

dae: Cylindrocopturus eatoni Buchanan, C. furnissi Buchanan, Pissodes approx¬

imate Hopkins, P. strobi (Peck), P. terminate Hopping. (Records of pulchri¬

pennis from Ips spp. (Overgaard, 1968) and from a combined rearing of

Dendroctonus frontalis Zimmermann and Ips spp. (Moser et al., 1971) probably

refer to Rhopalicus zolae\ see discussion under that species.)

Rhopalicus tutela (Walker)

(Fig. 24)

Cheiropachus tutela Walker, 1836b: 14-15, 6, 9. Lectotype 9, BMNH [examined].

Cleonymus maculifer Foerster, 1840:34, 9.

Pteromalus suspense Ratzeburg, 1844b: 189.

Pteromalus spinolae Ratzeburg, 1844b: 189, S, 9.

Pteromalus immaculate Ratzeburg, 1844b: 189, 205.

Pteromalus lunula Ratzeburg, 1848:193, 9 .

Pteromalus multicolor Ratzeburg, 1848:193 [n.n. for P. spinolae Ratzeburg 1844b:

189, nec Foerster, 1840:23].

Pteromalus aemulus Ratzeburg, 1848:203.

Rhopalicus annellus Thomson, 1878:42, 6, 9 .

Discussion.—Rhopalicus tutela is the largest (to 5 mm) and usually most easily

distinguished member of the genus based upon the infumate spot associated with

VOLUME 59, NUMBERS 1-4

the stigma (is is especially well developed in the males where it is associated with

an enlargement of the stigma. Fig. 24a). Females rarely have the spot reduced or

absent and the stigma is nearly always wider than high. In some specimens it may

be nearly as long as the stigmal vein (Fig. 24c), while in others it is slightly shorter

(Fig. 24b). The stigmal vein is definitely less than twice the length, however,

whereas in other species the stigmal vein is 3 or more times the length of the

stigma (Fig. 23). This species seems to be distantly related to others of the genus,

based upon the stigmal characters and those given in the key.

In 1934 approximately 1500 specimens of this species were introduced from

England into Canada for control of the Eastern spruce beetle, Dendroctonus ru-

fipennis (Kirby), (reported as piceaperda Hopkins: Scolytidae by McGugan and

Coppel, 1962); no “evidence of survival” was obtained. According to Britton

(1920), however, R. tutela (reported as suspensus Ratzeburg) was reared in 1914

from Pissodes strobi (Curculionidae) in New Haven, Connecticut. Specimens of

this rearing are now in the U.S. National Museum and confirm the fact that

Rhopalicus tutela occurred in the Nearctic before its release from Europe in 1934.

It is not known whether R. tutela could have been an accidental introduction of

earlier times.

Material examined. —In addition to the lectotype (BMNH), I have seen 95

specimens from Europe (BMNH, USNM) and 15 from the northeastern United

States (USNM).

Distribution.—Rhopalicus tutela is found from the northeastern United States

to Quebec and Ontario in Canada (Burks, 1979), is “widely distributed in Europe”

(Graham, 1969), and is reported from numerous localities in Japan (Kamijo,

1981).

Hosts. — In the Nearctic, this species has been reared from Ips pint (Say) (Sco¬

lytidae) and Pissodes strobi (Curculionidae). Hedqvist (1963) reported the biology

of this species in Europe and gave a complete host list which includes numerous

genera and species of Scolytidae as well as 3 species of Pissodes. Kamijo (1981)

stated that “in Japan this species is commonly found on trunks of pine and larch

infested by bark beetles, and to a lesser extent on those of spruce and fir.”

Unplaced Species

Rhopalicus pallipes Provancher

Rhopalicus pallpies [sic] Provancher, 1888:407, 2.

Burks (1964:1261) commented on the validity of this species. The type is lost,

and from indications cited by Burks this species is probably an eulophid. The

name was spelled pallipes in the index to Provancher’s work, and this is considered

to be the correct spelling.

Dinotiscus Ghesquiere

Dinotus Foerster, 1856:66, 70, 71. Preoccupied by Dinotus Guettard, 1770

( Vermes). Type-species: D. bidentulus Thomson, 1878; designated by Ashmead,

1904: 316. '

Dinotiscus Ghesquiere, 1946:370. New name for Dinotus Foerster (1856) nec

Guettard (1770).

PAN-PACIFIC ENTOMOLOGIST

This genus is presently known from 4 Palearctic species (Graham, 1969) and

6 Nearctic ones (Burks, 1979). In this paper, however, the number of previously

reported Nearctic species is reduced to 3 and a Palearctic species ( aponius Walker)

is reported as new to the Nearctic fauna. The genus now is made up of 6 species

with 2 Palearctic {colon, wichmanni), 2 Nearctic ( thornsoni, dendroctoni), and 2

Holarctic {aponius, eupterus).

Dinotiscus may be defined as follows: clypeus variable, either deeply emarginate

(with 2 well-defined lobes), with a slight emargination, or straight; venter of torulus

usually IX, and rarely 2X, own diameter above line connecting ventral margins

of eyes; thorax not similarly reticulate throughout, postalar plate vertically carinate

(Fig. 10; may appear elongately reticulate), epimeron usually smooth above, pro-

podeum nearly shiny or alutaceous medially; nucha merely a parallel sided rim

(Fig. 9), rarely intersected medially by a carina; supracoxal flange joined to the

nucha (as in Fig. 6); forewing with or without maculations; postmarginal and

marginal veins longer than stigmal, postmarginal longer than marginal, stigma

may be enlarged.

As with Rhopalicus, my concepts of the species are based upon Graham’s work

(1969), identification of Palearctic specimens based upon his key, and the types

of Nearctic species. Unfortunately there do not seem to be an abundance of specific

characters with which to separate species of the genus Dinotiscus. Although “typ¬

ical” forms of the stigmal shape are useful overall, the variability of small spec¬

imens (which tend to have the stigma disproportionately reduced) detracts from

the reliability of this character. I have included illustrations of typical stigmal

shapes to aid in recognition of the species (Figs. 26-28, 30-32).

Key to Female Dinotiscus

1. Apical free margin of clypeus deeply emarginate, forming 2 lobes laterally

(Fig. 34). 2

- Apical free margin of clypeus straight (Fig. 33) or feebly emarginate (Fig.

35) . 3

2. Stigma of forewing higher than wide (Fig. 26); basal vein and cell essen¬

tially bare (several setae may be present on basal vein); propodeum nar¬

rowed medially to ca. x h length of plica (Palearctic) . colon (Linnaeus)

- Stigma of forewing wider than high (Fig. 27); basal vein and cell with

some setae; propodeum medially at least % length of plica (Holarctic)

. aponius (Walker)

3. Flagellomeres 1 and 2 as long or longer than eye height or scape (each 3

to 4X longer than broad); upper epimeron sculptured much as lower

(Nearctic) . dendroctoni (Ashmead)

- Flagellomeres 1 and 2 shorter than eye height or scape (at most 2.5X as

long as broad); upper epimeron smooth. 4

4. Stigma of forewing wider than length of stigmal vein from marginal vein

to top of stigma (Fig. 28) (Palearctic) . wichmanni Boucek

- Stigma of forewing narrower than length of stigmal vein from marginal

vein to top of stigma (Figs. 30, 31). 5

5. Apical free margin of clypeus feebly emarginate (Fig. 35); prepectus not

evenly sculptured, dorsally with a shiny, usually pronounced, groove

VOLUME 59, NUMBERS 1-4

33 eupterus 34 colon

35 thomsoni

Figs. 25-35. Dinotiscus spp. 25, 29, parastigmal vein of forewing. 26-28, 30-32, stigma and stigmal

vein of forewing. 33-35, apical free margin of clypeus.

paralleling top border, surface of prepectus irregular, not in single plane

(Nearctic) . thomsoni (Crawford)

- Apical free margin of clypeus straight (Fig. 33); prepectus evenly sculp¬

tured and essentially in same plane (Holarctic) . eupterus (Walker)

Dinotiscus colon (Linnaeus)

(Figs. 25, 26, 34)

Sphex colon Linnaeus, 1758:571.

Dinotus calcaratus Thomson, 1878:40, 2.

Discussion. —Dinotiscus colon shares the deeply emarginate clypeus with apon-

ius from which it may be separated by characters in the key. Females of colon

tend to have an elongate abdomen which is about 3 times as long as the thorax,

whereas in aponius the abdomen is at most twice as long. Additionally in colon

the parastigma proximally forms a right angle with the submarginal vein (Fig.

25), but in aponius it gradually merges with the submarginal vein (Fig. 29).

Unfortunately the infumate spots associated with the wing veins are so variable

(even among a few specimens) as to be of little practical use. If developed, colon

has a spot at the parastigma and aponius does not.

Material examined.—I have seen 4 specimens of this species from Europe.

Distribution. — Britain, Sweden, Finland, Germany, and Czechoslovakia (Gra¬

ham, 1969).

PAN-PACIFIC ENTOMOLOGIST

Hosts. —Reared from Scolytidae as follows: Tomicus piniperda (Linnaeus), T.

minor (Hartig), and Ips acuminatus Gyllenhal (Hedqvist, 1963; reported as Din-

otiscus calcaratus).

Dinotiscus aponius (Walker)

(Figs. 27, 29)

Hetroxys aponius Walker, 1848:127, 215, 3, 9. Lectotype 3, BMNH [examined].

Pteromalus capitatus', Ratzeburg, 1848:196, pi. 3, fig. 7. (? misidentification of

capitatus Foerster, 1840).

Pteromalus patellatus Ratzeburg, 1848:196.

Dinotus bidentulus Thomson, 1878:39, 3, 9.

Discussion. — Graham’s (1969:411) citation of Pteromalus capitatus as being

described by Ratzeburg (1848) is in error. Ratzeburg correctly cited the specific

name as “32. P. capitatus Forst. (T. III. F. 7.).” However, under this species name,

Ratzeburg suggested that his specimens might represent a new species in which

case he proposed the name patellatus ; the description and diagnosis validate this

name. According to Graham (1969:411), the illustration in Ratzeburg (T. III. F.

7.) probably refers to what is currently called Dinotiscus aponius (Walker) and I

agree with this assessment. The wing venation is fairly diagnostic for the species.

As the specimens which Ratzeburg saw are presumably destroyed, the association

of names is provisional.

For comparative comments on this species see the key and discussion section

for D. colon. Dinotiscus aponius is apparently widespread in Europe and is re¬

ported for the first time in the Nearctic as a possible parasite of the smaller

European elm bark bettle (see hosts).

Material examined. — Forty-four Palearctic specimens (BMNH, USNM) deter¬

mined by Ruschka and Boucek; 3 female specimens from Detroit, Michigan

determined by Burks and confirmed by me. These latter specimens represent the

only record for this species in the Nearctic.

Distribution. —Northwestern and Central Europe (Graham, 1969), Japan (Kam-

ijo, 1981), and Michigan, U.S.A.

Hosts.— As summarized by Graham (1969), D. aponius in Europe attacks sco-

lytids as follows: Scolytus rugidosus Ratzeburg, S. multistriatus (Marsham), S.

ratzeburgi Jansson, and Hvlesinus varius (Fabricius) (reported as frax ini Panzer).

In the Nearctic, this species has been reared from elm infested with Scolytus

multistriatus (8 May 1973). According to Kamijo (1981) aponius appears to attack

scolytids on broad-leaved trees.

Dinotiscus dendroctoni (Ashmead)

(Fig. 32)

Cecidostiba dendroctoni Ashmead, 1874:337, 9. Lectotype 9, herein designated,

USNM.

Cecidostiba burkei Crawford, 1912:170, 9. Holotype 9, USNM [examined]. New

synonymy.

Discussion. —Ashmead’s dendroctoni was described from an unspecified num¬

ber of “specimens” in the U.S. National Museum. In the museum there are 3

specimens with red type labels. According to the type catalog, 2 specimens were

VOLUME 59, NUMBERS 1-4

originally logged as “types” in 1894 and a 3rd was labeled “type” by Gahan in

1928. As the species has had no official “type” designation, 1 have selected as

lectotype 1 of the 2 specimens originally registered in the type catalog. This

specimen has the head, wing, and antennae mounted on a slide. The specimen

labeled by Gahan as the 3rd type is one of a series on 9 pins (some with multiple

specimens) all bearing similar labels stating “Check list No. . . . , Det. by . . . .”

These may or may not actually have been examined by Ashmead and their status

is questionable. Whatever the status of these specimens, 2 were taken from the

series by Crawford (1912) and described as Cecidostiba ashmeadi which is a

synonym of D. eupterus. All the other specimens are eupterus as well.

Crawford (1912) separated his species burkei from Ashmead’s dendroctoni based

upon the supposedly longer postmarginal vein of the former. He had only the

holotype female for measurement. I have measured this specimen and the post¬

marginal is 1.3X the marginal, while in the lectotype of dendroctoni it is 1.2X. In

a reared series of 10 females from Colorado, the postmarginal ranges from 1.2 to

1.4X the marginal and burkei easily falls within this range. I consider them to be

synonyms, especially based upon the following autapomorphies which define the

species: the flagellomeres are all much longer than wide (ca. 3 to 4X), with FI

and F2 together longer than either the scape or the eye height (in other Dinotiscus

FI and F2 are shorter than the eye height or scape, and each flagellomere is rarely

up to 2.5X longer than wide); the upper and lower epimeron are similarly sculp¬

tured (upper epimeron polished in other species); and the last tergum is longer

than the postmarginal vein (noticeably shorter in other species). I have seen a few

specimens from Mexico (Chapingo; Coatepede) which agree with the above, but

the upper epimeron is sculptured only medially and is polished around this area.

These are the only specimens of Dinotiscus known from south of the United

States. I think additional specimens are needed to assess their status in relation

to typical dendroctoni.

Material examined. — In addition to the type series I have seen 80 specimens

of dendroctoni.

Distribution.— This species has been reported from British Columbia south to

New Mexico, Texas (and possibly Mexico), and east to Virginia (Burks, 1979).

Hosts.—Dinotiscus dendroctoni is known from the following hosts (Burks, 1979):

Cylindrocopturus furnissi Buchanan, Dendroctonus brevicomus LeConte, D. ru-

fipennis (Kirby), D. frontalis Zimmermann, D. ponderosae Hopkins, D. pseudo-

tsugae Hopkins, Ips grandicollis (Eichhoff), and Polygraphus rufipennis (Kirby).

The specimens from Mexico were associated with Pinus montezumae.

Dinotiscus wichmanni Boucek

(Fig. 28)

Dinotiscus wichmanni Boucek, 1967:635-636, 9, <3.

Discussion. — This species closely resembles eupterus and differs primarily by

the enlarged stigma (stigma wider than length of stigmal vein in wichmanni but

narrower in eupterus, cf. Figs. 28 and 30).

Material examined.— Forty-eight specimens from Yugoslavia, identified by

Boucek (BMNH).

Distribution.— Austria (Boucek, 1967) and Yugoslavia.

100

PAN-PACIFIC ENTOMOLOGIST

Hosts.— Known from Hylastinus obscurus (Marsham) (Scolytidae) on Cytisus

laburnum Linnaeus and Viburnum lantana Linnaeus, and from Hylastinus frank-

hauseri Reitter on Cytisus.

Dinotiscus thomsoni (Crawford)

(Figs. 9, 10, 31, 35)

Cecidostiba thomsoni Crawford, 1912:171, 2, 8. Lectotype 2, herein designated,

USNM.

Discussion.— This species Was described from “6 female and 6 male specimens

selected from a large series” now housed in the U.S. National Museum. From

among these, I have designated a female as lectotype. Dinotiscus thomsoni is

somewhat aberrant by virtue of its modified prepectus (i.e., fairly irregular in

surface contour and sculpture, with a smooth groove paralleling the dorsal margin)

and its slightly emarginate clypeus. Also, the stigma is produced angularly on the

inner ventral margin (Fig. 31) which is generally characteristic for the species.

The toruli are situated above the ventral margins of the eyes by a distance about

1.5 times their own diameter which is an additional character to separate thomsoni

from eupterus (situated less than own diameter above ventral eye margin).

Material examined. —In addition to the type series I have seen about 60 spec¬

imens of this species.

Distribution. — So far restricted to the northwestern United States.

Hosts.—Dinotiscus thomsoni is known from Pissodes sp. (Curculionidae),

Phloeosinus sp. (new record), and Scolytus ventralis LeConte (Scolytidae).

Dinotiscus eupterus (Walker)

(Figs. 30, 33)

Pteromalus eupterus Walker, 1836a:482, 2. Lectotype 2, BMNH [examined].

Pteromalus dimidiatus Walker, 1836b: 12, 2. Lectotype 2, BMNH [examined].

1 Pteromalus capitatus Foerster, 1840:21, 6.

Pteromalus lanceolatus Ratzeburg, 1848:204, <5, 2.

Dinotus clypealis Thomson, 1878:40, 8, 2.

Dinotus acutus Provancher, 1887:201, 2. New' synonymy.

Cecidostiba polygraphi Ashmead, 1894:338, 2, 8. Lectotype 2, herein designated,

USNM. New synonymy.

Cecidostiba ashmeadi Crawford, 1912:170-171, 2. Holotype 2, USNM [exam¬

ined].

Uriella pityogenis Ishii, 1939:189, 8, 2.

Discussion. — Burks (1964) saw the type of acutus and stated that ashmeadi

Crawford was a synonym. I have not seen the Provancher type, but I accept Burks’

placement. 1 have compared the known type material of polygraphi Ashmead (2

2, 1 8 syntypes) with ashmeadi Crawford (holotype and paratype) and find them

identical. Unfortunately only the paratype of ashmeadi is in good enough con¬

dition to be of value. I hereby designate the better of the 2 females of polygraphi

Ashmead as lectotype and have placed a label to this effect on the specimen.

This species and dendroctoni are the two most commonly encountered species

in the Nearctic. Both are widespread and parasitize numerous scolytid hosts on

VOLUME 59, NUMBERS 1-4

101

coniferous trees. The characters used to separate eupterus are given in the key

and an additional one is given in the discussion of thomsoni .

Material examined. — My concept of this species is based upon an examination

of about 80 specimens (USNM, BMNH), the lectotype series of P. eupterus

(BMNH), and the type series of C. polygraphi and C. ashmeadi (USNM).

Distribution.— This species is now considered to be Holarctic. The Nearctic

distribution includes Nova Scotia and Quebec south to Virginia and along the

northern United States to Oregon and California. The Palearctic records include

Great Britain, Sweden, Germany, and Central Europe (Graham, 1969), and Japan

(Kamijo, 1981).

Hosts. — In the Nearctic this species has been reared from Scolytidae as follows

(Burks, 1979): Dendroctonus ponderosae Hopkins and Polygraphus rufipennis

(Kirby). Palearctic hosts include scolytid species in the genera Polygraphus, Phtho-

rophloeus, Cryphalus, Pityophthorns, Pityogenes, and Dryocoetes (Graham, 1969),

and Ips (Kamijo, 1981).

Unplaced Species

Dinotus elongatus Ashmead

Dinotus elongatus Ashmead, 1888:175, 2. Holotype 2, USNM [examined].

I have examined the single holotype female specimen of this species and it is

definitely not a Dinotiscus. The pronotum lacks a transverse carina, and the

propodeum has a well developed, reticulate nucha. The type is fairly fragmentary,

but it should be placed near genera such as Dinarmus Thomson or Heteroschema

Gahan based on the structure of the propodeum.

Acknowledgments

I wish to thank Don Anderson (Systematic Entomology Laboratory, % U.S.

National Museum) for helping with current nomenclature in the Scolytidae and

Curculionidae. Zdenek Boucek (Commonwealth Institute of Entomology, British

Museum (National History)), Michael Schauff (Department of Entomology,

Smithsonian Institution), and Arnold Menke (Systematic Entomology Laboratory,

% U.S. National Museum) read the manuscript and offered numerous suggestions

for its improvement. Additionally, Mike Schauff prepared the scanning electron

micrographs (Figs. 7-12). I thank them for their help.

Literature Cited

Ashmead, W. H. 1888. On the chalcideous tribe Chiropachides. Can. Ent. 20:172-176.

-. 1894. Descriptions of new parasitic Hymenoptera. Trans. Amer. Ent. Soc. 21:318-344.

-. 1904. Classification of the chalcid flies. Mem. Carnegie Mus. 1 :i—xi, 225-551.

Berisford, C. W. [undated]. The southern pine beetle: Natural enemies and associated organisms,

pp. 30-52, Appendix tables 2, 3. In: R. C. Thatcher, J. L. Searcy, J. E. Coster, and G. D. Hertel.

U.S. Dept. Agnc., For. Ser., Sci. Educ. Admin. Tech. Bull. 1631, 267 pp.

Boucek, Z. 1967. New reared palearctic Pteromalidae. Acta Ent. Mus. Nat. Pragae 37:635-647.

Britton, W. E. 1920. The white pine weevil. Nineteenth report of the state entomologist for 1919.

Bull. Conn. Agric. Exp. Sta. No. 218:144-155.

Burks, B. D. 1964 (1963). The Provancher species of Chalcidoidea. Can. Ent. 95:1254-1263.

-. 1979. Pteromalidae, pp. 768-835. In: K. V. Krombein et al., eds., Catalog of Hymenoptera

102

PAN-PACIFIC ENTOMOLOGIST

in America north of Mexico. Vol. I. Symphyta and Apocrita (Parasitica). Smithsonian Institution

Press. Washington, D.C., 1198 pp.

Crawford, J. C. 1912. Descriptions of new Hymenoptera No. 5. Proc. U.S. Natl. Mus. 43:163-188.

Foerster, A. 1840. Beitrage zur Monographic der Pteromalinen, Nees. 1 Heft. Aachen, 47 pp., 1 pi.

-. Hymenopterologische Studien. 2 Heft. Chalcididae und Proctotrupii. Aachen, 152 pp.

Gahan, A. B. 1924. Some new parasitic Hymenoptera with notes on several described forms. Proc.

U.S. Natl. Mus. 65:1-23.

Ghesquiere, J. 1946. Contribution a l’etude des Microhymenopteres du Congo beige. X-XI. Rev.

Zool. Bot. Aff. 39:367-373.

Girault, A. A. 1916. New miscellaneous chalcidoid Hymenoptera with notes on described species.

Ann. Ent. Soc. Amer. 9:291-308.

Graham, M. W. R. de V. 1969. The Pteromalidae of Northwestern Europe. Bull. Brit. Mus. (Nat.

Hist.). Ent. Suppl. 16:1-908.

Hedqvist, K. J. 1963. Die Fiende der Borkenkafer in Schweden. I. Erzwespen (Chalcidoidea). Stud,

forestal. suec. no. 11:1-176.

Ishii, T. 1939. New chalcidoid and proctotiypoid wasps reared from forest insects by Dr. H. Kono.

Kontyu 13:187-191.

Kamijo, K. 1981. Pteromalid parasites of bark beetles from Japan, with descriptions of three new

species. Kontyu 49:86-95.

Linnaeus, C. 1758. Systema naturae (10th ed.). Vol. 1. Stockholm, 824 4- iii pp.

McGugan, B. M., and H. C. Coppel. 1962. A review of the biological control attempts against insects

and weeds in Canada. Part II. Biological control of forest insects, 1910-1958. Inst. Biol. Contr.

Comm. Trinidad, Tech. Comm. No. 2:35-216.

Moser, J. C., R. C. Thatcher, and L. S. Pickard. 1971. Relative abundance of southern pine beetle

associates in East Texas. Ann. Ent. Soc. Amer. 64:72-77.

O’Brien, C., and G. J. Wibmer. 1982. Annotated checklist of the weevils (Curculionidae sensu lato )

of North America, Central America, and the West Indies. Mem. Amer. Ent. Inst. 34:i-ix, 1-

382.

Overgaard, N. A. 1968. Insects associated with the southern pine beetle in Texas, Louisiana, and

Mississippi. J. Econ. Ent. 61:1197-1201.

Provancher, L. 1887-1888. Chalcidides, pp. 184-211 (1887), pp. 406-407 (1888). Additions et

corrections au volume II de la Faune Entomologique du Canada. C. Darveau, Quebec, 477 pp.

Ratzeburg, J. T. C. 1844a. Die Forst-Insecten. 3. Berlin, viii + 314 pp., 16 pis.

-. 1844b. Die Ichneumonen der Forstmsecten in entomologischer und forstlicher Beziehung.

1. Berlin, 224 pp., 4 pis.

-. 1848. Die Ichneumonen der Forstinsecten in entomologischer und forstlicher Beziehung. 2.

Berlin, vi + 238 pp., 4 tables, 3 pis.

Thomson, C. G. 1878. Hymenoptera Scandinaviae. 5. Pteromalus (Svederus) continuatio. Lund,

307 pp., 1 pi.

Walker, F. 1836a. Monographia Chalciditum. Ent. Mag. 3:465-496.

-. 1836b. Monographia Chalciditum. Ent. Mag. 4:9-23.

-. 1848. List of the specimens of hymenopterous insects in the collection of the British Museum.

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monograph. Great Basin Nat. Mem. No. 6:1-1359.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 103-107

A New Species of Bembix from Lower California

(Hymenoptera: Sphecidae)

Terry L. Griswold

USDA Bee Biology and Systematics Laboratory, Utah State University, Logan,

Utah 84322.

A collection of aculeate Hymenoptera obtained by William H. Clark and his

associates during ecological studies in central Lower California was received for

identification. Among this material was an undescribed species of Bembix Fa-

bricius which is here named after Richard M. Bohart, in recognition of his con¬

tribution to the taxonomy of Bembicini.

Bembix boharti Griswold, New Species

(Figs. 1, 2)

Male.— Length, 22 mm; forewing length, 14 mm. Black with extensive yellow

markings, except white postocularly, laterally on scutellum, medially on terga I-

V; pale markings as follows: scape; pedicel ventrally; lower half of frons with

linear extension dorsally along eye margin to level of midocellus, linear vertical

mark below midocellus; clypeus; labrum; mandible except teeth; posterior margin

of eye; pronotum except large quadrate dorsal mark and irregular mark at pronotal

angle; anterior half of tegula; anterior half of costal lamella; posterolateral corner

of scutum; mark on mesopleuron, vertical on upper half of sclerite then obliquely

angled ventrally toward anterior margin; basalar area of mesopleuron; basalar

lobe of metapleuron; most of metapleuron; lateral portion of propodeum; meso-

sternum medial to midcoxa; legs except coxae basally, trochanters dorsally, femora

at extreme base and central posterior region; tergum I with transverse mark

broadly interrupted medially; tergum II-V with transverse mark narrowly inter¬

rupted medially, submedially emarginate anteriorly; tergum VI with transverse

mark narrowly interrupted medially; tergum VII apically; sternum I except basal

inverted “W”-shaped area; sternum II except linear basomedial line and antero¬

lateral corner; sternum III-V basomedially, anterolaterally; basal third of sternum

VI; basal two-thirds of sternum VII; medial mark on dorsal surface of paramere.

Wings hyaline; veins reddish except subcosta black basally to junction with basal

vein. Body clothed with dense, silvery pubescence.

Midocellus a linear crescent; flagellomeres IV-VII with teeth, those on articles

VI, VII spinose; flagellomeres IV-X with tyloids; flagellomeres VIII-X expanded

ventrally, broader than preceding segments; flagellomere XI elongate, curved;

scape length greater than clypeal height; labrum without median angulation, length

more than twice clypeal height; mandible slightly curved apically, inner tooth

distinct; forebasitarsus with 7 rake setae; midfemur with small denticles ventrally

on apical four-fifths; midtibia with ventral flange on apical half (Fig. 1); length of

midtibial spur less than one-half length of midbasitarsus; first intersubmarginal

vein of forewing with strong, step-like double bend; tergum VII broadly rounded

apically, without lateral angle; spiracular lobe broad, rounded apically, length one-

104

PAN-PACIFIC ENTOMOLOGIST

Figs. 1, 2. Bembix boharti. 1, Portion of male midleg showing distally enlarged tibia. 2, Male

genital capsule (left side, ventral view; right side, dorsal view).

half that of tergum VII; sternum II with prominent, blunt, median keel; sternum

VI with broad, ventrally flattened median process notched apically, ventral surface

parallel to surface of sternum; sternum VII with submedian longitudinal carina

in addition to apically notched median carina; genitalia as in Fig. 2.

Female. — Length, 19-21 mm; forewing length, 13-14 mm. Markings as in male

except: white markings reduced to medial areas of terga I-IV; pale markings of

pronotum more extensive; scutum with anterolateral and discal stripes; meso-

pleural mark expanded posteroventrally to form rough triangle, mesopleuron with

additional narrowly isolated mark dorsoposteriorly; metapleuron entirely pale;

propodeum with lateral mark expanded to posterior face, additional marks along

posterior margin of propodeal triangle; terga II and sometimes III with markings

expanded anteriorly forming small enclosed black spots; tergum VI yellow except

basally; pale markings of sternum I expanded; sternum II pale except for oval

basomedial area; sternum VI pale except for narrow medial line; wing venation

darker.

Structures as in male except flagellomeres, midfemur, midtibia, sterna not mod¬

ified; ratio of maximum length of flagellomere I to maximum length of scape

between 1.04 and 1.11; ratio of length of flagellomere I to height of clypeus between

1.22 and 1.34; forebasitarsus with 7 rake setae, length of distal ones more than

twice width of basitarsus; pygidium slightly roughened, without lateral angle.

Type material.— Holotype 3, MEXICO, Baja California {Norte), 9 km NW

Rancho Santa Ines, 29°46 / N, 1 14°46'W, 10-VI-79, W. H. Clark. Paratypes: MEX¬

ICO: Baja California {Norte): 12 <$, 7 9, same data as holotype; 1 <5, 2 9, same

VOLUME 59, NUMBERS 1-4

105

except 11-VI-79; 1 2, same except 14-VI-79; 1 <5, 2 9, same except 17-VI-79; 1 9,

same except 19-VI-79; 6 <3, 20 9, same except E. R. Tinkham; 1 9, same except

11-VI-79, J. Hoch; 1 2 , same except 12-VI-79; I 6, 1 2 , 9.6 km (6 mi) NW Rancho

Santa Ines, 29°43'N, 114°43'W } 17-VI-79, J. Miles; 1 2, 1.6 km (1 mi) NE Santa

Catarina Ranch, 29°44 / N, 115°09 / W, 20-VI-79, W. EE Clark; 1 2, 8 km NW Santo

Tomas, 31°37 / N, 116°27'W, 200 m, 9-VI-79, W. H. & M. H. Clark; 1 6, El Marmol,

18-VI-38, Michelbacher & Ross. Baja California Stir: 1 <3, 4 2, La Paz and vicinity,

11/14-VI-75, H. E. Evans; 1 2, same except H. Evans, W. Rubink, & D. Gwynne;

1 <3, 1 2, Cape San Lucas, 19-III-28, T. Craig; 2 2, La Paz, 3-VI-21, E. P. VanDuzee;

1 2, same except 4-VI-21; 2 2, La Paz, 7.6 m (250, 12-V-69, S. C. Williams; 2 2,

4.8 km (3 mi) W San Miguel de Comondu, 457 m (15000, 21-IV-69, S. C.

Williams. Holotype will be deposited at the California Academy of Sciences,

paratypes at the Museum of Natural History, College of Idaho, Caldwell, Idaho,

California Academy of Sciences, Colorado State University, University of Cali¬

fornia at Davis, U.S. National Museum, USDA Bee Biology and Systematics

Laboratory, Logan, Utah, and the personal collection of E. R, Tinkham.

Additional material. — MEXICO, Baja California (Norte): 2 2, km 76.7 on road

to Sierra San Pedro Martir National Park, 21-VI1-77, D. Weissman &C. Mullinex;

1 2, same except km 76.5, 20-VII-77.

Discussion.— This species belongs to the amoena species-group of Evans and

Matthews (1968) which contains two other species, B. amoena Handlirsch and

B. sayi Cresson. B. boharti is close to B. sayi, and given the wide distribution

and considerable variation of B. sayi it might be questioned whether B. boharti

is distinct. However, I have studied 56 males and 51 females of B. sayi from

throughout its range and find there to be distinct and consistent differences between

the two species. Further, the range of B. sayi extends westward into California

only in the northern Mojave Desert, not entering either the Colorado Desert or

cismontane southern California. Thus, B. boharti, which appears to be restricted

to central and southern Lower California, is isolated from B. sayi by these inter¬

vening regions. The male of B. boharti runs to couplet 9 in Evans and Matthews’

key. It can be separated from the other two species in this group by the strong,

apically expanded ventral keel on the midtibia, the absence of a polished basal

welt on flagellomere XI, and the more evenly rounded apicolateral margin of the

paramere. It also differs from B. amoena in the short midtibial spur (less than

half length of midbasitarsus) and the broad spiracular lobe of tergum VII, and

from B. sayi by the median platform of sternum VI parallel to the surface of the

segment and sternum VII with strong lateral carinae. Males of B. boharti vary

slightly in the extent of the pale markings. Some possess faint white discal marks

on the scutum.

The female of B. boharti is easily distinguished from B. amoena by the short

midtibial spur (less than half length of midbasitarsus). It runs to B. sayi in the

key of Evans and Matthews (1968). The only consistent difference found between

female B. boharti and B. sayi is in the length of flagellomere I relative to the

maximum scape length (Fig. 3). The ratio of these lengths is greater in B. boharti

(flagellomere I length/maximum scape length: x = 1.07 ± 0.02, range = 1.04-

1.11, 7? = 54) than in B. sayi (x = 0.96 ± 0.03, range = 0.86-1.00, n = 51). Note

that the ranges of the ratios do not overlap. A Mann-Whitney U Test comparing

the ratios of these two species was significant (P < 0.0001). The length of flagello-

106

PAN-PACIFIC ENTOMOLOGIST

1 1.1 1.2 1.3 1.4 1.5 1.6

Maximum Length Scape (mm)

Fig. 3. Plot of length of flagellomere I against maximum scape length for female B. boharti and

B. sayi.

mere I in relation to the height of the clypeus is also usually greater in B. boharti

(flagellomere I length/median clypeal height: x = 1.27 ± 0.03, range = 1.22-1.34,

n = 54) than in B. sayi (x= 1.10 ± 0.06, range = 0.97-1.22, n = 51). A Mann-

Whitney U Test comparing these ratios was also significant (P < 0.0001). An

additional character that will serve to differentiate most specimens is the absence

of a complete longitudinal black stripe on the venter of the abdomen and the

presence of a fine medial black line on sternum VI. However, specimens of B.

boharti from near Sierra San Pedro Martir National Park have the black areas

more extensively developed and occasional specimens of both B. sayi and B.

arnoena have reduced black areas approaching the condition of B. boharti.

Evans (1976) recorded B. sayi from Baja California Sur, described aspects of

its nesting biology, and commented that it differed from populations at several

locations in the United States in its “mound-building behavior.” I have studied

these specimens and find them to be representatives of B. boharti. Thus, the

differences in nesting biology noted by Evans may be of species significance. Fox

(1923) also recorded B. sayi from Lower California. These records also refer to

B. boharti.

Acknowledgments

I wish to thank William Clark, Museum of Natural History, College of Idaho,

Caldwell, Idaho; Wojciech Pulawski, California Academy of Sciences; Howard

VOLUME 59, NUMBERS 1-4

107

Evans, Colorado State University; Richard Bohart, University of California at

Davis; and Ernest Tinkham, Indio, California for the loan of material, Frank

Parker for reviewing the manuscript, and Vincent Tepedino for assistance with

statistical details.

Literature Cited

Evans, H. E. 1976. Bembicini of Baja California Sur: Notes on nests, prey, and distribution. Pan-

Pac. Ent. 52:314-320.

Evans, H. E., and R. W. Matthews. 1968. North American Bembix, a revised key and suggested

grouping. Ann. Ent. Soc. Amer. 61:1284-1299.

Fox, C. L. 1923. Expedition of the California Academy of Sciences to the Gulf of California in 1921.

The Bembicini (digger wasps). Proc. Calif. Acad. Sci. (4)12:429-436.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 108-112

Three New Species of New Zealand Tar digrades

(Tardigrada: Echiniscidae)

Donald S. Horning, Jr. and Robert O. Schuster

(DSH) Macleay Museum, University of Sydney, New South Wales 2006, Aus¬

tralia; (ROS) Department of Entomology, University of California, Davis, Cali¬

fornia 95616.

Professor Walter Maucci (pers. comm.) indicated that the identification of

Pseudechiniscus lateromamillatus Ramazzotti (1964) in Horning, Schuster and

Grigarick (1978) was in error. A re-examination of the New Zealand specimens

considered to be variants of P. lateromamillatus led to the recognition of three

undescribed species.

We have retained these new species in Pseudechiniscus. The pseudosegmental

area of these species is too large for Echiniscus and hardly large enough for

Pseudechiniscus. These species could be placed in either genus, or a new genus.

The problem is confounded by questionable illustrations of named species and

the lack of a comparative study of the morphology of this part of the family. Eye

pigment is lacking, probably due to the fixation procedures.

The holotypes of the three new species are deposited in the National Museum

of New Zealand, Wellington (NMNZ). Paratypes are deposited in NMNZ and the

Department of Entomology Museum, University of California, Davis (UCD). The

number of paratypes from any sample is given parenthetically following the NZ

sample number. The sample numbers and their associated data have been listed

previously (Horning et al., 1978) and, unless otherwise noted, the collector was

D. S. Horning, Jr.

We dedicate this article to R. M. Bohart, our mentor for many years, on the

occasion of his 70th birthday.

Pseudechiniscus conversus Horning and Schuster, New Species

(Figs. 1, 2)

Length of dorsum excluding legs IV 188.5 /urn; including legs IV 203 /urn. Cuticle

with granular pattern; granules largest (1.3-2.5 /um diameter) on scapular plate,

median line and posterior margins of plates C and D, and terminal plate; gran¬

ulation of head plate slightly smaller, and of posterior of third median still small¬

er; remaining areas (stipple of Fig. 1) with granule diameter 0.3-0.4 u m; very

infrequent light spots (pores) randomly distributed. Head with simple buccal cirri,

interior 9.5 /um long, exterior 19 /um; buccal papilla 6 /um long, 4 fim wide; eye

pigment absent. Scapular plate with cirrus A 52 /um long, clava 6 /um long, 3 mm

wide. Lateral spines present at positions B, C, D and E, short, triangular, about

6 /um long. Leg I spine bluntly conical, 2.9 /urn long; leg IV papilla 3.3 /urn long;

inner claws of all legs with basal spur.

Discussion. — This species was the basis for the misidentification of P. latero¬

mamillatus Ramazzotti in Horning et al. (1978). It fits the description of that

VOLUME 59, NUMBERS 1-4

109

Figs. 1, 2. Pseudechiniscus conversus. 1, dorsal aspect. 2, detail of scapular cuticle.

species, but examination of a paratype revealed that lateromamillatus has pores

instead of granules as the major cuticular feature.

The cuticular pattern of diversely sized granules, the largest occurring in pre¬

cisely limited areas, and the near absence of pores are the most obvious recog-

nitional features of this species.

Etymology?. —Latin: conversus, opposite; noting the reverse of cuticular char¬

acteristics distinguishing the species from P. later omamillatus.

Types.— Holotype and 1 paratype: South Island : Canaan Road Terminus, Abel

Tasman National Park, 9-IV-1971, NZ 525 (holotype) (NMNZ—Arthropoda 10/

32) and NZ 526 (NMNZ).

Habitat. —The holotype was found on the foliose lichen Lobaria adscripta (Nyl.)

Hue on live Nothofagus sp. in a shaded beech forest. The paratype was found on

foliose lichens, mostly Pseudocyphellaria spp., on live Nothofagus in an open,

wind-exposed meadow. The sample contained 11 species of tardigrades, the high¬

est number of species found in 577 New Zealand tardigrade-positive samples.

Pseudechiniscus parvisentus Horning and Schuster, New Species

(Figs. 3, 4)

Length of dorsum excluding legs IV 157 jam; including legs IV 171 gm. Cuticle

of dorsal plates consists of granules and pores of generally uniform size; granules

110

PAN-PACIFIC ENTOMOLOGIST

Figs. 3, 4. Pseudechiniscus parvisentus. 3, dorsal aspect. 4, detail of scapular cuticle.

of about 0.3 diameter. Head with accuminate buccal cirri; interior cirrus 6.7

pm, exterior 10 ^.m long; buccal papilla 6 long, 4 pm wide; eye pigment absent.

Scapular plate with cirrus A 33 pm long; papilla 6 pm long, 3 pm wide. Lateral

spines 6 pm to 12 pm long, thick at base and tapered distally, present at positions

B, C, D and E. Leg I spine minute, about 2 pm long, 1 pm wide; leg IV papilla

3 pm long, 2 pm wide; internal claws of all legs with basal spur.

Discussion .—This species is similar to P. conversus in respect to size and ar¬

rangement of spines. However, the pattern of the dorsal plates is uniformly of

small granules and pores except that along the posterior margin of the terminal

plate the granules are slightly larger and pores are infrequent and obscure.

Etymology. — Latin: parvus, little; sentus, spine; in reference to the short lateral

spines.

Types .—Holotype and 5 paratypes: South Island : Otepatotu Bush, Banks Pen¬

insula, 3-XI-1970, NZ 392 (holotype) (NMNZ-Arthropoda 10/33), NZ 596 (1,

NMNZ). Additional paratypes: South Island, West Harpers Hut, 14 km S Bealey,

12-IV-1971, H. A. Best, NZ 539 (1, NMNZ), NZ 544 (1, UCD); Lewis Pass, 11-

V-1970, NZ 106 (1, NMNZ); Halpins Creek, Arthurs Pass National Park, 18-V-

1970, NZ 149 (2, NMNZ). North Island : Mountain House Road, Mt. Egmont

National Park, 23-V-1971, NZ 746 (1, NMNZ), NZ 752 (1, NMNZ); Ohakune

Scenic Reserve, 21-V-1971, NZ 711 (3, UCD).

Habitats.— All habitats were in wet Nothofagus forests. Pseudechiniscus par-

VOLUME 59, NUMBERS 1-4

111

Figs. 5-7. Pseudechiniscus perplexus. 5, dorsal aspect. 6, detail of scapular cuticle. 7, detail of

terminal cuticle.

visentus was found on mosses on trees and shaded rocks (five samples), foliose

lichens, Sticta spp., on trees (two samples) and one hepatic sample on a tree.

Pseudechiniscus perplexus Horning and Schuster, New Species

(Figs. 5-7)

Length excluding legs IV 193 pm, including legs IV 218 pm. Cuticle of dorsal

plates consists of granules generally of about 0.6 pm diameter, and frequent larger,

randomly spaced pores; granules larger on median line and posterior margins of

plates C and D. Head with accuminate buccal cirri, interior 11 pm long, exterior

23 pm long; buccal papilla 9 pm long, 7 ;um wide, eye pigment absent. Scapular

plate with cirrus A 58 pm long; clava 7 ^um long, 5 ^m wide. Lateral spines B, C,

D, and E present; approximate lengths of B 80 pm, C 140 pm, D 150 pm and E

170 pm. Leg I with acute spine 3 pm long; leg IV papilla about 4 ^,m long; inner

claws of all legs with basal spur.

Discussion .—The dorsal pattern of this species is pronounced in some areas,

similar to the pattern of P. conversus, and the presence of both granules and pores

is similar to the cuticle of P. parvisentus.

112

PAN-PACIFIC ENTOMOLOGIST

Etymology.— Latin: perplexus, intricate; for the beautifully intricate pattern of

bright and dark spots of the cuticle.

Types. — Holotype: South Island : Gouland Downs Hut, Heaphy Track, 12-IX-

1971, Margaret E. Miller, NZ 1109 (NMNZ—Arthropoda 10/34). Paratypes:

South Island : Halpins Creek, Arthurs Pass National Park, 12-X-1970, NZ 373

(1, NMNZ); Canaan Road Terminus, Abel Tasman National Park, 9-IV-1971,

NZ 526 (2, NMNZ; 3, UCD).

Habitats. — This species was found on foliose lichens, mostly Pseudocyphellaria

spp., on live trees and shrubs. It was also found with Pseudechiniscus conversus

(NZ 526).

Key to New Zealand Pseudechiniscus

1. Dorsal plates B, C, D and E with lateral spines . 2

- Dorsal plates B, C, D and E without lateral spines . 4

2. Dorsal plate pattern almost entirely granules, pores infrequent.

. P. conversus Horning and Schuster, n. sp.

- Dorsal plate pattern of both granules and frequent pores . 3

3. Lateral spines short (6 to 12 pm) .

. P. parvisentus Horning and Schuster, n. sp.

- Lateral spines long (80 to 150 ^m) ....

. P. perplexus Horning and Schuster, n. sp.

4. Pseudosegmental plate with simple posterior margin .

. P. s. suillus (Ehrenberg)

- Pseudosegmental plate with 2 broad, short spines on posterior margin

. P. n. novaezeelandiae (Richters)

Acknowledgments

Professor Walter Maucci kindly arranged the loan of a paratype specimen of

Pseudechiniscus lateromamillatus from the Istituto Italiano di Idrobiologia, Pal-

lanza, and Dr. Diane Nelson examined the specimens and illustrations and crit¬

ically read the manuscript.

Literature Cited

Horning, D. S., Jr., R. O. Schuster, and A. A. Grigarick. 1978. Tardigrada of New Zealand. New

Zealand J. Zoo. 5:185-280.

Ramazzotti, G. 1964. Tardigradi del Cile, III, con descrizione delle nuove specie Oreella minor e

Pseudechiniscus lateromamillatus. Atti Soc. Ital. Sc. Nat. 103:347-355.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 113-139

The boharti Species Group of the Genus Pherocera

(Diptera: Therevidae: Phycinae)

Michael E. Irwin

Illinois Natural History Survey, and the University of Illinois, 607 East Pea¬

body, Champaign, Illinois 61820.

Introduction

The published history of Pherocera is meager. The genus was described by Cole

(1923a) for three species: P. albihalteralis Cole, P.flavipes Cole, and P. signatifrons

Cole, with P. signatifrons designated as the type-species. At that time, Cole had

available for study only eight female specimens, six of which were P. signatifrons.

That same year, Cole (1923b) described a fourth species, Pherocera nigripes Cole,

based on a single female from Mexico. Except for passing mention by Hardy

(1927:385), Rodendorf (1964:86), Cole (1965:349), and Cole and Schlinger (1969:

170, 172), no reference was made of the genus Pherocera until the 1970’s.

An important factor in any account of Pherocera is its historical rarity in

entomological collections. Until the beginning of 1940 only 18 specimens were

known to exist in collections, and during the 1940’s only 34 were added. After

1952 the number of specimens of this genus increased greatly, due largely to the

collecting efforts of Evert I. Schlinger. I became interested in the group and added

considerably to the collections between 1963 and 1970. During the 1970’s spec¬

imens were gathered mainly through the efforts of Saul I. Frommer who main¬

tained a malaise trap in the P. L. Boyd Desert Research Center near Palm Desert,

Riverside County, California.

The genus Pherocera has a restricted distribution, being confined to western

North America. Its northern limit, according to specimens examined, is in Oregon

and Idaho; its southern limit is near Puebla, Mexico. It has been collected as far

east as Colorado, New Mexico, and western Texas. Its current center of distri¬

bution is southern California.

I completed a study of Pherocera and its relatives in 1971 (Irwin, 1971) and

published a paper on two closely related genera, Schlingeria Irwin and Para-

pherocera Irwin, six years later (Irwin, 1977). Previously I published on the com¬

parative morphology of the female terminalia and oviposition behavior in the

Therevidae, including a section on Pherocera (Irwin, 1976). A review of the

Nearctic genera of Therevidae (Irwin and Lyneborg, 1981a) contained a rede¬

scription of the genus Pherocera and included the following statement (p. 268):

“Thirty species of the genus Pherocera have been described but not formally

published.” My dissertation (Irwin, 1971) divided the 34 recognized species of

Pherocera into seven species groups, one of which included P. fiavipes and seven

additional species. Although designated the ‘ fiavipes group” in the dissertation,

I am renaming it the “ boharti group” in honor of Richard M. Bohart, mentor

and esteemed friend. This paper describes the seven unpublished species of the

boharti group and updates the description of P. fiavipes.

114

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Pherocera rupina, female (1725), lateral view.

Acknowledgments

Richard M. Bohart, more than any person, bolstered my early interest in insect

systematics. I strongly support the efforts of Arnold Menke and Eric Grissell in

dedicating this issue of Pan-Pacific Entomologist to Dick for his generous and

enriching fellowship. To Dick and his gracious wife, Margaret, I extend heartfelt

and eternal gratitude.

I earnestly thank the following for contributing to this manuscript: Evert I.

Schlinger, University of California at Berkeley; Saul I. Frommer, University of

California at Riverside; Donald W. Webb, David J. Voegtlin, and John K. Bouse-

man, Illinois Natural History Survey, for critical review of the manuscript; and

John P. Sherrod, Illinois Natural History Survey, for drawing Figs. 6-11. The

curators of several of the major entomological museums are gratefully acknowl¬

edged for loaning valuable material.

Support for this study was supplied in part by a National Defense Education

Act Fellowship, the University of California Regents Patent Fund, the University

of California Dry Lands Institute, the University of California Deep Canyon

Research Fund, the Illinois Natural History Survey, and the University of Illinois.

Methods and Procedures

Specimen number.—Each specimen has been assigned a unique number to

facilitate the association of data. The number appears below the specimen on a

yellow label bearing the following words: THEREVIDAE/M. E. IRWIN/SPEC¬

IMEN #. Numbers referring to specimens will not be found in the text even though

they have been used to incorporate ecological and label data associated with the

specimens into an automated data management system designed by Rauch (1970).

These numbers have been associated with individual specimens in the figures,

however. If ecological or depository information regarding individual specimens

is needed, please contact the author.

VOLUME 59, NUMBERS 1-4

115

Format of “Specimens examined” section. — To conserve space and include as

much information about each specimen as possible, the following layout, adopted

from Irwin (1977), was used in the “Specimens examined” section under each

species. Many of these terms are more fully explained by Stuclcenberg and Irwin

(1973): 1) LARGEST POLITICAL UNIT [country, or state within the United

States, in capital letters], 2) intermediate political unit [state or province, or county

within the United States, in boldface print], 3) smallest political unit [city or town

in regular print], 4) modifier of smallest political unit [distance (km) and direction,

or subunit, in regular print], 5) elevation [in meters (m) above sea level, in boldface

print], 6) year/month/day that the specimen was collected [year expressed by two

digits, the “19” has been omitted since all specimens examined were collected in

the 1900’s], 7) collectors) [acronym(s), not in parentheses, expanded below], 8)

number of specimens of each sex [6 or 2], and 9); [a semicolon indicates that data

for previous specimens terminate and data for the next series of specimens follow].

If subsequent specimen data are not included, data not repeated are the same as

those of preceding specimens.

Collectors.— AFH, Howland, A. F.; ALM, Melander, A. L.; ARG, Gillogly, A.

R.; ATM, MacClay, A. T.; CAT, Toschi, C. A.; CTB, Brues, C. T.; DEF, Fox,

D. E.; EIS, Schlinger, E. I.; EMP, Painter, E. M.; FDP, Parker, F. D.; FXW,

Williams, F. X.; GRB, Balmer, G. R.; HAH, Hunt, H. A.; HKC, Court, H. K.;

HRM, Moffit, H. R.; JAP, Powell, J. A.; JCB, Bradley, J. C; JCD, Downey, J.

C.; JCH, Hall, J. C.; JFL, Lawrence, J. F.; JRQ, Quezada, J. R.; JT, Turner, J.;

JW, Wilcox, J.; JWM, MacSwain, J. W.; KWB, Brown, K. W.; LAS, Stange, L.

A.; LFL, Lapre, L. F.; MEI, Irwin, M. E.; MTJ, James, M. T.; PAR, Rauch, P.

A.; PDH, Hurd, P. D.; PH A, Arnaud, P. H.; RAS, Stirton, R. A.; RCB, Bechtel,

R. C.; RES, Somerby, R. E.; RHP, Painter, R. H.; RMB, Bohart, R. M.; RMW,

Worley, R. M.; SIF, Frommer, S. L; SL, Larisch, S.; SLF, Frommer, S. L.; WAS,

Steffan, W. A.; WI, Icenogle, W.; WJH, Hanson, W. J.; WRM, Mason, W. R.

Depositories. — Specimens from this study have been deposited with the follow¬

ing museums: AMNH, American Museum of Natural History, New York; ANSP,

Academy of Natural Sciences, Philadelphia, Pennsylvania; ASU, Arizona State

University, Tempe; AMS, Australian Museum, Sydney; BMH, Bernice P. Bishop

Museum, Honolulu, Hawaii; BMNH, British Museum of Natural History, Lon¬

don, England; BYU, Brigham Young University, Provo, Utah; CAS, California

Academy of Sciences, San Francisco; CIS, California Insect Survey, University

of California, Berkeley; CMNH, Field Museum of Natural History, Chicago,

Illinois; CMP, Carnegie Museum, Pittsburgh, Pennsylvania; CNC, Canadian Na¬

tional Collection, Ottawa; CSDA, California State Department of Agriculture,

Sacramento; CSIR, Commonwealth Scientific Industrial Research Organization,

Canberra, Australia; CSU, Colorado State University, Fort Collins; CU, Cornell

University, Ithaca, New York; DEI, Deutsches Entomologisches Institut, Berlin,

East Germany; DSIR, Department of Scientific and Industrial Research, Nelson,

New Zealand; DZSA, Departamento de Zoologia Agricultura, Sao Paulo, Brazil;

EEA, Estacion Experimental Agronomica, Universidad de Chile, Maipu; FSCA,

Florida State Collection of Arthropods, Gainesville; IAS, Institute of Agricultural

Sciences, Tokyo, Japan; IE, Instituto di Entomologia, Bologna, Italy; IML, In¬

stitute Miguel Lillo, Tucuman, Argentina; INHS, Illinois Natural History Survey ,

Champaign; INIA, Instituto Nacional de Investigaciones Agricolas, Chapingo,

116

PAN-PACIFIC ENTOMOLOGIST

Fig. 2. Pherocera rupina, female (1725), head, frontal view. Arrows give linear measurements

corresponding to continuous character states in text: 2 = head width; 3 = distance from center of frons

to compound eye margin at dorsal margin of antennal socket; 4 = interocular distance at vertex; 5 =

width of frontal callosity; 6 = height of frontal callosity.

Mexico; IOC, Instituto Oswaldo Cruz, Rio de Janeiro, Brazil; IRSN, Institut Royal

des Sciences Naturelle de Belgique, Brussels; ISU, Iowa State University, Ames;

ITM, Instituto Technologico y de Estudios Superiores, Monterrey, Mexico; KSU,

Kansas State University, Manhattan; KUF, Kyushu University, Fukuoka, Japan;

LACM, Natural History Museum of Los Angeles County, California; LBSC, Cal¬

ifornia State University at Long Beach; MCZ, Museum of Comparative Zoology,

Harvard University, Cambridge, Massachusetts; MEI, M. E. Irwin Collection;

MHN, Museo de Historia Natural Javier Prado, Lima, Peru; MMB, Moravske

Museum, Brno, Czechoslovakia; MNH, Musei Nationalis Hungarici, Budapest,

Hungary; MNHN, Museum National d’Histoire Naturelle, Paris, France; MSU,

Michigan State University, East Lansing; NCSU, North Carolina State University,

Raleigh; NMB, Naturhistorisches Museum, Basel, Switzerland; NMP, Natal Mu¬

seum, Pietermaritzburg, South Africa; NMSU, New Mexico State University, Las

Cruces; NSDA, Nevada State Department of Agriculture, Reno; OSM, Ohio State

Museum, Columbus; OSU, Oregon State University, Corvallis; PAS, Polish Acad¬

emy of Sciences, Warsaw; RNHL, Rijkmuseum van Natuurlijke Historic, Leiden,

The Netherlands; SDAP, State Department of Agriculture, Harrisburg, Pennsyl-

VOLUME 59, NUMBERS 1-4

117

Figs. 3, 4. Thorax of Pherocera rupina female (1725). 3, Lateral view. 4, Dorsal view. Arrows give

linear measurements corresponding to continuous character states in text: 7 = length of thorax exclu¬

sive of scutellum; 8 = width of thorax at wing bases; 9 = length of scutellum; 10 = width of scutellum;

11 = distance between ridge above prothoracic spiracle and basal margin of forecoxa; 12 = length of

mesopleural suture separating katepisternum and katepimeron.

vania; SDCM, San Diego County Museum, California; SJSC, San Jose State

University, California; SMN, Staatlichen Museum furNaturkunde, Stuttgart, West

Germany; SWRS, Southwestern Research Station (AMNH), Portal, Arizona; TAM,

Texas Agricultural and Mechanical University, College Station; UA, University

Fig. 5. Wing of Pherocera rupina male (3185). Letters give landmarks between which linear mea¬

surements correspond to continuous character states in text: g-d (character 13) = length of vein M 3 -f-CuA,;

f-e (character 14) = length of Cu 2 -f2nd A; a-i (character 15) = distance from distal end of R 2+3 to

distal margin of 1st M 2 cell; h-b (character 16) = length of vein R 4 from fork of R 4I5 ; h-c (character

17) = length of vein R 5 from fork of R 4+5 ; b-c (character 18) = distance from distal end of vein R 4

to distal end of vein R 5 ; k-j (character 19) = longest length of 1st M 2 cell; h-i (character 20) = distance

from fork R 4+5 to closest point of 1st M 2 cell.

118

PAN-PACIFIC ENTOMOLOGIST

of Arizona, Tucson; UAC, University of Alberta, Canada; UCB, University of

British Columbia, Vancouver, Canada; UCD, University of California, Davis;

UCM, University of Colorado Museum, Boulder; UCR, University of California,

Riverside; UCVM, Universidad Central de Venezuela, Maracay; UG, University

of Georgia, Athens; UI, University of Idaho, Moscow; UK, University of Kansas,

Lawrence; UM, University of Minnesota, Saint Paul; UMA, University of Mich¬

igan, Ann Arbor; UNLP, Museo de Ciencias Naturales, Universidad Nacional de

La Plata, Argentina; USI, University of Southern Illinois, Carbondale; USNM,

United States National Museum, Washington, D.C.; USSR, Zoological Institute

USSR, Leningrad; USU, Utah State University, Logan; UTA, University of Tex¬

as, Austin; UTI, University of Tel Aviv, Israel; UU, University of Utah, Salt

Lake City; UW, University of Wisconsin, Madison; UZM, Universitetets Zool-

ogiske Museum, Copenhagen, Denmark; VNM, Naturhistorisches Museum, Vi¬

enna, Austria; WSU, Washington State University, Pullman; ZIB, Zoologisches

Institut, Berlin, West Germany; ZSI, Zoological Survey of India Collection, Cal¬

cutta, India.

Characters used—continuous character states.— The following characters were

measured for all female boharti group holotypes herein described (Table 1):

Overall length

Ch. 1. Measurements were taken from the posterior tip of the abdomen to

and including any protuberance on the frons, but not including the

antennae.

Head, measured as illustrated in Fig. 2.

Ch. 2. Head width.

Ch. 3. Distance from center of frons to compound eye margin at dorsal margin

of antennal socket.

Ch. 4. Interocular distance at vertex.

Ch. 5. Width of frontal callosity. The frontal callosity (= dark shiny area

between compound eyes of most females in the genus Pherocera ) is an

extremely important species specific character. Males in the boharti

group lack this callosity.

Ch. 6. Height of frontal callosity. This character was sometimes difficult to

measure on rubbed specimens because it was difficult to determine

where the callosity ended and the ocellar tubercle began.

Thorax, measured as illustrated in Figs. 3 and 4.

Ch. 7. Length of thorax exclusive of scutellum.

Ch. 8. Width of thorax at wing bases.

Ch. 9. Length of scutellum measured from posterior margin of scutellum,

through its center, to posterior margin of mesonotum.

Ch. 10. Width of scutellum, measured to include widest part of scutellum

excluding basal flanges.

Ch. 11. Distance between ridge above prothoracic spiracle and basal margin

of forecoxa.

Ch. 12. The length of the broad ventral section of the mesopleural suture sep¬

arating the katepistemum and katepimeron.

Wing, measured as illustrated in Fig. 5. This illustration gives landmarks, as letters

(a) through (k), and the following characters represent linear measurements

between landmarks:

VOLUME 59, NUMBERS 1-4

119

Ch. 13. g-d Length of vein M 3 +Cu,.

Ch. 14. f-e Length of Cu 2 +2nd A.

Ch. 15. a-i Distance from distal end of R 2+3 to distal margin of 1st M 2 cell.

Ch. 16. h-b Length of R 4 from fork of R 4+5 .

Ch. 17. h-c Length of R 5 from fork of R 4+5 .

Ch. 18. b-c Distance from distal end of R 4 to distal end of R 5 .

Ch. 19. k-j Longest length of 1st M 2 cell.

Ch. 20. h-i Distance from fork R 4+5 to closest point of 1st M 2 cell.

Characters used—discrete character states. — The characters listed below form

a matrix (Table 2) with the included species in the boharti group. It is hoped that

the matrix will prove useful in the separation and recognition of species, aug¬

menting to some degree the dichotomous key.

Ch. 21. Lower gena

a. entirely tomentose (Fig. 14).

b. with a definite bare stripe (Figs. 12, 13, 15-19). In one species no

bare area exists on the lower gena (P. tomentamala, state 21 a), while

a definite bare area, generally confined to a narrow stripe between

the frontobasal compound eye margin and the lower margins of the

oral cavity, is evident on the remaining species of the boharti group

(state 21b). Rubbed specimens sometimes have “bare” areas on the

genae, but these are generally less well defined than the true non-

tomentose stripe of state 21b.

Ch. 22. Cuticular coloration of thorax

a. orange to red

b. dark brown to black

Ch. 23. Cuticular coloration of abdomen

a. orange to red

b. light brown to tan

c. mostly dark brown to black

Ch. 24. Thoracic vittae

a. distinct, dark

b. indistinct, powdered over, but discernible

c. indiscernible

Ch. 25. Cuticular coloration of palp

a. light brown to tan

b. dark brown to black

Ch. 26. Cuticular coloration of hindfemur

a. mostly dark brown to black

b. mostly reddish-orange to tan

Ch. 27. Cuticular coloration of midfemur

a. mostly dark brown to black

b. mostly reddish-orange to tan

Ch. 28. Cuticular coloration of midtibia

a. mostly dark brown to black

b. mostly reddish-orange to tan

Ch. 29. Frontal callosity of female

a. reaching ocellar tubercle

b. not reaching ocellar tubercle

120

PAN-PACIFIC ENTOMOLOGIST

Table 1. Measurements in mm of holotype females of the boharti group (the P.flavipes specimen

is not the holotype).

Species and specimen number

Characters

bishop-

ensis

2804

boharti

1624

boydi

1537

flavipes

1330

nigragena

1578

rufoab-

dominalis

2626

rupina

1725

tomen-

tamala

1670

5.26

4.76

4.31

4.82

5.60

4.65

4.87

5.04

1.36

1.20

1.19

1.23

1.48

1.30

1.40

1.29

0.31

0.24

0.25

0.31

0.36

0.28

0.27

0.28

0.18

0.21

0.18

0.20

0.21

0.31

0.38

0.28

0.42

0.27

0.32

0.31

0.27

0.60

0.62

0.60

0.64

0.38

0.70

0.57

1.36

1.29

1.26

1.50

1.27

1.43

1.33

1.37

1.06

1.20

1.27

1.20

1.37

1.26

0.28

0.27

0.25

0.36

0.28

0.31

0.28

0.52

0.56

0.48

0.50

0.59

0.29

0.59

0.57

0.48

0.41

0.49

0.56

0.46

0.53

0.50

0.15

0.14

0.17

0.18

0.17

0.18

0.20

0.14

0.15

0.20

0.14

0.13

0.24

0.17

0.13

0.18

0.22

0.17

0.18

0.91

0.81

0.83

0.78

0.88

0.78

0.91

0.88

0.92

0.88

0.81

0.95

0.84

0.85

0.97

0.87

0.88

0.87

0.80

0.87

0.80

0.91

0.45

0.42

0.34

0.36

0.42

0.34

0.42

0.43

0.84

0.71

0.84

0.97

0.81

0.85

0.84

0.34

0.28

0.34

0.32

0.39

0.35

Ch. 30. Ventral gonocoxal spur of male terminalia

a. one-appendaged, molar-shaped or with minute teeth (Figs. 46-51)

b. one-appendaged, elongate molar-shaped, with rounded ventrolat¬

eral lobe (Fig. 45)

c. two-appendaged, one tapered, the other elongate, spine like (Fig.

44)

Ch. 31. Adults mainly occur in

a. desert canyon bottom environments

b. montane environments

c. dry wash environments

d. inland stable sand dune environments

Ch. 32. Principal landing substrate of adults is

a. rock

b. twig on ground

c. soil-leaf litter

Definition of special morphological terms. — Most terms used in this paper can

be found in Torre-Bueno (1950), Lyneborg (1968, 1972), Irwin (1973, 1976, 1977),

Irwin and Lyneborg (198 la, 1981b), and McAlpine et al. (1981); those that cannot

or are used in a different connotation or otherwise need clarification follow:

Frontal callosity: The area of the female frons usually devoid of tomentum.

Genal stripe: The stripe, devoid of tomentum, on the lower gena connecting

VOLUME 59, NUMBERS 1-4

121

Table 2. Discrete states of characters 21-32 explained in text for species of the boharti group.

Characters

Species

bishop¬

ensis

boharti

boydi

flavipes

nigragena

nifoab-

dominalis

rupina

tomen¬

tamala

a&b

the lower margin of the compound eye with the lower margin of the oral

cavity.

Ventral gonocoxal spur: An extension of the gonocoxite (Fig. 10, vgs), possibly

homologous with the ventral lobe of the gonocoxite of Lyneborg (1968) and

Irwin and Lyneborg (1981a).

Pile: Long, thin hair without a noticeable socket.

Tomentum: Short, scalelike hair densely covering exoskeleton; sometimes re¬

ferred to by other authors as pollen.

The boharti Group of Pherocera

Diagnosis. — Body usually dark brown to black ( rufoabdominalis and bishopensis

with red abdomens, boharti with reddish-orange thorax, tan abdomen); tomentum

usually silver to silver-gray, dense on face and thorax, sparse on abdomen.

Head: Tomentum dense, silver-gray on upper frons and vertex and silver on

lower frons and gena; facial pile white; palp white pilose; female dichoptic, facets

of uniform size; male holoptic, facets divided in most species, or facet size grading

from smaller ventrad to larger dorsad ( boycii, tomentamala)\ antennal flagellum

pear-shaped to elongate pear-shaped (Figs. 20-27); maxillary palp with distinct

terminal indentation, one-jointed (Figs. 28-35).

Thorax: Mesonotal vittae distinct {nigrigena, flavipes), indistinct but discernible

{bishopensis, boydi, flavipes, rufoabdominalis, rupina, tomentamala), or indiscer¬

nible {boharti)-, vein M 3 +CuA, reaching wing margin (Fig. 5, g-d). Halter knob

white; proximal % to 5 / 6 of foretibia with dense, white pile.

Abdomen: Male terminalia (Figs. 8, 9) with hypoproct fitted partially beneath

epandrium, variously shaped (Figs. 36-43); ventral gonocoxal spurs (vgs), gonostyli

(g) (Fig. 10) variously shaped (Figs. 44-51), and aedeagus (Fig. 11) complex (Figs.

52-59). Female terminalia (Figs. 6, 7) simple; lurca with two membranous areas,

through the anterior of which passes the coalesced spermathecal duct (Fig. 7).

Relationships.— The boharti group has a sister-group relationship with the al-

bihalteralis group, for which P. albihalteralis was formally described (Cole, 1923a),

but can be distinguished from it by the ventral gonocoxal spur being one-appen-

122

PAN-PACIFIC ENTOMOLOGIST

Figs. 6, 7. Female terminalia of Pherocera rufoabdominalis (2506). 6, Lateral view. 7, Dorsal view

with tergite 8 removed. Scale = 0.2 mm.

daged, one molar-shaped or rounded and with minute teeth (Figs. 46-51), one-

appendaged, elongate molar-shaped with a lateroventral rounded lobe (Fig. 45),

or two-appendaged, one molar-shaped, the other spinelike (Fig. 44), while mem¬

bers of the albihalteralis group have spurs two-appendaged, sharp, spinelilce, equal

or unequal in length or one-appendaged, sharp, elongate or fanglike and elongate.

VOLUME 59, NUMBERS 1-4

123

Figs. 8, 9. Male terminalia of Pherocera rufoabdominalis (2432). 8, Lateral view. 9, Dorsal view.

Scale = 0.2 mm.

The bo hart i group is also characterized by having a bare stripe on each lower gena

between the frontobasal compound eye margins and the lower margins of the oral

cavity (except P. tomentamala).

Within the bo hart i group the species may be further grouped on the basis of

their ventral gonocoxal spurs and gonostyli. Two related species, bishopensis and

124

PAN-PACIFIC ENTOMOLOGIST

Figs. 10, 11. Male genitalia of Pherocera rufoabdominalis (2511). 10, Dorsal view with epandrium

removed. 11, Aedeagus, dorsolateral view. Scale = 0.1 mm.

rufoabdominalis, are distinct in having hatchet-shaped gonostyli and cuspidlike

ventral gonocoxal spurs. P. boydi is distinct in having spinelike and exaggeratedly

lobed ventral gonocoxal spurs. P. boharti is characterized by an elongate, cuspid¬

shaped ventral gonocoxal spur and a gonostylus with an elongate lateral spine. P.

rupina, tomentamala, nigragena, and flavipes appear closely related, having elon¬

gate, sometimes double-tipped, gonostyli and single ventral gonocoxal spurs that

are molar-shaped in appearance.

Included species.—P. bishopensis Irwin, sp. nov.; P. boharti Irwin, sp. nov.; P.

boydi Irwin, sp. nov.; P. flavipes Cole, 1923a; P. nigragena Irwin, sp. nov.; P.

rufoabdominalis Irwin, sp. nov.; P. rupina Irwin, sp. nov.; and P. tomentamala

Irwin, sp. nov.

VOLUME 59, NUMBERS 1-4

125

Key to the Species of the Boharti Group

1. Palps dark brown to black . 2

- Palps yellow to light brown. 5

2. Hindfemur dark brown to black; ventral gonocoxal spur and gonostylus

(<$) as in Fig. 49; frontal callosity (2) as in Fig. 16 .. nigragena Irwin, sp. nov.

- Hindfemur mostly orange to tan . 3

3. Lower gena without a bare stripe; ventral gonocoxal spur and gonostylus

(6) as in Fig. 46; frontal callosity (2) as in Fig. 14.

. tomentamala Irwin, sp. nov.

- Lower gena with bare stripe . 4

4. Ventral gonocoxal spur (<5) bilobed, one elongate, spinelike (Fig. 44); fron¬

tal callosity (2) as in Fig. 12. Mesonotal vittae indistinct, powdered over

. boydi Irwin, sp. nov.

- Ventral gonocoxal spur (<$) molar-shaped (Fig. 48); frontal callosity (2) as

in Fig. 17. Mesonotal vittae usually distinct. flavipes Cole

5. Abdomen red; thorax dark brown to black . 6

- Abdomen tan or brown; thorax dark brown to tannish-orange . 7

6. Frontal callosity (2) with central tomentose zone (Fig. 19); ventral gon¬

ocoxal spur and gonostylus (<$) as in Fig. 51 . bishopensis Irwin, sp. nov.

- Frontal callosity (2) without tomentose island (Fig. 18); ventral gonocoxal

spur and gonostylus (3) as in Fig. 50. rufoabdominalis Irwin, sp. nov.

7. Thorax orangish-tan; frontal callosity (2) as in Fig. 13; ventral gonocoxal

spur and gonostylus (3) as in Fig. 45. boharti Irwin, sp. nov.

- Thorax black or dark brown; frontal callosity (2) as in Fig. 15; ventral

gonocoxal spur and gonostylus (<$) as in Fig. 47 . rupina Irwin, sp. nov.

Pherocera bishopensis Irwin, Sp. Nov.

(Figs. 19, 26, 34, 43, 51, 58)

Derivation of name.— Bishop = type locality; -ensis (Latin) = of or from.

Diagnosis.—P. bishopensis can be distinguished from all other species of the

boharti group by the following combination of characters: mesonotal vittae in¬

distinct, abdominal coloration red, frontal callosity of female not reaching ocellar

tubercle and having a tomentose area within the bounds of the frontal callosity

(Fig. 19); male terminalia with hypoproct as in Fig. 43 and ventral gonocoxal

spur and gonostylus as in Fig. 51.

Description, female holotype.—See Table 1 for holotype measurements, Table

2 for discrete character states.

Head: Dark brown. Face and frontal callosity as in Fig. 19. Tomentum silver-

gray on vertex and upper frons, brownish above frontal callosity, silver on lower

frons and gena. Pile lacking on ocellar tubercle; sparse, short, white dorsolaterad

of antennal socket; sparse, longer on lower gena and palp. Ratio of antennal scape :

pedicel: flagellum, 15:9:32; scape reddish-tan with two black setae ventrad, red¬

dish-tan pile sparsely over entire segment: pedicel dark brown with short, white

pile forming ring distally; flagellum (Fig. 26) dark brown, bare. Palp (Fig. 34) light

brown, nearly translucent. Proboscis dark brown.

Thorax: Dark brown; mesonotal vittae indistinct but discernible; tomentum

gray-brown, moderately dense on mesonotum, very dense on scutellum; meso-

126

PAN-PACIFIC ENTOMOLOGIST

16 nigragena 17 flavipes 18 rut oabdominalis 19 bishopensis

Figs. 12-19. Heads of females of Pherocera boharti group, frontal views. 12, ( 2794)] 13, ( 2800)]

14, (1670)] 15, (2808)] 16, (2796)] 17, (2831)] 18, (2806)] 19, (2804). Scale = 0.5 mm.

notal pile very short, light colored, sparse, present only posteriorly; pile short,

sparse on scutellum; pleural region sparsely silver tomentose, even sparser on

anepimeron and katatergite; coxae mostly red with proximal Vi of hindcoxa and

most of forecoxa dark brown; legs mostly red with forefemur, distal l / 6 of foretibia,

and all tarsi dark brown; pile on legs sparse, white.

Abdomen: Cuticular coloration red; tomentum almost nonexistent, gray, sparse

dorsally and centrally; pile short, evenly covering abdomen, white anteriorly and

ventrally to black posteriorly; tergites I-IV fasciate.

Male. — Same as female holotype except as follows: pile short, black on ocellar

tubercle; frons silver tomentose, pile on mesonotum white, short anteriorly to

long posteriorly, even longer on scutellum; scape dark brown; coxae dark brown;

distal sixth of middle and hindtibiae dark brown; distal third of foretibia dark

brown; abdomen with a single mid-dorsal dark brown blotch on each of tergites

I-IV; tergites I—III, faintly IV, fasciate; aedeagus as in Fig. 58; hypoproct as in

Fig. 43; ventral gonocoxal spur and gonostylus as in Fig. 51.

Variation. — Overall length 3.6 mm to 5.1 mm <3; 4.1 mm to 6.3 mm 2; tergites

I—II to I-IV fasciate; one specimen has a more intensely yellow palp and more

intensely orange abdomen than other examined specimens; some have dark blotches

on dorsal midline of tergites.

Distribution and ecological placement.—P. bishopensis appears to be restricted

to dry washes and desert canyon bottoms where Populus spp. abound in the Owens

Valley (Inyo and Mono Counties) and the Mojave Desert (Los Angeles and Kern

Counties), California.

Specimens examined. — Holotype: 2, Irwin specimen number 2804, CAS type

no. 10454; 4.8 kilometers south of Bishop (label reads Bishop), Inyo Co., Cali¬

fornia; EIS; July 20, 1960; under Populus trichocarpa Torrey & Gray. Paratopo-

VOLUME 59, NUMBERS 1-4

127

boharti

boydi

tomentamala

rupina

f lavipes

nigragena

bishopensis ruf oabdominalis

Figs. 20-27. Antennal flagella of males of Pherocera boharti group, lateral view. 20, (7655); 21,

(7525); 22, (7669); 23, (7507); 24, (7792); 25, (7594); 26, (2774); 27, (2453). Scale = 0.1 mm.

types: 85 <3; 64 2. Other specimens: 1 6; 2 2. The following specimens were ex¬

amined:

CALIFORNIA: Inyo Co.: Bishop, 4.8 km S, 1265 m, 60/07/20, EIS, 85 6, 65

9; Kern Co.: Walker Pass, 1585 m, 58/08/19, RHP EMP, 1 <3; Los Angeles Co.:

Lancaster, 12.9 km E, 56/09/06, EIS, 1 9; Mono Co.: Benton Station, 50/07/20,

HAH, 1 2.

Commerce—Morphological separation of some males of P. bishopensis and P.

ruf oabdominalis is difficult. Female frontal callosity, ecological associations, and

distribution patterns all help separate this species from P. rufoabdominalis, its

sister species.

Pherocera boharti Irwin, Sp. Nov.

(Figs. 13, 20, 28, 37, 45, 52)

Derivation of name .—Named in honor of Dr. Richard M. Bohart.

Diagnosis. — This species can be distinguished from all others in the boharti

group by the following combination of characters: frontal callosity (2) as in Fig.

13; thorax orange; mesonotal vittae indistinguishable; male terminalia with hy-

poproct as in Fig. 37 and ventral gonocoxal spur and gonostylus as in Fig. 45.

Description, female holotype. — See Table 1 for holotype measurements, Table

2 for discrete character states.

Head: Face and frons similar to Fig. 13, slightly less tomentum on frons. Light

brown from midfrons down, dark brown above. Tomentum dense, silver through¬

out; pile lacking on ocellar tubercle; white, moderately long dorsolaterad of an¬

tennal socket; sparse, moderately long, white posterior of genal stripe. Ratio of

antennal scape: pedicel: flagellum, 17:7:34; scape with one short, black seta ven-

trad, white pilose; pedicel with distal ring of white and black pile; flagellum (Fig.

20) bare; antennae dark brown; palp (Fig. 28) light brown; proboscis dark brown.

Thorax: Orange; mesonotal vittae indiscernible; tomentum extremely sparse,

gray, not obliterating ground color; pile black, sparse over mesonotum and scu-

tellum; pleural region sparsely silver tomentose on anepisternum, katepisternum,

katepimeron and metanepisternum; lacking on anepimeron and katatergite; an¬

episternum sparsely short, white tomentose; katatergite with sparse, tan and elon¬

gate tomentum; coxae orangish-tan, silver tomentose, white pilose; legs orange

128

PAN-PACIFIC ENTOMOLOGIST

with proximal V 2 of hindtibia light brown and distal V 2 dark brown, forecoxa and

femur brown, distal V 2 of midtibia brown, proximal % of foretibia white, distal x h>

dark brown; all tarsi dark brown; wing veins brown.

Abdomen: Tan, darker brown dorsally; tomentum absent; pile short, black,

appressed mesally; tergites I—II, faintly IV, fasciate.

Male. — Same as female holotype except as follows: antennal scape with two

black setae ventrad; pile short, black on ocellar tubercle; tomentum white on

anterolateral margins of mesonotum; tomentum golden on scutellum; hindcoxa

and trochanter dark brown; midtrochanter dark brown; abdomen dark brown;

abdominal pile long, white, erect laterally, shorter, black, appressed ventrally and

dorsally; aedeagus as in Fig. 52; hypoproct as in Fig. 37; ventral gonocoxal spur

and gonostylus as in Fig. 45.

Variation. —Overall length 3.6 mm to 5.0 mm <3; 4.2 mm to 5.2 mm 2; one

female with black pile on ocellar tubercle; abdomen of female sometimes darker.

Distribution. —P. boh art i has been collected from the transverse mountain range

including the San Gabriel Mountains in Los Angeles County and the San Ber¬

nardino Mountains in San Bernardino County, California, in association with

oak-woodland leaf litter. A small series was collected by RMB near Warner Springs,

San Diego County, California. It was subsequently learned (R. M. Bohart, pers.

comm.) that the collector had just recently come from the San Gabriel Moun¬

tains and had collected the specimens from inside the car. It is possible that these

specimens were transported by car from the Tanbark Flat area to Warner Springs.

Specimens examined.— Holotype: 2, Irwin specimen number 1624, CAS type

no. 10455; Wildwood Canyon, 8 km east of Calimesa, San Bernardino Co., Cal¬

ifornia, MEI; July 12, 1965; on wet sand near watering trough. Paratopotypes: 1

<3; 23 2. Other paratypes: 4 <3; 5 2. Other specimens: 6 2. The following specimens

were examined:

CALIFORNIA: Los Angeles Co.: Big Dalton Canyon, 52/07/23, ATM, 1 <3, 1

2; Glendale, 54/08/25, EIS, 1 <3; 56/07/29, 1 2; San Francisquito Canyon, 56/07/

15, LAS, 2 2; Tanbark Flat, 56/07/14, RMB, 1 <3, 1 2. San Bernardino Co.:

Calimesa, 8 km E at Wildwood Canyon, 57/07/09, HRM, 1 2; 57/07/13, JCH, 1

<3; HRM, 2 2; 58/07/07, HRM, 2 2; 65/07/10, MEI, 2 2; 65/07/12, MEI, 13 2; 68/

07/24, MEI, 4 2; Camp Baldy, 4.8 km W, 65/09/09, MEI, 1 <3; Mountain Home,

44/08/24, ALM, 2 2. San Diego Co.: Warner Springs, 14.5 km S (Sic! see text),

56/07/08, RMB, 4 2.

Pherocera boydi Irwin, Sp. Nov.

(Figs. 12, 21, 29, 36, 44, 53)

Derivation of name. —Named in honor of Mr. Philip L. Boyd who donated land

that now forms the Deep Canyon Desert Research Center of the University of

California, Riverside campus.

Diagnosis.— P. boydi can be distinguished from all other members of the boharti

group by the following combination of characters: frontal callosity (2) as in Fig.

12; mesonotum with indistinct vittae, heavily powdered over with silver to silver-

gray tomentum; male terminalia with large hypoproct (Fig. 36) and unique-shaped

ventral gonocoxal spur (Fig. 44).

Description, female holotype. —See Table 1 for holotype measurements, Table

2 for discrete character states. Body medium brown.

VOLUME 59, NUMBERS 1-4

129

Head: Face similar to Fig. 12, frontal callosity fuller, rounder, extending through

ocellar tubercle; tomentum gray near vertex, silver-gray on upper frons, silver on

lower frons and gena; pile absent on ocellar tubercle, short, white dorsolaterad of

antennal socket, absent on gena except short, white posterior to genal stripe; ratio

of antennal scape: pedicel: flagellum, 15:7:35; scape with two long black setae

ventrad, light brown pile overall; pedicel with distal ring of light tan pile; flagellum

similar to Fig. 21, bare; antenna dark brown, brown tomentose; palp (Fig. 29)

and proboscis dark brown.

Thorax: Mesonotal vittae indistinct but discernible, densely powdered over

with silver-gray tomentum; tomentum covering scutellum; pile very short, sparse

over mesonotum and scutellum; pleural region densely silver-gray tomentose on

anepisternum, katepisternum, katepimeron, and metanepisternum, lacking on

anepimeron and katatergite; anepisternum and katatergite with sparse, long, white

pile; coxae reddish brown, silver-gray tomentose, white pilose; middle and hind¬

legs mostly orange or tannish-orange; foreleg black except proximal % of tibia

whitish.

Abdomen: Tomentum lacking; pile short, white (some black), more or less

erect, covering entire abdomen; tergites I-IV fasciate.

Male. — Same as female holotype except as follows: light tan pile on ocellar

tubercle; pile posterior to genal stripe longer and whiter; mesonotum more densely

silver tomentose, short, white pilose; white pile on scutellum longer; white pile

on abdomen much longer on tergites; tergites I—II, III faintly fasciate; aedeagus

as in Fig. 53; hypoproct as in Fig. 36; ventral gonocoxal spur and gonostylus as

in Fig. 44.

Variation.— Overall length 3.4 mm to 4.5 mm <3; 4.0 mm to 5.4 mm $; female

frontal callosity variable, more bulbous on lower frons, narrowing on upper frons;

coxae and trochanter of male often black.

Distribution and ecological placement. — P. boydi seems restricted to desert can¬

yon bottom environments, often in association with rock substrates under desert

willow, Chilopsis linearis (Cavanilles). It has been collected from the San Jacinto,

Santa Rosa, and Little San Bernardino Mountains, Riverside County; from the

Daggett Hills, San Bernardino County; from the eastern foothills of the White

Mountains, Inyo County, California; from Gila County, Arizona; from Hidalgo

County, New Mexico; and from the eastern slope of the Sierra Juarez, Baja Cal¬

ifornia Norte, Mexico.

Specimens examined. — Holotype: 2, Irwin specimen number 1537, CAS type

no. 10456; P. L. Boyd Deep Canyon Desert Research Center, 6.4 km south of

Palm Desert, Riverside Co., California; EIS; July 1, 1964; under Chilopsis in

lower part of canyon. Paratopotypes: 18 6; 17 2. Other specimens: 8 <3; 8 2. The

following specimens were examined:

MEXICO: Baja California Norte: Guadalupe Canyon, 365 m, 67/07/11, MEI,

1 6 .

USA-ARIZONA: Gila Co.: San Carlos, 18/05/12, JCB, 1 2. Maricopa Co.:

Canyon Lake, 58/05/30, JCH, 1 <3. CALIFORNIA: Inyo Co.: Antelope Springs,

60/06/19, HKC, 1 6, 6 2. Westgard Pass, 6.4 km W, 61/06/19, JAP, 1 Riverside

Co.: Massacre Canyon, 64/07/29, MEI, 1 <3; P. L. Boyd Deep Canyon Desert

Research Center, 6.4 km S of Palm Desert, 320 m, 63/05/15, EIS, 1 <3; 63/06/21,

EIS, 2 <3, 1 2; 64/05/19, MEI, 1 2; 64/06/24, EIS, 2 <3, 2 2; 64/07/01, GRB, 1 6;

130

PAN-PACIFIC ENTOMOLOGIST

28 boharti

Figs. 28-35. Palps of males of Pherocera boharti group, lateral view. 28, ( 1633); 29, (1523); 30,

(1669); 31, (1807); 32, (1409); 33, (1594); 34, (2714); 35, (2453). Scale = 0.1 mm.

EIS, 1 2; 65/06/08, MEI, 1 67/05/19, MEI, 1 6; 69/05/18, MEI, 1 2; 69/05/21,

MEI, 1 2; 69/05/24, MEI SIF, 4 3, 4 2; 69/06/15, SIF RMW, 3 2; 69/06/23, SIF

RMW, 1 3; 69/06/28, SIF RMW, 1 3; 69/06/29, SIF RMW, 1 2; 69/07/03, SIF

RMW, 2 2; 69/07/05, SIF RMW, 1 3; 65/06/08, MEI, 1 3; 373 m, 65/06/08, MEI,

1 2; 381 m, 69/06/29, SIF, 1 3; Whitewater Canyon, 63/07/26, EIS, 2 3. San

Bernardino Co.: Daggett, 55/05/22, WRM, 1 2. NEW MEXICO: Hidalgo Co.:

Rodeo, 8 km N, 1380 m, 65/06/01, MEI, 1 3.

Pherocera flavipes Cole

(Figs. 17, 24, 32, 41, 48, 56)

Pherocera flavipes Cole, 1923a, Proc. U.S. Natl. Mus. 62(4):22.

Derivation of name.—flavus (Latin) = yellow; pes (Latin) = foot.

Diagnosis. —P. flavipes can be distinguished from all other species in the boharti

group by the following combination of characters: female frontal callosity occu¬

pying nearly all of frons (Fig. 17); cuticular coloration usually black; mesonotal

vittae usually distinct; palp dark brown; middle and hindfemora and tibiae usually

orange; antennal flagellum as in Fig. 24; palp as in Fig. 32; male terminalia with

aedeagus as in Fig. 56, ventral gonocoxal spur and gonostylus as in Fig. 48, and

hypoproct as in Fig. 41.

Description, female. — (Comments on holotype): This species was adequately

described by Cole (1923a, pp. 22-23) except as follows: length 5.3 mm; both

antennal styli are entire; coxae partly orange, partly brown.

VOLUME 59, NUMBERS 1-4

131

Male. — Similar to female holotype (see Cole, 1923a) except as follows: pile

short, black on ocellar tubercle, white dorsolaterad of antennal socket, longer,

white posterior to genal stripe; ratio of antennal scape : pedicel: flagellum, 18:3:

39; scape with four black setae ventrad and white pile; pedicel with distal ring of

white pile; flagellum bare; palp and proboscis dark brown; mesonotal vittae dis¬

tinct, black, silver tomentose; mesonotal pile long, white; tarsi dark brown; ab¬

domen black, white pilose, not tomentose; aedeagus as in Fig. 56; hypoproct as

in Fig. 41; ventral gonocoxal spur and gonostylus as in Fig. 48.

Variation.— Overall length 3.7 mm to 5.3 mm <3; 4.1 mm to 5.9 mm 2; the

Wildwood Canyon, San Bernardino County, California population has the me¬

sonotal vittae discernible but not distinct. The shape of the antennal flagellum

varies from pear-shaped to elongate pear-shaped. P. flavipes is a variable species

with rather indistinct and usually inseparable populations.

Distribution and ecological placement.— This species inhabits montane and des¬

ert canyon bottom environments where adults are often found on rock substrates.

It has been collected from Arizona, California, Idaho, Nevada, New Mexico, and

Utah in the United States; Baja California Norte and Baja California Sur in

Mexico. P. flavipes has the widest distribution of all species of Pherocera.

Specimens examined. — Holotype: 2, USNM: Higley (not Highley as reported

by Cole, 1923a), Superstition Mountain, Pinal Co., Arizona; EGH; June 13, 1917.

Other specimens: 132 <3; 227 2. The following specimens were examined:

MEXICO: Baja California Norte: La Rumorosa, 14.5 km S, 1370 m, 67/07/

08, MEI, 1 2 . Baja California Sur: Isla Cerralvo, Gordas Point, 53/03/20, PHA,

1 2. Guadalupe, 100 km NW of La Paz, 107 m, 68/04/20, MEI, 6 3, 1 2. Loreto,

14.5 km S, 18 m, 68/04/17, MEI, 1 2. San Ignacio, 4.8 km E, 170 m, 68/04/14,

MEI, 4 <3, 2 2. Santa Rosalia, 40 km S, 61 m, 68/04/15, MEI, 1 3, 1 2.

USA-ARIZONA: Pima Co.: Alamo Canyon, Oregon Pipe National Monument,

47/04/16, ALM, 6 2. CALIFORNIA: Inyo Co.: Antelope Springs, 61/06/14, CAT,

1 6; 61/07/01, JAP, 1 5; 66/07/09, HKC, 1 2; Batchelder Spring, 67/07/21, MEI,

1 (3, 7 2; Mountain Springs Canyon, Argus Mountains, 59/06/27, JCD, 4 3, 6 2;

WCF, 1 <3. Lassen Co.: Hallelujah Junction, 49/07/04, JWM, 1 2; 54/07/12, RMB,

1 2. Los Angeles Co.: Crystal Lake, 56/06/28, RCB, 1 2. Napa Co.: Monticello

Dam, 62/09/15, MEI, 1 <3. Riverside Co.: Massacre Canyon, 64/07/29, MEI, 1 2.

Millard Canyon, 63/06/20, EIS, 1 2. P. L. Boyd Deep Canyon Desert Research

Center, 6.4 km S of Palm Desert, 63/04/13, WAS, 1 3; 64/05/19, MEI, 2 3; 69/

06/07, SIF, 1 3; 69/06/13, SIF, 1 3; 320 m, 64/05/19, MEI, 1 3; 64/05/31, MEI,

1 2; 65/05/24, MEI SIF, 2 3, 10 2; 65/06/08, MEI, 1 3; 69/05/16, MEI, 3 3; 69/

05/17, JRQ, 1 3; 69/05/18, MEI, 13 3, 4 2; 69/06/10, SIF, 1 2; 69/06/15, SIF

RMW, 1 3; 343 m, 64/05/19, MEI, 6 3, 2 2; 64/05/31, MEI, 1 2; 65/04/28, MEI,

1 3; 67/05/02, MEI, 5 2; 350 m, 65/06/08, MEI, 1 3; 366 m, 67/05/19, MEI, 1 3,

4 2; 373 m, 65/04/28, EIS, 1 3; 65/06/08, MEI, 1 3; 67/05/02, MEI, 1 2; 381 m,

63/05/03, EIS, 3 3; 63/05/17, EIS, 1 3; 63/06/21, EIS, 2 3; 65/06/08, MEI, 1 3;

67/05/19, MEI, 1 2; 400 m, 63/05/03, EIS, 6 2; 63/05/30, EIS, 1 2; 63/06/21, EIS,

2 2; 64/05/19, MEI, 5 3, 1 2; 64/05/31, MEI, 4 3, 1 2; 65/04/28, EIS, 1 3; MEI,

3 3, 3 2; 67/05/02, MEI, 1 3; 420 m, 65/04/28, EIS, 1 3; 67/05/19, MEI, 1 3; 1036

m, 65/06/1 1, MEI, 3 3, 8 2; EIS, 1 2; Palm Springs, Andreas Canyon, 55/05/03,

WRM, 1 2; Pinon Flat, 48/05/26, AFH, 1 3; Thousand Palms Canyon, Willis

Oasis, 55/04/12, WRM, 1 3; Whitewater Canyon, 63/06/07, EIS, 1 2; Winchester,

67/06/20, WI, 1 2; 67/06/23, WI, 2 2; 68/06/20, WI, 1 3, 1 2. San Bernardino

132

PAN-PACIFIC ENTOMOLOGIST

Figs. 36-43. Hypoprocts of male genitalia of Pherocera boharti group, dorsal view. 36, (7579); 37,

(1635)- 38, (7669); 39, (7757); 40, (7594); 41, (7792); 42, (5209); 43, (2639). Scale = 0.1 mm.

Co.: Calimesa, 8 km E at Wildwood Canyon, 57/07/09, HRM, 2 2; 57/07/13,

JCH, 4 5, 1 2; HRM, 2 2; 65/07/10, MEI, 4 3, 59 2; RES, 10 2; 65/07/12, MEI,

17 6; 65/09/12, MEI, 1 67/06/04, MEI PAR, 1 2; 68/07/24, MEI, 4 2; Camp

Baldy, 4.8 km W, 65/09/09, MEI, 1 <3; Daggett, 55/05/22, WRM, 4 6, 6 2; He-

lendale, 55/05/21, WRM, 1 2; San Diego Co.: Borrego Springs, 45/05/03, ALM,

1 2; 55/04/27, PDH, 1 2; 57/04/18, EIS, 2 3, 3 2; 57/04/19, JCH, 2 1 2; RMB,

5 2; 60/04/21, JFL, 1 2; Culp Canyon, Anza-Borrego State Park, 58/06/12, EIS,

3 5, 8 2; JCH, 10 2; HRM, 5 2. Ventura Co.: Dry Canyon, 14.5 km E, 57/07/26,

RAS, 1 2. IDAHO: Twin Falls Co.: Buhl, 32/07/08, DEF, 1 <3. NEVADA: Clark

Co.: Charleston Mountains at Willow Creek, 54/07/01, EIS, 2 3, 2 2; JWM, 1 <3,

1 2. NEW MEXICO: Hidalgo Co.: Rodeo, 24 km N, 1380 m, 65/06/01, MEI, 1

<3, 1 2; Rodeo, 8 km N, 1380 m, 65/06/01, MEI, 1 2. UTAH: Grand Co.: Arch

Canyon, 35/06/23, CTB, 1 2; Browns Park, W of, 46/06/23, MTJ, 1 3; Castle

Valley, 63/06/09, WJH, 1 3.

Pherocera nigragena Irwin, Sp. Nov.

(Figs. 16, 25, 33, 40, 49, 57)

Derivation of name. — nigra (Latin) = black; gena (Latin) = cheek.

Diagnosis.—P. nigragena can be distinguished from all other species of the

boharti group by the following combination of characters: female frontal callosity

as in Fig. 16; mesonotum with vivid and distinct vittae; cuticular coloration of

both mid and hindfemora mostly black or dark brown; midtibia mostly orange;

cuticular coloration of palp dark brown; male terminalia with hypoproct as in

Fig. 40 and ventral gonocoxal spur and gonostylus as in Fig. 49.

Description, female holotype. — See Table 1 for holotype measurements, Table

2 for discrete character states. Body black, silver tomentose.

VOLUME 59, NUMBERS 1-4

133

Head: Face and frons similar to Fig. 16; frontal callosity narrow, tapering

gradually from base to anterior ocellus. Pile absent on ocellar tubercle; sparse,

white dorsolaterad of antennal socket; short, white posterior to genal stripe; white,

longer, on palp. Ratio of antennal scape : pedicel: flagellum, 20:9:38; scape with

three large, black setae lateroventrad, white pilose; pedicel with distal ring of short

black pile; flagellum similar to Fig. 25; bare, antenna black; palp (Fig. 33) and

proboscis dark brown.

Thorax: Mesonotal vittae distinct; mesonotal pile short, black; pleural region

tomentose except for katatergite and anepimeron; anepisternum and katatergite

densely long, white pilose; coxae tomentose, white pilose; legs black with distal

l / 4 of femora and proximal V 3 of tibiae of mid and hindlegs orange; distal V 10 of

femur and proximal V 10 of tibia of foreleg orange; remainder of foretibia white

and densely white pilose; legs predominantly black pilose.

Abdomen: Tomentum absent; tergite 1 with long, white pile; rest of abdomen

short, black pilose; tergites I—III fasciate.

Male. — Same as female holotype except as follows: abdominal pile long, white

laterad; shorter, dark, dorsad and ventrad. Aedeagus as in Fig. 57; hypoproct as

in Fig. 40; ventral gonocoxal spur and gonostylus as in Fig. 49.

Variation. — Overall length 4.2 mm to 5,5 mm 5; 4.2 mm to 6.2 mm 2; legs of

some specimens with slightly more orange to almost no orange.

Distribution and ecological placement. —P. nigragena was collected in dry mon¬

tane environments characterized by cold winters and cool summers; most of the

specimens collected by the author were associated with rock substrates under

willow trees ( Salix spp.). It has been collected in the more northern areas of

California (Inyo, Mono, and Modoc Counties; with a single specimen from Vic¬

torville, Los Angeles County) and Nevada (Mineral and Washoe Counties).

Specimens examined. — Holotype: 2, Irwin specimen number 1578, CAS type

no. 10470; Batchelder Spring, Inyo Co., California; MEI; July 21, 1967; under

Salix on rock. Paratopotypes: 3 2. Other paratypes: 3 5, 18 2. Other specimens:

10 5, 10 2. The following specimens were examined:

CALIFORNIA: Inyo Co.: Antelope Springs, 1675 m, 60/06/19, HKC, 4 5 , 14

2; 60/06/20, HKC, 1 5 , 2 2; 65/07/02, MEI, 3 5 , 4 2; 67/07/21, MEI, 1 5 ; Batchelder

Spring, 67/07/10, RMB, 1 2; 67/07/21, MEI, 3 2; Big Pine, 57/07/06, JW, 1 5 ;

Lone Pine, 1.6 km N, 64/06/12, MEI, 1 2; Westgard Pass, 9.7 km E, 65/07/01,

MEI, 1 2. Modoc Co.: Canby, NW of, 60/06/24, EIS, 1 2. Mono Co.: Topaz Lake,

57/06/26, JWM, 2 5 . San Bernardino Co.: Victorville, 44/05/30, ALM, 1 5 . NE¬

VADA: Mineral Co.: Basalt, 6.4 km S, 65/07/01, MEI, 5 2. Washoe Co.: Patrick,

64/06/30, FDP, 1 5 .

Pherocera rufoabdominalis Irwin, Sp. Nov.

(Figs. 6-11, 18, 27, 35, 42, 50, 59)

Derivation of name. — rufus (Latin) = red or reddish; abdomen (Latin) = belly.

Diagnosis. — P. rufoabdominalis can be distinguished from all other species of

the boharti group by the following combination of characters: abdomen red;

mesonotal vittae indistinct; female frontal callosity as in Fig. 18, not reaching

ocellar tubercle and being entirely bare (tomentose area only within callosity of

P. bishopensis females); male terminalia with hypoproct as in Fig. 42 and ventral

gonocoxal spur as in Fig. 50.

134

PAN-PACIFIC ENTOMOLOGIST

46 47

tomentamala rupina

Figs. 44-51. Ventral gonocoxal spurs on outer lateroventral portions of gonocoxites (light shading)

and gonostyli (dark shading) of male genitalia of Pherocera boharti group, ventrolateral view. 44,

(1519); 45, (1635); 46, (1669); 47, (1787); 48, (1192); 49, (1594); 50, (3209); 51, (2639). Scale = 0.1

mm.

Description, female holotype. —See Table 1 for holotype measurements, Table

2 for discrete character states. Also see diagnosis of boharti group.

Head: Black; face and frons similar to Fig. 18; tomentum silver-gray on vertex

and upper frons, silver on central and lower frons and gena. Pile lacking on ocellar

tubercle; sparse, short, white dorsolaterad of antennal socket and lower gena,

longer on palp. Ratio of antennal scape : pedicel: flagellum, 14:7:34; scape with

two black setae ventrad; sparse, white pile over entire surface; pedicel with short,

white pile forming distal ring; flagellum similar to Fig. 27, bare; entire antenna

dark brown with whitish to light brown tomentum; palp (Fig. 35) light brown,

translucent; proboscis dark brown.

Thorax: Black; mesonotal vittae indistinct but easily discernible; tomentum

gray to gray-brown, denser on scutellum than mesonotum; mesonotal pile not

evident on anterior portion, very sparse, white posteriorly, sparse on scutellum;

pleural region densely silver tomentose except for lcatatergite and anepimeron;

fine white pile on anepisternum and katatergite; coxae dark brown, silver tomen¬

tose and white pilose except for red cuticular coloration on distal % of middle

and distal Va of hindcoxae; legs red with femur, distal Va of tibia (proximal 3 A of

tibia white), tarsus of foreleg, tarsus of midleg, and distal V 6 of tibia and tarsus

of hindleg dark brown. Pile on legs white; wing veins brown.

Abdomen: Red; tomentum gray, almost nonexistent; pile short, white anteriorly

to black posteriorly; tergites I—III fasciate.

Male. — Same as female holotype except as follows: pile short, black on posterior

edge of ocellar tubercle; frons silver tomentose; mesonotum short, white pilose;

pile longer, denser on scutellum; palp medium brown; coxae black; distal V 6 of

middle tibia and distal Va of hindtibia dark brown. Aedeagus as in Fig. 59; hy-

poproct as in Fig. 42; ventral gonocoxal spur and gonostylus as in Fig. 50; aedeagus

as in Fig. 59.

VOLUME 59, NUMBERS 1-4

135

Variation. — Overall length 3.5 mm to 5.4 mm <5; 3.7 mm to 5.8 mm 2; with

some brown tomentum above and lateral to female frontal callosity; sometimes

tergite IV also fasciate.

Distribution and ecological placement.—P. rufoabdominalis is generally found

within the confines of desert canyon bottom environments, closely associated with

rock substrates and, when present, is often found under desert willow ( Chilopsis

linearis). It is found along the eastern side of the San Jacinto and Santa Rosa

Mountains, Riverside County, California, the southeastern San Bernardino Moun¬

tains, San Bernardino County, California, the Santa Ana Mountains, Orange

County, California, and the eastern slope of the Sierra Juarez Mountains, Baja

California Norte, Mexico.

Specimens examined. — Holotype: 2, Irwin specimen number 2626, CAS type

no. 10475; P. L. Boyd Deep Canyon Desert Research Center, 6.4 km south of

Palm Desert, Riverside Co., California; MEI; July 17, 1965; at Willow Pool

(Marker O ) on rock under willow tree ( Salix sp.) in filtered light. Paratopotypes:

129 <3; 153 2. Other topotypes: 119 <3; 31 2. Other specimens: 4 <3; 4 2. The following

specimens were examined:

MEXICO: Baja California Norte: La Rumorosa, 14.5 km S, 1372 m, 67/07/

08, MEI, 1 5.

USA-CALIFORNIA: Kern Co.: Walker Pass, 1585 m, 58/08/19, RHP EMP,

1 <3. Orange Co.: Irvine Lake, 63/09/11, MEI, 1 <3. Riverside Co.: Cottonwood

Canyon, 69/07/30, SIF LFL, 1 2; P. L. Boyd Deep Canyon Desert Research Center,

6.4 km S Palm Desert, 63/06/21, EIS, 14 2; 63/07/11, EIS, 9 2; 63/07/26, EIS, 4

6, 8 2; 63/08/09, EIS, 1 <3; 63/08/23, EIS, 4 <3, 6 2; 63/09/23, MEI EIS, 1 2; 63/

09/24, EIS, 3 5, 5 2; 63/10/09, EIS, 1 2; MEI, 3 2; 64/07/01, KWB, 1 2; 64/09/

02, PAR, 5 3, 3 2; ARG, 1 2; 64/09/10, MEI, 1 3, 5 2; 69/06/07, SIF, 4 2; 69/06/

13, SIF, 4 3, 1 2; 320 m, 64/06/24, EIS, 7 3, 5 2; 65/06/08, MEI, 1 3; 67/10/17,

MEI, l 3; 68/11/08, MEI, 6 3; 68/11/17, MEI, 3 3; 69/05/16, MEI, 1 3; 69/05/

18, MEI, 7 3; 69/05/21, MEI, 1 3; 69/05/24, MEI SIF, 4 3, 1 2; 69/06/10, SIF, 2

2; 69/06/15, SIF RMW, 4 3, 6 2; 69/06/19, SIF RMW, 2 3, 1 2; 69/06/20, SIF

RMW, 1 3; 69/06/23, SIF RMW, 2 3, 1 2; 69/05/25, SIF RMW, 4 3, 4 2; 69/06/

27, SIF RMW, 2 3; 69/06/28, SIF RMW, l 3; 69/06/29, SIF RMW, 2 3; 69/07/

05, SIF RMW, 6 3, 1 2; 69/07/10, SIF RMW, 5 3; 69/07/13, SIF RMW, 1 3; 69/

07/14, SIF RMW, 1 3; 69/07/15, SIF RMW, 1 3; 69/07/17, SIF RMW, 2 3; 69/

07/18, SIF RMW, 6 3, 1 2; 69/07/22, SIF RMW, 1 3; 69/07/23, SIF RMW, 1 3;

69/07/25, SIF RMW, 3 3; 69/07/27, SIF RMW, 1 3; 69/07/29, SIF RMW, 2 2;

69/08/01, SIF RMW, 2 3, 2 2; 69/08/03, SIF RMW, 4 3, 3 2; 69/08/05, SIF RMW,

2 3, 1 2; 69/08/06, SIF RMW, 1 3; 69/08/09, SIF RMW, 1 3; 69/08/12, SIF RMW,

2 3, 1 2; 69/08/16, SIF RMW, 1 3; 69/08/17, SIF RMW, 1 3; 69/08/20, SIF RMW,

3 3; 69/08/22, SIF RMW, 6 3; 69/08/24, SIF RMW, 5 3, 1 2; SIF JT, 1 2; 69/08/

25, SIF JT, 1 3; 69/08/26, SIF JT, 1 3; 69/09/01, SIF JT, 4 3; 69/09/05, SIF JT,

16 3, 1 2; 69/09/07, SIF MEI, 60 3, 16 2; 335 m, 64/09/10, EIS, 5 3; 366 m, 64/

09/02, MEI, 1 3; 64/09/10, MEI, 1 3; EIS, 2 2; 67/05/19, MEI, 1 3; 373 m, 64/

06/24, EIS, 2 2; 64/09/02, MEI, 8 3, 6 2; 64/09/10, MEI, 10 3, 3 2; 65/06/08,

MEI, 1 2; 69/08/25, SIF, 2 3, 2 2; 381 m, 64/09/02, MEI, 1 3, 6 2; 64/09/10, MEI,

1 3; 69/06/13, SIF, 1 3; 69/06/29, SIF, 1 2; 400 m, 64/05/31, MEI, 1 2; 64/07/

08, EIS, 1 2; 65/07/17, MEI, 3 3, 3 2; 64/09/02, MEI, 1 3, 7 2; 64/09/10, MEI, 1

136 PAN-PACIFIC ENTOMOLOGIST

52 boharti 53 boydi 54 tomentamala 55 rupina

56 flavipes 57 nigragena 58 bishopensis 59 rufoabdominalis

Figs. 52-59. Aedeagi of male genitalia of Pherocera boharti group, dorsal (left) and lateral (right)

views. 52, ( 1635 ); 53, (7579); 54, (7669); 55, (7757); 56, (7792); 57, (1594)- 58, (2639); 59, (3209).

Scale = 0.1 mm.

2; 410 m, 64/07/01, EIS, 1 6; 64/07/08, EIS, 1 <3; 64/09/02, MEI, 4 <3, 8 9; 420 m,

64/07/23, MEI, 1 2; 434 m, 64/09/02, MEI, 4 <3, 11 2; 64/09/10, EIS, 2 <3, 4 2;

1036 m, 65/06/11, MEI, 2 2. San Bernardino Co.: Morongo, 44/09/26, ALM, 2

2. San Diego Co.: Culp Canyon, Anza-Borrego State Park, 58/06/12, EIS, 1 2;

junction of highways 78 and S2, 65/08/07, SIF SLF SL, 1 < 3 .

Pherocera rupina Irwin, Sp. Nov.

(Figs. 1-5, 15, 23, 31, 39, 47, 55)

Derivation of name. —rupina (Latin) = rocky chasm.

Diagnosis.— P. rupina can be distinguished from all other species of the boharti

group by the following combination of characters: Palp light brown to tan; female

frontal callosity as in Fig. 15; legs almost entirely dark brown to black; male

terminalia with hypoproct as in Fig. 39 and ventral gonocoxal spur and gonostylus

as in Fig. 47.

Description, female holotype {Fig. 1). — See Table 1 for holotype measurements,

Table 2 for discrete character states. Black; silver tomentose; white pilose.

Head: Face and frons similar to Fig. 15; frontal callosity slightly less pointed

ventrally; pile absent on ocellar tubercle, short, sparse dorsolaterad of antennal

socket, short posterior to genal stripe, longer on palp; ratio of antennal scape:

pedicel: flagellum, 16:8:35; scape with two black setae ventrad, pilose; pedicel

VOLUME 59, NUMBERS 1-4

137

sparsely pilose; flagellum similar to Fig. 23, smoother, bare; antenna dark brown;

palp (Fig. 31) tan; proboscis black.

Thorax: Mesonotal vittae indistinct but discernible; mesonotal pile short, sparse;

pleural region tomentose except for katatergite and anepimeron; anepisternum

and katatergite long pilose; coxae tomentose, pilose; legs white pilose, dark brown

with proximal % of foretibia white; wing veins light brown.

Abdomen: Tomentum absent dorsally, relatively dense ventrally; tergites I—III

relatively long pilose and fasciate; rest of abdomen with shorter, sparse pile.

Male. — Same as female holotype except as follows: pile dorsolaterad of antennal

socket longer, denser; legs brown or black with foretibia white; pile relatively long

on abdominal tergites; aedeagus as in Fig. 55; hypoproct as in Fig. 39; ventral

gonocoxal spur and gonostylus as in Fig. 47.

Variation .—Overall length 3.1 mm to 4.7 mm <3; 3.5 mm to 5.3 mm 9; genal

stripe occasionally lightly powdered with tomentum; the populations from Baja

California Sur, Mexico, and some southern California specimens have three or

four black setae on antennal scape.

Distribution and ecological placement. — P. rupina was collected repeatedly on

rock substrates in desert canyon bottom environments in Riverside County, Cal¬

ifornia and in Baja California Sur and Norte, Mexico.

Specimens examined. — Holotype: 9 , Irwin specimen number 1725, CAS type

no. 10476; P. L. Boyd Deep Canyon Desert Research Station, 6.4 km south of

Palm Desert, Riverside Co., California; MEI; June 8, 1965; at marker 56.8 on

rock in filtered light under Chilopsis, not near water, in sunny area in light wind.

Paratopotypes: 188 < 3 ; 94 9 . Other topotypes: 9 < 3 . Other specimens: 48 6, 13 9 .

The following specimens were examined:

MEXICO: Baja California Norte: San Quintin, inland from, 55/05/25, FXW,

1 9. Baja California Sur: Guadalupe, 100 km NW of La Paz, 107 m, 68/04/20,

MEI, 7 <3; Loreto, 8 km W, 15 m, 68/04/17, MEI, 1 <3, Santa Rosalia, 40 km S,

10.7 m, 68/04/15, MEI, 2 5.

USA-CALIFORNIA: Riverside Co.: Massacre Canyon at highway 79, 64/07/

29, MEI, 25 6 , 1 9 ; PAR, 5 6 , 7 9 ; 64/07/30, MEI, 7 3 ; MacCoy Springs, 63/04/

08, JCH, 1 9 ; P. L. Boyd Desert Research Center, 6.4 km S of Palm Desert, 63/

05/03, EIS, 7 < 3 , 6 9 ; 63/05/17, EIS, 5 < 3 , 2 9 ; 63/05/30, EIS, 3 9 ; 63/06/21, EIS,

1 3 ; 63/07/11, EIS, 3 6 , 1 9 ; 63/07/21, EIS, 5 5 ; 63/07/26, EIS, 3 5 ; 63/09/05, EIS,

1 9 ; 63/09/24, EIS, 8 < 3 ; 64/05/09, MEI, 1 < 3 ; 64/07/01, KWB, 3 6 , 1 9 ; 69/06/07,

SIF, 4 3 ; 320 m, 64/05/09, MEI, 2 6 ; 64/05/25, MEI, 2 6 ; 64/06/24, EIS, 7 9 ; 64/

07/01, EIS, 1 9 ; 64/07/23, MEI, 3 6 ; 64/09/10, MEI, 1 6 ; 65/06/08, MEI, 3 < 3 , 5

9 ; 65/07/17, MEI, 1 6 ; 67/10/17, MEI, 2 6 ; 68/09/08, MEI, 4 2 9 ; 69/05/18,

MEI, 8 < 3 , 3 9 ; 69/05/21, MEI, 1 6 , 2 9 ; 69/05/24, MEI SIF 10 6 , 7 9 ; 69/06/10,

SIF, 2 9 ; 69/06/15, SIF RMW, 2 6 , 9 9 ; 69/06/19, SIF RMW, 3 9 ; 69/06/20, SIF

RMW, 1 9 ; 69/06/23, SIF RMW, 1 < 3 , 1 9 ; 69/06/25, SIF RMW, 1 9 ; 69/06/27,

SIF RMW, 1 < 3 ; 69/06/29, SIF RMW, 1 < 3 , 1 9 ; 69/07/01, SIF RMW, 2 < 3 ; 69/07/

02, SIF RMW, 1 < 3 ; 69/07/05, SIF RMW, 2 6 , 1 9 ; 69/07/10, SIF RMW, 1 < 3 ; 69/

07/13, SIF RMW, 1 6 ; 69/07/15, SIF RMW, 1 6 ; 69/07/22, SIF RMW, 1 3 ; 69/

08/24, SIF RMW, 1 3 ; 69/09/07, SIF MEI, 2 3 ; 335 m, 65/06/08, MEI, 2 3 ; 343

m, 64/05/19, MEI, 7 9 ; 64/06/24, EIS, 30 3; 64/07/01, EIS, 12 3 ; 64/09/10, MEI,

1 3 ; 65/06/08, MEI, 4 3 ; 350 m, 64/09/10, MEI, 2 3 ; 65/06/08, MEI, 2 3 ; 358 m,

65/06/08, MEI, 1 3 ; 366 m, 63/09/10, EIS, 1 3; 64/05/31, MEI, 4 3; 64/09/02,

138

PAN-PACIFIC ENTOMOLOGIST

MEI, 4 <3; 64/09/10, MEI, 2 6; 65/06/08, MEI, 1 <3, 1 9; 373 m, 64/05/19, MEI, 3

9; 64/05/31, MEI, 1 9; 64/09/02, MEI, 2 9; 64/09/10, MEI, 2 3 9; 65/06/08,

MEI, 1 <3; 381 m, 64/09/02, MEI, 1 <3; 400 m, 64/05/09, MEI, 8 <3; 64/05/19, MEI,

4 9; 64/06/24, EIS, 1 9; 64/07/08, EIS, 7 6, 2 9; 64/09/02, MEI, 2 9; 65/04/28,

MEI, 1 3; 65/06/08, MEI, 2 9; 65/07/17, MEI, 3 3; 412 m, 64/05/31, MEI, 5 <3;

64/07/01, EIS, 1 6; 64/07/08, EIS, 6 3; 420 m, 67/05/19, MEI, 2 9; 434 m, 64/

09/02, MEI, 1 9; 64/09/10, EIS, 1 9. Winchester, 68/06/20, WI, 1 9. San Diego

Co,: Culp Canyon, Anza-Borrego State Park, 58/06/26, EIS, 1 9.

Pherocera tomentamala Irwin, Sp. Nov.

(Figs. 14, 22, 30, 38, 46, 54)

Derivation of name. — tomentum (Latin) = wooly hairs; mala (Latin) = jaw.

Diagnosis.—P. tomentamala can be distinguished from all other species of the

boharti group by the following combination of characters: female frontal callosity

as in Fig. 14; mesonotum with indistinct vittae; genal stripe lacking; male ter-

minalia with hypoproct as in Fig. 38 and ventral gonocoxal spur as in Fig. 46.

Description, female holotype. — See Table 1 for holotype measurements, Table

2 for discrete character states. Dark brown; tomentum silver; pile white.

Head: Face and frons as in Fig. 14; pile lacking on frons, very sparse on posterior

portion of lower gena, elongate on palp; ratio of antennal scape : pedicel: flagellum,

15:7:39; scape with two elongate black setae ventrad and white pilose overall;

pedicel very sparsely pilose; flagellum similar to Fig. 22; bare; antenna, palp (Fig.

30) and proboscis brown.

Thorax: Mesonotal vittae indistinct but discernible; mesonotal pile short, sparse;

pleural region tomentose except for katatergite and anepimeron; anepisternum

and katatergite pilose; coxae tomentose, white pilose; legs orange with ventral

proximal % of femora, distal V 5 of tibiae and tarsi of mid and hindlegs black;

foreleg entirely black except for proximal % of tibia white and joint between femur

and tibia orange; legs predominantly white pilose, distal V 3 of foretibia short, black

pilose; wing veins light brown.

Abdomen: Tomentum absent dorsally, sparse ventrally; pile moderate, covering

entire abdomen; tergites I—II fasciate.

Male. — Same as female holotype except as follows: abdominal pile longer; ae-

deagus as in Fig. 54; hypoproct as in Fig. 38; ventral gonocoxal spur and gonostylus

as in Fig. 46.

Variation. — Overall length 4.9 mm to 5.2 mm <3; 5.0 mm 9.

Distribution and ecological placement.—P. tomentamala has been collected in

association with dead twigs under Cercidium trees on inland stable sand dunes

and in cotton fields. Although only eight specimens are presently known, they

indicate a wide distributional range from Imperial, Riverside, and San Bernardino

Counties in California and Maricopa County in Arizona.

Specimens examined. — Holotype: 9, Irwin specimen number 1670, CAS type

no. 10480; 10 km north of Glamis, Algadones Dunes, Imperial Co., California

at 100 m; MEI; May 21, 1967; under Cercidium tree. Paratypes: 3 <3, 1 9. Other

specimens: 2 <3, 1 9. The following specimens were examined:

ARIZONA: Maricopa Co.: Gila Bend, 11.3 km NW, 215 m, 61/04/10, RHP

EMP, 1 <3. CALIFORNIA: Imperial Co.: Calexico, 57/07/11, collector unknown,

VOLUME 59, NUMBERS 1-4

139

1 2; Glamis, 10 km N, 100 m, 67/05/21, MEI, 1 2. Holtville, nr., 75/08/10, GRB,

1 <3, 1 2. Riverside Co.: Indio, Keosegan Ranch, 70/07/15, MEI, 1 <3; 70/07/26,

MEI, 1 <3. San Bernardino Co.: Oro Grande, 54/06/16, collector unknown, 1 <3.

Literature Cited

Cole, F. R. 1923a. A revision of the North American two-winged flies of the family Therevidae.

Proc. U.S. Natl. Mus. 62:1-140.

-. 1923b. Expedition of the California Academy of Sciences to the Gulf of California in 1921.

Diptera from the islands and adjacent shores of the Gulf of California. II. General Report.

Proc. Calif. Acad, of Sci., Series 4, 12:457-481.

-. 1965. Family Therevidae, pp. 348-354. In: A. Stone, C. W. Sabrosky, W. W. Wirth, R. H.

Foote, and J. R. Coulson, eds., A catalog of the Diptera of America north of Mexico. U.S.

Dept. Agr. Handbook 276. Washington, D.C., 1696 pp.

Cole, F. R., and E. I. Schlinger. 1969. The flies of western North America. Univ. Calif. Press, Berkeley

& Los Angeles, 693 pp.

Hardy, G. H. 1927. On the phylogeny of some Diptera Brachycera. Proc. Linn. Soc. New South

Wales 52:380-386.

Irwin, M. E. 1971. Ecology and biosystematics of the pherocerine Therevidae. Ph.D. Dissertation

in Entomology, Univ. of Calif., 256 pp.

-. 1973. A new genus of the Xestomyza- group from the western coast of South Africa, based

on two new species with flightless females (Diptera: Therevidae). Ann. Natal Mus. 21:533-556.

-. 1976. Morphology of the terminalia and known ovipositing behavior of female Therevidae

(Diptera: Asiloidea), with an account of correlated adaptations and comments on phylogenetic

relationships. Ann. Natal Mus. 22:913-935.

-. 1977. Two new genera and four new species of the Pherocera- group from western North

America, with observations on habitats and behavior (Diptera: Therevidae: Phycinae). Proc.

Ent. Soc. Wash. 79:422-451.

Irwin, M. E., and L. Lyneborg. 1981a. The genera of Nearctic Therevidae. Bull. Ill. Natur. Hist.

Surv. 32:191-277.

-. 1981b. Family Therevidae, Chapter 37, pp. 513-523. In: J. F. McAlpine, B. V. Peterson,

G. E. Shewed, H. J. Teskey, J. R. Vockeroth, D. M. Wood, coordinators, Manual of Nearctic

Diptera. Agr. Canada Monogr. 27. Ottawa, Ontario, Vol. 1, 674 pp.

Lyneborg, L. 1968. A comparative description of the male terminalia in Thereva Latr., Dialineura

Rond., and Psilocephala Zett. (Diptera, Therevidae). Ent. Medd. 36:546-559.

-. 1972. A revision of the Xestomyza-group of Therevidae (Diptera). Ann. Natal Mus. 21:297-

376.

McAlpine, J. F., B. V. Peterson, G. E. Shewed, H. J. Teskey, J. R. Vockeroth, and D. M. Wood. 1981.

Manual of Nearctic Diptera. Agr. Canada Monogr. 27. Ottawa, Ontario, Vol. 1, 674 pp.

Rauch, P. A. 1970. Electronic data processing for entomological museums, an economical approach

to an expensive problem. Ph.D. Dissertation in Entomology, LIniv. of Calif, 78 pp.

Rodendorf, B. B. 1964. Historical development of two-winged insects [in Russian]. Proc. Acad. Sci.

U.S.S.R. (Paleontological Institute) 100:1-311.

Stuckenberg, B. R., and M. E. Irwin. 1973. Standards for entomological labels. Bud. Ent. Soc.

Amer. 19:164-168.

Torre-Bueno, J. R. de la. 1950. A glossary of entomology. Brooklyn Ent. Soc., Brooklyn, N.Y.,

336 pp.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 140-147

Review of the Euchroeine Chrysidids

(Hymenoptera: Chrysididae)

Lynn S. Kimsey

Department of Entomology, University of California, Davis, California 95616.

The Euchroeini contains some of the more highly modified and unusual-looking

genera in the Chrysididae. The taxonomy of this group has long been confused

and as many as 20 different generic names have been used. Included in the

Euchroeini are the largest chrysidids, members of the genus Stilbum, reaching up

to 25 mm in length. Although Euchroeini is primarily an Old World group, 2

species of Spinolia are found in the New World, and Neochrysis is strictly neo¬

tropical.

I am following the higher classification proposed by Bohart and Kimsey (1980)

and Kimsey and Bohart (1981). Members of the Euchroeini are distinguished by

their characteristic wing venation: the forewing radial sector extends over half the

length of the marginal cell and bends slightly away from the coastal margin. Based

on this characteristic 5 genera are included: Euchroeus, Stilbichry>sis, Spinolia,

Stilbum and Neochrysis.

This study is intended as a preliminary reorganization of the generic taxonomy

of these chrysidids in preparation for a more detailed cladistic analysis of the

chrysidid genera.

Under each genus I give a preliminary list of included species. These lists are

based on Linsenmaier (1959, 1968), Mocsary (1889), original descriptions and

examination of types, as indicated by an asterisk. Some names in the lists may

prove to be synonyms as I have not seen all of the types. Host and distributional

data are summarized in Table 1 and Fig. 1, respectively. Male genitalia are not

particularly useful in this group at the generic level, except to distinguish Neo¬

chrysis.

The following abbreviations are used: RS = radial sector, 1A = anal vein, Cu =

cubital vein, T = gastral tergum, S = gastral sternum, and MOD = midocellus

diameter.

This paper is dedicated to Richard M. Bohart who has done more to straighten

out the taxonomy of the Chrysididae than any other systematist living or dead.

Key to Genera of Euchroeini

1. Metanotum produced posteriorly into large spoon-like projection (Fig.

5); least frontal width less than eye width in front view (Fig. 2); T-III

with 4 symmetrical apical teeth.. Stilbum Spinola

- Metanotum not produced into spoon-like projection, least frontal width

usually greater than eye width, T-III variable. 2

2. Hindwing with 3 sclerotized veins and M+Cu forked; scutellum elevated

above level of scutum and metanotum; metanotal surface irregular and

denticulate .. Stilbichrysis Bischoff

- Hindwing with 2 or fewer sclerotized veins and M+Cu, not forked or

VOLUME 59, NUMBERS 1-4

141

StiJbum

Euchroeus

Stilbichrysis

= Spinolia

Neochrysis

Fig. 1. Distribution map of euchroeine genera.

absent; scutellum not elevated above level of scutum and metanotum;

metanotal surface relatively even and smooth . 3

3. Mesopleuron with 2 large teeth pointing ventrally, similar to Fig. 6; T-III

dentate, with 8 or more asymmetrical teeth or more rarely 4 white teeth

. Euchroeus Latreille

- Mesopleuron without 2 large teeth pointing ventrally; T-III with 2, 4 or

6 symmetrical, non-white teeth (as in Fig. 8) or edentate . 4

4. T-III with 4 or 6 apical teeth . Neochrysis Linsenmaier

- T-III apical rim edentate, with small irregularities or with 2 small lateral

teeth. Spinolia Dahlbom

Euchroeus Latreille

Chrysis Linnaeus, in part.

Euchroeus Latreille, 1809:578. Type species: Chrysis purpurata Fabricius, 1787.

Brugmoia Radoszkowski, 1877:26. Type species: Brugmoia pellucida Radosz-

kowski, 1877, orig. desig.

Brugmoja Radoszkowski, 1877:pl. 2, fig. 12. Misspelling of Brugmoia.

Polyodontus Radoszkowski, 1877:25. Type species: Polyodontus stchurovsky Ra¬

doszkowski, 1877, orig. desig.

Euchroeides Nurse, 1904:23. Type species: Euchroeides oblatus Nurse, 1904, by

monotypy.

Afrospinolia Linsenmaier, 1968:42. Type species: Euchroeus katanganus Linsen¬

maier, 1968, orig. desig. (treated as subgenus of Euchroeus). New synonymy.

Diagnosis.— Malar space 1.0 MOD long or usually longer; face with transverse

frontal carina (Fig. 3); tongue as long as eye height or longer; mesopleuron with

transverse medial groove and 2 or more sharp teeth pointing ventrally; metanotum

evenly rounded; T-I anterior corners rounded; T-III apically with more than 8

142

PAN-PACIFIC ENTOMOLOGIST

Table 1. Hosts of euchroeine species.

Euchroeine species

Host species

Family

Author

Stilbum cyanurum

Sceliphron destillatorium Illig.

Sphecidae

Grandi, 1961

Eumenes unguicidatus Villers

Eumenidae

Sceliphron sp.

Sphecidae

Zimmermann, 1937

Eumenes sp.

Eumenidae

Odynerus sp.

Rhynchium sp.

Chalcidoma sp.

Megachile sp.

Megachilidae

S. viride

Eumenes maxillosus DeGeer

Eumenidae

Spinolia neglectus

Odynerus spinipes (Linnaeus)

Odynerus reniformis (Gmelin)

Linsenmaier, 1959

S. humboldti

Paravespa grandis Morice

S. dallatorreanus

Pterocheilus bembeciformis

S. lamprosomus

Odynerus spiricornis (Spinola)

Neochrysis sp.

Podium sp.

Sphecidae

Summarized in

Trypoxylon sp.

Bohart and Kim-

Sceliphron sp.

sey, 1980

irregular teeth or denticles (Fig. 9) (except singularis ), and apical rim and teeth

often at least partly white or transparent; female T-V and S-V unmodified (as in

Figs. 11, 12).

Distribution (Fig. 1).— Europe and the Middle East, south to South Africa.

Included species.—artifrons Edney 1947*, aurovirens ( Mocsary) 1913 ( Chrysis ),

binodatus Edney 1947*, candens Dahlbom 1845, chrysidiformis Magretti 1898,

doursi Gribodo 1875, egregius Buysson 1887, eos (Semenov) 1954 (Chrysis),

hellenicus ( Mocsary) 1913 (Chrysis)*, jordanicus Linsenmaier 1968, katanganus

Linsenmaier 1968, limbatus Dahlbom 1854*, mongo liens Tsuneki 1947, moricei

Buysson 1896*, ob/atus (Nurse) 1904 (Euchroeides)*, oculatissimus Buysson 1898,

pellucidus ( Radozskowski) 1877 (Brugmoia), purpuratus (Fabricius) 1787 (Chry¬

sis)*, rugulosus (Mocsary) 1909 (Chrysis)*, singularis (Spinola) 1838 (Chrysis)*,

stchurovsky (Radozskowski) 1877 (Polyodontus).

Discussion. — Members of Euchroeus can be readily distinguished from other

chrysidids by the dentate mesopleuron and numerous irregular teeth along the

apical rim of T-III. One exception to this is the species singularis, which has

roughly 4 symmetrically arranged white teeth. Other characteristics of this species

including the mesopleural teeth, long malar space and unmodified female T-V

and S-V, definitely place singularis in Euchroeus. Another distinctive feature that

occurs in several species, including singularis and candens, is the white coloration

on the apical rim of T-III. This characteristic also occurs in Neochrysis (Ipsiura),

Argochrysis and Spintharosoma, the latter 2 belonging in the Chrysidini.

Euchroeus species are rarely encountered; purpuratus, a European wasp, is

probably the most commonly collected.

Neochrysis Linsenmaier

Chrysis Linnaeus, in part.

Neochrysis Linsenmaier 1959:74. Type species: Chrysis punctatissima Spinola,

VOLUME 59, NUMBERS 1-4

143

2 Stilbum calens

4 Spinolia dournovi

— —

6 Stilbichrysis biselevata

5 Stilbum calens

8 Neochrysis carina

7 Neochrysis panamensis

Spino'lia termini

tev e vli

■t l c * •£/■»&

9 Euchroeus purpurata

10 Stilbum

cyanurum

Spinolia lamprosoma

Figs. 2-14. 2, 3, Male face, front view. 4, Female head, lateral view: eye height (EH), malar space

(MS), tongue length (TL). 5, Scutellum, metanotum and propodeum, dorsal view. 6, Male body, lateral

view: radial sector (RS), cubital vein (Cu), mesopleural teeth (MT). 7, Tergum I, dorsal view. 8, 9,

Male tergum III, apical rim. 10, Ovipositor, lateral view. 11-14, Ovipositor segments. 11,13, Tergum

V, dorsal view. 12, 14, Sternum V, ventral view.

144

PAN-PACIFIC ENTOMOLOGIST

1840, orig. desig., nec Villers 1789 = carina Brulle, 1846 (treated as subg. of

Chrysis).

Neochrysis Linsenmaier, Bohart 1966 (elevated to genus).

Pleurocera Guerin 1842:149.Preocc.byP/e , wroc^raRafinesque, 1818. Type species:

Pleurocera viridis Guerin, 1842 (nec Oliver, 1790) = Chrysis bruchi Brethes,

1902, by monotypy.

Pleurochrysis Bohart 1966:144, n.n. for Pleurocera Guerin (treated as subg. of

Neochrysis by Bohart and herein).

Exochrysis Bohart 1966:141. Type species: Chrysis panamensis Cameron, 1888,

orig. desig. (treated as subg. of Neochrysis by Bohart and herein).

Tpsiura Linsenmaier 1959:74. Type species: Chrysis marginal is Brulle, 1846, orig.

desig. (treated as subg. of Neochrysis by Bohart and herein).

Diagnosis. — Malar space 1.5 MOD long or less, tongue length usually shorter

than eye height, mesopleuron without medial groove, T-I anterior comers sharply

angled (Fig. 7), T-III with 4 or 6 symmetrical apical teeth (Fig. 8), female T-V

and S-V unmodified or T-V with small ridge-like annulations, male paramere

divided into a distinct, articulated telomere and basomere.

Distribution (Fig. 7). —Western Hemisphere, the southern half of the United

States south to Chile and Argentina.

Discussion. —See Bohart and Kimsey (1980) and Kimsey and Bohart (1981) for

discussion of generic and subgeneric taxonomy and included species. The male

genitalia of Neochrysis is unique in the Euchroeini due to the articulated telomere.

Spinolia Dahlbom

Chrysis Linnaeus, in part.

Euchroeus Latreille, in part.

Holochrysis of Mocsary, in part.

Pseudochrysis of Balthasar and Trautmann, in part.

Spinolia Dahlbom 1854:363. Type species: Chrysis magnifica Dahlbom, 1854,

orig. desig. (= lamprosoma Forster, 1853).

Gonochrysis Lichtenstein 1876:227. Type species: Chrysis albipennis Dahlbom,

1854 (= unicolor Dahlbom, 1831).

Achrysis Semenov 1892:486. Type species: Chrysis unicolor Dahlbom, 1831 (treated

as subg. of Pseudochrysis).

Spinolaia Schulz 1906:154, emendation of Spinolia.

Pseudospinolia Linsenmaier 1951:31. Type species: Chrysis uniformis Dahlbom,

1854, orig. desig. (treated as subg. of Euchroeus). New synonymy.

Neospinolia Linsenmaier 1968:39. Type species: Chrysis tertrini Buysson, 1897,

orig. desig. (treated as subg. of Euchroeus). New synonymy.

Pseudospinolia Linsenmaier, Bohart and Kimsey, 1980, elevated to genus.

Diagnosis. — Malar space 1.0 MOD long or shorter (Fig. 4); tongue as long as

eye height or longer (Fig. 4) (except tertrini ); mesopleuron with transverse medial

groove and without teeth or knobs; metanotum evenly rounded; T-I anterior

corners rounded; T-III apical rim smooth or with slight irregularities and rounded,

some species with 2 small lateral teeth.

Distribution (Fig. 1). — Palearctic, Africa, 1 species reaching North America

(neglectus) and 1 in Chile (tertrini).

VOLUME 59, NUMBERS 1-4

145

Included'species. — ardens (Mocsary) 1902 ( Chrysis)*, aureicollis (Abeille) 1878

( Chrysis )*, bouvieri (Buysson) 1897 (C/zrysw)*, chalcites (Mocsary') 1890 (Chry¬

sis)*, chobauti (Buysson) 1891 (Chrysis)*, dallatorreanus (Mocsary) 1896 (Chrysis)*,

dournovi (Radozskowski) 1866 (Chrysis)*, gestroi (Gribodo) 1874 (Chrysis), gra-

tiosus (Mocsary) 1889 (Chrysis), herodianus Morice 1909, humboldti (Dahlbom)

1845 (Chrysura)*, ignithorax (Balthasar) 1853 (Pseudochrysis), incrassatus (Spi-

nola) 1838 (Chrysis), insignis (Lucas) 1849 (Chrysis), lamprosomus (Forster) 1853

(Chrysis)*, marqueti (Buysson) 1887 (CArys/s)*, morawitzi (Mocsary) 1889 (Chry¬

sis)*, neglectoides Linsenmaier 1959 NEW COMB., neglectus (Shuckard) 1837

(Chrysis)*, pulawskii Linsenmaier 1968, rogenhoferi (Mocsary) 1889 (Chrysis)*,

rugosa Buysson 1900, tertrini (Buysson) 1897 (Chrysis)*, there side (Buysson) 1900

(Chrysis)* NEW COMB., tmnsversus (Dahlbom) 1854 (Chrysis), tricoloricornis

Linsenmaier 1968 NEW COMB., unicolor (Dahlbom) 1831 (Chrysis)*, uniformis

(Dahlbom) 1854 (Chrysis)*, vogti (Trautmann) 1926 (Pseudochrysis).

Discussion. — Linsenmaier (1959, 1968) placed the euchroeine species without

apical teeth on T-III in 6 subgenera, Spinolid, Pseudospinolid, Stilbichrysis, Pri-

mdchroeus, Neochrysis, Prospinolid and Neospinolid, which he included in Eu-

chroeus. Primdchroeus actually belongs in the Chrysidini, based on the position

of the RS vein. The remaining subgenera were distinguished by the structure of

the mesopleuron, length of the malar space, presence or absence of a transverse

frontal carina on the face and the structure of T-II and T-III. In the case of

Neochrysis and Stilbichrysis this approach is too conservative as these groups

differ sufficiently to require generic status. After examining additional species

from Africa it has become apparent that there are no clear-cut differences between

the remaining subgenera. The elevation of Pseudospinolid to generic status by

Bohart and Kimsey (1980) was premature. Characteristics such as the mesopleural

sculpture, transverse frontal carina, pronotal carina and modification of the female

ovipositor segments (T-V and S-V) vary too inconsistently between species to

distinguish subgenera (see Table 2). As a result, I am synonymizing these names

under Spinolid Dahlbom, which has priority.

Stilbichrysis Bischoff

Stilbichrysis Bischoff 1910:445. Type species: Stilbichrysis biselevdtd Bischoff,

1910, orig. desig.

Didgnosis. —Body, lateral view (Fig. 6), face with frontal carina, tongue longer

than eye height, malar space about 1.0 MOD long, mesopleuron with 2 teeth and

strong medial groove, hindwing with well developed RS, M+Cu and 1A veins,

scutellum elevated, metanotum elevated with medial tooth, T-I anterior corners

rounded, T-III apical rim smooth and rounded; female T-V and S-V apically with

large ridge-like annulations (as in Fig. 10).

Distribution (Fig. 1).— Africa: Zimbabwe, Mozambique, South Africa and Tan¬

zania.

Included species. —biselevdtd Bischoff 1910*, serruldtd (Edney) 1947 (Chrysis)*

NEW COMB.

Discussion. — Both of these species are rarely collected; they superficially resem¬

ble Stilbum but can be distinguished from it and other euchroeine genera by: T-III

rim edentate, metanotum denticulate, malar space short, mesopleuron dentate

and grooved and hindwing with sclcrotized M+Cu and 1A veins.

146

PAN-PACIFIC ENTOMOLOGIST

Table 2. Morphological characteristics of selected species of Spinolia representing 3 subgenera

according to Linsenmaier (1959, 1968).

Species

Sub¬

genus

Neo-

Subgenus Subgenus spino-

Spinolia s.s. Pseudospinolia lia

Characteristics

Face with transverse frontal carina

—

- +

— —

Pronotum with lateral carina

—

— —

—

— —

—

Malar space longer than 1 MOD

—

— —

—

— —

Mesopleuron with medial groove

- +

—

— —

Mesopleuron with U-shaped carina

Female T-V and S-V annulate

+ -

(Fig. 10)

Female T-V and S-V dentate

—

- +

(Figs. 13, 14)

Female T-V and S-V unmodified

—

(Figs. 11, 12)

—

— —

—

+ -

Tongue shorter than eye height

—

— —

—

— —

Stilbum Spinola

Chrysis, Linnaeus, in part.

Stilbum Spinola 1806:9. Type species: Chrysis calens Fabricius, 1781, orig. desig.

Diagnosis.— Face with frontal carina, malar space 2.0 MOD or more long (Fig.

2), tongue about as long as eye height, mesopleuron with strong medial groove

and 3 or more teeth or knobs, scutellum elevated, metanotum enlarged into a

backward projecting spoon-like structure (Fig. 5), T-I anterior comers rounded,

T-III with 4 apical teeth, female T-V and S-V apically pointed with large ridge¬

like annulations (Fig. 10).

Distribution (Fig. 1).— Southern Europe, Africa, Madagascar, India to the Aus¬

tralasian Region.

Included species (after Linsenmaier, 1959).—calens (Fabricius) 1781 (Chrysis),

chrysocephalum Buysson 1897, cyanurum Forster 1771, pici Buysson 1891, viride

Guerin 1842.

Discussion.—Stilbum includes the largest chrysidids and certainly the most

commonly encountered species in Africa, parts of Asia and Australia. The exact

number of species in this genus is controversial; unlike Linsenmaier (1959), Zim-

mermann (1937) considered Stilbum to contain only 2 species, cyanurum and

viride, and he synonymized the remainder under cyanurum. The question of the

number of species in Stilbum needs further study. Members of this genus can be

distinguished by the unusually long narrow face (Fig. 2), mesopleural sculpture,

metanotal projection and T-III with 4 teeth apically. Stilbum commonly para¬

sitizes wasps that build mud nests on a variety of surfaces including man-made

structures, which may account for the commonness and wide distribution of this

genus.

VOLUME 59, NUMBERS 1-4

147

Literature Cited

Bischoff, H. 1910. Die Chrysididen des koniglichen zoologischen Museums zu Berlin. Mitt. Zool.

Mus. Berlin 4:427-493.

Bohart, R. M. 1966. The genus Neochrysis in America north of Mexico. Bull. Brooklyn Ent. Soc.

63:139-144.

Bohart, R. M., and L. S. Kimsey. 1980. A generic synopsis of the Chrysididae of America north of

Mexico. J. Kansas Ent. Soc. 53:137-148.

-. 1982. A synopsis of the Chrysididae in America north of Mexico. Mem. Amer. Ent. Inst.

33:1-266.

Dahlbom, A. G. 1854. Disposito methodica specierum Hymenopterorum, etc. Berlinianis, Lund,

20 pp.

Grandi, G. 1961. Studi di un entomologo sugli imenotteri superiori. Boll. Inst. Ent. Univ. Bologna

25:1-659.

Guerin-Meneville, F. E. 1842. Description de quelques Chrysidides nouvelle. Rev. Mag. Zool. (1)

5:144-150.

Kimsey, L. S., and R. M. Bohart. 1981 (1980). A synopsis of the chrysidid genera of neotropical

America. Psyche 87:75-91.

Latreille, P. H. 1809. Genera crustaceorum et insectorum. Vol. 4. A. Koenig, Parisiis et Argentorati,

397 pp.

Lichtenstein, J. 1876. Note sur le genre Chrysis. Petites Nouv. Ent. 2:227.

Linsenmaier, W. 1951. Die europaischen Chrysididen. Mitt. Schweiz. Ent. Ges. 24:1-110.

-. 1959. Revision der Familie Chrysididae. Mitt. Schweiz. Ent. Ges. 32:1-232.

-. 1968. Revision der Familie Chrysididae. Zweiter Nachtrag. Mitt. Schweiz. Ent. Ges. 41:1 —

144.

Mocsary, A. 1889. Monographia Chrysididarum orbis terrestris universi. Budapest, Academia Scien-

tarum Hungarica, 643 pp.

Nurse, C. G. 1904. New species of Indian Hymenoptera. J. Bombay Nat. Hist. Soc. 16:19-26.

Radoszkowski, O. 1877. Chrysidiformis, Mutillidae and Sphecidae, pp. i-ii, 1-42. In: A. Fedtschen-

ko, Reise in Turkestan, vol. 2 (Zool. Theil), part 5.

Schulz, W. A. 1906. Spolia Hymenopterologica. A. Pape, Paderbom, 355 pp.

Semenov, A. 1892. De genere Pseudochrysis. Horae Soc. Ent. Rossicae 26:480-491.

Spinola, M. 1806. Insectorum Liguriae species novae aut rariores, etc., Vol. 1. A. Koenig, Parisiis

et Argentorati, xvii + 160 pp.

Zimmermann, H. 1937. Uber die Verbreitung und Formenbildung der Gattung Stilbum. Arch.

Naturges. (Leipzig) 6:645-662.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 148-151

Bohartiellus, a New Genus of Doryctinae from

South America (Hymenoptera: Braconidae)

Paul M. Marsh

Systematic Entomology Laboratory, IIBIII, Agricultural Research Service, U.S.

Department of Agriculture, Beltsville, Maryland 20705.

Throughout my undergraduate study in entomology at the University of Cal¬

ifornia, Davis, I was undecided about an area of specialization. During the summer

of my senior year I enrolled in the summer held course, Ent. 49, which was

required for graduation. That year the six-week course was held at the Southwest

Research Station in Portal, Arizona, under the direction of Richard M. Bohart.

Those six weeks were a memorable experience and I probably learned more about

insects during that short amount of time than during the previous four years.

Dick’s enthusiasm about insects and his interest in the students made it clear to

me that taxonomy was the field I would pursue.

When I began graduate work, the major question became which group of insects

to study. Since Dick was my advisor, I expected he would suggest a group of

aculeate wasps, his first love. During our first discussion, however, I recall Dick

saying to me, “Paul, if I were starting now, I would take up the Braconidae.” He

must have sensed the emerging concern about the effects of insecticides on the

environment and the resulting need to emphasize biological control, at the heart

of which is knowledge of natural enemies of insect and mite pests, particularly

the parasitic wasps. I took his advice and launched into a thesis research problem

on the systematics of Braconidae. Shortly before I completed my Ph.D. degree in

1964, Rachel Carson published “Silent Spring” and biological control took on a

new and important meaning. Three days after handing in my thesis, I reported

to Washington, D.C., to begin my career as a specialist on the Braconidae for the

USDA’s Systematic Entomology Laboratory.

I owe an extreme debt of gratitude to “Doc” Bohart for his influence on my

career and his training and encouragement during my student days at Davis. To

this end I dedicate the remarkable genus of Braconidae described below to a

remarkable man.

Bohartiellus Marsh, New Genus

(Braconidae: Doryctinae)

Diagnosis. — Head cuboid, occipital carina present, clypeus semi-circularly

emarginate and together with mandibles forming a circular mouth opening; fore¬

tibia with row of 6-8 stout spines along anterior edge; propodeum medially at

base with a laterally flattened lamella which projects dorsally (Fig. 1); forewing

(Fig. 3) with two cubital cells, recurrent vein ending at first cubital cell, nervulus

present, first brachial cell closed at apex well before recurrent vein; hind wing (Fig.

3) without nervellus; male abdomen elongated, often more than twice as long as

thorax, due to telescoping and extension of segments 4-7.

Type of genus. — Bohartiellus cornutus Marsh, new species.

VOLUME 59, NUMBERS 1-4

149

Figs. 1, 2. Bohartiellus cornutus. 1, Propodeum, lateral view. 2, Head and thorax, dorsal view.

Etymology.— Named for Richard M. Bohart, mentor and friend.

This genus is similar in wing venation and elongation of the male abdo¬

men to Aivalykus Nixon but differs from Aivalykus and from all known western

hemisphere Doryctinae by the unusual lamella projecting dorsally from the pro¬

podeum (Fig. 1). In this regard it is similar to Spathioplit.es Fischer from Chad

150

PAN-PACIFIC ENTOMOLOGIST

Fig. 3. Bohartiellus cornutus, fore and hindwings.

(French Equatorial Africa) but the lamella-like structure of Spathioplites arises

from the postscutellum rather than from the propodeum as in Bohartiellus. Fur¬

thermore, Bohartiellus has two cubital cells in the forewing whereas Spathioplites

has three.

Bohartiellus cornutus Marsh, New Species

(Figs. 1-3)

Female .—Length of body, 2.5-3.0 mm; ovipositor, 2.0-2.5 mm. Color: head

dark honey yellow; antenna honey yellow, apical 3-4 flagellomeres brown; thorax

and abdomen dark brown, median mesonotal lobe light brown anteriorly; legs

brown, fore tarsus yellow, hind tibia yellow at base; forewings banded (Fig. 3).

Head: vertex (Fig. 2) and frons strigate 1 , temples smooth, face strigate-rugose with

median raised smooth area; malar space about Vs eye height; ocell-ocular distance

about 3 times diameter of lateral ocellus; antenna 17-segmented (broken in ho-

lotype). Thorax: median mesonotal lobe strigate-rugose, anterior corners produced

into large horn-like structures (Fig. 2), lateral mesonotal lobes smooth; notauli

not distinct, indicated only by carinate lines; scutellar furrow with 6-8 cross

carinae; scutellum swollen, smooth; mesopleuron smooth, subalar groove weakly

scrobiculate; sternaulus absent; propodeum strigate-rugose dorsally, the strigae

emanating from median raised lamella, propodeum smooth laterally. Abdomen:

first tergum about 2.5 times as long as wide at apex, strigate-rugose; rest of terga

smooth and polished; ovipositor as long as abdomen and thorax combined.

Male. — Essentially as in female except for sexual differences.

Types.— Holotype 2: BOLIVIA: Cavinas, Rio Beni, Wm. M. Mann, Feb. 1922,

deposited in the National Museum of Natural History (USNM). Paratypes: 1 2,

same data as holotype; 1 2, 2 66, BRASIL: Rio Caraguala, 21°48' B., 52°27' L.,

Fritz Plaumann, April 5-7, 1953, deposited in the USNM and the Canadian

National Collection, Ottawa (CNC).

1 All terms for sculpturing are based on Harris, 1979.

VOLUME 59, NUMBERS 1-4

151

This species differs from Bohartiellusplaumanni, new species, by the large horn¬

like anterior corners of the mesonotum, the shorter malar space which is Vi the

eye height, the swollen scutellum, and the smooth mesopleuron.

The specific name is from the Latin cornutus which means horned in reference

to the horn-like projections on the anterior corners of the mesonotum.

Bohartiellus plaumanni Marsh, New Species

Female. — Length of body, 2.5 mm; ovipositor, 2.0 mm. Color: entire body and

legs brown except basal antennal segments and all tarsi which are honey yellow;

forewing banded as in cornutus. Head: vertex and frons strigate, temples smooth,

face strigate-rugose with median raised smooth area; malar space about x h eye

height; ocell-ocular distance about 3 times diameter of lateral ocellus; antenna

17-segmented. Thorax: mesonotal lobes coriaceous, anterior corners of meso¬

notum without large horn-like projections as in cornutus ; notauli distinct and

strongly scrobiculate; scutellar furrow with 6-8 cross carinae; scutellum flat, smooth;

mesopleuron strigate, subalar groove weakly scrobiculate; sternaulus absent; pro-

podeum strigate-rugose dorsally, the strigae emanating from the median lamella,

propodeum smooth laterally. Abdomen: first tergum about 1.5 times as long as

wide at apex, strigate-rugose; rest of terga smooth and polished; ovipositor as long

as abdomen and thorax combined.

Male. — Essentially as in female except for sexual differences.

Types.— Holotype 2: BRASIL: Rondon, 24°38' B., 54°0 1' L., Fritz Plaumann,

VIII-1952, deposited in the CNC. Paratypes: 1 2, 2 $6, same data as holotype,

deposited in USNM and CNC.

This species differs from Bohartiellus cornutus by the reduced anterior corners

of the mesonotum, the longer malar space which is Vi the eye height, the flat

scutellum, and the strigate mesopleuron.

This species is named after its collector Fritz Plaumann.

Literature Cited

Harris, R. A. 1979. A glossary of surface sculpturing. Calif. Dept. Food Agric., Ent. Occas. Pap. No.

28, 31 pp.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 152-162

Biology of a New Try poxy Ion that Utilizes Nests of

Microstigmus in Costa Rica

(Hymenoptera: Sphecidae)

Robert W. Matthews

Department of Entomology, University of Georgia, Athens, Georgia 30602.

In the Neotropics, spider-hunting wasps of the genus Trypoxylon are extraor¬

dinarily diverse (Bohart and Menke, 1976). The subject of this report was first

noted in 1968 (Matthews, 1968:37), but no adults were collected until 1980.

Professor O. W. Richards and Dr. Arnold S. Menke kindly examined the wasp,

declared it to be a new species, and very graciously offered to describe it. My

notes on nesting biology follow the species description.

Trypoxylon latro Menke & Richards, New Species

(Figs. 1-13)

Holotype male. — Color: Black. Scape, pedicel pale brown above, whitish yellow

below; clypeal margin, labrum and mandible pale brown; palpi, pronotal lobe,

precoxal lobes, coxae apically, trochanters and foretarsus pale yellow; forefemur

brownish above, brownish yellow below; midfemur brownish; fore and midtibiae

brownish yellow; midtibial spur and basal half of tarsomere I pale yellow, rest of

midtarsus brownish; basal third of hindtibia and spurs pale yellow; hindtarsus

brownish.

Vestiture: Clypeus and lower frons, including eye emarginations, with dense

appressed silver setae; gena, pronotal collar, scutum laterally, metanotum laterally,

mesopleuron, propodeal dorsum and hindface with sparser, suberect silver setae;

rest of body covered with fine, pale setae except metapleuron and propodeal side

glabrous.

Structure: Flagellomere VIII abruptly expanded apically in profile, IX-XI of

same width (Fig. 13); flagellomeres III-IV and VII-VIII with narrow polished tyli

(those on III-IV very difficult to detect even at 75X); flagellomere IV flattened

and with small apical tooth-like process; comparative lengths of flagellomeres I-

II and XI: 7:5.5:17; greatest length of flagellomere XI slightly more than combined

lengths of VIII-X; frontal carina evanescent, represented by short, polished fine

above transverse supra-antennal carina; frons minutely granulate, dull, shallowly

punctate, punctures separated by about two diameters; least interocular distance

at vertex nearly twice that at clypeus (21:12); ratio of ocellocular distance to

hindocellus diameter to distance between hindocelli: 2:6:4.5; clypeal free margin

as in Fig. 2, edge thickened at middle third as seen from below; labrum consisting

of two narrow, fingerlike lobes; occipital carina a complete circle, separated from

hypostomal carina by slightly less than a hindocellus diameter (4.5:6); scutal and

scutellar sculpture and punctation similar to frons but weakly shining; propodeal

dorsum with pair of shallow, arcuate furrows which are evanescent posterad,

resulting enclosure with broad, shallow, median longitudinal depression which is

cross carinulate, carinae strongest and converging at base of enclosure; propodeal

VOLUME 59, NUMBERS 1-4

153

side highly polished, impunctate except for zone of dense setae posteriorly above

hindcoxa, side delimited dorsad by arcuate shelf-like ridge, dorsal rim of petiole

socket without a backward projecting lamella; posterolateral corner of propleuron

with roughly egg-shaped platelike area which is delimited inwardly by a carina;

mesopleuron finely, shallowly punctate, depression around scrobe about equal to

hindocellus diameter; metapleuron highly polished, impunctate except above

hindcoxa; metapleural flange lamellate, its margin translucent, not down curved;

intercoxal carina slightly arcuate; dorsal carina of hindcoxa not reaching coxal

apex; gaster claviform, segment I gradually expanding toward II, I not quite twice

as long as wide (46:28) in dorsal view; sternum VIII as in Fig. 8; genitalia as in

Fig. 11, aedeagal shaft arcuate in lateral profile, penis valve head with sharp lateral

process.

Length: 6 mm.

Female. — Color: As in male except scape yellow brown, pedicel brown above,

yellow brown beneath; foreleg whitish yellow except upper surface of femur with

brownish stripe and coxa yellowish brown; midfemur brownish but with pale

yellow area at base; midtibia pale yellow behind, basal fourth of gastral segments

II—III ringed by pale yellow.

Vestiture: Same as male.

Structure: As in male except: flagellum simple, comparative lengths of flagello-

meres I—II and X: 13.5:11:15.5; flagellomere X length equal to combined lengths

of VIII-IX; least interocular distance at vertex more than twice that at clypeus:

23.5:10.5; ratio of ocellocular distance to hindocellus diameter to distance between

hindocelli: 1.5:7.5; clypeal margin less sinuate than male (Fig. 5); inner margin

of mandible simple as in male (Fig. 5); length of tergum I a little less than twice

width (range: 58:35-55:32).

Length: 6-6.5 mm.

Discussion. — The clavate gaster of T. latro is suggestive of the monotypic genus

Pisoxylon Menke (1968), and both have a bilobed labrum and somewhat similar

male antennae. But Pisoxylon has a much more compact gaster, the occipital

carina runs into the hypostomal carina, the propodeal side curves to the dorsum

without interruption, the metapleural flange is not lamellate, the hindcoxa has

only a suggestion of a dorsal carina, sternum VIII is roundly acuminate, the

gonostyle is not bilobate and the penis valve lacks spines or other processes. In

T. latro the occipital carina is a complete circle which is separated from the

hypostomal carina, the propodeal side is highly polished, glabrous, and sharply

delimited dorsally by a shelf-like ridge, the metapleural flange is broadly lamellate,

the hindcoxal carina is strongly indicated, male sternum VIII is truncate apically

and bispinose, the gonostyle is bilobed and the penis valve is adorned with spinose

processes. These characters negate a close relationship with P. xanthosoma, and

they indicate that T. latro belongs in the fabricator group (see Bohart and Menke,

1976:344, and Richards, 1934:280) of Trypoxylon. In fact, latro is very similar

to the North American species T. johnsoni Fox and T. clarkei Krombein as

demonstrated by the labrum, the male antenna, sternum VIII, the male genitalia

and the general habitus.

T. latro differs from johnsoni in the following ways: In both species the pro¬

podeal side just above the hindcoxa is densely covered by setae which rather

sharply delimit this zone from the remainder of the side. In johnsoni a ridge

VOLUME 59, NUMBERS 1-4

155

extends dorsad from the hindcoxal articulation partially delimiting this setal zone.

This ridge is usually paralleled anterad by several shorter ridges. The least inter¬

ocular distance is comparatively shorter in females of latro. When the interocular

distance at the vertex is compared to that at the clypeus and the length of fla-

gellomere I the resulting ratios are obtained: latro : 23.5:10.5:13.5, johnsoni: 26:

18:16. The female clypeus of latro (Fig. 5) lacks the two prominent, rounded lobes

(Fig. 4) found in johnsoni. The inner margin of the female mandible is simple in

latro (Fig. 5), but there is a weak tooth in johnsoni (Fig. 4). In both sexes of latro

the hindocelli are separated by less than an ocellus diameter (5:7.5), while in

johnsoni they are separated by an ocellus diameter or more (7:7). The male

antennae are quite similar in the two species, but flagellomere VIII of johnsoni

lacks a tyloid. Furthermore, flagellomere IV in johnsoni has a semicircular depres¬

sion which accentuates the apical toothlike process. Tergum I in latro is about

twice as long as wide or slightly less in the female, while in johnsoni it is almost

three times as long as wide. In the male of latro I is less than twice as long as

wide, while in johnsoni it is slightly more than twice as long as wide. The male

genitalia are very different (compare Figs. 10, 11). In latro the gonostyle ends in

two elongate, broad flat lobes which overlay each other dorsoventrally (Fig. 11).

In johnsoni the gonostyle ends in two narrow lobes which lie side by side, the

outermost being narrower and attenuate apically (Fig. 10).

T. latro is immediately separable from clarkei by the complete occipital carina.

In clarkei the carina is incomplete below, although the gap between the ends is

narrow in some males. The propodeal side of clarkei is less densely setose pos¬

teriorly above the hindcoxa than in latro, and the shelf-like dorsal ridge is weaker,

but these are features that require specimens of both species to appreciate. The

intercoxal carina is straight in clarkei, arcuate in latro. In the female of clarkei

the least interocular distance at the vertex is about twice that at the clypeus, the

distance at the latter equalling the length of flagellomere I (29:14:14). In latro the

ratios are 23.5:10.5:13.5. The female clypeus of clarkei is produced into a truncate

156

PAN-PACIFIC ENTOMOLOGIST

lobe, and there is a weak tooth on the inner margin of the mandible (Fig. 6). In

both sexes of clarkei the ocellocular distance is equal to half (female) or two-thirds

(male) a hindocellus diameter. In both sexes of latro the ocellocular distance is

less than half a hindocellus diameter. As in latro, the apical lobes of the gonostyle

of clarkei are broad and overlay each other, but ventrally there is a dense “basket”

of long curved setae at their base (Fig. 12, also illustrated by Sandhouse, 1940:

Fig. 48 —misidentified as johnsoni). The gonostyle of latro lacks the basal “basket”

and is sparsely setose in general (Fig. 11).

Etymology’. — Latro is a Latin noun meaning robber which refers to the habit

of using nests of Microstigmus for its own.

Types.— Holotype 6: COSTA RICA, Puntarenas Prov., Corcovado Natl. Park,

Sirena Station, nest 71, March 8, 1981, R. W. Matthews (USNM type #100374).

Paratypes: 4 <3, 10 $, same place as type, various dates in February and March

1981, and Oct. 1980, all R. W. Matthews or Matthews and C. K. Starr, deposited

in U.S. National Museum, British Museum (Natural Ftistory), Univ. of Georgia,

Univ. of Costa Rica, and Univ. of Sao Paulo.

Biological Notes

Study method.— This study was conducted between October 1980 and Septem¬

ber 1982 at Sirena, in Corcovado National Park on the Osa Peninsula, Guanacaste

Province, Costa Rica. In the primary forests along the Ollas Trail, the eusocial

sphecid wasp Microstigmus comes nests abundantly on the palm Crysophila gua-

gara. Monthly censuses of all M. comes nests in the study site were taken, so that

complete records of individual nest histories were obtained for over 700 nests,

many of which were appropriated by T. latro (Fig. 14). Trypoxylon latro nests

were collected during 4 extended visits to the study site. Dissections of these nests,

rearings, and incidental observations produced the information reported here.

Nest architecture. —Trypoxylon latro utilizes only the empty upper portion or

“attic” of the M. comes nest. There appears to be no modification of existing M.

comes cells or the nest entrance other than the application of irregular splotches

of mud to the floor and sides of the nest interior. I could find no trace of old M.

comes cells or their contents. Petioles of nests appropriated by Trypoxylon often

lose their distinctive tight coil and become “wavy” (Fig. 14), a useful field cue.

Presumably this loss of coil tightness reflects a lack of the maintenance normally

given by Microstigmus adults, rather than modification by T. latro females.

Dissections of 86 nests from 4 collections are summarized in Table 1. Nests

contained an average of 1.3 cells (range 1-3). There was a marked seasonal dif¬

ference in the average nest size. Nests constructed in the wet season (August-

October) averaged 1.19 cells (n = 57), whereas those constructed in the dry season

(February-March) were larger, averaging 1.62 cells (n = 21). This difference is

highly significant (Mann-Whitney Test, z = 3.56, P < 0.0002). Similarly, the

number of nests with 2 cells was significantly greater in the dry season (x 2 — 9.64,

P < 0.01). When 2 cells were constructed, a weak mud partition of about 0.5 mm

thickness separated the lower (first-made) cell from the upper one. Due to the

pear-like shape of an M. comes nest, the upper cell was invariably smaller in size

and volume. In the single nest with 3 cells, the lower space was partitioned in

half. Empty “vestibule” cells between the upper cell and the sealed nest entrance

VOLUME 59, NUMBERS 1-4

157

Fig. 14. A Trypoxylon latro female carrying prey into the entrance of a Microstigmus comes nest.

Constructed of the waxy pubescence found on the underside of Crysophila guagara palms and bound

together with silk, the nest now lacks the strong petiole coil characteristic of an active Microstigmus

colony (cf. Fig. 4 in Matthews, 1968). The petiole is about 14 mm long.

occurred in 7% of all nests, all from the wet season. Four nests had a final closure

but were entirely empty.

Life history.— The egg, about 1.2 mm long and 0.4 mm wide, was attached

obliquely to the anterior lateral surface of the spider’s abdomen. Two eggs ob¬

served in the laboratory hatched within 3 days of collection.

Larval growth was rapid. In the single case observed, a newly hatched larva

completed its full growth and constructed a cocoon in 4 days. Such rapid devel-

158

PAN-PACIFIC ENTOMOLOGIST

Table 1. Contents of 86 Trypoxylon latro nests from 4 collections (1 dry season, 3 wet seasons)

taken at Sirena, Corcovado National Park, Costa Rica.

Nests without cells

Nests with cells

Nests

Mud-

Empty

but

with

Total

cells/nest

Vesti¬

bule

Provisioning

Coccoon

Collection date

sample

lined

only

final

closure

cell

present

Incom¬

plete

Scav¬

enged

Egg

Un¬

emerged

Emerged

October 1980

Feb.-March

1981

September

1981

August 1982

10 1

1 Three individuals of an unidentified chloropid fly were reared from 2 nests.

opment may explain the absence of nests with larvae among those summarized

in Table 1.

Rounded at both ends, cocoons were attached basally to a bit of the mud in

the cell bottom and stood upright or obliquely in the cell (Fig. 15A). Dull gray

and of a grainy and brittle texture, cocoons measured an average of 6.8 mm long

(range 5.3-7.5, n = 20) and 2.1 mm wide (range 1.8-2.3, n = 20). Four female

cocoons averaged 7.0 mm long (range 6.3-7.5) and a single male cocoon was 6.5

mm long. Overall, cocoons from nests with 2 cells averaged slightly smaller than

those from single-celled nests, for those from the upper cells were all at the small

end of the size range.

Duration of the pupal stage could not be precisely documented, but eclosions

from collected nests continued over a period of at least 2 months. No diapause

or quiescent stage was detected.

Overall the sex ratio of the reared adults was 18 females to 8 males. Flowever,

emergences from the 14 single-celled nests were strongly female-biased (11 fe¬

males : 3 males). Five of the 6 two-celled nests from which both adults emerged

yielded one of each sex. Presumably the males came from the smaller cocoons of

the upper cells. Upon emergence, adults characteristically chewed directly through

the side wall of the nest bag rather than through the mud closure plug.

Prey.—Cells were mass provisioned with spiders (Fig. 15B). A sample of 133

spiders from 8 cells was preserved. The number of prey per cell averaged 16.6

(range 9-28). Dr. H. W. Levi of the Museum of Comparative Zoology, Harvard

University, identified them as Leucauge and Dolichognatha in the Tetragnathidae.

Leucauge was uncommon in the sample, with only 6 juveniles represented. How¬

ever, at least 2 species of Dolichognatha were represented. Nearly half (48%) of

the individuals were juveniles, 13.7% were males, and 38.3% were females. Ac¬

cording to Dr. Levi, Dolichognatha constructs small orb webs at the base of trees.

Neotropical tetragnathids are largely unstudied, but recent revisions of Nearctic

genera are available (Levi, 1980, 1981).

Natural enemies. — As shown in Table 1, nest cells attacked by scavenger fly

larvae were found in all 3 wet season collections but not in the dry season col¬

lection. In the wet season 12.5-32.1% of total cells were infested. In August 1982,

3 unidentified chloropid fly adults were successfully reared from scavenged prey

VOLUME 59, NUMBERS 1-4

159

Figs. 15 A, B. A, cocoon of T. latro attached by its posterior end to a spot of mud in the bottom

of the nest. Cocoon is 7.0 mm long. B, Dolichognatha spider prey and small larva of T. latro in an

M. comes nest.

in 2 nests. An unidentified ichneumonid wasp was reported as emerging from a

cocoon in a mud-plugged nest in 1968 from a different locality (Matthews, 1968),

but none were taken in the present study.

Adult behavior. — On 7 March 1981, a female was observed carrying prey into

160

PAN-PACIFIC ENTOMOLOGIST

an M. comes nest in which, 6 days earlier, a single M. comes female had been

present. The nest was less than 4 weeks old according to census records. Whether

T. latro usurped the nest or took over after it had become abandoned is unde¬

termined, but the evidence suggests that the nest may have been usurped.

Provisioning was first noted at 1110 AM CST when the female T. latro was

seen to enter with prey. During the next 116 minutes she brought in 10 additional

spiders. Time away from the nest ranged 3-42 minutes (n = 10), whereas the time

inside the nest to store the prey was always brief, ranging 5-18 seconds. After a

lapse of 81 minutes I returned to the nest at 1645 PM. The female returned with

a spider at 1647, and remained inside for 49 seconds before departing. Presumably

oviposition occurred at this time, for 2 minutes later she returned carrying mud,

then made 5 more trips for mud in rapid succession, each requiring less than 1

minute. All of this mud was applied inside the nest, with the wasp spending 46-

70 seconds each time at this task. During these periods of mud application I was

unable to detect any audible buzz like that produced by the subgenus Trypargilum

during mud application. Presumably the wasp was closing olf the lower cell at

this time.

Impact on host population.— At my study site, monthly census of the status of

all M. comes nests from February, 1981 to May, 1982 revealed that T. latro

utilized a relatively small but more or less constant proportion of about 5% of

the potentially available nests. The average number that became Try poxy Ion nests

each month was 9.8 (range 5-21) of an average of 187 active M. comes nests.

Two other Sirena samples of nests from outside of the census area gave com¬

parable rates of T. latro nest utilization. The first involved 64 M. comes nests

collected at night on 8 March 1981 (dry season). Five (7.8%) proved to be occupied

by Trypoxylon. A second night-collected sample, made on 4 September 1982 (wet

season), involved 65 nests, of which 6 (9.2%) were Trypoxylon. Of these, 4 were

in nests known to be constructed by M. comes more than 3 months earlier, and

2 were in nests constructed within the past 3 months. According to the nest census

records from the study site, nests of all ages are vulnerable to appropriation by

T. latro. No preference for nests of a particular age could be detected. Overall,

activity of T. latro accounted for 24% of the total M. comes mortality in the study

site.

Discussion. —Trypoxylon is a huge genus with over 450 species and a great

diversity of behavior. It includes two subgenera, Trypoxylon and Trypargilum.

The revision of the American Trypoxylon by Richards (1934) remains the defin¬

itive study of the genus in the New World. Coville’s (1982) revision of the subgenus

Trypargilum in North America is a recent contribution to the taxonomy of that

group. The widespread use of preformed cavities for nesting, including the aban¬

doned cells of other wasps, has allowed researchers to use the trap-nest technique

(Coville and Coville, 1980; Coville, 1981; Coville and Griswold, in prep.) to

expand our knowledge of Neotropical Trypargilum. Species of the subgenus Try¬

poxylon are less well known, and probably many undescribed species exist. Many

also take trap-nests and at least 2 species, T. johnsoni and T. clarkei, character¬

istically nest in preexisting cavities (Krombein, 1967). However, members of the

fabricator group, to which T. latro, johnsoni, and clarkei belong, typically con¬

struct free mud nests.

The use of Microstigmus nests by T. latro is unique. Building a mud nest within

VOLUME 59, NUMBERS 1-4

161

the small, flexible silk-lined nest sacs of these primitively social wasps would seem

rather analogous to building a house inside a laundry bag. Moreover, abandoned

nest sacs are rather delicate and rarely persist for long before the petiole breaks

or becomes detached from the plant leaf.

In the Neotropics, insects, particularly ants, compete intensely for cavities suit¬

able for nesting. Ants have exploited a great variety of seemingly unlikely cavities,

from hollow thorns to galls. Virtually every hollow twig that one breaks open is

occupied by ants. The vast majority of trap-nests that I have placed in the field

in various localities in Costa Rica have been quickly exploited by ants. However,

of over 700 Microstigmus nests monitored over a 16 month period, only one was

found to be occupied by an ant colony (a species of Tapinoma). Thus, Microstig¬

mus nest bags are apparently either being overlooked by searching ants or are in

some manner unacceptable to them. By exploiting this unusual habitat, T. latro

minimizes competition from ants. Furthermore, since no parasites were reared

from any nests in this study, T. latro also appears to be avoiding certain other

sources of mortality, such as chrysidid wasps, that regularly parasitize related

species nesting in more “typical” habitats.

Yet despite the abundance of potential nest sites, T. latro is a relatively rare

species. Due to the small size of the Microstigmus nest bag, nests rarely contain

more than 2 cells. While T. latro nests continuously throughout the year, it clearly

is more successful during the dry season. Dry season nests contained significantly

more cells and suffered no losses to scavenger flies. These differences probably

reflect differences in prey abundance, humidity, and available time for nesting.

Trypoxylon latro is unusual in that its brittle grainy cocoons are unlike “typical”

Trypoxylon cocoons, which are papery and tan-colored. However, at least one

other Trypoxylon species, T. johnsoni, constructs a similar cocoon and also nests

in preformed cavities (Krombein, 1967). As Menke and Richards note in their

description, T. latro is closely related to T. johnsoni morphologically. Dr. Krom¬

bein kindly loaned me cocoons of T. johnsoni for comparison and they are virtually

identical to those of T. latro, which is additional evidence for the close relationship

of these 2 species.

Acknowledgments

It is a pleasure to dedicate this paper to Dr. Richard M. Bohart, whose con¬

tributions to our knowledge of wasps have been truly monumental. I thank Dr.

Rollin E. Coville for initially determining that T. latro was undescribed and Prof.

O. W. Richards and Dr. Arnold Menke for describing it. Dr. K. V. Krombein

kindly loaned cocoons of T. johnsoni for comparison. Dr. H. W. Levi determined

the spider prey; voucher specimens are deposited in the Museum of Comparative

Zoology, Harvard University. Dr. Christopher K. Starr, now at Visayas State

Agricultural College in Baybay, Leyte, Philippines, assisted in data collection. I

thank Servicio de Parques Nacionales, San Jose, Costa Rica for permission to

work in Corcovado Park. This work was suported by NSF grant BMS7925708.

Literature Cited

Bohart, R. M., and A. S. Menke. 1976. Sphecid wasps of the world. Univ. Calif. Press, Berkeley,

ix + 695 pp.

Coville, R. E. 1981. Biological observations on three Trypoxylon wasps in the subgenus Trypargilum

162

PAN-PACIFIC ENTOMOLOGIST

from Costa Rica: T. nitidum schulthessi, T. saussurei, and T. lactitarse (Hymenoptera: Spheci-

dae). Pan-Pac. Ent. 57:332-340.

-. 1982. Wasps of the genus Trypoxylon subgenus Trypargilum in North America (Hymenop¬

tera: Sphecidae). Univ. Calif. Pub. Ent. 97:1-147.

Coville, R. E., and P. L. Coville. 1980. Nesting biology and male behavior of Trypoxylon ( Trypar¬

gilum) tenoctitlan in Costa Rica (Hymenoptera: Sphecidae). Ann. Ent. Soc. Amer. 73:110-119.

Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests and associates. Smithsonian

Press, Washington, D.C., 570 pp.

Levi, H. W. 1980, The orb-weaver genus Mecynogea, the subfamily Metinae and the genera Pachy-

gnatha, Glenognatha and Azilia of the subfamily Tetragnathinae North of Mexico (Araneae:

Araneidae). Bull. Mus. Comp. Zool. 149:1-74.

-. 1981. The American orb-weaver genera Dolichognatha. and Tetragnatha north of Mexico

(Araneae: Araneidae, Tetragnathinae). Bull. Mus. Comp. Zool. 149:271-318.

Matthews, R. W. 1968. Nesting biology of the social wasp Microstigmus comes (Hymenoptera:

Sphecidae, Pemphredoninae). Psyche 75:23-45.

Menke, A. S. 1968. New genera and species of wasps of the tribe Trypoxylonini from the Neotropical

region (Hymenoptera: Sphecidae, Larrinae). Contrib. Sci. Los Angeles Co. Mus. (135): 1-9.

Richards, O. W. 1934. The American species of Trypoxylon. Trans. R. Ent. Soc. Lond. 82:173-362.

Sandhouse, G. A. 1940. A review of the Nearctic wasps of the genus Trypoxylon (Hymenoptera:

Sphecidae). Amer. Mid. Nat. 24:133-176.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 163-175

A Review of the Genus Polemistus in the New World

(Hymenoptera: Sphecidae)

Arnold S. Menke and David L. Vincent

(ASM) Systematic Entomology Laboratory, IIBIII, ARS, USDA, % U.S. Na¬

tional Museum, Washington, D.C. 20560; (DLV) Beneficial Insect Introduction

Laboratory, IIBIII, ARS, USDA, Beltsville, Maryland 20705.

Dick Bohart has had a life long love of wasps. While most of us choose other

pursuits in our free time, Dick’s devotion to his wasps is so intense that they have

assumed the status of a hobby with him—he works on them day and night! Few

people achieve the broad knowledge and experience in different families of insects

possessed by Dick. His work has spanned many aculeate families, and his influence

will be felt for many generations to come. The Sphecidae has been one of his

favorite groups, and he has published more comprehensive works on this family

than any other. We take great pleasure in dedicating our short treatise on Pole¬

mistus to him.

The cosmopolitan genus Polemistus contains 19 described species (Bohart and

Menke, 1976). Two of these are poorly known Neotropical species, one of which,

the Mexican pusillus, was incorrectly reported by Bohart and Menke to occur in

the southwestern United States. Our studies of approximately 100 specimens of

Polemistus from the New World indicate that three species occur in the United

States, all of which are undescribed. The three North American species are newly

described here, as is a fourth new species discovered in Central America. A key

to the six known New World species is provided. Each of us has assumed sole

authorship of two of the new species.

Material for this study was borrowed from the following institutions:

Academy of Natural Sciences, Philadelphia (ANSP).

American Entomological Institute, Ann Arbor.

American Museum of Natural History, New York (AMNH).

British Museum (Natural History), London (BMNH).

California Academy of Sciences, San Francisco (CAS).

Cornell University, Ithaca (CUI).

Museum d’Histoire Naturelle, Geneva (Geneva).

National Museum of Natural History, Washington DC (USNM).

Naturhistorisches Museum, Vienna (Vienna).

University of Arizona, Tucson (UA).

University of California, Davis (UCD).

The holotype of P. yoda was borrowed from the Mississippi Entomological

Museum, Mississippi State University. Richard L. Brown, director, has graciously

allowed us to deposit the type in the USNM on indefinite loan. We would also

like to thank Norita Chaney, Mike Schautf and Gene Taylor for taking the scanning

electron photographs used here.

The New World species of Polemistus represent at least two, and possibly three,

164

PAN-PACIFIC ENTOMOLOGIST

species groups. In the pusillus group {pusillus, dickboharti, yoda ) the clypeus is

densely, evenly covered by appressed silver setae; the clypeal outline, especially

in the female, is similar among the species, and there is a narrow malar space in

both sexes (Figs. 8-10, 14, 15); a hypersternaulus is usually present (sometimes

very short in yoda) (Figs. 5, 6); and terga I—II lack a lateral carina, or it is weak.

Polemistus stieglmayri appears to belong in the pusillus group based on clypeal

characters, the presence of a hypersternaulus and the setation of the male abdom¬

inal venter, but the frontal swellings (Fig. 23), and the strong lateral carina on

terga I—II set the species apart. The remaining species, chewbacca and vaderi, are

closely related to the Palearctic species abnormis (Kohl). The abnormis group is

characterized by having a sparsely or non-silvered female clypeus; a large, deep

clypeal emargination bounded by large, blunt teeth (Figs. 12, 13) in the female;

no malar space in the female and at most a very narrow one in the male; a strong

lateral carina on terga I—II; and by the absence of a hypersternaulus (Fig. 7).

In contrast to the ornately sculptured head and thorax of many Old World

species, the New World members of Polemistus are rather plain, and species

characters are few. In the male the presence or absence of antennal tyli and their

form, when present, are diagnostic. The presence or absence of long sternal setae

on the male abdominal venter is useful. The outline of the free margin of the

clypeus can be distinctive in both sexes but unfortunately the mandibles usually

have to be spread or the marginal clypeal setae scraped away in order to view

this structure. The shape of the labrum is sometimes distinctive. On the scutum

the form of the notaulus pits, and the presence or absence of ridges or foveae

adjacent to the scutellum are important. Species group characters are used in

conjunction with these. Body sculpture and punctation differ little in New World

species, and except in stieglmayri, we found no apparent differences on the ab¬

domen. Male genitalia were not examined. The known New World species have

a slender spine in the scapal basin just above the antennal sockets. Generally in

New World species the integument surface has an imbricate microsculpture (sensu

Harris, 1979) as seen at 75X, and the epistemal sulcus, omaulus, hypersternaulus,

and notaulus are pitted or foveate (the notaulus sometimes obscurely so). In the

New World species the sculpture of the propodeal dorsum is coarse and varies

from areolate to rugose to rugulose (Harris, 1979). The sculpture changes to finer,

parallel, oblique ridging on the propodeal side.

Key to New World Species of Polemistus

Note: Antennal characters are best seen by using high magnification (75X) and

good lighting. Male is unknown for yoda.

1. Male: Antenna with 13 articles, gaster with 7 visible segments . 2

- Female: Antenna with 12 articles, gaster with 6 visible segments . 6

2. Flagellomeres I-VII without projections or ridges ventrally . 3

- Flagellomeres II-IV with projections or ridges ventrally (Figs. 2-4) ... 4

3. Frons without pair of obliquely transverse swellings at upper margin of

scapal basin; pronotal collar without sharp transverse carina; hindmar-

gin of gastral sterna IV-V each with pair of long, separated setae; sw.

U.S., nw. Mexico . dickboharti Menke

- Frons with pair of obliquely transverse swellings at upper margin of

scapal basin (Fig. 23); pronotal collar with sharp transverse carina; hind-

VOLUME 59, NUMBERS 1-4

165

1 stieglmayri

4 chewbacca

Figs. 1-4. Polemistus species. 1, Left profile of female abdominal petiole and first tergum. 2-4,

Profile of male flagellomeres I-VI.

margin of gastral sterna IV-V each with transverse row of long, pale

setae; Brasil . stieglmayri (Kohl)

4. Hypersternaulus present (Fig. 5); flagellomeres II-V (or VI) with rounded

projections apicoventrally (Fig. 2); hindmargin of gastral sterna IV-V

each with transverse row of long, pale setae; central Mexico.

. pusillus Saussure

- Hypersternaulus absent (Fig. 7); flagellomeres I-V or II-VI with low

ridges ventrally (Figs. 3, 4); sterna IV-V without transverse rows of

setae; sw. U.S

166

PAN-PACIFIC ENTOMOLOGIST

Figs. 5-7. Left mesopleuron of Polemistus species, female. 5, pusillus. 6, dickboharti. 7, vaderi.

5. Flagellomere I with long ridge, II-IV or V with progressively shorter

ridges (Fig. 3); clypeal outline as in Fig. 17 . vaderi Vincent

- Flagellomere I at most with weak apical process, II-VI with long, prom¬

inent ridges (Fig. 4); clypeal outline as in Figs. 18, 19 . . chewbacca Menke

6. Hypersternaulus absent (Fig. 7); clypeus arcuately uni- or bi-emarginate

(Figs. 12, 13), its surface with few silver setae only near apex . 7

- Hypersternaulus present as row of large pits (Fig. 6) (if weak, following

clypeal characters diagnostic); clypeus sinuately emarginate (Figs. 8-11),

its surface uniformly, moderately covered by appressed silver setae ... 8

7. Labrum triangular apically (Fig. 12); clypeal outline as in Fig. 12; Utah,

Ariz., New Mexico . vaderi Vincent

- Labrum truncate apically (Fig. 13); clypeal outline as in Fig. 13; se. Ariz.

. chewbacca Menke

8. Notaulus ending posterad in large circular depression whose diameter

equals that of hindocellus (Fig. 20), depression usually sharply rimmed

cephalad when viewed from rear; central Mexico . pusillus Saussure

- Notaulus not ending posterad in large circular depression, or if so, it is

vague and no more than % diameter of hindocellus (Figs. 21, 22) .... 9

9. Frons with pair of obliquely transverse swellings at upper margin of

scapal basin (Fig. 23); middle lobe of clypeus exceeded by lateral teeth

(Fig. 11); Brasil . stieglmayri (Kohl)

- Frons without swellings at upper margin of scapal basin; middle lobe

of clypeus on line drawn between lateral teeth (Figs. 9, 10) . 10

10. Scutal surface next to scutellum with short, longitudinal or oblique ru-

gulae (Fig. 21); hypersternaulus often extending posterad beyond level

of scrobe, pits of hypersternaulus well defined (Fig. 6); sw. U.S., nw.

Mexico . dickboharti Menke

- Scutal surface without rugulae posteriorly; hypersternaulus not extend¬

ing beyond level of scrobe, often shorter, pits of hypersternaulus be¬

coming indistinct posterad; El Salvador, Honduras, Nicaragua .

. yoda Vincent

Polemistus pusillus Saussure

(Figs. 2, 5, 8, 14, 20)

Polemistus pusillus Saussure, 1892:567. Holotype male, “Mexico calida (Cor-

VOLUME 59, NUMBERS 1-4

167

8 pusillus 9 dickboharti 10 yoda

11 stieglmayri 12 vaderi 13 chewbacca

14 pusillus 15 dickboharti 16 stieglmayri

17 vaderi 18 chewbacca 19 chewbacca

Figs. 8-19. Clypeal and labral outlines of Polemistus species. 8-13, Females. 14-19, Males (19 is

holotype).

doba)” (Geneva). Bohart and Menke, 1976:185 (listed). Passaloecus pusillus,

Rau, 1943:648 (biology).

Diagnosis .—The sawtooth-like projections of the tyli on male flagellomeres II-

V or VI (Fig. 2), and the large, deep last notaular pit (diameter equal to hindocellus)

in the female (Fig. 20) are diagnostic for pusillus. The anterior side of the last

notaular pit usually is sharply margined by a ridge which may extend sinuately

between both notauli. A number of parallel microridges usually precede it. The

bottom of the last pit is smooth and polished. Males also have the large notaular

pit but sometimes it is smaller than the hindocellus.

The labrum is triangular in pusillus. It and the clypeus are shown in Figs. 8,

14. The scutum is unridged posteriorly except in one female where three weak

rugulae can be seen. The hypersternaulus extends posterad to the level of the

scrobe in males, slightly beyond in females (ends about half way to mesopleural

168

PAN-PACIFIC ENTOMOLOGIST

pit. Fig. 5). Terga I and II lack a lateral carina. In the male sterna IV-V have

transverse rows of long, whitish setae posteromedially which stand out against

the short vestiture of the rest of the venter.

Saussure identified his type as a female, but it is a male and the abdomen is

now missing.

Distribution.— This species is known only from central Mexico. Records of

pusillus from the United States (Bohart and Menke, 1976:185; Krombein, 1979:

1606) are based on misidentified material of dickboharti.

Material studied.— MEXICO, Guerrero, Acapulco, 1939-40, biological note

#2480, P. Rau, 2 9, 2 6 (USNM). Jalisco, Guadalajara, July 21, McClendon, 1 <3

(ANSP). Morelos, Cuernavaca, 5500', May 22, June 18, 1959, H. E. and M. A.

Evans, 2 9 (CUI); 5 mi. e. Cuernavaca, March 29, 1962, F. D. Parker, 1 9 (UCD).

Vera Cruz, Cordoba, type 6 (Geneva).

Biolog}’. — Rau (1943) found pusillus nesting in vacant mud cells of a Trypoxylon

species. Each cell was provisioned with 6-8 aphids of the genera Aphis and Mac-

rosiphum, and the entrance to the nest was sealed with a “transparent, glasslike

substance.” Some cells were partitioned with the same material. Chrysidid clepto-

parasites of the genera Omalus and Chrysis, and a torymid parasitoid of the genus

Monodontomerus were found in some pusillus cells.

Polemistus dickboharti Menke, New Species

(Figs. 6, 9, 15, 21)

Holotype male. — Black except as follows: pale yellow: palpi, tegula; brownish

yellow: ventral surface of scape, pedicel and basal flagellomeres; fore and midlegs

except coxae and posterior surface of midfemur; hindtrochanter, basal half of

hindtibia, hindtarsus. Tegula brown.

Clypeus and lower frons densely covered with appressed silver setae which

obscure surface. Sterna IV-V each bearing a pair of separated, long, erect setae

on posterior margin.

Scapal basin crossed by many fine, irregular ridges, the lower of which curve

downward laterally; frons above scapal basin finely punctate (75X), punctures 2-

3 diameters apart; inner orbit paralleled by crenulate carina opposite scapal basin;

upper two thirds of outer orbit with similar non-crenulate carina; flagellomeres

without tyli ventrally; free edge of clypeus with lateral tooth, weakly bilobate

mesally (Fig. 15); labrum triangular apically (Fig. 15). Scutum densely micro-

punctate (75X), punctures separated by less than puncture diameter, scutum with

longitudinal rugulae next to scutellum which are oblique laterally (similar to Fig.

21); last pit of notaulus largest, its diameter approximately equal to one half

diameter of hindocellus; hyperstemaulus extending posterad nearly to mesopleural

pit; scrobal sulcus represented by horizontal crest between scrobe and episternal

sulcus. Terga I—II without lateral carina. Length 4.5 mm.

Variation in males. — The fore and midfemora are sometimes suffused with

brown except at their apices. The tegula varies from brown to translucent yellow.

The last notaular pit varies in size. It may be no larger than the other pits, and

in one specimen it is as large as a hindocellus. The pleuron between the scrobal

sulcus and hyperstemaulus sometimes is polished and lacks microsculpture. In

VOLUME 59, NUMBERS 1-4

169

Figs. 20-23. Polemistus species. 20-22, Dorsal view of female scutum and scutullem. 20, pusillus\

21, dickboharti\ 22, stieglmayri. 23, Face of stieglmayri showing transverse swellings below ocelli.

the single male from California, the carina along the inner orbit is not crenulate,

and the scutum does not have rugulae at its posterior border.

Female .—As in male except: labrum and mandible brownish, flagellum usually

dark brown except for pale venter of flagellomeres I—II. Tegula translucent yellow.

Occasionally all legs yellow, but usually hind leg brownish except for trochanter

and base of tibia.

Clypeus and lower frons moderately, uniformly covered with appressed silver

170

PAN-PACIFIC ENTOMOLOGIST

setae, surface partially obscured; clypeal outline as in Fig. 9. Last pit of notaulus

usually about same size as others, but not exceeding two thirds diameter of

hindocellus; scutum lacks rugulae posteriorly in specimen from Aduana, Mexico;

hyperstemaulus extends posterad two thirds to three fourths distance to meso-

pleural pit (Fig. 6). Length 4-5 mm.

Discussion.— This species is similar to pusillus. Females have the same clypeal

outline, but the very large last pit of the notaulus (equal to diameter of hindocellus)

in pusillus (Fig. 20) contrasts with the smaller last pit in dickboharti (no more

than two thirds diameter of hindocellus and often much smaller, Fig. 21). Also

females of dickboharti nearly always have rugulae along the hind margin of the

scutum (Fig. 21). In most pusillus females (and those of yoda also) the scutum is

plain posteriorly. Males of dickboharti separate easily from those of pusillus on

the basis of the simple flagellomeres. The basal flagellomeres in pusillus males

have prominent, sawtooth-like tyli ventrally (Fig. 2). The lateral tooth of the male

clypeus is much larger in dickboharti (Fig. 1 5) than in pusillus (Fig. 14). Males of

dickboharti lack the transverse rows of long setae found in males of pusillus ;

instead there are two long, separated setae on sternum IV-V.

Etymology. — With great affection I dedicate this species to Dick Bohart, a dear

friend, colleague, and mentor.

Distribution.— Arizona, southern Utah, southern California, and Sonora, Mex¬

ico.

Types. — Holotype 6\ ARIZONA, Cochise Co., Portal, 5000', Sept. 5, 1959, H.

E. Evans (CUI). Fifteen <3, 22 2 paratypes as follows: ARIZONA, Cochise Co.,

same place as type, on honey dew on walnut, Sept. 2-12, 1959, H. E. Evans, 4 <3

(CUI); 1 mi. ne. Portal, on walnut, Sept. 7, 1959, J. R. Powers, 1 2 (CAS); 3 mi.

ne. Portal, on Agave, July 22, 1964, G. D. Butler, 1 2 (UCD); Southwest Research

Station, 5400', 5 mi. w. Portal, on honey dew on Populus, Aug. 3-Sept. 12, 1959,

H. E. Evans, 9 <3, 2 2 (CUI); same place, July 3-13, 1963, J. Rozen, D. Oliver, A.

Moldenke, J. Woods, 5 2 (AMNH, UCD); same place, July 11-14, 1965, V. Roth,

2 2 (AMNH, UCD); Huachuca Mtns., June 26, 1940, R. A. Rock, 1 2 (UCD);

Miller Canyon, 6000', 1 mi. w. parking area, July 16, 1964, M. Noller, 1 <3 (UCD).

Graham Co., 0.9 mi. along road to Marijilda Canyon from highway 666, 3860',

Aug. 3, 1965, H. B. Leech, 1 2 (CAS). Gila Co., 3 mi. sw. Christmas, June 5,

1962, malaise trap by Condalia, F. Werner, 1 2 (UCD). Navajo Co., Carrizo, June

21, 1957, Butler and Werner, 1 2 (UCD). Pima Co., Tucson, April 14, 1940, R.

H. Crandall, 1 2 (UGD); 27 mi. se. Tucson, May 29, 1965, M. E. Irwin, 1 2 (UCD);

Sabino Canyon, Santa Catalina Mtns., May 11, 1961, R. and E. Painter, 1 2

(UCD). Santa Cruz Co., Patagonia, on Cessia leptocarpa, July 29, 1962, M. Noller,

P. Johnson, 1 2 (UCD); Canelo, Aug. 3, 1956, G. Butler, 1 <3 (UCD). UTAH,

Garfield Co., Bryce Canyon, June 18, 1947, G. Bohart, 1 2 (UCD). Washington

Co., St. George, June 15, 1930, E. W. Davis, 1 2 (USNM). MEXICO, Sonora,

Nogales, 1 2 (BMNH).

The following nonparatypic material has been seen: ARIZONA: Cochise Co.,

Douglas, Sept. 19, 1933, W. W. Jones, 1 2 (USNM); Southwest Research Station,

5 mi. w. Portal, 5400', Sept. 16, 1963, V. Roth, 1 2 (AMNH). CALIFORNIA,

San Diego Co., Palm Canyon, Borego, May 3, 1945, A. Melander, 1 <3 (USNM).

MEXICO, Sonora, Aduana, March 15, 1962, L. Stange, 1 2 (UCD).

VOLUME 59, NUMBERS 1-4

171

Polemistus yoda Vincent, New Species

(Fig. 10)

Holotype female.— Black except as follows: yellow: scape ventrally, mandible

(apex brownish), palpi, pronotal lobe; yellow brown: labrum, ventral surface of

pedicel and flagellum (last few flagellomeres darker), foreleg (except outer surface

of tibia yellow), midleg (except femur black with extreme apex brownish), hind-

trochanter and tarsus; hindtibial base yellow. Tegula translucent yellow anteriorly,

dark brown posteriorly.

Clypeus, lower frons, propleuron posteriorly and forecoxa ventrally densely

covered by appressed silver setae which obscure surface.

Scapal basin irregularly, finely cross carinulate; frons above scapal basin finely

punctate (75X), punctures 1-2 diameters apart; inner orbit closely paralleled by

fine carina opposite scapal basin, upper two thirds of outer orbit with similar

carina; clypeal outline as in Fig. 10; labrum triangular apically (Fig. 10). Scutum

densely micropunctate (75X), punctures separated by less than puncture diameter;

notaulus pits sharply formed, uniform in width; hypersternaulus ending posterad

at level of scrobe, pits of hypersternaulus smaller, shallower than those of epi-

sternal sulcus; scrobal sulcus represented by horizontal crest between scrobe and

episternal sulcus. Terga I—II with weak lateral carina. Length 4.5 mm.

Male. — Unknown.

Variation in female.—The hypersternaulus is weakly indicated in both para-

types. It does not reach the level of the scrobe and only the first pit is large and

clearly formed.

Discussion. — Females of P. yoda are most similar to those of dickboharti, but

the ridging along the posterior margin of the scutum in the latter (Fig. 21) contrasts

with the plain scutum of the former. Polemistus yoda has weak lateral carinae on

terga I—II, but these are absent in dickboharti and pusillus. Also these last two

species lack the dense appressed silver setae found on the propleuron and forecoxa

of yoda.

Etymology. —The name yoda, a noun in apposition, is based on a character in

the movie “The Empire Strikes Back.”

Distribution. — El Salvador, Honduras, and Nicaragua.

Types. — Holotype 2: NICARAGUA, 14 km. sw. Camoapa, swept from door-

yard Gossypium hirsutum, Dec, 1, 1976, W. H. Cross (USNM). Two $ paratypes

as follows: HONDURAS, Tegucigalpa, June 12, 1918, F. J. Dyer (USNM). EL

SALVADOR, 2 and one half mi. w. Quezaltepeque, July 12, 1961, M. E. Irwin

(UCD).

Polemistus stieglmayri (Kohl)

(Figs. 1, 11, 16, 22, 23)

Passaloecus stieglmayri Kohl, 1905:359. Holotype female, “Brasilien (Rio Grande

do Sul)” (Vienna). Kohl, 1905:529 (listed). Polemistus stieglmayri, Bohart and

Menke, 1976:185 (listed).

Diagnosis.— The pair of oblique swellings on the frons above the scapal basin

are unique to stieglmayri (Fig. 23). The absence of tyli on the male flagellum and

the long clypeal teeth (Figs. 11, 16) are also distinctive. The labrum is triangular

172

PAN-PACIFIC ENTOMOLOGIST

apically. In both sexes the scutum usually has a pair of longitudinally elongate

foveae posteromedially which are narrowly separated by a septum (Fig. 22). In

some specimens these two foveae are broad, roughly triangular depressions which

fade laterally and may contain rugulae. The notaulus pits are uniform in size. The

hypersternaulus extends posterad to or just beyond the level of the scrobe. Terga

I and II have a lateral carina. Tergum I sometimes has a strong transverse subapical

impression which, when viewed in lateral profile, results in a humped outline

(Fig. 1). This impression is quite weak in two thirds of the specimens at hand.

The posterior margin of sterna IV-V in males of stieglmayri bear transverse rows

of long, pale setae, a feature shared with pusillus.

Distribution.— Known only from southern Brasil.

Material examined.— BRASIL, Rio Grande do Sul, no specific locality, Stie-

glmayr, 2 type (Vienna). Santa Catarina, Nova Teutonia, Nov.-Feb. 1964-65,

Fritz Plaumann, 8 2, 1 $ (UCD).

Polemistus vaderi Vincent, New Species

(Figs. 3, 7, 12, 17)

Holotype male.— Black except as follows: pale yellow: scape and pedicel ven-

trally, palpi, pronotal lobe, tegula, fore and midtibiae and tarsi, basal half of

hindtibia; brownish yellow: fore and midtrochanter ventrally and hindtrochanter;

closing face of fore and midfemora yellow, brown above; hindtarsi brownish.

Clypeus and lower frons densely covered by appressed silver setae which obscure

surface.

Scapal basin crossed by many fine, irregular ridges which curve downward

laterally; frons above scapal basin finely punctate (75X), punctures 2-3 diameters

apart; inner orbit closely paralleled by fine carina opposite scapal basin; upper

two thirds of outer orbit with similar carina; flagellomeres I-IV with low ridgelike

tyli ventrally, those on I—II extending the length of each flagellomere, those on

III-IV shorter, occupying apical one half and one third, respectively (similar to

Fig. 3); free edge of clypeus as in Fig. 17; labrum roundly triangular apically (Fig.

17). Scutum densely micropunctate (75X), punctures separated by less than punc¬

ture diameter; notaulus pits shallow, ill defined laterad; propodeal side with many

fine, parallel, oblique ridges which reach metapleural sulcus; scrobal sulcus rep¬

resented by horizontal crest between scrobe and episternal sulcus; hypersternaulus

absent although episternal sulcus pits enlarged at that level of pleuron. Terga I-

II with lateral carina. Length 4.5 mm.

Variation in males.— In some specimens the fore and midlegs are entirely yellow

except the coxae, and the basal flagellomeres are yellowish beneath. Tyli occur

on the first six flagellomeres in some specimens, but those on III-VI are repre¬

sented by very short apical tooth-like elevations. The tylus on fiagellomere II

sometimes is restricted to the apical two thirds or one half of the article (Fig. 3).

The notauli are sometimes weakly impressed and the pitting obscure.

Female.— As in male except: mandible apex, labrum, and fore and midtro¬

chanters brownish black; forefemur brownish black except yellow on apical one

fourth; midfemur yellow only at extreme apex. Clypeus with few appressed silver

setae near apex, surface not obscured; appressed silver setae on frons between eye

and scapal basin only partially obscure surface; scapal basin shining, without

ridges; frons between scapal basin and eye with feeble, vertical microridges; carina

VOLUME 59, NUMBERS 1-4

173

along inner orbit short, weak; outer orbit without such carina; free edge of clypeus

with large, deep, semicircular emargination containing a weak central tooth (Fig.

12); clypeal surface shining, broadly concave; labrum triangular apically. Length

4.5-6 mm.

Discussion. — The broad, deep, semicircular clypeal emargination and triangular

labrum of the female (Fig. 12), and the presence of a long tylus on the first male

flagellomere (Fig. 3) are diagnostic for vaderi . The absence of a hypersternaulus

(Fig. 7) and the absence in the female of dense silver clypeal setae relate vaderi

to chewbacca, but the latter has a truncate labrum in the female (Fig. 13), and

flagellomere I has at most a short apical tylus in the male (Fig. 4). The clypeal

outlines also differ between these species (compare Figs. 12, 13).

Etymology .—This species is named after Darth Vader, a character in the movies

“Star Wars” and “The Empire Strikes Back.”

Distribution.— Utah, Arizona and New Mexico.

Types .—Holotype <3: UTAH, Beaver Co., Beaver Mountain, 6000-7000', rear¬

ing code BM 6-B, Aug. 1972, D. Vincent (USNM). Seven <3 and 3 2 paratypes as

follows: ARIZONA, Cochise Co., Southwest Research Station, 5 mi. w. Portal,

5400', July 8, 1963, J. Rozen, D. Oliver, A. Moldenke & J. Woods, 1 2 (AMNH);

same location, Aug. 29, 1959, H. E. Evans, 1 8 (CUI); same location, Sept. 13,

1959, on honey dew on Populus, H. E. Evans, 1 <3 (CUI). NEW MEXICO, Lincoln

Co., Capitan Mountains, May 26, 1907, ex galleries of Dendroctonus convexifrons,

Hopkins #5470b, J. L. Webb, 1 <3 (USNM); San Miguel Co., 5 mi. nw. Las Vegas,

reared from galls of Andricus ruginosus Bassett, April 29, 1918, Hopkins #15602,

L. H. Weld, 3 8 (USNM). UTAH, same data as holotype, rearing codes BM 6-C,

BM 5-B, BM 3-B, 1 <3, 2 2 (USNM).

Biology ?.—Four trap nests from the holotype locality were examined. The nests

were constructed in 2.0 X 32.0 mm trap-nest borings in soft pine blocks. The

trap-nest blocks were placed at the site in June 1972 and recovered in August

1972. Partitions and closures ranged from 0.5 to 2.0 mm thick and w-ere made

of pale yellow translucent resin. Three nests contained one brood cell each, and

one nest contained two brood cells. Brood cells ranged from 12.0 to 20.0 mm

long. Each nest had a single vestibular cell which ranged in length from 6.0 to

15.5 mm. Four of the five developing larvae spun a single, vestigial cocoon

consisting of a delicate, transverse, silken partition immediately anterior to the

larva’s head. The larva of the female from the same nest as the holotype spun

two of these silken partitions a few millimeters apart. Fecal pellets were deposited

at the rear of each brood cell. Aphids were the prey but the number used per cell

could not be determined.

Polemistus chewbacca Menke, New Species

(Figs. 4, 13, 18, 19)

Holotype male. — Black except as follows: yellow: ventral surface of antenna

(yellow changes to brownish yellow toward apex), trochanters ventrally, apex of

fore and midfemora, fore and midtibiae, basal half of hindtibia, all tarsi; pale

yellow: palpi, pronotal lobe and tegula.

Clypeus and lower frons densely covered by appressed silver setae which obscure

surface.

Scapal basin nearly smooth, with scattered, weak, fine, transverse ridges; ex-

174

PAN-PACIFIC ENTOMOLOGIST

tremely fine scattered punctures visible (75X) on frons below ocelli, punctures 2-

3 diameters apart; no carina paralleling inner or outer orbits; flagellomere I with

weak toothlike tylus apicoventrally, flagellomeres II-VI with low ridgelike tyli

ventrally, that on II visible only on apical one fourth, those on III-V extending

length of each flagellomere, that of VI occupying outer three fourths of flagellomere

(Fig. 4); free edge of clypeus as in Fig. 18; labrum arcuate apically, weakly indented

medially (Fig. 18). Scutum with extremely fine punctures anteriorly (75X), punc¬

tures 1-2 diameters apart; notaulus weakly impressed, not clearly pitted; pro-

podeal side smooth, shining adjacent to metapleuron but changing to oblique

parallel ridging and areolation dorsad; scrobal sulcus represented by crest between

scrobe and epistemal sulcus; hypersternaulus absent. Terga I—II with lateral carina,

but carina on I weak. Length 4.5 mm.

Variation in male. — The antennal tyli occupy only the apical one half to three

fourths of flagellomeres II-VI in one specimen, the apical three fourths on V-VI

in three specimens, and in one of these last the tyli on II-IV are full length. In

one specimen the propodeal side is ridged to the metapleuron and the dorsum is

broadly rugose. The angle at the center of the clypeal notch is more prominent

in the paratypes (Fig. 19).

Female .—As in male except: mandible apex and labrum brownish, trochanters

sometimes completely yellow. Clypeus with few appressed silver setae near apex,

surface not obscured. Free edge of clypeus deeply bi-emarginate, its lateral lobes

reflexed (Fig. 13); clypeal surface shiny; labrum truncate (Fig. 13); scutal punc-

tation obscure at 75X. Terga I—II with strong lateral carina. Length 4.5-5 mm.

Discussion .—The bi-emarginate female clypeus (Fig. 13), the rounded male

labrum, and the absence of a long tylus on male flagellomere I (Fig. 4) are diagnostic

for chewbacca. The absence of a hypersternaulus and the sparsely silvered female

clypeus relate chewbacca to vaderi, but the latter has a semicircular clypeal emar-

gination in the female (Fig. 12), the labrum is triangular in the female (Fig. 12),

and flagellomere I has a long tylus in the male (Fig. 3). The male clypeal and

labral outlines also differ between these two species (compare Figs. 17-19).

Etymolog}?. — The name chewbacca, a noun in apposition, is based on a rather

wild looking but friendly beast in the movies “Star Wars” and “The Empire

Strikes Back.”

Distribution. — Southeastern Arizona.

Types. — Holotype 8: ARIZONA, Cochise Co., Southwest Research Station, 5

mi. w. Portal, 5400', Sept. 9, 1959, H. E. Evans (CUI). Four 8 and 3 9 paratypes

as follows: ARIZONA, same locality as type, Aug. 20, 23, 29, Sept. 9, 13, 1959

(one on honey dew on Populus ), H. E. Evans, 2 8, 3 9 (CUI); Huachuca Mts.,

5000', June 14, 1920, A. A. Nichol, 1 9 (USNM); Miller Canyon, 6000', 1 mi. w.

parking area, Huachuca Mts., July 16, 1964, M. Noller, 1 8 (UA).

Literature Cited

Bohart, R. M., and A. S. Menke. 1976. Sphecid wasps of the world. Univ. Calif. Press, Berkeley,

ix + 695 pp.

Harris, R. A. 1979. A glossary of surface sculpturing. Occ. Papers Ent., Calif. Dept. Food Agric.,

no. 28, 31 pp.

Kohl, F. F. 1905. Hymenopteren aus der neotropischen Fauna. Verhandl. zool.-bot. Ges. Wien 55:

338-366.

VOLUME 59, NUMBERS 1-4

175

-. 1905. Zur Kenntnis der Hymenopterengattung Passaloecus Shuck. Verhandl. zool.-bot. Ges.

Wien 55:517-529.

Krombein, K. V. 1979. Superfamily Sphecoidea, pp. 1573-1740. In: Krombein et al., Catalog of

Hymenoptera in America north of Mexico, vol. 2. Smithsonian Institution Press, Washington

DC.

Rau, P. 1943. The nesting habits of certain sphecid wasps of Mexico, with notes on their parasites.

Ann. Ent. Soc. Amer. 36:647-653.

Saussure, H. de. 1892. Histoire naturelle des Hymenopteres, pp. 177-590 in vol. 20. In: Grandidier,

Histoire physique, naturelle, et politique de Madagascar. Paris.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 176-187

The Classification of the Lithurginae

(Hymenoptera: Megachilidae ) 1

Charles D. Michener

Department of Entomology, University of Kansas, Lawrence, Kansas 66045.

The Lithurginae is a small world-wide subfamily of Megachilidae, with its

maximum diversity in the arid temperate parts of South America. Its position

relative to related groups of bees is indicated in Fig. 1. The synapomorphies shown

in the figure and in Table 1 are not an exhaustive list but only enough to establish

the cladistic relationships and to characterize the major taxa related to the Li¬

thurginae. Peters (1972) has provided a similar list and come to conclusions similar

to mine, although he did not consider the diversity of Lithurginae here discussed.

The Fideliidae are considered as a subfamily of Megachilidae by Rozen (1977)

and this position for that group may be quite appropriate. My placement of it

here as a family does not indicate a strong conviction as to its status.

The definition of the Lithurginae given by Michener (1944) is not correct, as

noted by Moure (1949). (In 1944 I had not seen South American material of the

subfamily, and my comments were based on a limited sample of the species of

the genus Lithurge.) Thus the jugal lobe of the posterior wing, while generally

longer in Lithurginae than in Megachilinae, is only half as long as the vannal lobe

in Trichothurgus colloncurensis Ogloblin. The hindtibiae lack spicules in many

males and the spicules are weak, especially in Trichothurgus. The face of the

female is often not elevated below the antennal sockets in Trichothurgus. The

epistomal suture is often complete, even in some species of Lithurge. Finally, in

Trichothurgus the lowest rather than the middle mandibular tooth is often longest

and the middle one is not always advanced in front of the others, as in Lithurge

and Microthurge.

Characteristic features of Lithurginae include the following: Synapomorphies:

Characters 8 to 14, Table 1. Plesiomorphies: Jugal lobe of hindwing half to three

fourths as long as vannal lobe. Basitibial plate usually defined along posterior

margin and at apex in female, but not evident in several species of Trichothurgus.

Metasoma of male not curled under at apex as in most megachilids; sixth tergum

without transverse carina, teeth, and the like; seventh tergum fully exposed and

directed posteriorly. Six sterna fully exposed. Pygidial plate of female represented

by a longitudinal, dorsal, bare zone on sixth tergum, extended posteriorly as flat

or concave dorsal surface of apical tergal projection or spine, sometimes expanded

at apex; of male a broader plate, margined laterally by carinae, often pointed

apically and quite slender, almost like an apical tergal spine.

A common feature of the subfamily found in females of nearly all Lithurge, all

1 Contribution number 1839 from the Department of Entomology, University of Kansas, Lawrence,

Kansas 66045.

This paper is dedicated with pleasure and enthusiasm to Dr. Richard M. Bohart, in recognition not

only of his seventieth birthday but of his life-long work as a hymenopterist and as a teacher of systematic

entomology. A new species, Microthurge boharti, is named in his honor.

VOLUME 59, NUMBERS 1-4

177

Lithurginae

Fig. 1. Cladogram showing the position of Lithurginae. Numbers represent the apomorphies listed

in Table 1.

Microthurge, and a few Trichothurgus, and in males of a few species of Lithurge

(e.g., L. fortis Cockerell) and Microthurge, is a facial prominence. This is an

elevation near the median segment of the epistomal suture. Sometimes the suture,

if discernible, is largely above it so that the prominence is on the upper part of

the clypeus or largely so [T. osmioides (Friese), wagenknechti (Moure)]; L. rubri-

catus Smith). Sometimes the suture is below it, so that the prominence is on the

supraclypeal area or largely so [T. dubius (Sichel), Lithurgopsis, Microthurge ]. In

still others both areas are involved, so that the suture goes across the prominence

( L. sparganotes Schletterer, atratus Smith, scabrosus Smith, etc.).

Other features useful in recognizing the subfamily are the tridentate mandibles,

with the middle tooth the longest except in Trichothurgus. Arolia are absent or

nearly so in all females and some males.

Another interesting feature of the subfamily is the structure of the apex of the

labial palpus. Attached to the end of the very long second segment is the small,

flattened third segment, which continues in the same direction as the second and

is often somewhat pointed. Only the fourth segment, when present, diverges from

the axis of the preceding segments. In most long-tongued bees both third and

178

PAN-PACIFIC ENTOMOLOGIST

Table 1. Synapomorphies indicated in Fig. 1, with explanations or plesiomorphies in parentheses.

1. Labrum longer than broad. (The labrum is broader than long in other bees, with the exception

of some Nomadinae which must have evolved the elongate labrum independently.)

2. Scopa on metasomal sterna, not on legs. (No other bees have this feature although some short-

tongued bees have sternal in addition to leg scopal hairs.)

3. Hairy larval bodies. (Most other bees have bare larvae or larvae with only a few minute spicules.

Exceptions are allodapine anthophorids which must have evolved hairs independently.)

4. Hindbasitarsus with long hairs used in flicking soil away from nest entrance. (Not found in other

bees.)

5. Two submarginal cells. (Three is widespread, although reduction to two occurs in many bees.)

6. Seventh metasomal sternum of male reduced, without large apical lobes. (Seventh metasomal

sternum produced to paired thin apical lobes, a widespread feature of short-tongued and some

long-tongued bees.)

7. Volsellae lost or fused to form lobes on gonocoxites; opposable digitis and cuspis not recognizable

or at least not moveable. (Although volsellar reduction is common, volsellae are separate sclerites,

each with digitis and cuspis, in most bees.)

8. Proboscis elongate, in repose often reaching metasoma. (With few exceptions other megachilids

have shorter proboscides.)

9. Third segment of labial palpus flattened, on same axis as second. (Third segment directed laterally

in nearly all other long-tongued bees.)

10. Outer surfaces of tibiae, except hindtibiae of some males, with coarse hairless spicules. (Spicules

if present bearing an apical hair.)

11. Hindbasitarsi slender, almost cylindrical. (The hindbasitarsi are flattened in most bees, at least

in females.)

12. First metasomal tergum small, flattened, posterior margin rounded. (In other bees this tergum is

more convex in profile with anterior and dorsal surfaces, and has the posterior margin transverse.)

13. Posterior margin of sixth tergum of female with strong sublateral tooth, completely hidden in

dense hair in Trie hot hurgus. (Such teeth are absent in other megachilids.)

14. Male genitalia and hidden sterna extraordinarily small. (In other bees these structures are larger

and commonly more heavily sclerotic as well as more ornate.)

15. Basitibial plate completely lost. (Such loss is also common in other bees, although the plate is

widespread and presumably plesiomorphic.)

16. Jugal lobe of hindwing less than half as long as vannal lobe. (The jugal lobe is half or commonly

much more than half as long as the vannal lobe in many bees although reduction also occurs in

many groups other than megachilids.)

17. Pygidial plate and pygidial fimbria absent. (Both are present in most bees, although independently

lost in various groups; in Fideliidae they may also be lost although the condition found in that

group may result from great expansion of the plate.)

fourth segments diverge, although Prochelostoma in the Megachilinae resembles

the Lithurginae in this feature. The fourth segment of the labial palpus is some¬

times absent. It is certainly easily broken off, but seems to be actually absent in

males of Trichothurgus laticeps (Friese), in all specimens of Microthurge exam¬

ined, and in some Old World species of Lithurge (see below).

Key to Genera of Lithurginae

1. Labrum longer than clypeus, often much longer. Hindtibia of female

rather uniformly hairy on outer and anterior and posterior surfaces, spic¬

ules relatively inconspicuous among hairs. Lower mandibular tooth long¬

er than middle tooth or in some females, lower and middle teeth equal.

. Trichothurgus

VOLUME 59, NUMBERS 1-4

179

- Labrum about as long as clypeus. Hindtibia of female with hairs of broad,

longitudinal outer zone shorter and sparser than those of anterior and

posterior surfaces, spicules large and conspicuous in outer zone. Lower

mandibular tooth conspicuously shorter than middle tooth, which is long¬

est mandibular tooth. . 2

2. Body small, slender, heriadiform; claws of female bifid; stigma of mod¬

erate size, broadest at base of vein r, sides converging toward base; max¬

illary palpi 2-segmented. Microthurge

- Body broad, megachiliform; claws of female simple; stigma small, sides

basal to vein r parallel or nearly so; maxillary palpi 3- or 4-segmented.

. Lithurge

Moure (1949) placed Trichothurgus (with Lithurgomma ) in a separate tribe,

the Trichothurgini. While it is the most distinctive genus of Lithurginae, I see no

need to segregate it as a tribe, and follow Snelling (in press) in not doing so. Most

of its characters are plesiomorphies or probable plesiomorphies. The large and

elongate labrum, however, appears to be a synapomorphy uniting the species of

Trichothurgus , which can be regarded as a sister group to the other two genera

combined. The sparsely hairy wing membrane is probably also a synapomorphy

although this character arises independently in various other bees.

Lithurge and Microthurge agree in several synapomorphies: the reduced lower

mandibular tooth such that the middle tooth is longest as well as more anterior

than the others, the coarse tibial spicules in areas of short and sparse hairs, the

low profile of the posterior part of the thorax and especially of the propodeum,

the median apical pygidial spine of the female, the strong tooth lateral to this

spine on the posterior margin of tergum VI. Compared to Trichothurgus with its

many plesiomorphies, Lithurge and Microthurge have few, the most noteworthy,

perhaps, being the ordinary sized labrum and the moderate to high density of

hairs on the wing membrane.

Microthurge has few apomorphies. These include the short maxillary palpi and

perhaps the larger stigma (plesiomorphic for Apoidea as a whole, no doubt, but

perhaps derived in connection with small body size in this genus as well as in

some related small megachilids such as Chelostoma and Heriades). Bifid claws

in females (Fig. 5) are considered a plesiomorphy by Peters (1972) in other mega¬

chilids, probably correctly, but could be an apomorphy in Microthurge. Genes

for this character must be retained in all species, because the claws of males are

cleft. A minor regulatory change could therefore cause their reactivation in fe¬

males, and would be apomorphic. The broad pygidial plate of males of Micro¬

thurge (Fig. 8) seems to be plesiomorphic, as compared to all other megachilids,

but one must question this idea in view of the slender produced plate in both

Trichothurgus and Lithurge (Fig. 7). If this similarity is due to homology, then

the broad plate of male Microthurge is a reversion toward the more primitive

apoid condition, but for this genus it would be an apomorphy.

For Lithurge the situation is equally confusing. If the stigmal, claw, and pygidial

characters listed above are plesiomorphic for Microthurge, then the alternative

characters are apomorphic for Lithurge. Regardless of such matters, there is enough

phenetic difference between Lithurge and Microthurge to justify generic status for

them both.

Figs. 2-6. 2-4, sixth metasomal terga of females; 2, Trichothurgus wagenknechti, 3, Lithurge apicalis,

and 4, Microthurge pharcidontus. 5, 6, claws of females; 5, Microthurge pharcidontus and 6, Lithurge

apicalis.

The long, slender arolia in males of Lithurginae are of interest. No doubt their

presence is plesiomorphic; they occur in both sexes of most bees, but are absent

in female lithurgines. They are present in males of Trichothurgus, in the Western

Hemisphere subgenus Lithurgopsis of Lithurge, and in one Australian species,

Lithurge ( Lithurge ) rubricatus Smith. They are absent in the rest of the genus

Lithurge and in Microthurge.

Genus Trichothurgus Moure

(Fig. 2)

Trichothurgus Moure, 1949:240. Type species: Megachile dubia Sichel, by original

designation.

Lithurgomma Moure, 1949:277. New synonym. Type species: Lithurgomma wa¬

genknechti Moure, by original designation.

Commonly robust, hairy bees without metasomal hair bands. Facial promi¬

nence commonly absent, present in females of T. dubius (Sichel) (clypeal); os-

mioides (Friese), pseudocellatus (Moure), and wagenknechti (Moure) (all supra-

clypeal). Epistomal suture complete. Labrum longer than clypeus, sometimes as

long as distance from clypeal apex to anterior ocellus. Mandible with lower tooth

longer than median tooth or in females of some species, these two teeth subequal.

Maxillary palpus 3-segmented; labial palpus 4-segmented or fourth segment some¬

times missing. First flagellar segment over twice as long as greatest breadth [less

than twice as long in osmioides (Friese)], second much shorter than first and

usually distinctly broader than long. Tibiae with anterior, outer, and posterior

VOLUME 59, NUMBERS 1-4

181

surfaces rather uniformly hairy; fore and middle tibiae each with row of small

spines or spicules extending basad on posterior outer surface from each of the

two apical tibial spines, these rows limited to distal part of tibia and absent in

some males; hindtibia with similar spicules scattered among hairs of outer surface

in females but not in males. Arolia of male distinct, long, slender (as in Lithur-

gopsis). Claws of female simple. Hindbasitarsus of female subequal to tibia or

usually shorter. Stigma small, slender, sides basal to vein r parallel. Wing mem¬

brane bare or with sparse hairs. Profile of first metasomal tergum with basal

concavity less horizontal than dorsal surface. Pygidial plate of female a dorsal

strip on tergum VI, usually slightly expanded at apex, not projecting as a spine;

of male a dorsally flat or concave, broad, blunt projection or robust spine, similar

to that of Lithurge but often hidden by long hair from adjacent parts of tergum.

Tergum VI of female with apical tooth at each side small and hidden in the dense

hairs (Fig. 2).

This genus is known from Chile, Argentina, and Peru. At least some species

visit flowers of Cactaceae for pollen. They range from very large (up to 21 mm

long, T. dubius (Sichel)) to rather small (7 mm long, T. colloncurensis Ogloblin).

The following species are placed in this genus: albiceps (Friese), 1908; lalpestris

(Friese), 1923; aterrimus (Cockerell), 1926; colloncurensis Ogloblin, 1957; dubius

(Sichel), 1867; herbsti (Friese), 1905 (= muticus (Herbsl), 1918); holomelan (Moure),

1949; laticeps (Friese), 1906; neoqueenensis (Friese), 1910; osmioides (Friese),

1910; pseudocellatus (Moure), 1949; shajovskoyi Ogloblin, 1957; wagenknechti

(Moure), 1949.

The name Lithurgomma was proposed for certain species in which there is a

defined impunctate area in front of the anterior ocellus. The size and sharpness

of definition of this area varies and intergradation to typical Trichothurgus thus

seems to occur. Moreover, other features do not show close relationship among

the forms having such an area. I therefore see no justification for recognition of

Lithurgomma as a genus or even as a subgenus, a viewpoint supported by Snelling

(in press).

Microthurge Michener, New Genus

(Figs. 4, 5, 8)

Type species: Lithurgus pharcidontus Moure.

Small, slender species having the form of a large Heriades or one of the small

species of Chelostomoides; metasomal terga with apical pale hair bands. Facial

prominence present in females, weakly so in some males, supraclypeal. Labrum

as long as clypeus. Mandible with middle tooth longest. Maxillary palpus ex¬

tremely short, 2-segmented; labial palpus 3-segmented (fourth segment absent).

First flagellar segment less than twice as long as greatest width but over twice as

long as second segment which is at least twice as broad as long. Tibiae, especially

hindtibia, with shorter hairs on outer surface than on anterior and posterior

surfaces and with strong spicules arranged as in Lithurge, present in males but

more strongly developed in females. Arolia of male absent. Claws of female bifid.

Hindbasitarsus of female about as long as tibia. Stigma larger than in other genera,

sides basal to vein r diverging toward apex. Wing membrane with dense short

hairs. Profile of first metasomal tergum as in Lithurge. Pygidial plate of female

represented by flat upper surface of short apical spine of tergum VI; of male a

182

PAN-PACIFIC ENTOMOLOGIST

broad triangle, nearly pointed apically, not or but little extending beyond rest of

tergum VII. Tergum VI of female with strong apical, marginal tooth at each side,

not hidden in dense hairs.

Distribution.— This genus occurs in Argentina, Boh via, and probably southern

Brazil. The following species are included: boharti new species, pharcidontus

(Moure), 1948; pygmaeus (Friese), 1908; and probably corumbae (Cockerell),

1901.

Discussion. —Microthurge is more similar to Lithurge than to Trichothurgus, as

shown by the labral size, the large median mandibular tooth of both sexes, the

tibial vestiture and spiculation, the densely hairy wings, the profile of the first

tergum, and the apical spine and sublateral apical teeth of tergum VI of the female.

It differs not only from Lithurge but also from Trichothurgus, however, by its

small, slender form, the short two-segmented maxillary palpus, the cleft claws of

the female, the larger pterostigma, the short apical spine of tergum VI of the

female, and the broad rather than apically produced pygidial plate of the male.

From other Western Hemisphere Lithurginae (except the possibly adventive Li¬

thurge huberi Ducke), Microthurge differs by the lack of arolia in the male.

Etymology.— The generic name, formed in the manner of Padre J. S. Moure in

naming Trichothurgus, is from micro, small, plus part of the generic name Li¬

thurge.

Genus Lithurge Latreille

(Figs. 3, 6, 7, 9-11)

Robust but with metasoma somewhat flattened, commonly with hair bands.

Facial prominence almost always present in females, absent in most males (present

in males of L. fortis, etc.). Epistomal suture present or absent in females between

tentorial pits. Labrum about as long as clypeus. Mandible with middle tooth

longest. Maxillary palpus 3- to 4-segmented; labial palpus 4-segmented except in

certain Old World species that lack fourth segment. First and second flagellar

segments variable. Tibiae, especially posterior one, with hairs shorter on outer

surface than on posterior surface. Fore and middle tibiae in female each with two

longitudinal rows of coarse spicules extending basad on posterior part of outer

surface from each of the two apical tibial spines, these rows outlining a channel

on each tibia between rows which extends to the middle of the tibia or beyond

nearly to base; sometimes some spicules between the rows; in male spicules small¬

er, rows shorter, sometimes absent. Hindtibia of female with coarse spicules

scattered over short-haired outer surface; spicules smaller in males, absent in

many species, present but small in Lithurgopsis. Arolia of male present or absent.

Claws of female simple. Hindbasitarsus of female slightly shorter than to longer

than tibia. Stigma small, sides basal to vein r parallel or nearly so. Wing membrane

hairy, often densely so. Profile of hrst metasomal tergum with dorsal surface almost

a direct continuation in a straight line from surface of basal concavity. Pygidial

plate of female a flat or concave dorsal surface of long apical spine of tergum VI;

of male a dorsally concave, broad, blunt projection or robust spine. Tergum VI

of female with strong apical, marginal tooth at each side, not hidden in dense

hairs; pygidial plate of male a broad flat-topped apical pointed projection.

This genus is world-wide in distribution in tropical and warm to moderate

VOLUME 59, NUMBERS 1-4

183

Figs. 7-11. 7, 8, seventh metasomal terga of males; 7, Lithurge apicalis and 8, Microthurge phar-

cidontus. 9-11, Lithurge apicalis, male; 9, genitalia (dorsal side on left), 10, eighth sternum, 11, seventh

sternum.

temperate zones, except that in the Americas it may be largely absent from the

wet tropics. Two subgenera are commonly recognized, as follows.

Subgenus Lithurge Latreille, s. str.

Lithurge Latreille, 1825:463. Type species: Andrena cornuta Fabricius (mono¬

basic).

Lithurgus Berthold, 1827:467. An emendation of Lithurge, hence type species

automatically the same.

Arolia absent or rudimentary in both sexes (see Cockerell, 1905) except for L.

rubricatus (from Australia) in which arolia of the male are as large and long as in

male Lithurgopsis. Facial prominence of female involving upper part of clypeus

and usually also part of supraclypeal area. First flagellar segment about twice as

long as broad, more than twice as long as second, which is much broader than

long.

This subgenus is found in Eurasia, Africa, Australia and intervening islands

and includes all of the Old World species of the genus. In Africa it reaches the

Cape, in Australia, New South Wales but is apparently absent from southern parts

of the continent, while in Eurasia it extends north to Japan, China, Hungary,

Spain, etc. To the east it occurs as far as Tahiti, perhaps having been carried by

184

PAN-PACIFIC ENTOMOLOGIST

primitive boats. One species, L. huberi Ducke, occurs in Brazil; it is probably the

same as one of the Old World species and adventive in Brazil (Snelling, in press).

Another species, L. chrysurus Fonscolombe, has been recorded by Roberts (1978)

as introduced presumably from the Mediterranean region into New Jersey.

L. rubricatus Smith does not agree with Lithurgopsis except in the presence of

long arolia in the male. Such arolia may be a primitive feature that survived in

Australia and the Americas while being replaced in the rest of the world by forms

lacking such arolia in both sexes.

While some species of this subgenus collect pollen from Malvaceae which have

coarse pollen like that of the Cactaceae used by the subgenus Lithurgopsis, other

species collect fine pollen of Compositae and perhaps other flowers. There may

be morphological correlates with the type of flowers visited for pollen. Thus most

species have coarse (malvaceous?) pollen on them and have the proboscis long,

with the fourth segment of the labial palpus present. A few species have fine

(composite?) pollen on them and have the proboscis shorter, with the fourth

segment of the labial palpus absent. Such species are L. chrysurus Fonscolombe,

capensis Friese (= ovatus Cameron), and fuscipennis Lepeletier.

Latreille’s original spelling (1825) of the genus and subgenus name has been

used here, although it has been regarded by many authors as French vernacular

(e.g., Fox, 1902). Such authors use the emended Latinized form Lithurgus, intro¬

duced by Berthold (1827) when he translated Latreille’s work into German. The

spelling Lithurge, as noted by Fox (1902), was used by Lepeletier (1828:795) but

by few subsequent authors. However, it was shown as the correct name by Sand-

house (1943) in her treatment of apoid generic names, and has since been used

by Michener (1944, 1951) and Hurd (1979) in major studies and catalogues, as

well as in shorter papers by various authors.

Subgenus Lithurgopsis Fox

Lithurgus (Lithurgopsis ) Fox, 1902:138. Type species: Lithurgus apicalis Cresson

(original designation).

Male with arolia. Facial prominence of female entirely supraclypeal (absent in

one species being described by Snelling, in press). First flagellar segment not or

little longer than broad, slightly longer than to shorter than second, which is nearly

as long as broad to longer than broad.

This subgenus is limited to the Western Hemisphere, where it ranges from

South Dakota to Argentina, but is absent from broad areas in the tropics. It

collects pollen from Cactaceae. Snelling (in press) has reviewed the North Amer¬

ican species.

Appendix

Below I establish a lectotype for a name in Trie hot hurgus, and present descrip¬

tions of the species of Microthurge available to me.

Trichothurgus herbsti (Friese)

Moure (1949) apparently correctly placed Lithurgus muticus Herbst, 1918, in

the synonymy ot “Megachile” herbsti Friese, 1905. Two males and two females

of Lithurgus muticus Herbst are in the Museum of Comparative Zoology, Harvard

VOLUME 59, NUMBERS 1-4

185

University, and evidently are cotypes, no holotype or lectotype having been des¬

ignated. All are labelled Chile, Rio Blanco, P. Herbst; one female XII-10-17, one

male and one female XII-11-17, and one male XII-26-17. The females and the

male taken on XII-10 all bear the label “in floribus Echinocactus ceratites Otto,”

while the remaining male, probably taken on another flower, is labelled “Her¬

barium No. 46.” Each also bears Herbst’s label “ Lithurgns muticus P. Herbst,

Type” (the sex also being indicated on this label) and a red label “MCZ Type

17207.” Herbst described the female first and in greater detail than the male; I

therefore designate and have labelled the female taken on XII-10 as the lectotype.

Microthurge pygmaeus (Friese), New Combination

Lithurgus pygmaea Friese, 1908:62.

Female.— Upper margin of base of mandible just below lateral part of anterior

clypeal margin with large, dorsoventrally flattened projection extending antero-

ventrally (projection described as clypeal by Friese, 1908); apex of facial protu¬

berance a transverse, shining carina over one third as long as width of face at that

level and much nearer to antennal bases than is anterior ocellus; mesoscutum

punctate, finely and closely so around margins, somewhat more coarsely and less

closely so on disc.

Male.— Upper margin of clypeus with interspaces between punctures broader

than elsewhere, forming a shining transverse band, sometimes weak; supraclypeal

area slightly elevated and with a small tubercle on each side; shiny transverse

basal zone of labrum uniform in convexity.

This species was described from Mendoza and is widespread in Argentina, as

indicated by the following collecting data from specimens in the Snow Entomo¬

logical Museum, University of Kansas. Buenos Aires Prov.: Villa Elias, Feb., 1955

(J. Foerster); Ing. R. Otamendi, Nov., 1954 (F. H. Walz); Tigre, Nov. 3-10, 1956;

Dique, Lujan, Nov., 1954 (F. H. Walz). Entre Rios Prov.: Villaguay, Nov. 20,

1951 (J. Foerster). Misiones Prow: Obera, Oct. 17, 1950 (M. Senkute). Tucuman

Prov.: 20 km W. of S. M. de Tucuman, Dec. 10, 1971 (D. J. Brothers).

Microthurge pharcidontus (Moure), New Combination

Lithurgus pharcidontus Moure, 1948:321.

Female.— Mandibular projection absent; apex of facial protuberance shining,

transverse, slightly emarginate medially seen from above, less than one third as

wide as face at that level, almost as far below antennal bases as median ocellus

is above; mesoscutum finely and closely punctate around margins, but broad

median transverse zone between tegulae coarsely, transversely rugosopunctate,

several transverse rugae particularly strong.

Male.— Unknown (see below).

This species was described from Tarancas, San Pedro de Colalao, Tucuman

Prov., Argentina. Specimens from that locality, collected in December and Feb¬

ruary by P. J. M. Arnau and J. Foerster, are at hand. Additional specimens are

from: ARGENTINA: Tucuman Prov.: El Cadillal; Oct., 1951 (J. Foerster). BO¬

LIVIA: Cochabamba Prov.: Feb., 1952 (F. H. Walz).

A male that might be this species is from Guayaramerin, El Beni, Bolivia, Dec.

6, 1956 (M. Fritz). It is similar to that of M. pygmaeus but the upper part of the

186

PAN-PACIFIC ENTOMOLOGIST

clypeus is densely punctate; the shiny transverse basal zone of the labrum is

distinctly prominent medially, not uniformly convex; the eyes, unlike those of

other known species, are closer together below, separated by little more than the

maximum ocular width; and the facial hairs are more golden.

Microthurge boharti Michener, New Species

Female. — Length 8 mm (varying to 6 mm among paratypes); forewing length

6 mm (varying to 5 mm). Mandibular projection absent; apex of facial projection

bilobed, shining laterally but punctate medially, less than one third as wide as

face at that level, about two-thirds as far below antennal bases as median ocellus

is above; mesoscutum similar to that of M. pygmaeus but disc slightly more

coarsely and less closely punctate.

Male.— Length 7 mm, forewing length 6 mm. Upper margin of clypeus densely

punctate like adjacent areas; supraclypeal area as described for M. pygmaeus ;

shiny transverse basal zone of labrum slightly prominent medially (sometimes

only feebly so), not uniformly convex.

Types. —Holotype 2, allotype 6, and seven 2 and one <3 paratypes: ARGENTINA:

Cordoba Prov., Isla Martin Garcia, Jan., 1943 (M. J. Viana). One 2 paratype:

Cordoba Prov., Valle Hermoso (M. J. Viana). One <3 and one 2 paratype: Cordoba

Prov., Huerta Grande, Jan., 1941 (F. M. Walz). One 2 paratype: labelled merely

“Argentina.” A series of specimens of both sexes in the Los Angeles County

Museum of Natural History is from ARGENTINA, Catamarca Prov., Andalgala,

(J. L. Neff). Some or all were taken on flowers of Abutilon virgatum on March

14, 1973, and January 16, 1974. The holotype and allotype are in the Museo

Argentina de Ciencias Naturales “Bernardino Rivadavia” (Buenos Aires), other

type material is in the Snow Entomological Museum, University of Kansas.

Etymology?. — Microthurge boharti is named in honor of R. M. Bohart.

Discussion. —This species could be Microthurge corumbae (Cockerell), new com¬

bination, the type of which I have not been able to locate. The description of

corumbae, however, indicates a species with more abundant white hair and with

the femora and first two metasomal segments dark ferruginous (Cockerell, 1901).

These areas are black in M. boharti. Moreover, M. corumbae is from Corumba,

Mato Grosso, Brazil, which is in a moist tropical region, while M. boharti is from

desertic areas farther south.

Micorthurge sp.?

Two small males (the smallest only 5 mm long) from Pocitos, Salta, Argentina,

Nov. and Dec., 1956 (M. Fritz) may be conspecific with the male tentatively

placed with M. pharcidontus. The supraclypeal area, however, lacks the two tu¬

bercles characteristic of that species.

Acknowledgments

I am indebted to R. J. McGinley of the Museum of Comparative Zoology for

the loan of lithurgine material, and to R. R. Snelling of the Los Angeles County

Museum of Natural History not only for the loan of material but for sending me

a copy of his unpublished paper on North American Lithurge.

This paper is a product of National Science Foundation grant DEB 77-23035.

VOLUME 59, NUMBERS 1-4

187

Literature Cited

Berthold, A. A. 1827. Latreille’s naturliche Familien des Thierreichs mit Anmerkungen und Zusatzen.

Weimar, 8 + 602 pp.

Cockerell, T. D. A. 1901. Descriptions of new bees collected by Mr. H. H. Smith in Brazil—II. Proc.

Acad. Nat. Sci. Philadelphia 53:216-222.

-. 1905. Notes on some bees in the British Museum. Trans. Amer. Ent. Soc. 31:309-364.

Fox, W. J. 1902. Lithurgopsis, a new genus of bees. Ent. News 13:137-140.

Friese, H. 1908. Die Apidae (Blumenwespen) von Argentina nach den Reisenergebnissen der Herren

A. C. Jensen-Haarup und P. Jorgensen in den Jahren 1904-1907. Flora og Fauna 10:1-94.

Hurd, P. D. 1979. Apoidea, pp. 1741-2209. In: K. V. Krombein, P. D. Hurd, D. R. Smith, and B.

D. Burks, Catalog of Hymenoptera in America north of Mexico, vol. 2. Smithsonian Institution

Press, Washington, D.C.

Latreille, P. A. 1825. Families naturelles du regne animal .... Paris, 570 pp.

Lepeletier de St. Fargeau, A. [1828]. Encyclopedic Methodique. Insectes, vol. 10, pp. 345-832. Paris.

Michener, C. D. 1944. Comparative external morphology, phylogeny, and a classification of the

bees. Bull. Amer. Mus. Nat. Hist. 82:151-326.

-. 1951. Megachilidae, pp. 1136-1186. In: C. F. W. Muesebeck, K. V. Krombein, and H.

Townes, Hymenoptera of America north of Mexico. Synoptic Catalog. U.S. Dept. Agric., Agric.

Monog. 2.

Moure, J. S. 1948. Notas sobre algumas abelhas de Tacanas, Tucuman, Argentina. Rev. Ent. 19:

313-346.

-. 1949. Las especies chilenas de la sub-familia Lithurginae. Arq. Mus. Paranaense 7:265-286.

Peters, D. S. 1972. Uber die Stellung von Aspidosmia Brauns 1926 nebst allgemeinen Erorterungen

der phylogenetischen Systematik der Megachilidae. Apidologie 3:167-186.

Roberts, R. B. 1978. The nesting biology, behavior and immature stages of Lithurge chrysurus, an

adventitious wood-boring bee in New Jersey. J. Kansas Ent. Soc. 51:735-745.

Rozen, J. G., Jr. 1977. The ethology and systematic relationships of fideliine bees, including a

description of the mature larva of Parafidelia. Amer. Mus. Novitates no. 2637:1-15.

Sandhouse, G. A. 1943. The type species of the genera and subgenera of bees. Proc. U.S. Natl. Mus.

92:519-619.

Snelling, R. (In press) The North American species of the bee genus Lithurge.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 188-217

Key to North and Central American Species of the Mealybug

Genus Hypogeococcus (Homoptera: Coccoidea: Pseudococcidae)

with Descriptions of Four New Species and

Redescription of the Type Species

Douglass R. Miller

Systematic Entomology Laboratory, IIBIII, ARS, USDA, Beltsville, Maryland

20705.

I wish to express my gratitude to Dick and Margaret Bohart for their unselfish

efforts in my behalf while I attended the University of California at Davis. I take

great pleasure in dedicating this article to them as a token of my sincere appre¬

ciation for their guidance, example, and many kindnesses.

Introduction

The genus Hypogeococcus was described by George Rau in 1938 and included

only H. barbarae Rau. Ferris (1953) included a second species, and later three

more were added to the genus (Miller and McKenzie, 1971; Williams, 1973;

Willink, 1981). In this paper four additional species are described, and the type

species is redescribed.

Hypogeococcus is known in the New World only and occurs from New York

to Argentina. Species have been collected on such diverse hosts as composites,

cacti, orchids, and the unusual monocotyledonous genus Eriocaulon. Several species

are of economic importance: Hypogeococcus boharti n. sp., H. gilli n. sp., H.

othnius Miller and McKenzie, and H. spinosus Ferris often are encountered at

ports-of-entry in the United States and are excluded from the U.S. because they

are considered to be potential pests of cacti and orchids. Hypogeococcus festerianus

(Lizer y Trelles) was introduced into Queensland, Australia as a biological control

agent of the weedy cactus Eriocereus martinii. McFayden (1979) reported that

the mealybug causes deformed and distorted growth to the cactus, and growth

and seed development are greatly reduced. McFayden and Tomley (1978) pre¬

dicted that successful biological control of the cactus in Australia likely would be

achieved by the mealybug in combination with an introduced cerambycid.

Materials and Methods

Although all specimens in the type series of the new species were examined in

detail, a total of 10 specimens was used to determine ranges and means. The 10

specimens were selected from as many localities and hosts as possible. Averages

were rounded off to the nearest whole number or to the nearest tenth of a micron.

Specific geographic localities are not always given because many collections were

taken at LLS. ports-of-entry and specific data are unknown. Specimens were stud¬

ied using a Zeiss, phase contrast compound microscope at magnifications of 160,

400, and 1000X.

Terminology is the same as that used by Miller (1975) with the exception of

abdominal segmentation of the female. I have followed the numbering system

VOLUME 59, NUMBERS 1-4

189

adopted by Beardsley (1965), with the first visible abdominal segment numbered

I. This system, unlike the one used by Ferris (1950), is consistent with the seg¬

mentation of the males (Theron, 1958). The term cisanal seta is applied to all

of the setae on the venter of segment IX not just the posterior setae as suggested

by Ezzat and McConnell (1956) and MacGillivray (1921). This was done to avoid

giving each seta on this segment a separate name.

Depositories of types and their abbreviations are as follows: Auburn University,

Auburn, Alabama (AUA); British Museum (Natural History), London (BMNH);

California Department of Food and Agriculture, Sacramento (CDAS); Florida

State Collection of Arthropods, Gainesville (FSCA); Institute of Zoology, Aca¬

demia Sinica, Beijing, People’s Republic of China (IZAS); Museo de Historia

Natural de la Ciudad de Mexico, Mexico, D.F. (MCM); Museum National d’His-

toire Naturelle, Paris (MNHP); Shanghai Institute of Entomology, Academia Sin¬

ica, People’s Republic of China (SIE); University of California, Davis (UCD);

University of Georgia, Experiment (UG); University of Hawaii at Manoa, Ho¬

nolulu (UH); United States National Museum of Natural History, Washington,

D.C. (USNM); Virginia Polytechnic Institute and State University, Blacksburg

(VPI); and Zoological Institute, Academy of Sciences of USSR, Leningrad (ZIL).

Hypogeococcus Rau

Hypogeococcus Rau, 1938:159. Type species: Hypogeococcus barbarae Rau, 1938,

by monotypy.

Diagnosis of adult females. — Dorosomedial setae on posterior abdominal seg¬

ments enlarged, often conical, similar to cerarian setae; without trilocular pores;

cerarii on abdomen only, with 4-6 pairs; posterior cerarii each with more than 2

conical setae; circulus small and oval, not divided; unusually large anal ring for

pseudococcid.

Notes.— No single character can be used to distinguish species of Hypogeococcus

from all other pseudococcid genera. Members of the genus can be separated by

use of a combination of the above mentioned characters; individual species possess

a majority but not all of these diagnostic features. In fact, the character states

mentioned in the diagnosis are absent on one or more of the included species.

As in previous studies (Ferris, 1953; Miller and McKenzie, 1971), the mono-

phyletic nature of Hypogeococcus remains questionable. There appear to be several

groups of species within the confines of what currently is considered to be Hy¬

pogeococcus. These are: 1. boharti and othnius (without trilocular pores and dor-

somedial conical setae, translucent pores absent from hind coxa, anal ring with

2 or 3 rows of pores, first instar with triloculare only and more than 2 cerarii, on

orchids); 2. festerianus, pungens, and spinosus (without trilocular pores, dorso-

medial conical setae abundant on posterior 4 or 5 abdominal segments, anal ring

broad with 2 or 3 rows of pores, translucent pores on hind coxa, first instar with

triloculars and 1 or 2 pairs of cerarii, primarily on cactus); 3. hamoni and mar-

garetae (without trilocular pores, dorsomedial conical setae uncommon on pos¬

terior 4 or 5 abdominal segments, anal ring broad, with 6-8 rows of pores, trans¬

lucent pores on hind coxa, first instar with triloculars and more than 2 pairs of

cerarii, on Eriocaulon ); 4. barbarae (with trilocular pores, dorsomedial conical

setae abundant on posterior 4 or 5 abdominal segments, anal ring narrow with 2

190

PAN-PACIFIC ENTOMOLOGIST

rows of pores, translucent pores on hind coxa, first instar unknown, on Compos-

itae); 5. gilli (with trilocular pores, dorsomedial conical setae abundant on pos¬

terior 4 or 5 abdominal segments, anal ring broad with 4 or 5 rows of pores,

translucent pores on hind coxa, first instar unknown, on orchids).

Groups 2 and 3 each possess enlarged leg setae, and the basal segment of the

labium is closely appressed to the second segment. These characters, which prob¬

ably are synapomorphies, may suggest a close relationship between these groups.

Douglas J. Williams, Commonwealth Institute of Entomology, London, En¬

gland and Christina Grana de Willink, Universidad Nacional de Tucuman, Ar¬

gentina, are continuing their studies of South American species of the genus.

According to Doug Williams (pers. corresp.) several problems remain unresolved

in regard to the Argentine species. I hesitate to instigate any generic rearrangements

until these problems are resolved, until more species are described, and until first

instars and adult males have been studied more completely.

Hypo geococcus contains the following species:

1. barbarae Rau, 1938:159 (New York) (Compositae).

2. boharti Miller, n. sp. (Central America, Mexico) (Orchidaceae).

3. festerianus (Lizer y Trelles), 1942:24 (Argentina) (Cactaceae) (see Williams,

1973).

4. gilli Miller, n. sp. (Costa Rica) (Orchidaceae).

5. hamoni Miller, n. sp., (Florida) (Eriocaulon- Eriocaulaceae).

6. margaretae Miller, n. sp. (Florida) (Eriocaulon-Eriocaulaceae).

7. othnius Miller and McKenzie, 1971:578 (Central America, northern South

America, Mexico) (Orchidaceae).

8. pungens Willink, 1981:61 (Argentina) (Alter nanthera-Polygon&czdLt).

9. spinosus Ferris, 1953:382 (California, Texas) (Cactaceae).

Pseudococcus mirabilis Brain (1915), an African species, was placed in Hypo-

geococcus by De Lotto (1974). It is clear that mirabilis does not belong in the

latter genus since it has only slightly enlarged dorsomedial setae, has trilocular

pores, has only 2 conical setae in each posterior cerarius, lacks a circulus, and has

a narrow, weakly developed anal ring.

Key to Adult Females of

Hypogeococcus from North and Central America

1. Trilocular pores present. 2

- Trilocular pores absent. 3

2(1) Oral-collar tubular ducts absent from medial area of dorsum; dorsal

area of posterior abdominal segment sclerotized throughout; without a

circulus .. gilli Miller, n. sp.

- Oral-collar tubular ducts present over dorsal surface; dorsal area of

posterior abdominal segment with sclerotization restricted to cerarian

area; usually with a circulus . barbarae Rau

3(2) Without conical or enlarged setae on dorsomedial areas of segments VI

and VII, these areas with bristle-shaped setae only . 4

- At least 1 conical or enlarged seta on dorsomedial areas of either or

both of segments VI and VII . 5

4(3) Posterior 4 pairs of cerarii each with 3 or more conical setae .

. boharti Miller, n. sp.

VOLUME 59, NUMBERS 1-4

191

- Posterior 4 pairs of cerarii each with 2 conical setae .

. othnius Miller and McKenzie

5(3) Oral-collar tubular ducts on dorsomedial areas of thorax ... spinosus Ferris

- Oral-collar tubular ducts absent from dorsomedial areas of thorax ... 6

6(5) Femora of hind 2 pairs of legs with enlarged setae; dorsal setae pre¬

dominately enlarged; antennae 5-segmented, occasionally with third

segment partially divided; penultimate cerarius with same number or

more conical setae than anal-lobe cararius . margaretae Miller, n. sp.

- Femora of hind 2 pairs of legs without enlarged setae; dorsal setae

predominately unenlarged; antennae 6-segmented, or 7-segmented;

penultimate cerarius with fewer conical setae than anal-lobe cerarius

. hamoni Miller, n. sp.

Hypogeococcus barbarae Rau

Hypogeococcus barbarae Rau, 1938:159; Ferris, 1953:380; McKenzie, 1967:201;

Miller and McKenzie, 1971:578; Williams, 1973:567, De Lotto, 1974:109;

Cruttwell and Fidalgo, 1974:1; McFadyen, 1979:284; Willink, 1981:61.

Etymology’.— The species epithet was named in honor of Mrs. Barbara Rau,

the mother of George Rau.

Type data.—In the original description Rau stated that the holotype was de¬

posited in his private collection. I have been unable to locate Mr. Rau or his

collection. None of the type series is part of the holdings of the American Museum

of Natural History, Cornell University, or the New York State Museum, Albany.

A single slide with data identical to that given by Rau in the original description

has been located in the collection of the University of California at Davis. It is

labelled in Rau’s handwriting as follows: “Hypogeococcus/barbarae Rau/Aster

sp./Saratoga Springs,/Sept. 11, 1936./Rau.” The slide contains 3 specimens, and

although no type designation is on the slide, it clearly is part of the paratype series.

Two additional slides each containing 3 specimens are labelled in Ferris" hand¬

writing; they probably are part of the type series, but they lack a specific date in

September. Another slide from the type series is deposited in the collection of the

British Museum (Natural History). It contains 2 specimens and was collected a

year later; it is labelled in Rau’s handwriting as follows: “Hypogeococcus/barbarae

Rau/On Aster sp./Saratoga Springs,/N.Y. S/6/31 ./Paratypes/Rau.” Rau appar¬

ently tried to recollect the species again in 1938. Specimens collected at Saratoga

Springs, on Aster sp., in June and late August are also in the collection at Davis,

but these specimens are not Hypogeococcus barbarae and are not part of the type

series. Three topotype specimens on 2 slides deposited in the U.S. National

Museum are labelled as Hypogeococcus barbarae. These specimens were collected

in late September 1936 and 1938 but are not H. barbarae; they are similar to

species of Humococcus by virtue of the form of their anal ring, but their correct

identity is uncertain.

Adult Female (4th Instar)

(Fig. 1)

This description is based on the 5 paratype specimens labelled by Rau and the

6 specimens labelled by Ferris mentioned above.

Description. — Mounted 0.9-1.4 (1.2) mm long, 0.6-0.8 (0.7) mm wide. Body

elongate oval.

192

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Adult female paratype of Hypogeococcus barbarae Rau. Saratoga Springs, New York, IX-

11-36, on Aster sp.

Dorsum with 4-6 (6) pairs of cerarii. Anal-lobe cerarius with little or no scler-

otization, no bristle-shaped setae, 5-9 (7) conical setae, 5-12 (9) trilocular pores,

0-1 (0) discoidal pores. Penultimate cerarius with 4-9 (7) conical setae, small

cluster of trilocular pores. Antepenultimate cerarius with 3-7 (5) conical setae,

occasional basal trilocular pore. Fourth cerarius with 1-5 (3) conical setae, without

basal trilocular pores. Fifth cerarius with 0-2 (2) conical setae, without associated

VOLUME 59, NUMBERS 1-4

193

trilocular pores. Sixth cerarius with 0-2 (1) conical setae, without associated

trilocular pores. With 2 kinds of setae; posterior setae conical, not as large as

cerarian setae, gradually becoming thinner anteriorly; setae at juncture of thorax

and abdomen thin, not bristle-shaped, present over remainder of abdomen; longest

seta on segment VII, excluding those in surrounding cerarian cluster, 20-25 (22)

li long, segment IV with 30-41 (34) setae excluding those in cerarii. Trilocular

and discoidal pores scattered over surface, discoidal pores less abundant. Mul-

tilocular pores absent from 1 of 11 specimens, when present, with 1-8 (4) pores

on 1 or all of segments VI-VIII. Oral-collar tubular ducts of 2 distinct sizes,

scattered over surface, without conspicuous collar. Ostioles obvious.

Anal ring on dorsal surface touching apex of abdomen, with 2 rows of pores;

anal-ring setae with apex slightly rounded or swollen, longest seta 1 16-165 (135)

ix long, longest seta 1.9-2.4 (2.1) times longer than greatest diameter of anal ring.

Venter with 1 kind of seta; bristle-shaped setae with longest seta on segment

VII 27-37 (30) ix long; with 3 pairs ofcisanal setae, longest seta 28-47 (37) n long;

longest anal-lobe seta 123-138(131)^ long; length of longest anal-ring seta/length

of longest anal-lobe seta 0.8-1.2 (1.0). Multilocular pores in medial areas near

anterior margin of segments VI or VII-IX, in medial areas near posterior margin

of segments III, IV, or V-IX, in lateral areas on each abdominal segment rare or

absent on III and/or II, multilocular pores present on thorax in clusters surround¬

ing mouthparts, spiracles, and legs. Trilocular and discoidal pores scattered over

surface. Oral-collar tubular ducts of same 2 sizes as on dorsum, present over

surface. Ventral anal lobe weakly sclerotized.

Circulus absent from 4 of 11 specimens, when present, transversely oval, width/

length 1.0-1.7 (1.3), 12-37 (28) fx wide. Hind legs with 42-112 (79) translucent

pores on dorsal surface of hind coxa, 49-86 (66) on ventral surface; with 3 trans¬

lucent pores on 1 femur of 1 specimen, absent from femora of remaining speci¬

mens; hind tibiae with translucent pores on 9 of 10 specimens, when present,

with 4-17 (8) pores arranged in 2 clusters on dorsal surface. Hind femora 101—

114 (108) ix long; tibiae 73-91 (84) ix long; tarsi 52-62 (57) \x long; length offemur/

tibia 1.2-1.5 (1.3); length of tibia/tarsus 1.4-1.6 (1.5). Tibiae with 5-7 (6) setae,

those on lateral surface (as drawn) very small. Leg setae bristle shaped; digitules

usually broken, tarsal digitules apparently reaching tip of claw, digitules of all 3

pairs of legs each apparently with 1 thick seta with capitate apex and 1 thin seta

with acute apex; claw digitules extending beyond apex of claw, with conspicuous

club, about equal in size. Antennae 7-segmented, 168-217 (200) /x long, with

bristle-shaped setae. Labium with basal segment separate, 67-81 (74) ix long. Eye

without associated discoidal pores.

Comparison. — Adult females of Hypogeococcus barbarae and H. gilli superfi¬

cially arc similar by having trilocular pores. In addition to the characters used in

the key, H. barbarae differs by having the anal-lobe cerarius with 5-9 (7) conical

setae, 5-12 (9) trilocular pores, and 0-1 (0) discoidal pores, by having dorsal

multilocular pores absent or present in the dorsomedial areas of the posterior

abdominal segments, 2 distinct sizes of oral-collar tubular ducts, longest anal-ring

seta 116-165 (135) /x long, 3 pairs of cisanal setae, longest anal-lobe seta 123—

138 (131) ix long, multilocular pores present around mouthparts, hind coxae with

42-112 (79) translucent pores on the dorsal surface, hind tarsi each 52-62 (57) /x

long, hind tibiae each with 5-7 (6) setae, antennae each 168-217 (200) /x long.

194

PAN-PACIFIC ENTOMOLOGIST

Hypogeococcus gilli has the anal-lobe cerarii each with 9-14 (12) conical setae,

17-21 (19) trilocular pores, and 1-3 (2) discoidal pores, has the dorsal multilocular

pores absent from the medial areas, 1 size of oral-collar tubular duct, longest anal-

ring seta 205-217 (213) g long, 2 pairs of cisanal setae, longest anal-lobe seta

163-187 (177) g long, multilocular pores absent from area around mouthparts,

hind coxae each with 15-21 (18) translucent pores on the dorsal surface, hind

tarsi each 87-103 (97) g long, hind tibiae each with 9-10 (9) setae, antennae each

246-273 (256) g long.

Hypogeococcus boharti Miller, New Species

Etymology. — The species epithet is given in recognition of the many scientific

accomplishments of Dr. Richard M. Bohart and of the many hours that he spent

teaching me about insect systematics. Dick’s never ending enthusiasm and energy

for acquiring new knowledge about the taxonomy of insects has been an important

inspiration to my career as a systematist.

Although most know him as an authority of the Hymenoptera, he also has

contributed to the field of coccidology. In 1942 he published a paper on the life

history of the armored-scale pest Diaspis boisduvalii Signoret, and it remains the

most important work on the biology of the species. In 1958 he collected an

undescribed species of mealybug that later was named in his honor ( Rhizoecus

boharti McKenzie) by McKenzie (1960). It was through Dick Bohart’s adminis¬

trative efforts that Howard L. McKenzie was hired at the University of California

at Davis and the Gordon F. Ferris collection of scale insects was obtained and

incorporated in the Entomology Department’s Collection. Dick’s encouragement

and advice had major impact on the highly productive, later years of Howard

McKenzie’s research career. Therefore, this species is named in honor of Dick

Bohart for his several contributions to the field of scale-insect systematics. It

seems especially appropriate that the epithet “boharti” be given to this orchid

infesting species since his 1942 paper was about a pest of Cattleya orchids.

Type data. — Holotype adult $, right specimens of 3 on slide, with left label

“Hypogeococcus/DRAW/On Orchidaceae/leaf Mexico/(Coyutla Vera Cruz)/at

Hidalgo 1359 1-1-1974 75-6363/D. R. Riley coir.”; right label has a map showing

position of holotype and “Hypogeococcus/boharti/Miller/HOLOTYPE” (USNM).

In addition to the holotype there are 44 paratype specimens on 22 slides. The

paratype data are as follows: BELIZE—On orchid, III-6-70, J. C. Buff. CANAL

ZONE-On orchid, IV-7-75, H. L. Rubin. EL SALVADOR-On Oncidium or-

nithorhynchwn, 11-25-69, B. E. Davis, A. J. Benton. GUATEMALA—On On¬

cidium cristagolli, VI-10-65, R. Reasoner; on orchid, V-30-80, F. Matthews.

HONDURAS—On orchid, III-7-77, W. Forster; on orchid, 1-5-81, J. Torres; on

orchid, VIII-26-81, F. Matthews; on Oncidium sp., 11-18-75, F. Matthews; on

Orchidaceae, VII-31-68, L. R. Gillogly. MEXICO—On Odontoglossum bicto-

niense, VI-18-73, G. L. Piro; on orchid, V-21-51, Cary; on Sobralia sp., III-29-

38, R. H. Buffham; Coyutla, on Orchidaceae, 1-1-74, D. R. Riley; Oaxaca, on

orchids, XII-6-65, L. R. Gillogly; Vera Cruz, on Coffea arabica, II-1-67, G. K.

Browne; Vera Cruz, on orchid, 1-1-74, D. R. Riley. Paratype slides have been

deposited in BMNH, CDAS, FSCA, MCM, MNHP, UCD, UG, UH, USNM,

VPI, and ZIL.

VOLUME 59, NUMBERS 1-4 195

Adult Female (4th Instar)

(Fig. 2)

Description. — Holotype mounted, 1.5 mm long (paratypes 1.6-2.7 (2.0) mm),

0.7 mm wide (paratypes 0.8-1.6 (1.1) mm). Body elongate oval.

Dorsum with 5 and 6 pairs of cerarii (paratypes with 5-7 (6) cerarii). Anal-lobe

cerarius, including ventral area with conical setae, with basal sclerotization, 7

bristle-shaped setae, 9 and 8 conical setae, 6 multilocular pores, 3 discoidal pores

(paratypes with 5-7 (6) bristle-shaped setae, 3-9 (7) conical setae, 1-6 (4) mul¬

tilocular pores, 1-5 (3) discoidal pores). Penultimate cerarius with basal sclero¬

tization, 11 conical setae (paratypes with 7-14 (10) conical setae) and several

bristle-shaped setae, oral-collar tubular ducts, multilocular and discoidal pores.

Antepenultimate cerarius with slight basal sclerotization, 7 and 8 conical setae

(paratypes with 5-11 (8) conical setae), and 1 or more bristle-shaped setae, oral-

collar tubular ducts, multilocular and discoidal pores. Fourth cerarius with or

without weak basal sclerotization, 6 and 7 conical setae (paratypes with 3-7 (6)

conical setae), and 1 or more oral-collar tubular ducts and discoidal pores. Fifth

cerarius with 4 and 5 conical setae (paratypes with 3-7 (4) conical setae), and 1

or more oral-collar tubular ducts and discoidal pores. Sixth cerarius absent on 1

side with 2 conical setae on other (paratypes with 0-3 (2) conical setae) and 1 or

more discoidal pores. Seventh cerarius absent (present on 6 of 10 paratypes, with

0-3 (1) conical setae). With 1 kind of seta; bristle-shaped setae longer posteriorly;

longest seta on segment VII, excluding those surrounding cerarian cluster, 55 p

long (paratypes 47-75 (57) /u); segment IV with 33 setae, excluding those in cerarii

(paratypes with 27-35 (31) setae). Trilocular pores absent. Discoidal pores scat¬

tered over surface; small, irregular sclerotized spots also scattered over surface.

Multilocular pores present over surface, least abundant near posterior apex. Oral-

collar tubular ducts with inconspicuous collar, short, in marginal or submarginal

areas of head to segment VII, in mediolateral areas of all segments except pro¬

thorax, mesothorax, and segment VIII (paratypes with oral collars absent from

mediolateral areas of segment VIII and any or all of prothorax, mesothorax,

metathorax, or segment I), in medial areas on all segments except segment VIII

and prothorax to segment 2 (paratypes with medial oral collars absent from

prothorax to segment I, II, III, or IV, and absent from segment VIII). Ostioles

small.

Anal ring dorsal, touching apex of abdomen (3 of 10 paratypes with anal ring

bent around posterior apex of abdomen), with 2 or 3 rows of pores; anal-ring

setae with narrow, acute apices, largest seta 240 p long (paratypes 215-245 (228)

A), longest seta 2.7 times longer than greatest diameter of ring (paratypes 2.3-3.1

(2.6) times).

Venter with bristle-shaped setae only, excluding setae in cerarii; with 3 pairs

of cisanal setae, longest cisanal seta 40 p long (paratypes 27-43 (38) ju); anal-lobe

setae 157 and 153 p long (paratypes 110-173 (138) /u); length of longest anal-ring

seta/length of longest anal-lobe seta 1.5 (paratypes 1.3-2.0 (1.7) units). Multilocu¬

lar pores scattered over surface except anterior margin of medial areas of segments

I-VI (paratypes with multiloculars absent from anterior margin of segments I, II,

or III to V or VI), posterior margin of medial areas of segments I-IV (paratypes

196

PAN-PACIFIC ENTOMOLOGIST

Fig. 2. Adult female holotype of Hypogeococcus boharti n. sp. Coyutla, Vera Cruz, Mexico, 1-1-

74, on Orchidaceae.

with multiloculars absent from posterior segments I, II, or III to IV or V). Tri-

locular pores absent. Discoidal pores and irregular sclerotized pores of same nature

and distribution as on dorsum. Oral-collar tubular ducts of same size and shape

as on dorsum, on marginal or submarginal areas from head to segment VII (para-

types often with oral collars also on segment VIII), in medial and mediolateral

VOLUME 59, NUMBERS 1-4

197

areas of segments IV-YI (paratypes with oral collars from II, III, or IY to VI or

VII). Ventral anal-lobe area broadly sclerotized, with 1 or 2 conical setae, bristle¬

shaped setae, multilocular and discoidal pores.

Circulus nearly round, width/length 1.1 (paratypes 0.9-1.3 (1.0) units), 35 p

wide (paratypes 30-60 (40) /x). Hind legs with 35 and 29 translucent pores on

dorsal surface of each femur arranged in cluster near distal apex of anterior margin

(as drawn) and in cluster along posterior margin (paratypes with 13-35 (26) pores,

occasionally without distal cluster); tibia with 27 and 48 pores on dorsal surface

scattered along outer margin (as drawn) (paratypes with 26-42 (33) pores). Hind

femora 155 and 153 fx long (paratypes 140-165 (151) /x); tibiae 107 and 113 /x

long (paratypes 104-123 (112) /*); tarsi each 67 /x long (paratypes 61-73 (67) /x );

length of femur/tibia 1.4 (paratypes 1.3-1.4 (1.3) units); length of tibia/tarsus 1.6

and 1.7 (paratypes 1.5-1.7 (1.7) units). Tibiae with 8 and 9 setae (paratypes with

9-10 (9) setae). Leg setae bristle shaped; tarsal digitules not quite reaching tip of

claw, digitules of hind 2 pairs of legs approximately equal in length, 1 noticeably

more slender than other including apical club, digitules of front legs unequal, 1

without apical club, other similar to digitules of hind tarsi; claw digitules extending

beyond apex of claw, equal in size, slightly larger than largest tarsal digitule.

Antennae 6- and 7-segmented (paratypes 6- and 7-(7-)segmented), 6-segmented

antenna with third segment partially divided, 263 and 250 ix long (paratypes 245-

310 (275) A), with bristle-shaped setae. Labium with basal segment separate, 90

fx long (paratypes 93-113 (99) p). Eye with 2 and 3 loosely associated discoidal

pores (paratypes with 1-3 (2) pores).

Third-Instar Female

(Fig. 3)

Description.— Mounted, 1.1 and 1.2 mm long, 0.5 and 0.6 mm wide. Body

elongate.

Dorsum with 5-6 (5) pairs of cerarii. Anal-lobe cerarius, including ventral area

with conical setae, with basal sclerotization, 2-3 (3) bristle-shaped setae, 4-5 (4)

conical setae, 1-2 (1) multilocular pores, 0-1 (0) discoidal pores. Penultimate

cerarius with basal sclerotization, 5-7 (6) conical setae, and 1 or more bristle¬

shaped setae, discoidal pore, and oral-collar tubular duct. Antepenultimate ce¬

rarius with slight basal sclerotization, 3-5 (4) conical setae. Fourth cerarius with

or without weak basal sclerotization, 3-4 (3) conical setae. Fifth cerarius with 3-

4 (3) conical setae. Sixth cerarius with 0-2 (1) conical seta. With 1 kind of seta;

bristle-shaped setae longer posteriorly; longest seta on segment VII, excluding

those surrounding cerarian cluster, 40 and 43 /x long; segment IV with 18 setae,

excluding those in cerarii. Trilocular pores absent. Discoidal pores scattered over

surface; small, irregular sclerotized spots also scattered over surface. Multilocular

pores present over surface, least abundant medially. Oral-collar tubular ducts of

same structure as on adult, in marginal and subma rginal areas of head to segment

VII, in mediolateral areas of all segments except prothorax to segment I and

segment VIII, in medial areas of all except prothorax to segment III and segment

VIII on 1 specimen, absent from medial area on other specimen. Ostioles small.

Anal ring bent around posterior apex of abdomen, with 2 rows of pores; anal-

ring setae with acute apices, longest seta 145 and 170 ix long, longest seta 1.9 and

2.9 times longer than greatest diameter of ring.

198

PAN-PACIFIC ENTOMOLOGIST

Fig. 3. Third-instar female paratype of Hypogeococcus boharti n. sp. Flonduras, VIII-26-81, on

“orchid leaf.”

Venter with bristle-shaped setae only, excluding setae in cerarii; longest cisanal

seta 27 p long; anal-lobe setae 103 and 115 p long; length of longest anal-ring

seta/length of longest anal-lobe seta 1.3 and 1.7. Multilocular pores scattered over

surface except medial areas of abdomen and mediolateral areas of segments I—III

on 1 specimen and segment II on other. Trilocular pores absent. Discoidal pores

and irregular sclerotized pores scattered over surface. Oral-collar tubular ducts of

VOLUME 59, NUMBERS 1-4

199

same size and shape as on dorsum, on marginal and submarginal areas of head

to segment VII in mediolateral areas of segment V on 1 specimen, V and VI on

other, absent medially. Ventral anal-lobe area broadly sclerotized, with bristle¬

shaped setae, multilocular and discoidal pores.

Circulus nearly round width/length 1.1 and 1.3, 20 and 25 p wide. Hind legs

with traces of translucent pores on femora and tibiae (these pores may be from

the legs of the adult before shedding). Hind femora 105-117 (110)/x long; tibiae

73-80 (76) /x long; tarsi 67-80 (68) p long; length of femur/tibia 1.4-1.5 (1.4);

length of tibia/tarsus 1.1. Tibiae with 9 setae. Leg setae bristle-shaped; digitules

same as on adult. Antennae 6-segmented, 1 antenna with third segment partially

divided, 203-230 (219) p long, with bristle-shaped setae. Labium with basal

segment separate, 75 and 73 ^ long. Eye with 0-2 (1) loosely associated discoidal

pores.

First Instar

Specimens examined.—The following description is based on 42 embryos from

the following 6 localities (complete data given under “Type Data”): Belize; Gua¬

temala (1965 and 1980); Honduras (1-5-1981); Mexico (1938 and 1974).

Description.—O nly salient features are included because of the poor condition

of the available material.

Mounted 326-440 (391) p long, 175-225 (198) p wide. Body elongate.

Dorsum with 4-5 (4) pairs of cerarii that have at least 1 conical seta, with up

to 14 paired, enlarged bristle-shaped setae along each body margin including those

with conical setae; cerarii usually without associated pores, without basal scler-

otization. Anal-lobe cerarii with 2 conical setae, 1 trilocular pore between cerarian

setae; penultimate and antepenultimate cerarii each with 2 conical setae; fourth

cerarii with 1-2 (1) conical setae, 0-1 (0) bristle-shaped setae; fifth cerarii with

0-1 (0) conical setae, 1-2 (2) bristle-shaped setae. With 1 kind of seta; bristle¬

shaped setae scattered over dorsum, arranged in 4 longitudinal lines on most of

abdomen, shortest setae in center of body; longest seta on segment VII 17-27

(23) p long, segment IV with 4 setae excluding those in cerarii. Trilocular pores

scattered over surface, on abdomen usually with 1 pore between each cerarius

and mediolateral seta, and 1 pore between each mediolateral seta and medial seta,

located between anterior and posterior margin of each segment, forming 4 lon¬

gitudinal lines of trilocular pores on abdomen. Discoidal and multilocular pores

absent. Tubular ducts absent. Ostioles present.

Anal ring apical, with 2 rows of pores; anal-ring setae with acute apices, longest

setae 54-73 (63) p long.

Venter with 1 kind of seta, forming 6 longitudinal lines of setae on abdomen;

bristle-shaped setae scattered over surface, longest seta 16-37 (25) p long; longest

anal-lobe seta 50-55 (52) p long. Trilocular pores associated with submarginal

and mediolateral lines of setae on abdomen. Multilocular pores absent. Discoidal

pores associated with submarginal line of setae. Tubular ducts absent. Ventral

anal-lobe area not sclerotized.

Circulus oval, not divided, 8-11 (10) p wide. Legs unclear; tibia with 7-8 (7)

setae; leg setae bristle shaped; tarsal digitules extending beyond tip of claw, dig¬

itules of hind 2 pairs of legs equal in length, 1 noticeably more slender than other,

digitules of front legs unequal, 1 without apical club, other about same as larger

200

PAN-PACIFIC ENTOMOLOGIST

digitule on hind legs; claw digitules extending beyond tip of claw, 1 digitule slightly

thinner than other. Antennae unclear. Labium details unclear. Eye with 1 asso¬

ciated discoidal pore.

Comparison.— Adult females of H. boharti and H. othnius are similar; both

lack dorsomedial conical setae, have 2 or 3 rows of pores in the anal ring, have

apices of the anal-ring setae slightly rounded or capitate, have the length of the

longest anal-ring seta/the greatest diameter of the anal ring about 2.6, have the

longest anal-ring seta about 220 y long, have translucent pores on the femur and

tibia, usually have a discoidal pore associated with the eye, and occur on orchids.

Hypogeococcus boharti differs by having more than 2 conical setae in each of the

posterior 4 pairs of cerarii, and circulus oval, rarely longer than wide; H. othnius

has 2 conical setae in each of posterior 4 pairs of cerarii (1 specimen has 1 cerarius

with 3 conical setae), and circulus elongate, usually longer than wide. The above

mentioned species share several similarities with H. gilli', all have the apex of the

anal-ring setae rounded or capitate, usually have 7-segmented antennae, and have

9 or 10 setae on each hind tibia. Hypogeococcus gilli is easily separated by having

trilocular pores, by lacking a circulus, by having many pores near the base of the

cerarian conical setae in the anal-lobe cerarius, and by having translucent pores

on the hind coxa; H. boharti and H. othnius lack trilocular pores, have a circulus,

lack pores near the base of the cerarian conical setae in the anal-lobe cerarius,

lack translucent pores on the hind coxa.

A comparison of H. boharti and H. margaretae, is given in the species treatment

of the latter species.

Hypogeococcus gilli Miller, New Species

Etymology .—The species epithet is named in honor of Mr. Raymond J. Gill,

California Department of Food and Agriculture, Sacramento, not only for pro¬

viding the type series of this species, but also for his dedication and many im¬

portant contributions to the study of scale-insect systematics.

Type data.— Holotype adult 9, single specimen on slide, with left label “Hy-

pogeogoccus/gilli Miller/HOLOTYPE/Det. by CDA/Vn-62/C 4 N 8 0 AF + LP-B.”;

right label “sp. 13033/No. 62G20-55 Cal. Dept. Agr./Q. fr./Loc. Costa Rica at

San/Pedro, Cahfomia/17-VII-1962/ex. orchid leaf/Draw/J. Lindsay coll” (USNM).

In addition, there are 2 adult female paratypes on 2 slides with the same data

(CDAS and BMNH).

Adult Female (4th Instar)

(Fig. 4)

Description.— Holotype, mounted, 1.7 mm long (paratypes 1.8 and 2.0 (1.9)

mm) 0.8 mm wide (paratypes each 1.0 mm). Body elongate oval.

Dorsum with 5 pairs of cerarii, sixth pair reduced to single conical seta on 1

side of specimen, with 2 such setae on other side (1 paratype with definite sixth

cerarius with basal sclerotization). Anal-lobe cerarius part of sclerotized plate on

dorsum of segment VIII, with 11 conical setae, 17 trilocular pores, 2 discoidal

pores (paratypes with 9-14 (12) conical setae, 17-21 (19) trilocular pores, and 1-

3 (2) discoidal pores). Penultimate cerarius defined by basal sclerotization ex¬

tending on to ventral surface with 11 and 12 conical setae with 2 larger than

others (paratypes with 11-18 (14) conical setae), with cluster of basal trilocular

VOLUME 59, NUMBERS 1-4 201

Fig. 4. Adult female holotype of Hypogeococcus gilli n. sp. Costa Rica (quarantined at San Pedro,

California), VII-17-62, on “orchid leaf.”

pores and discoidal pores. Antepenultimate cerarius with 8 and 11 conical setae

with 2 larger than others (paratypes with 5-9 (7) conical setae), basal sclerotization,

cluster of trilocular pores. Fourth cerarius with 3 and 5 conical setae with 2 larger

than others (paratypes with 2-5 (3) conical setae), basal sclerotization, few triloc¬

ular pores. Fifth cerarius with 2 conical setae (paratypes with 2-3 (2) such setae),

202

PAN-PACIFIC ENTOMOLOGIST

basal sclerotization, 1 or 2 trilocular pores. With 2 kinds of setae; conical setae

in medial and mediolateral areas of segments IV-VII, in mediolateral areas only

of segments III and VIII; remainder of dorsum with bristle-shaped setae; longest

seta on segment VII, excluding those surrounding cerarian cluster, 25 /x long

(paratypes 25 and 27 fx), segment IV with 29 setae excluding those in cerarii

(paratypes with 35 and 37). Trilocular and discoidal pores scattered over surface,

discoidals less abundant. Multilocular pores in marginal and submarginal areas

of segment II-VII on 1 side, II, IV-VII on other side (paratypes with these pores

from metathorax, segments I, II, or III-VII). Oral-collar tubular ducts incon¬

spicuous, with unusually short duct and darkly staining collar in marginal or

submarginal areas of prothorax, segments I-VII on 1 side, II-VII on other side

(paratypes without dorsal oral-collar tubular ducts on thorax). Ostioles obvious.

Anal ring on dorsal surface touching apex of abdomen, with 4 or 5 rows of

pores; anal-ring setae broken at apex except 1, apex of unbroken seta rounded

(paratypes with apices of anal-ring setae rounded), longest. 217 y long (paratypes

205 and 216 /x ), longest seta 2.2 times longer than greatest diameter of anal ring

(paratypes each 2.2 times).

Venter with 2 kinds of setae; conical setae restricted to marginal and submarginal

areas of abdominal segments; bristle-shaped setae with longest seta on segment

VII 45 ju long (paratypes 30 n long); with 2 pairs of cisanal setae, longest seta 50

/u long (broken on paratypes); longest anal-lobe seta 187 long (paratypes 163

and 180 /x long); length of longest anal-ring seta/length of longest anal-lobe seta

1.2 (paratypes 1.2 and 1.3). Multilocular pores in medial and mediolateral areas

of segments V-VIII (paratypes VI-VIII), in marginal and submarginal areas of

segments 1I-IV, on thorax anterior of each spiracle, represented by 1, 2, or 3 pores

in each cluster (paratypes with thoracic clusters each represented by 1-5 (3) pores).

Trilocular and discoidal pores scattered over surface, least abundant near center

of surface. Oral-collar tubular ducts of same size and shape as on dorsum generally

in same areas as multiloculars. Ventral anal-lobe area broadly sclerotized.

Circulus absent. Hind legs with 22 translucent pores on dorsal surface of each

coxa (paratypes 15-21 (18) pores), 23 and 25 pores on ventral surface of each

coxa (paratypes 11-29 (21) pores); tibia with 3 and 6 pores on dorsal surface

(paratypes 2-9 (5) pores), absent from ventral surface. Hind femora 125 and 127

g long (paratypes 113-135 (127) g)\ tibiae 87 and 93 ju long (paratypes 93-103

(97) fi)\ tarsi 73 and 75 /x long (paratypes 70-75 (73) ju); length of femur/tibia 1.4

(paratypes 1.2-1.7 (1.4) units); length of tibia/tarsus 1.2 (paratypes 1.2-1,4 (1.3)

units). Leg setae bristle shaped except 2 setae at distal apex of inner side of tibia;

tibia with 9 and 10 setae (paratypes 9-10 (9) setae); tarsal digitules extending

beyond apex of claw, apical clubs of unequal size; claw digitules extending beyond

apex of claw, equal in size, about equal in size to largest tarsal digitule. Antennae

7-segmented, 260 ix long (paratypes 246-273 (256) g). Labium with basal segment

separate, 75 y long (paratypes 87 and 100 ix ). Eye without associated pores.

Comparison. —Hypogeococcus gilli is similar to H. barbarae by having trilocular

pores. For a comparison of these species see the description of the latter. A

comparison of H. gilli is also made in the description of H. boharti.

Hypogeococcus hamoni Miller, New Species

Etymology. — The species epithet is named in honor of Dr. Avas B. Hamon,

VOLUME 59, NUMBERS 1-4

203

Florida Department of Agriculture and Consumer Services, Gainesville, for pro¬

viding specimens of many interesting mealybugs from Florida including this species.

Type data. — Flolotype adult 2, single specimen on slide, with left label “Hy-

pogeococcus/hamoni/Miller/HOLOTYPE , ’; right label “Palatka, Fla./17 VI 1980/

on/Eriocaulon/compressum/DPI# 126354/draw imm. USNM/Balsam” (USNM).

In addition, there are 2 adult female paratypes on 2 slides with the same data

(FSCA and BMNH).

Adult Female (4th Instar)

(Fig. 5)

Description.— Holotype, mounted 1.6 mm long (paratypes 1.6 and 1.8 mm),

0.9 mm wide (paratypes 0.8 and 1.0 mm). Body elongate.

Dorsum with 4 and 5 pairs of cerarii (paratypes with 5). Anal-lobe cerarius,

including ventral area with conical setae, with basal sclerotization, 0 and 1 bristle¬

shaped setae, 10 and 11 conical setae, no pores (paratypes with 10-11 (10) conical

setae, 0-2 (1) bristle-shaped setae, no multilocular or discoidal pores). Penultimate

cerarius with weak basal sclerotization, 5 conical setae, without pores (paratypes

with 3-4 (4) conical setae). Antepenultimate cerarius without basal sclerotization,

2 and 3 conical setae, no associated pores (paratypes with 2 conical setae). Fourth

cerarius without basal sclerotization, 2 conical setae, no pores (paratypes also with

2 conical setae). Fifth cerarius absent from 1 side, with 1 conical seta on other,

without pores (paratypes with 1-2(1) conical setae). With 2 kinds of setae; basally

enlarged bristle-shaped setae restricted to medial areas of segments VI and VII;

bristle-shaped setae scattered elsewhere, all about same size except larger near

anterior and posterior apices of body; longest seta on segment VII, excluding those

surrounding cerarian cluster, 40 p long (paratypes 40 and 45 p), segment IV with

18 setae, excluding those in cerarii (paratypes with 16 and 19 setae). Trilocular

pores absent. Discoidal pores scattered over surface. Multilocular pores present

over surface, least abundant in center of body and near posterior apex. Oral-collar

tubular ducts with inconspicuous collar, short, in marginal or submarginal areas

of head to segment VII, in medial and mediolateral areas of segments III-VII

(paratypes with medial or mediolateral oral collars on segments IV and V on 1

specimen, on III-VII on other). Ostioles small and inconspicuous.

Anal ring dorsal, touching apex of abdomen, with 7-9 rows of pores; anal-ring

setae with acute apices, longest seta 143 p long (paratypes 143 and 140 p ), longest

seta 1.4 times longer than greatest diameter of ring (paratypes 1.5 and 1.4 times).

Venter with bristle-shaped setae only, excluding setae in cerarii; longest seta on

segment VII 25 p long (paratypes 37 p)\ with 2 pairs of cisanal setae, longest

cisanal seta 40 p long (paratypes 40 and 30 p)\ anal-lobe setae broken on all

specimens. Multilocular pores scattered over surface, most abundant near pos¬

terior margin of segments V-VII, sparse in medial areas of segments I—III. Triloc¬

ular pores absent. Discoidal pores in small numbers over surface. Oral-collar

tubular ducts of same size and shape as on dorsum; in marginal or submarginal

areas of prothorax to segment VIII, in medial or mediolateral areas of segments

V-VII (paratypes with marginal and submarginal oral collars from head or pro¬

thorax to segment VII, medial oral collars absent from 1 specimen, on segments

VI and VII on other). Ventral anal-lobe area broadly sclerotized, with several

bristle-shaped setae.

204

PAN-PACIFIC ENTOMOLOGIST

Fig. 5. Adult female holotype of Hypogeococcus hamoni n. sp. Palatka, Florida, VI-17-80, on

Eriocaulon compressa.

Circulus oval, width/length 1.3 (paratypes 1.4 units) not divided, 29 p wide

(paratypes 31 and 33 p). Hind legs with 14 and 17 translucent pores on dorsal

surface of each coxa (paratypes 11-19 (15) pores), 9 and 11 pores on ventral

surface of each coxa (paratypes 10-21 (17) pores); tibia without pores 1 side with

7 pores on other (paratypes 0-4 (2) pores). Hind femora 177 and 180 p long

VOLUME 59, NUMBERS 1-4

205

(paratypes 187-195 (191) ju); tibiae 150 and 160 y long (paratypes 157-167 (164)

y); tarsi 83 and 95 y long (paratypes 93-103 (99) a 0; length of femur/tibia 1.1 and

1.2 (paratypes 1.1-1.2 (1.2) units); length of tibia/tarsus 1.7 and 1.8 (paratypes

1.6-1.8 (1.7) units). Tibia with 7 setae (paratypes with 5-7 (6) setae). Leg setae

bristle shaped; tarsal digitules not quite reaching tip of claw, digitules of hind 2

pairs of legs unequal in length, with apical clubs of approximately same size,

digitules of front legs unequal, 1 without apical club, other similar to digitule of

hind tarsi; claw digitules extending beyond apex of claw, equal in size, about equal

to largest tarsal digitule. Antennae 6-segmented (paratypes 6-segmented and

7-segmented), with third segments partially divided, 217 y long (paratypes 210-

230 (222) y), with bristle-shaped setae. Labium with basal segment fused with

second segment, 85 y long (paratypes 85 and 87 y). Eyes without associated pores.

Comparison. — Adult females of Hypogeococcus hamoni and H. margaretae are

similar; both lack multilocular pores associated with anal-lobe cerarius, have the

anal ring with 7-9 rows of pores, have translucent pores on the hind coxa and

tibia, have the hind femora longer than 177 y and the hind tibia longer than 140

y, and have the basal labial segment closely appressed to the second segment.

Hypogeococcus hamoni differs by having 10-11 (10) conical setae in each anal-

lobe cerarius, having few or no enlarged bristle-shaped setae on the dorsum, having

16-19 (18) dorsal setae on segment IV excluding those in the cerarii, having short

setae (longest seta on dorsum of segment VII 40-45 y long, longest seta on venter

of segment VII 25-37 y long, longest cisanal seta 30-40 y long), having 5-7 (6)

setae on each hind tibia, and having 6- or 7-(6-)segmented antennae; H. mar¬

garetae has 24-40 (26) conical setae in each anal-lobe cerarius, has numerous,

conspicuous enlarged bristle-shaped setae over the dorsum, has 33-48 (42) dorsal

setae on segment IV excluding those in cerarii, has long setae (longest seta on

dorsum of segment VII 57-80 (68) y long, longest seta on venter of segment VII

35-50 (42) y long, longest cisanal seta 60-123 (85) y long), has 8-9 (9) setae on

each hind tibia, and has 5- or 6-(5-)segmented antennae. The above mentioned

species share with H. spinosus the occurrence of enlarged leg setae, short anal-

ring setae, and the basal labial segment which is closely appressed with the second

segment. Hypogeococcus spinosus is readily separated by having dorsomedial

conical setae on abdominal segments II and III, having 2 or 3 rows of pores in

the anal ring, having 4 pairs of cisanal setae, having 7-segmented antennae, and

having hind length of tibia/tarsus about 1.3; H. hamoni and II. margaretae differ

by having dorsomedial conical setae absent from abdominal segments II and III,

having 7-9 rows of pores in the anal ring, having 3 pairs of cisanal setae, usually

having less than 7-segmented antennae, and having hind length of tibia/tarsus

about 1.7 or 1.8.

Hypogeococcus margaretae Miller, New Species

Etymology. —The species epithet is formed in honor of Mrs. Margaret E. Russell

Bohart, who has been a major contributor to the scientific achievements of the

Bohart regime at Davis. It is clear that many of the research and administrative

achievements of Dick Bohart could not have been accomplished without Mar¬

garet’s devotion to science and support of many, necessary behind-the-scenes

activities. Therefore, it is appropriate that Margaret Bohart be honored along with

her husband.

206

PAN-PACIFIC ENTOMOLOGIST

Type data .—Holotype adult 2, single specimen on slide, with left label “Hy-

pogeococcus/m< 2 r < gflrpZflp/Miller/HOLOTYPE”; right label “nr. Port St. Lucie,/

0.3 mi N. Hiway 609/on Hiway 709, St./Lucie Co., Fla./l 7-XII-1981/K. Hibbard/

on Eriocaulon/decangulare/DPI-126357” (USNM). In addition there are 11 para-

types on 11 slides from the same locality collected on the same date, 25 paratypes

on 25 slides from the same locality collected 28-VII-1982, and 4 paratypes on 2

slides from Jackson Co., Mississippi, V-2-76, B. J. Muse, on Eriocaulon sp.

Paratype slides have been deposited in AUA, BMNH, CD AS, FSCA, IZAS, MCM,

MNHP, SIE, UCD, UG, UH, USNM, VPI, and ZIL.

Adult Female (4th Instar)

(Fig. 6)

Description. — Holotype, mounted, 1.8 mm long (paratypes 1.4-2.6 (2.1) mm),

0.9 mm wide (paratype 0.8-1.3 (1.0) mm). Body elongate.

Dorsum with 4 and 6 pairs of cerarii (paratypes with 4-6 (5) cerarii). Anal-lobe

cerarius including ventral area with conical setae, with basal sclerotization, 25

and 26 conical setae, no pores (paratypes with 24-30 (26) conical setae, 0-3 (0)

enlarged, bristle-shaped setae, no multilocular pores, 0-1 (0) sclerotized discoidal

pores). Penultimate cerarius defined by basal sclerotization extending onto ventral

surface with 13 and 15 conical setae, without pores (paratypes with 12-16 (14)

conical setae). Antepenultimate cerarius with basal sclerotization, 5 and 8 conical

setae, no associated pores (paratypes with 4-8 (6) conical setae). Fourth cerarius

with basal sclerotization, 3 and 4 conical setae, no pores (paratypes with 3-6 (4)

conical setae). Fifth cerarius without basal sclerotization, with 1 and 2 conical

setae, without associated pores (paratypes with or without basal sclerotization,

with 1-5 (2) conical setae). Sixth cerarius absent from 1 side, on other side without

basal sclerotization and associated pores, with 2 conical setae (paratypes with

sixth cerarius on 6 of 10 specimens, when present, without basal sclerotization

and associated pores, with 1-2 (1) conical setae). With 2 kinds of setae; conical

setae on medial areas of segments VI and VII (on paratypes on any or all of

segments IV-VII), on mediolateral areas of segments V-VII (on paratypes on any

or all of segments III-VII), on submarginal areas of segment VII (paratypes usually

with these setae on 1 or more of segments III-VII); enlarged, bristle-shaped setae

scattered over remainder of surface, largest posteriorly and anteriorly, smallest in

center of body ; longest seta on segment VII, excluding those surrounding cerarian

cluster, 63 p long (paratypes 57-80 (68) p), segment IV with 42 setae, excluding

those in cerarii (paratypes with 33-48 (42) setae). Trilocular pores absent. Dis¬

coidal pores represented by small, irregular sclerotized spots, distributed in same

relative abundance as multilocular pores. Multilocular pores scattered over sur¬

face, least abundant on posterior portion of abdomen, becoming numerous on

abdominal segments I or II, numerous on thorax and head. Oral-collar tubular

ducts with inconspicuous collar, unusually short for mealybug, in marginal or

submarginal areas of segments V-VII (paratypes with ducts absent or on 1, and

some, or all of segments IV-VII). Ostioles small and usually inconspicuous.

Anal ring dorsal, usually touching apex of abdomen, with 7-9 rows of pores;

anal-ring setae with acute apices, longest seta 133 p long (paratypes 125-150 (139)

p), longest seta 1.4 times longer than greatest diameter of ring (paratypes 1.3-1.7

(1.5) times).

VOLUME 59, NUMBERS 1-4

207

Fig. 6. Adult female holotype of Hypogeococcus margaretae n. sp. near Port St. Lucie Co., Florida,

XII-17-81, on Eriocaulon decangulare.

Venter with 3 kinds of setae; conical setae restricted to marginal and submarginal

areas of abdomen; enlarged, bristle-shaped setae in marginal and submarginal

clusters except on anterior apex of head, medial setae on segments III-V slightly

enlarged; bristle-shaped setae in medial and mediolateral areas of abdomen, thorax,

and head, excluding segments III-V, also on anterior apex of head; longest bristle-

208

PAN-PACIFIC ENTOMOLOGIST

shaped seta on segment VII 45 p long (paratypes 35-50 (42) ^u); with 2 pairs of

cisanal setae, longest seta 77 g long (paratypes 60-123 (85) g)] longest anal-lobe

seta 170 g long (paratypes 163-205 (182) g ); length of longest anal-ring seta/length

of longest anal-lobe seta 0.8 (paratypes 0.7-0.9 (0.8) units). Multilocular pores

abundant over surface, absent or sparse in medial areas of segments II-IV or V.

Trilocular pores absent. Discoidal pores of same nature and distribution as on

dorsum. Oral-collar tubular ducts of same size and shape as on dorsum, on

marginal or submarginal areas of segments II or III-VII (paratypes often with

fewer ducts on fewer segments). Ventral anal-lobe area broadly sclerotized, some¬

what indefinite anteriorly, apparently with conical setae, bristle-shaped setae, and

multilocular pores.

Circulus oval, width/length 1.5 (paratypes 1.3-1.7 (1.5) units) not divided, 25

A* wide (paratype 20-30 (25) g). Hind legs with 30 and 35 translucent pores on

dorsal surface of each coxa (paratypes 30-46 (38) pores), 18 and 26 pores on

ventral surface of each coxa (paratypes 20-44 (27) pores); tibia without pores

(paratypes without pores on 5 of 10 specimens, when present, located on dorsal

surface at distal apex, with 1-6 (3) pores). Hind femora 210 and 213 g long

(paratypes 195-217 (205) ^); tibiae 167-170 g long (paratypes 147-170 (159) g);

tarsi 93 and 90 g long (paratypes 80-90 (87) g); length of femur/tibia 1.3 (paratypes

1.2-1.4 (1.3) units); length of tibia/tarsus 1.8 (paratypes 1.7-1.9 (1.8) units). Tibia

with 9 setae (paratypes 8-9 (9) setae). Leg setae bristle shaped except for some

setae on trochanter, femur, and tibia as follows: hind trochanter with enlarged

bristle-shaped seta on lower-distal portion (as drawn) (paratypes sometimes with

2 enlarged at distal end); hind femur with enlarged, bristle-shaped setae on ventral

surface at distal end of segment and posterior edge (as drawn) (paratypes often

with setae on anterior edge also enlarged); hind tibia with enlarged, bristle-shaped

setae on ventral surface at distal end of segment and along anterior margin (as

drawn) (paratypes occasionally with enlarged seta on posterior margin near middle

of segment); front legs with few enlarged, bristle-shaped setae on femur (paratypes

often with many more enlarged setae on front femur); tarsal digitules usually not

quite reaching tip of claw, digitules of hind 2 pair of legs unequal in length, with

apical clubs of approximately same size (1 or 2 paratypes with digitules apparen tly

of same length), digitules of front legs unclear (paratypes with front digitules

unequal, 1 without apical club, other similar to digitule of hind tarsi); claw digitules

extending beyond apex of claw, equal in size, about equal to largest tarsal digitule.

Antennae 5-segmented (3 of 10 paratypes with third segment partially divided),

255 and 260 g long (paratypes 225-260 (245) /u); with enlarged, bristle-shaped

setae on anterior margin (as drawn) of segments 1-5 (paratypes on segments 1-

3, 4, or 5). Labium with basal segment closely attached to second segment, 87 g

long (80-100 (89) g). Eye without associated pores.

Comparison.—A comparison is given of the adult females of Hypogeococcus

margaretae and H. hamoni in the treatment of the latter species.

Third-Instar Female

(Hg. 7)

Specimens examined.—The following description is based on 3 specimens col¬

lected near Port St. Lucie, Florida on 28-VII-1982.

Description.— Mounted, 1.1-1.2 (1.2) mm long, 0.5-0.6 (0.5) mm wide. Body

elongate.

VOLUME 59, NUMBERS 1-4

209

Fig. 7. Third-instar female paratype of Hypogeococcus margaretae n. sp. near Port St. Lucie,

Florida, VII-28-82, on Eriocaulon decangulare.

Dorsum with 4 or 5 (5) pairs of cerarii. Anal-lobe cerarius with basal sclero-

tization, 12-14 (13) conical setae, 1-3 (2) enlarged, bristle-shaped setae, 1 spec¬

imen with 3 discoidal pores, remaining specimens without pores. Penultimate

cerarius with basal sclerotization, 6 or 7 conical setae, 1-3 (2) enlarged, bristle¬

shaped setae, no pores. Antepenultimate cerarius with basal sclerotization, 4-5

210

PAN-PACIFIC ENTOMOLOGIST

(4) conical setae, 0-1 (1) enlarged, bristle-shaped setae, no pores. Fourth cerarius

with basal sclerotization, 1-5 (3) conical setae, 0-2 (1) enlarged, bristle-shaped

setae, no pores. Fifth cerarius, when present, with 1-3 (2) conical setae, 1-2 (1)

enlarged, bristle-shaped setae, no pores. With 1 or 2 kinds of setae; 1 conical seta

present on 1 specimen, located dorsomedially on segment VII, absent from re¬

maining specimens; enlarged, bristle-shaped setae scattered over surface; longest

seta on segment VII, excluding those surrounding cerarian cluster, 40-49 (45) g

long, segment IV with 19-22 (20) setae, excluding those in cerarii. Trilocular pores

absent. Discoidal pores represented by small, irregular sclerotized spots, distrib¬

uted in same relative abundance as multilocular pores. Multilocular pores present

over surface, most abundant on thorax. Oral-collar tubular ducts in marginal or

submarginal areas on any or all of segments 11-VII. Ostioles small.

Anal ring dorsal, touching apex of abdomen, with 5 or 6 (6) rows of pores; anal-

ring setae with acute apices, longest seta 111-119 (115) g long, longest seta 1.5-

1.7 (1.6) times longer than greatest diameter of ring.

Venter with 2 kinds of setae; abdominal segments each with 10 or more setae;

enlarged, bristle-shaped setae in marginal and submarginal areas except on head,

with 3-5 setae cluster near each lateral margin of each segment; bristle-shaped

setae scattered over remainder of surface; longest bristle-shaped seta on segment

VII 32-47 (41) g long; with 2 pairs of cisanal setae, longest seta 52-62 (58) g long;

longest anal-lobe seta 143-148 (145) g long; length of longest anal-ring seta/length

of longest anal-lobe seta 0.8. Multilocular pores most abundant on thorax, usually

absent from medial areas of segments I or II to VI or VII. Trilocular pores absent.

Discoidal pores of same nature and distribution as on dorsum. Oral-collar tubular

ducts on marginal and submarginal areas of metathorax and all or some of seg¬

ments III to VII. Ventral anal-lobe area broadly sclerotized, with several enlarged,

bristle-shaped setae, sometimes with 1 or 2 conical setae, 1 bristle-shaped seta,

several multilocular pores.

Circulus oval, width/length 1.1-1.3 (1.2), 17-20 (18) g wide. Hind legs with

trace of translucent pores on coxa; hind femora 138-146 (142) g long; tibiae 101 —

109 (105) g long; tarsi 77-79 (78) g long; length of femur/tibia 1.3-1.4 (1.3); length

of tibia/tarsus 1.3-1.4 (1.4). Tibia with 7 setae. Leg setae bristle shaped except

for some enlarged, bristle-shaped setae on trochanter, femur, and tibia; tarsal

digitules of hind 2 pairs of legs unequal in length, with apical clubs of approxi¬

mately same size, digitules of front legs unequal, 1 without club, other similar to

digitules of hind tarsi; claw digitules extending beyond apex of claw, equal in size.

Antennae 5-segmented, third segment sometimes partially divided, 180-203 (192)

g long; enlarged, bristle-shaped setae on anterior margin (as drawn) of segments

1-3 or 4. Labium with basal segment closely attached to second segment, 77-79

(78) g long. Eyes without associated pore.

Comparison.— Third instar of Hypogeococcus margaretae and H. boharti differ

by the former having 12-14(13) conical setae in each anal-lobe cerarius, enlarged,

bristle-shaped setae scattered over dorsum, oral-collar tubular ducts absent from

head and medial areas, anal ring with 5 or 6 rows of pores, longest anal-ring seta

111-119 (115) g long, longest cisanal seta 52-62 (58) g long, longest anal-lobe

seta 143-148 (145) g long, hind femur 138-146 (142) g long, tibia/tarsus ratio

1.3-1.4 (1.4), tibiae each with 7 setae, some leg setae enlarged, antennae 5-seg-

mented, eye without associated pores; H. boharti has 4-5 (4) conical setae in each

VOLUME 59, NUMBERS 1-4

211

anal-lobe cerarius, no enlarged, bristle-shaped setae, oral-collar tubular ducts on

head and dorsomedial areas of abdomen, anal ring with 2 rows of pores, longest

anal-ring seta 145-170 g, longest cisanal seta 27 g long, longest anal-lobe seta

103-115 g long, hind femur 105-117 (110) g long, length of tibia/tarsus ratio 1.1,

tibiae each with 9 setae, no enlarged leg setae, antennae 6-segmented, eye with

associated pores.

Third-instar females of H. margaretae can be separated from second-instar

females of the same species by having more dorsal setae (segment IV with about

20 setae), more cerarian setae (anal-lobe cerarius with about 13 conical setae),

longer anal-lobe setae (about 145 g long), and length of tibia/tarsus about 1.4;

second-instar females have 7 setae on segment IV, 4 or 5 conical setae in each

anal-lobe cerarius, anal-lobe setae about 128 g long, and length of tibia/tarsus

about 1.1.

Second-Instar Female

(Fig. 8)

Specimens examined .—The following description is based on 2 specimens col¬

lected near Port St. Lucie, Florida on 28-VII-1982.

Description.— Mounted, 0.9 mm long, 0.4 mm wide. Body elongate.

Dorsum with 5 pairs of cerarii. Anal-lobe cerarius with basal sclerotization, 4

or 5 conical setae, 0 or 1 enlarged, bristle-shaped seta, no pores. Penultimate

cerarius with basal sclerotization, 3 conical seta, 0 or 1 enlarged, bristle-shaped

seta, no pores. Antepenultimate cerarius with basal sclerotization, 1 or 2 conical

setae, 0 or 1 enlarged, bristle-shaped seta, no pores. Fourth and fifth cerarii each

with basal sclerotization, l conical seta, 1 enlarged, bristle-shaped seta, no pores.

With 1 kind of seta; enlarged, bristle-shaped setae scattered over surface, largest

posteriorly and anteriorly, smallest in center of body; longest seta on segment VII

37 g long, segment IV with 7 setae, excluding those in cerarii. Trilocular pores

absent. Discoidal pores represented by small, irregular sclerotized spots, present

near multilocular pores. Multilocular pores scattered over surface, most abundant

anteriorly. Oral-collar tubular ducts in marginal areas on any or all of segments

II-VII. Ostioles small.

Anal ring dorsal, touching apex of abdomen, with 4 or 5 rows of pores; anal-

ring setae with acute apices, longest seta 89 and 99 g long, longest seta 1.5 and

1.6 times longer than greatest diameter of ring.

Venter with 2 kinds of setae, abdominal segments each with 6-10 (8) setae;

enlarged, bristle-shaped setae in marginal areas on segments III or IV to VII. with

1 seta near each lateral margin of each segment; bristle-shaped setae scattered

over remainder of surface; longest bristle-shaped seta on segment VII 25 g long;

with 2 pairs of cisanal setae, longest seta 25 and 32 g long; longest anal-lobe seta

126 and 131 /u, long; length of longest anal-ring seta/length of longest anal-lobe

seta 0.7 and 0.8. Multilocular pores most abundant on thorax, absent from medial

areas of segments IV or V to VIII. Trilocular pores absent. Discoidal pores of

same nature and distribution as on dorsum. Oral-collar tubular ducts on marginal

or submarginal areas of metathorax and all or any of segments V-VII. Ventral

anal-lobe area sclerotized, with enlarged, bristle-shaped setae.

Circulus oval, width/length 1.2 and 1.3, 12^ wide. Hind legs without translucent

pores. Hind femora 103 g long; tibiae 74 /u long; tarsi 67 and 69 g long; length

212

PAN-PACIFIC ENTOMOLOGIST

Fig. 8. Second-instar female paratype of Hypogeococcus margaretae n. sp. near Port St. Lucie,

Florida, VII-28-82, on Eriocaulon decangulare.

of femur/tibia 1.4; length of tibia/tarsus 1.1. Tibia with 7 setae. Leg seta bristle

shaped except for some enlarged, bristle-shaped setae on femur, tibia, and some¬

times trochanter and tarsus; tarsal digitules reaching tip of claw, digitules of hind

2 pairs of legs subequal in length, with apical clubs of approximately same size,

VOLUME 59, NUMBERS 1-4

213

digitules of front legs unequal, 1 without club, other similar to digitule of hind

tarsi; claw digitules extending beyond apex of claw, equal in size. Antennae

5-segmented, third segment partially divided, 160 and 168 g long; enlarged, bris¬

tle-shaped setae on anterior margin (as drawn) of segments 1-3 or 4. Labium with

basal segment closely attached to second segment, 67 g long. Eyes without as¬

sociated pores.

Comparison. — Second instars of H. margaretae can be separated from first

instars of the same species by having 4 or 5 conical setae in each anal-lobe cerarius,

4 dorsal setae on segment IV, no trilocular pores, many multilocular pores, and

marginal oral-collar tubular ducts; first instars have 2 conical setae in each anal-

lobe cerarius, 7 dorsal setae on segment IV, many trilocular pores, and no mul¬

tilocular pores or oral-collar tubular ducts.

For a comparison of second-instar males and females of H. margaretae see the

comparison section of the second-instar male. For a comparison of third-instar

and second-instar females, see the comparison section of the third-instar female.

Second-Instar Male

Specimen examined.— The following description is based on 1 specimen col¬

lected near Port St. Lucie, Florida on 28-VII-1982.

Description. — Second-instar males are very similar to second-instar females.

Only differences are given here.

Dorsum with 4 or 5 pairs of cerarii. Anal-lobe cerarius with 2 conical setae, 3

enlarged, bristle-shaped setae. Penultimate cerarius with 2 conical setae, no bristle-

shaped setae. Remaining 2 or 3 cerarii each with 1 conical seta and 1 enlarged,

bristle-shaped seta. Longest seta on segment VII 35 g long. Oral-collar tubular

ducts in marginal and mediolateral areas of most of surface.

Anal-ring with longest seta 79 g long, longest seta 1.5 times longer than greatest

diameter of ring.

Venter with longest bristle-shaped seta on segment VII 25 g long; longest cisanal

seta 30 g long; longest anal-lobe seta 148 g long; length of longest anal-ring seta/

length of longest anal-lobe seta 0.5. Oral-collar tubular ducts on marginal areas

from head to thorax.

Circulus width/length 1.2, 12 g wide. Hind femora 99 g long; tibia 72 g long;

tarsus 62 g long; length of tibia/tarsus 1.2. Antennae 6-segmented, 151 g long.

Labium 64 g long.

Comparison. — Male and female second instars can be distinguished by the

former having fewer conical setae in the hind 2 pairs of cerarii, more oral-collar

tubular ducts scattered over most of the dorsum, and by having 6-segmented

antennae.

First Instar

(Fig. 9)

Specimen examined.— The following description is based on 1 first instar para-

type from near Port St. Lucie, Florida collected 28-VII-1982 and 8 embryos that

are inside 3 of the 4 adult female paratypes collected in Jackson Co., Mississippi.

Description. — Only salient features are included because of the poor condition

of the available material.

Mounted, 380-632 (445) g long, 187-304 (219) g wide. Body elongate oval.

214

PAN-PACIFIC ENTOMOLOGIST

Fig. 9. First-instar paratype of Hypogeococcus margaretae n. sp. near Port St. Lucie, Florida, VII-

28-82, on Eriocaulon decangulare.

Dorsum with 4-7 (5) pairs of cerarii that have at least 1 conical seta, with about

12 paired, enlarged bri stle-shaped setae along each body margin in form of cerarii

including those with conical setae; cerarii without associated pores, with basal

sclerotization. Anal-lobe cerarii with 2 conical setae; penultimate cerarii with 1-

2 (2) conical setae and 0-1 (0) enlarged, bristle-shaped setae; antepenultimate and

VOLUME 59, NUMBERS 1-4

215

fourth cerarii each with 1-2(1) conical setae and 0-1 (0) enlarged, bristle-shaped

setae; fifth cerarii with 0-1 (1) conical setae and 1-2 (1) enlarged, bristle-shaped

setae; sixth and seventh cerarii each with 0-1 (0) conical setae and 1-2 (2) enlarged,

bristle-shaped setae; remaining cerarii each with 2 enlarged, bristle-shaped setae.

With 2 kinds of setae; conical setae restricted to cerarii except 1 specimen has 1

such seta in medial area of segment 5; enlarged, bristle-shaped setae scattered

over dorsum, arranged in 4 longitudinal lines on most of abdomen, excluding

setae in cerarii, with extra setae on thoracic segments, shortest setae in center of

body; longest seta on segment VII, excluding those in cerarii, 23-32 (27) /x long,

segment IV with 4 setae including those in cerarii. Trilocular pores scattered over

surface most abundant on thorax, on abdomen usually with 6 longitudinal lines

of pores with transverse row of pores located near anterior and posterior margin

of each anterior abdominal segment, forming 6 or 8 longitudinal lines of trilocular

pores on abdomen. Discoidal and multilocular pores absent. Tubular ducts absent.

Ostioles present.

Anal ring apical, with 3 or 4 rows of pores; anal-ring setae with acute apices,

longest seta 63-77 (71) /x long.

Venter with 2 kinds of setae, forming 6 longitudinal lines of setae on abdomen;

enlarged, bristle-shaped setae restricted to submarginal areas of posterior 2 or 3

abdominal segments; bristle-shaped setae on rest of surface; longest bristle-shaped

setae on segment VII about 20 /x long; with 2 pairs of cisanal setae; longest anal-

lobe seta 87-151 (124) ju long. Trilocular pores associated with submarginal and

mediolateral lines of setae on abdomen, scattered on thorax and head. Multilocular

and discoidal pores absent. Tubular ducts absent. Ventral anal-lobe area sclero-

tized.

Circulus oval, not divided, 7-11 (9) /x wide. Legs generally unclear on embryos,

on single available crawler hind femora each 79 ix long; tibiae each 59 /x long;

tarsi 62 and 59 ju long; length of femur/tibia 1.3; length of tibia/tarsus 1.0. Tibia

with 7 setae; leg setae bristle shaped except for some enlarged setae on trochanter,

femur and tibia; tarsal digitules of hind 2 pairs of legs about equal in size, extending

beyond apex of claw, of front pair of legs, each tarsus apparently with 1 long,

capitate digitule and 1 short, apically acute digitule; claw digitules equal, extending

beyond tip of claw. Antennae unclear on embryos, on single crawler 6-segmented,

each 133 /x long; with enlarged, bristle-shaped setae on anterior margin (as drawn)

of segments 1-3 or 4. Labium with basal segment closely attached to second

segment, about 50 /x long. Eye without associated pores.

Comparison. — First instars of Hypogeococcus margaretae and H. boharti differ

as follows: H. margaretae lacks a trilocular pore between the conical setae of the

anal-lobe cerarius, has enlarged, bristle-shaped setae, has 2 transverse rows of

trilocular pores on anterior abdominal segments, lacks ventral discoidal pores,

has enlarged, bristle-shaped setae on the femora and tibiae, and lacks a discoidal

pore associated with the eye; H. boharti has a trilocular pore between the conical

setae of the anal-lobe cerarius, lacks enlarged, bristle-shaped setae, has only 1

transverse row of trilocular pores on abdominal segments, has a line of discoidal

pores on the venter near the base of the submarginal setae, lacks enlarged setae

on the legs, and has a discoidal pore associated with each eye. For a comparison

of second instars of H. margaretae with first instars of the same species see the

comparison section of the second-instar female.

216

PAN-PACIFIC ENTOMOLOGIST

Discussion

During the course of this study special emphasis was devoted to several char¬

acters that previously were given little or no detailed study. These characters

included numbers of leg setae (i.e., number of setae on hind tibia), numbers of

setae on the abdomen (i.e., number of setae on the dorsum of segment IV), lengths

of the abdominal setae (i.e., lengths of dorsal and ventral setae on segment VII

and of the cisanal setae), numbers of cisanal setae, and relative sizes of the claw

and tarsal digitules. Each of these characters has proven to be of value in distin¬

guishing among species of Hypogeococcus and may be useful in other groups of

mealybugs. Characteristics of the digitules were especially interesting. Features of

digitules have been used in the genus Rhizoecus (Hambleton, 1976) and seem to

characterize genera of the tribe Phenacoccini (Miller, 1975). In all of the species

of Hypogeococcus, and in Ferrisia virgata (Cockerell), Nipaecoccus nipcie (Mas¬

ked), Parcidoxococcus mcdanieli McKenzie, P/anococcus citri (Risso), Pseudococ¬

cus maritimus (Ehrhorn), Tnonymus winnemucae McKenzie, and probably most

mealybugs, the front legs have asymmetrical tarsal digitules, with one digitule on

each leg capitate and with the other digitule bristle shaped and without an enlarged

apex. Both tarsal digitules of each of the mesothoracic and metathoracic legs

usually are capitate, although one usually is much more robust than the other.

The claw digitules usually are larger than the tarsal digitules and frequently are

equal in size on all three pairs of legs. The study of embryos has proven to be of

considerable value in learning about the morphology of the first instars of 2

Hypogeococcus species and may prove to be of use in other pseudococcid studies.

Embryos have been used in armored scales (Miller et al., in press) and have the

added advantage of giving a clear association with an adult female.

Studies of immature forms continue to provide interesting and valuable infor¬

mation relative to the formulation of a usable mealybug classification. The oc¬

currence of trilocular pores on the first instars of Hypogeococcus is somewhat

surprising in light of the fact that Heterococcus, another genus of mealybug that

lacks trilocular pores as adults, has only quinquelocular pores in first instars

(Miller, 1975). Study of second instars of Hypogeococcus support the hypothesis

that in most mealybugs, the second-instar male has more oral-collar tubular ducts

than does the second-instar female.

It is clear that after current research on South American species is complete

and adult males and first instars have been studied, the composition of Hypo¬

geococcus will need to be reevaluated. It is likely that Hypogeococcus eventually

will be divided into two or three genera.

Acknowledgments

Special appreciation is extended to Dr. Manya B. Stoetzel and Dr. E. Eric

Grissell, Systematic Entomology Lab., Agric. Res. Serv., U.S. Dept. Agric. and

to Dr. Douglas J. Williams, Commonwealth Institute of Entomology, % British

Museum (Nat. Hist.), London, England, for their valuable comments and criti¬

cisms of the manuscript. To Mr. Kenneth L. Hibbard, Division of Plant Industry,

Florida Department of Agriculture and Consumer Services, Port Pierce, I give

special thanks for his considerable efforts to collect and recollect material of

Hypogeococcus margaretae. I am grateful to Ms. Helen Proctor, Systematic Ento¬

mology Lab., for typing the various drafts of the paper and to Ms. Pamela Hollyoak

VOLUME 59, NUMBERS 1-4

217

for preparing most of the illustrations. I also acknowledge the assistance of Dr.

Jennifer Cox and Dr. Douglas J. Williams (BMNH), Mr. Raymond J. Gill and

Mr. Robert O. Schuster (UCD), and Dr. Avas B. Hamon (FSCA) for loaning

specimens for this study.

Literature Cited

Beardsley, J. W. 1965. Notes on the pineapple mealybug complex, with descriptions of two new

species. Proc. Hawaii. Ent. Soc. 19:55-68.

Bohart, R. M. 1942. Life history of Diaspis boisduvalii and its control on Cattleya with calcium

cyanide. J. Econ. Ent. 35:365-368.

Brain, C. K. 1915. The Coccidae of South Africa.—I. Trans. Roy. Soc. S. Afr. 5:65-194.

Cruttwell, R. E., and P. Fidalgo. 1974. Studies of the biology and host-specificity of the cactus

mealybug Hypogoecoccus festerianus (Lizer y Trelles), attacking Eriocereus martinii in Argen¬

tina. Report Commonw. Inst. Biol. Control, S. Amer. Substa., 11 pp.

De Lotto, G. 1974. On two genera of mealybugs. J. Ent. Soc. S. Afr. 37:109—115.

Ezzat, Y. M., and H. S. McConnell. 1956. A classification of the mealybug tribe Planococcini. Univ.

Md., Agr. Exp. Sta. Bull. A-84, 108 pp.

Ferris, G. F. 1950. Atlas of the scale insects of North America. Vol. V. The Pseudococcidae (Part

I). Stanford Univ. Press, Calif., pp. 1-278.

-. 1953. Atlas of the scale insects of North America. Vol. VI. The Pseudococcidae (Part II).

Stanford Univ. Press, Calif., pp. 279-506.

Hambleton, E. J. 1976. A revision of the New World mealybugs of the genus Rhizoecus. U.S. Dept.

Agr. Tech. Bull. no. 1522, 88 pp.

Lizer y Trelles, C. A. 1942. Pedronia festeriana n. sp. Revista Fac. Agron. Vet. Univ. B. Aires 10:

24-28.

MacGillivray, A. D. 1921. The Coccidae. Scarab Co., Urbana, Ill., 502 pp.

McFadyen, R. E. 1979. The cactus mealybug Hypogeococcus festerianus an agent for the biological

control of Eriocereus martinii in Australia. Entomophaga 24:281-287.

McFadyen, R. E. and Tomley, A. J. 1978. Preliminary indications of success in the biological control

of Harrisia cactus ( Eriocereus martinii Lab.) in Queensland. Proc. 1 st Conf. Council Aust. Weed

Sci. Soc., pp. 108-112,

McKenzie, H. L. 1960. A new subterranean Rhizoecus mealybug from Arizona. Pan-Pac. Ent. 36:

139-141.

-. 1967. Mealybugs of California with taxonomy, biology, and control of North American

species. Univ. Calif. Press, Berkeley, 525 pp.

Miller, D. R. 1975. A revision of the genus Heterococcus Ferris with a diagnosis of Brevennia Goux.

U.S. Dept. Agr. Tech. Bull. No. 1497, 61 pp.

Miller, D. R., J. A. Davidson, and M. B. Stoetzel. A taxonomic study of the armored scale Pseudi-

schnaspis Hempel. Proc. Ent. Soc. Wash, (in press).

Miller, D. R., and H. L. McKenzie. 1971. Sixth taxonomic study of North American mealybugs,

with additional species from South America. Hilgardia 40:565-602.

Rau, G. J. 1938. Four more species of mealybugs from New York State. Canad. Ent. 70:157-165.

Theron, J. G. 1958. Comparative studies on the morphology of male scale insects. Ann. Univ.

Stellenbosch 34:1-71.

Williams, D. J. 1973. Two cactus-feeding mealybugs from Argentina. Bull. Ent. Res. 62:565-570.

Willink, C. G. de. 1981. Neuva especie de Hypogeococcus Rau de Tucuman, Republic Argentina.

Neotropica 27:61-65.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 218-228

Habitat Differences Among Container-Breeding Mosquitoes in

Western Puerto Rico (Diptera: Culicidae)

Chester G. Moore 1

Division of Vector-Borne Viral Diseases, Center for Infectious Diseases, Centers

for Disease Control, Public Health Service, U.S. Department of Health and Hu¬

man Services, P.O. Box 2087, Fort Collins, Colorado 80522-2087.

Aedes aegypti (Linn.), the vector of dengue virus in the Caribbean, has assumed

major importance in recent years because of continued dengue epidemics (Neff

et al., 1967; Likosky et al., 1973; Von Allmen et al, 1979).

From 1964 to 1968, an intensive Ae. aegypti Eradication Program was carried

out in the southeastern United States, Puerto Rico, and U.S. Virgin Islands. During

that program, there was widespread removal of container habitats in the domestic

and peridomestic environment and insecticidal treatment of remaining habitats.

McClelland (1967) suggested that this type of treatment might profoundly affect

the behavior of a “weedy” species, such as Ae. aegypti, by applying strong selective

pressure to change breeding site selection. Intensive control would eliminate

panmictic breeding, leading to higher inbreeding rates, which would further en¬

hance rapid evolutionary change. According to McClelland (1967), any repellent

quality of the insecticides used to treat larval habitats would also increase selection

by inducing ovipositing females to seek untreated containers, presumably those

not considered the usual types or those at some distance from houses. Moore

(1977) subsequently found that Abate® 2 and malathion, two widely used larvi-

cides, repelled ovipositing Ae. aegypti.

Kellett and Omardeen (1957) found Ae. aegypti in Trinidad breeding in tree

holes as high as 10 meters above ground level. They suggested this behavior might

be related to repellent action of benzene hexachloride (BHC) used for larval

control. Fox et al (1960) found Ae. aegypti breeding in coraline rock holes in

western Puerto Rico. Larvae from rock holes were found to be highly resistant to

DDT, dieldrin, and lindane. Weinbren and O’Gower (1966) and Weinbren and

Weinbren (1970) reported Ae. aegypti breeding in the Luquillo rain forest in

eastern Puerto Rico. However, Haber and Moore (1973) concluded that the rain

forest infestation resulted from introduction of eggs during a long-term radiation

ecology experiment, and no evidence of permanent establishment was found.

The foregoing reports of habitat and behavior changes prompted a survey of

Ae. aegypti distribution with respect to container habitat and distance from houses

in western Puerto Rico. The study also provided an opportunity to evaluate

possible competition between Ae. aegypti and the local Ae. ( Gymnometopa) me-

diovittatus (Coq .) and to assess the degree of association between the various

container-breeding mosquito species.

1 Work performed while the author was on the faculty of the Department of Biology, University of

Puerto Rico, Mayaguez, P.R.

2 Use of trade names is for identification only and does not constitute endorsement by the Public

Flealth Service or by the U.S. Department of Health and Human Services.

VOLUME 59, NUMBERS 1-4

219

Fig. 1. Location of study area in Puerto Rico. Municipalities surveyed were Aguada, Aguadilla,

Anasco, Cabo Rojo, Hormigueros, Isabela, Lajas, Las Marias, Maricao, Mayaguez, Moca, Rincon,

Sabana Grande, San German, and San Sebastian.

Materials and Methods

This study was carried out concurrently with collections of local mosquito

species for the “Mosquitoes of Middle America” (MMA) project (Belkin et al.,

1965a, b). Egg, larval, pupal, and adult mosquito collections were made from

artificial (man-made) and natural (e.g., tree holes, rock holes, plant axils) water¬

holding containers or holes around rural dwellings in 15 municipalities in western

Puerto Rico (Fig. 1), from January through June, 1971. Collecting methods were

as described by Belkin et al. (1965b), and collection data were recorded on the

standardized MMA collection form. Distance from the nearest house was also

estimated and noted on the collection form. Collections were returned to the

laboratory for identification. Scrapings and other collections of dried material

were flooded with water and subjected to vacuum for 15 to 30 minutes to induce

hatching. Samples that did not produce larvae were dried and then flooded a

second time. Samples producing no larvae after the second flooding were consid¬

ered to be negative.

Larvae were reared in enamel pans (18 X 30 X 5 cm) containing approximately

1 liter of 0.01 M NaCl solution in place of tap water and were fed finely ground

rabbit food. Larvae were identified in the fourth instar and, when appropriate,

samples were preserved for the MMA project. When less common species were

collected, individual rearings were carried out to preserve larval and pupal skins

with the associated adults (Belkin et al., 1965b). Representative samples of the

material collected were deposited in the MMA project collection (Belkin and

Heinemann, 1975), which is now located at the Smithsonian Institution.

Habitats were grouped by type and distance from the nearest human habitation.

Container habitats were grouped according to source (artificial or natural) and

size. Later, several groups were pooled to obtain reasonable sample sizes. The

final categories were: 1) artificial containers—a) containers of 20-liter capacity or

greater and b) containers of less than 20-liter capacity, and 2) natural containers

of all sizes. The number of containers decreased as distance from houses increased.

Most houses were grouped together and not separated from other houses by great

distances; also, inspectors were less likely to discover all containers farther away

220

PAN-PACIFIC ENTOMOLOGIST

Table 1. Availability of distance data, type of sample and positivity of samples for 772 artificial

and natural container habitats in western Puerto Rico, January-June 1971.

Type of sample

Larvae/pupae

Scrapings

Total

Positive samples

Distance known

383

391

Distance unknown

209

221

Total

592

612

Negative samples

Distance known

Distance unknown

Total

155

160

Total samples

597

175

772

from houses. Thus, there was a highly skewed distribution of containers over

distance from houses. To obtain a small number of distance classes with approx¬

imately equal numbers of collections, a logarithmic scale using four distance

classes was based upon a 13X expansion, beginning at 0.5 (-1.70, 0.81, 1.93, . . .

on the logarithmic scale) (Williams, 1964).

Data for species pairs were tested for positive or negative association by chi-

square or Fisher’s exact test and by calculation of Cole’s (1949) coefficient of

interspecific association (C 7 ), as described by Southwood (1966). Temperature

and rainfall data were obtained from 19 weather stations scattered throughout

the study area (NOAA, 1972). Changes in mosquito abundance during the study

were examined for possible correlation with temperature and rainfall variations.

Results

Twelve species of mosquitoes were collected during the study: Aedes aegypti

(Linnaeus), Ae. mediovittatus (Coquillett), 3 Anophcdes grabhamii Theobald. Culex

antillummagnorum Dyar, Cx. atraius Theobald, Cx. habilitator Dyar & Knab,

Cx. nigripalpus Theobald, Cx. p. quinquefasciatus Say, 4 Cx. secutor Theobald,

Toxorhynchites portoricensis (Roeder), Uranotaenia socialis Theobald, and Wyeo-

myia sp. In addition, a chaoborid gnat (Diptera, Chaoboridae) was collected in

rock hole habitats. Both the Wyeomyia and Corethrella spp. are apparently un¬

described species (Belkin and Heinemann, 1975). Because the larval stages of

chaoborids and culicids are morphologically and ecologically similar, the chao-

borids are included within the Culicidae following Belkin et al. (1970).

Table 1 shows the number of positive and negative samples of different types.

It was not always possible to determine distance to the nearest house, so that

distance information was available for only 61% of the collections. Nearly all

(97%) of the negative samples were scrapings from dry container habitats, mostly

tree and rock holes. All negative samples were discarded from the analysis, since

3 Generic abbreviations for Culicidae follow Reinert (1975); Co. — Corethrella.

4 Sirivanakam (1976) elevated Cx. p. quinquefasciatus to full specific status. I have elected to retain

the subspecific designation for this taxon because elevation only serves to further confuse an already

complex biosystematic problem.

VOLUME 59, NUMBERS 1-4

221

Fig. 2. Number of species per sample in 772 collections from various artificial and natural water¬

holding containers, western Puerto Rico, January-June 1971.

true negativity (absence of viable mosquito eggs) could not be differentiated from

methodological problems in the hatching technique.

Distance from human habitation and species per sample.—Of the 160 negative

samples, 155 were collections of material removed from dry containers while 5

were collections of water thought to contain larvae or pupae. Of 612 positive

samples, 488 (79.7%) contained a single mosquito species, 106 (17.3%) contained

2, 14 (2.3%) contained 3, and 4 (0.7%) contained 4 species (Fig. 2). No sample

contained more than 4 mosquito species.

The number of species per sample was related to proximity to houses. In the

383 positive larval/pupal collections of known distance, single species collections

were more frequent near houses, while multi-species collections were more fre¬

quent away from houses. The relationship was significant (P < 0.05, chi-square),

regardless of the geometric class intervals (i.e., 3X, 5X, 1IX, 13X, 17X, 23X) used

to group the collections (Williams, 1964).

Collections from containers less than 1 meter from houses (i.e., inside, attached

to, or resting against the house) accounted for 34% of all positive collections. Over

50% of all positive collections were made within 4 meters of a house, and 96%

222

PAN-PACIFIC ENTOMOLOGIST

Table 2. Percent distribution of seven container-breeding mosquito species in relation to distance

to nearest human residence, Puerto Rico (n = 383 samples).

Species

Distance from houses (meters)

Total

samples

1-6

7-84

85-1098

Ac. aegypti

44.0

37.0

18.1

0.8

243

Ae. mediovittatus

14.8

13.6

51.1

20.5

Co. sp.

0.0

10.7

42.9

46.4

Cx. antillummagnorum

6.7

26.7

33.3

Cx. p. quinquefasciatus

30.8

19.2

50.0

0.0

Cx. secutor

48.7

10.3

35.9

5.1

Tx. portoricensis

9.1

18.2

63.6

9.1

Samples per class

132

105

115

383

were no farther than 120 meters away from a house. In the areas under study,

there were relatively few localities where houses were more than 240 meters apart.

Table 2 shows the distribution of the 7 most common species with respect to

distance from houses. Aedes aegypti was most common near houses. Aedes me-

diovittatus was more common in collections away from houses, but nearly 30%

of collections of this species were within 6 meters of a house. Culex p. quinque-

fasciatus and Cx. secutor, which share fairly similar container habitats, were

somewhat similar in spatial distribution.

Importance of different kinds of container habitat.— Table 3 shows the relative

importance of major container categories as breeding sites for the 7 most common

mosquito species. Natural container habitats made up 23.8% (140/592) of all

positive larval/pupal collections. Of those, 51.4% were rock holes, 32.1% were

leaf axils, 7.1% were tree holes, and 9.3% were miscellaneous container types

(snail shell, coconut shell, palm spathe, etc.). Because of the small numbers of

collections from each container type, it was not possible to analyze the preference

of a given species for specific kinds of natural containers.

Several species (e.g., Ae. aegy’pti, Ae. mediovittatus, Cx. p. quinquefasciatus, Cx.

secutor) were broadly scattered among the different categories, while others (Co-

rethrella sp., Cx. antillummagnorum) were highly selective and occurred only in

a few kinds of container habitats. Several collections, such as Cx. antillummag¬

norum from rock holes, may be due to larvae being washed from a more common

habitat into another habitat by heavy rain.

Culex secutor is primarily an inhabitant of pools in mountain streams (Belkin

et al., 1970), but as shown by these data, it has a broad range of larval habitats.

It is also possible that the Cx. secutor of Puerto Rico consists of more than one

species (Belkin et al., 1970).

Zavortink (1972) recorded Ae. mediovittatus from all of the larval habitats

shown in Table 3, plus ground pools, which were not sampled in this study.

The rock hole area previously studied by Fox et al. (1960) was located through

the assistance of Mr. A. H. Boike. Aedes aegypti was still found breeding in the

rock holes 12 years after their original survey. The rock hole area is located, as

noted by Fox et al., about 25 to 90 meters (80 to 300 ft) from the nearest houses.

The area had been used as an unauthorized dump for some years, and by 1971,

large numbers of artificial containers were present to serve as additional breeding

VOLUME 59, NUMBERS 1-4

223

Table 3. Percent of breeding of 7 mosquito species in different container types, western Puerto

Rico, 1971 (n = 592 samples positive for one or more species, adjusted for differing numbers of

containers within types).

Species

Drum

Tire

Misc.

2:20 liters <20 liters

Rock hole

Misc.

natural

Ae. aegypti

29.7

22.7

13.0

17.7

7.8

9.1

Ae. mediovittatus

4.9

14.7

18.9

13.3

37.1

11.0

Co. sp.

0.0

2.2

8.2

3.1

86.5

0.0

Cx. until/iimmagnorum

0.0

4.1

7.1

10.3

5.0

73.4

Cx. p. quinquefasciatus

10.5

19.7

43.8

14.9

8.8

2.3

Cx. secutor

9.7

14.4

28.2

22.8

13.3

11.6

Tx. portoricensis

2.6

29.6

17.2

18.9

0.0

31.8

Total containers

178

137

sites. Fox et at. (1960) concluded that rock hole specimens of Ae. aegypti repre¬

sented the spillover of oviposition due to population increases in other, more

normal habitats, and no evidence was found in the present study to suggest

otherwise. Additional coraline rock hole breeding sites were found near the town

of Lajas.

Interspecific association. — Seven species were collected in sufficient numbers to

permit calculation of Cole’s (1949) coefficient of interspecific association (C 7 ).

Aedes aegypti was negatively associated with all of the other 6 species (Table 4).

Aedes mediovittatus was positively associated with Corethrella sp. All other species

pairings were random (i.e., species were not associated).

Sufficient data were available for Ae. aegypti and Ae. mediovittatus to compute

C 7 ’s for either container type or distance from houses and, to a certain extent, for

both factors simultaneously. There was a strong negative association between the

species in terms of the type of container in which they occurred (Table 5). Table

3 shows the major differences in types of containers utilized. Water storage drums

and tires were the most frequent sources of Ae. aegypti, while rock holes and

miscellaneous large containers were the most frequent sources of Ae. mediovittatus.

Joint occurrences of Ae. aegypti and Ae. mediovittatus were random with respect

to distance from houses at 1-6 meters and 85-1098 m, but C 7 ’s were significantly

negative at distances of less than 1 meter and from 7-84 meters (Table 5). Only

28 percent of all Ae. mediovittatus collections occurred within 6 meters of a house,

while 81% of all Ae. aegypti were found within that distance (Table 3). Lack of

significance in C 7 ’s at 1-6 and 85-1098 meters may have been due to small

numbers of one species in these two distance categories, or even a zone of overlap

in the 1- to 6-meter distance class.

Aedes aegypti and Cx. p. quinquefasciatus, both widely distributed domestic

species, were negatively associated. The separation in this case appeared to be

due to type of container and not to distance from houses. Most Cx. p. quinque¬

fasciatus were found in tires and miscellaneous large containers (Table 4). Different

preference for organic content of the container habitat probably also limits co¬

habitation.

Aedes mediovittatus and Corethrella sp., a predator, were positively associated

(C 7 = 0.40 ± 0.10, Table 4), and Ae. mediovittatus may form a major part of the

Table 4. Coefficients of interspecific association (C 7 , Cole, 1949) between container-breeding mosquitoes in western Puerto Rico (n = 383 samples). C 7 ± SE

significant (chi-square or Fisher’s exact test) at P < 0.05 (*), 0.01 (**), or 0.001 (***). NS = non-significant (random) association between species.

Ae. mediovittatus

Co. sp.

Cx. antillum¬

magnorum

Cx. p. quinque¬

fasciatus

Cx. secutor

Tx. portoricensis

Ae. aegypti

Ae. mediovittatus

Co. sp.

Cx. antillummagnorum

Cx. p. quinquefasciatus

Cx. secutor

-.71 ± .07***

-.89 ± .14***

.40 ± .10***

-.89 ± .19***

-.13 ± .46 NS

-1.00 ± .90 NS

-.48 ± .14**

-.03 ± .34 NS

-.49 ± .66 NS

-.05 ± .92 NS

-.68 ± .12***

.10 ± .08 NS

-.65 ± .54 NS

-1.00 ± .75 NS

-.27 ± .55 NS

-.71 ± .22**

-.60 ± .54 NS

-1.00 ± 1.06 NS

-1.00 ± 1.47 NS

-1.00 ± 1.08 NS

.09 ± .10 NS

Table 5. Effect of container type and distance from houses on coefficient of association (C 7 ) between Ae. aegypti and Ae. mediovittatus in western Puerto Rico

(n = 383 samples). C 7 ± SE significant (chi-square or Fisher’s exact test) at P < 0.05 (*), 0.01 (**), or 0.001 (***). NS = non-significant (random) association.

Distance (m)

Container type

1-6

7-84

85-1098

All distances

Large, artificial

-.62 ± .12***

1.00 ± 2.30 NS

-1.00 ± .40*

-1.00 ± .41 NS

-.66 ± .08***

Small, artificial

-.63 ± .32 NS

-.40 ± .16*

-.69 ± .26*

—

-.61 ± .15***

Natural

—

-.23 ± .49 NS

-.76 ± .20***

—

-.78 ± .17***

All containers

-.62 ± .13***

-.22 ± .11 NS

-.77 ± .15**

-1.00 ± .58 NS

-.71 ± .07***

224 PAN-PACIFIC ENTOMOLOGIST

VOLUME 59, NUMBERS 1-4

225

100

■J 0

Tx. portoricensis

-i ■ i •-

30|-

= 201-

1 iol- A

F M A M

Hot - dry

5* 0

y " I i

t r . -H-r r —l

M A M J

30 h

Warm - moist

-r

F M A M J

Month

Fig. 3. Temperature, rainfall, and monthly abundance of container-breeding mosquito species in

western Puerto Rico, January-June 1971 (X ± SD shown for temperature and rainfall).

predator’s diet. Moore and Haber (unpublished data) found that late instars of

this Corethrella sp. consumed as many as 2.2 Ae. mediovittatus larvae per 24 h

in laboratory experiments.

Weather pattern and temporal relationships between species.— The study area

was not homogeneous with respect to temperature and rainfall. Average (January

through June 1971) monthly temperature and rainfall for 11 stations in the study

area fell into 4 distinct zones: a) hot-dry (26°C, 7 cm rain/month), b) warm-moist

(24°C, 14 cm/month), c) cool-moist (21°C, 16 cm/month), and d) warm-wet (24°C,

21 cm/month). Approximately 95% of all collections came from the hot-dry or

warm-moist zones.

January and February were the coolest months, and temperatures rose gradually

and uniformly from March through June, when the study ended (Fig. 3). Rainfall

was lowest in January. As is usual for the western part of Puerto Rico (Calvesbert,

1970), rainfall increased in February but was diminished again in March. Rainfall

increased in April and May but was unusually low in June. June precipitation

records were from 5 to 11 cm below normal for most parts of the island.

Some of the observed negative associations could have been due to temporal

separation of population peaks, even though the study encompassed only a 6-month

period (Fig. 3). Except for Ae. aegypti and Cx. secutor, the temporal occurrence

of the species was fairly constant. There was a suggestion that several species may

226

PAN-PACIFIC ENTOMOLOGIST

have shown seasonal changes or shifts in utilization of different container types,

but there were insufficient data to test that possibility. Aedes aegypti gradually

declined in frequency over the course of the study. Although aegypti abundance

appeared to be inversely related to environmental temperatures, island-wide stud¬

ies by Moore et al. (1978) indicate that this is not the case. Culex secutor abun¬

dance was highly correlated with rainfall (Kendall rank correlation coefficient, r =

0.71, P = 0.001), with a lag effect of about 15 days.

Discussion

There was no evidence of behavioral change in Aedes aegypti populations in

western Puerto Rico as a result of the Aedes aegypti Eradication Program. Over

99% of all positive collections were made at distances of less than 85 meters from

the nearest house, and no Ae. aegypti were found more than 100 m from houses.

Similarly, natural container breeding sites made up only 17% of the total. Reports

of unusual behavior (Fox et. al, 1960; Weinbren and Weinbren, 1970) were shown

to be due to spillover from nearby domestic breeding sites or to transitory estab¬

lishment following introduction of Ae. aegypti into an isolated focus of intense

human activity (Haber and Moore, 1973).

Whenever Aedes aegypti has been introduced into new regions of the tropics,

it has tended to displace the indigenous container-breeding Aedes of that region

(Bailey and Bohart, 1952; Macdonald, 1956; Gilotra et al, 1967; Chan et al.,

1971). Laboratory studies have similarly demonstrated the ability of Ae. aegypti

to exclude other Aedes species (Wilton, 1968; Moore and Fisher, 1969; Peters et

al, 1969).

Aedes aegypti was probably introduced into Puerto Rico soon after the arrival

of slaves from Africa in the early 16th century (Pico, 1969). Thus, Ae. aegypti

has had more than 400 years and perhaps 5000 to 13,000 generations in which

to evolve an optimal division of the container environment with local species.

The highly negative association between Ae. aegypti and the other major con¬

tainer-breeding species suggests that, if competitive displacement occurred, it did

so some time ago. There appear to be major differences in oviposition sites (e.g.,

shade, organic content, pH), not measured in this study, that influence gravid

females in selecting containers not attractive to Ae. aegypti. Conversely, Ae. aegypti

seems able to utilize certain container habitats more effectively than can other

species. This appears to be an example of an “included niche” (Schoener, 1974),

as the other species have a broader niche than Ae. aegypti (either on the container

type or distance axes, or both).

Additional studies are needed to clearly define the parameters that reduce or

prevent competition among container-breeding mosquitoes. Some parameters

that should be included in future studies are shade, height, density and species of

surrounding vegetation; organic content (quality and quantity), pH, salinity, and

source (rain or human activity) of the water; and orientation of the container.

Future studies should also be carried on long enough to demonstrate any seasonal

separation between species.

Acknowledgments

This paper is dedicated to Dr. Richard M. Bohart who, with the late Dr. S. B.

Freeborn, provided the resources and encouragement for my first mosquito ecol-

VOLUME 59, NUMBERS 1-4

227

ogy studies. The late Dr. J. N. Belkin and the Mosquitoes of Middle America

Project provided supplies for this study. The assistance of W. A. Haber, L. Ace¬

vedo, T. Feliberty-Ortiz, H. Nieves-Caro, C. Ortiz-Baez, W. Rodriguez and L.

Bonilla-Navarro is gratefully acknowledged. I thank R. E. Bailey for advice on

statistical analyses.

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Ent. 45:947-952.

Belkin, J. N., and S. J. Heinemann. 1975. Collection records of the project “Mosquitoes of Middle

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296.

Belkin, J. N., S. J. Heinemann, and W. A. Page. 1970. Mosquito studies (Diptera, Culicidae). XXI.

The Culcidae of Jamaica. Contr. Amer. Ent. Inst. 6:1-458.

Belkin, J. N., R. X. Schick, P. Galindo, and T. H. G. Aitlcen. 1965a. Mosquito studies (Diptera,

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Amer. Ent. Inst. 1(2): 1-17.

Belkin, J. N., C. L. Hogue, P. Galindo, T. H. G. Aitken, R. X. Schick, and W. A. Powder. 1965b.

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trib. Amer. Ent. Inst. 1(2): 19-78.

Calvesbert, R. J. 1970. Climate of Puerto Rico and U.S. Virgin Islands. Climatography of the United

States, No. 60-52. U.S. Dept, of Commerce.

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Singapore City. 4. Competition between species. Bull. Wld. Hlth. Org. 44:643-649.

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Gilotra, S. K., L. E, Rozeboom, and N. C. Bhattacharya. 1967. Observations on possible competitive

displacement between populations of Aedes aegypti Linnaeus and Aedes albopictus Skuse in

Calcutta. Bull. Wld. Hlth. Org. 37:437-446.

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one-year survey. Mosq. News 33:576-578.

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Trinidad, B. W. I. W. Ind. Med. J. 6:179-188.

Likosky, W. H., C. H. Calisher, A. L. Michelson, R. Correa-Coronas, B. E. Henderson, and R. A.

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Parasit. 50:385-398.

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PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 229-232

A New Perdita from Utah’s San Rafael Desert

(Hymenoptera: Andrenidae)

F. D. Parker

Bee Biology & Systematics Laboratory, ARS, USDA, Utah State University,

UMC 53, Logan, Utah 84322.

Bees of the genus Perdita are a large and diversified assemblage of species that

are principally western North American in distribution. The number of species

and subspecies described is well over 600, and most of our knowledge of Perdita

has been through the important papers of P. H. Timberlake, now deceased. Many

of our native flowering shrubs and forbs have one or more species of Perdita

associated in some manner with the plant. For most species of Perdita, however,

the nature of this bee-plant relationship has not been investigated. Perhaps their

small size (most species are 2-4 mm long) has contributed to the lack of pollination

studies of this interesting group of bees.

The systema tics of Perdita is not easily understood because the genus is speciose,

and the existing keys are difficult to use without a representative collection for

comparisons. Although there are a number of unnamed Perdita in our collection,

I am describing here only one, a rather bizarre species that was recently collected

in southern Utah. This new bee is easily separable from all known Perdita by its

unusually large size (10 mm) and sternal modifications.

This Perdita was collected in a 2500 2 -mile Utah desert that has been almost

completely isolated by surrounding mountains since the late Pliocene (Hunt,

1974), and which has more endemic plant species than any other area in the

Intermountain Region (Cronquist et al., 1972).

Perdita bohartorum Parker, New Species

(Figs. 1-7)

Holotype male. — Black; head, thorax dark blue-green; scutum, scutellum with

coppery reflections; orange-red on mandible apex, mouth parts, apical margin of

tergum 7; yellowish markings on mandible, labrum, clypeus, laterally on supra-

clypeal area, beneath antenna, stripes laterally on crest of pronotum, pronotal

lobe except medially, forefemoral apex, anterior stripe on fore and midtibiae;

tergum 1 with only lateral stripe, terga 2-6 with lateral and apical stripes that

become wider on apical terga, tergum 7 all yellow, sterna 1-7 mostly yellow except

basally; tegula hyaline, wings hyaline, veins white; forebasitarus whitish. Pubes¬

cence white, not dense on head, thorax; subapical thin hair band on sterna, terga.

Punctation on body not uniform; head mostly shiny with punctures separated by

more than their diameter, surface polished, upper frons, behind interocellar area

dull; punctures closer on thorax and legs but surface shiny; microstriae medially

on dorsal surface of propodeum; abdomen shiny with shallow punctures. Clypeus

with median truncation (V 3 width of clypeus); mandibles as long as width of

clypeus, tapering to point; labrum twice as wide as long, small; mouthparts ex¬

tending beyond forecoxae in repose; galea 1.6X longer than labrum; maxillary

VOLUME 59, NUMBERS 1-4

231

palpal segments of equal length; basal labial palpal segment 4X longer than apical

ones; hypostomal carina raised flange-like; head measurements as follows: wider

(1.2X) than long (Fig. 6); antennocular distance 1.1X wider than interantennal

distance; distance between median ocellus and antennal base shorter than distance

from median ocellus to hind margin of vertex (1.5X); distance between lateral

ocelli equal to ocellocular distance; distance between lateral ocelli greater (2.5X)

than distance between median and lateral ocelli; distance between ocular and hind

margin of vertex greater (1.4X) than ocellocular distance; postocular area (Fig. 7)

wider (2X) than width of eye; malar space as wide as diameter of median ocellus;

clypeus not protuberant in lateral profile; antenna nearly as long as length of eye;

flagellomeres I, XI 1.4X longer than other flagellomeres, I somewhat flattened;

scape short, as wide as 2nd submarginal cell; facial fovea small, oval, about size

of median ocellus; hind-femur 5X as long as wide, hindtibia about 9X as long as

wide; hindbasitarsus 6X as long as wide; fore, midlegs shorter; hindtrochanter

normal; pronotum laterally with wide, flat depression, collar slightly indented

medially, rounded laterally; scutum 1.2X wider than long, scutellum as long as

horizontal area of propodeum, 1.3X wider than metanotum; 1st submarginal cell

slightly longer (1.1X) than marginal cell; mesopleuron without sulcus connecting

subalar pit and base of wings; abdomen slightly wider 1.05X than thorax; broad

hyaline band apically on terga, sterna, band wider than length of subapical hair

band; sternum 1 with shallow median longitudinal groove, sternum 2 with prom¬

inent U-shaped, keel-like median projection (Fig. 1); sterna 3-5 with lateral low

swellings forming shallow median basin; sternum 7 with median notch; sternum

8 (Fig. 3); tergum 7 with lateral U-shaped notch (Fig. 5); tergum 8 (Fig. 4); genitalia

as in Fig. 2; length 10 mm.

Female. — Like male except: head, propodeum with coppery reflection; man¬

dible black; terga 2-5 mostly yellow, black apically, basally on some; pubescence

on tergum 5 long, covering most of segment; scopal hairs on hind legs sinuate,

as long as forebasitarsus; terga 3-5 with white patch of bristles ventrolaterally;

clypeus longer, extending just beyond base of mandible; protuberant in lateral

profile; mandible bent apically; face more oval; inner eye margins converging

below; closed mandible nearly attaining opposite mandibular base; facial fovea

narrow, nearly touching inner margin of eye, extending just beyond mid length

of eye, as wide as antennal base; distance between median ocellus and antennal

base equal to distance from median ocellus to hind margin of vertex; distance

between lateral ocelli greater (1.3X) than ocellocular distance; distance between

ocular and hind margin of vertex equal to ocellocular distance; postocular area

slightly wider than width of compound eye; malar space narrower; antenna longer

than eye; segments of hindleg shorter; sterna without projections; pygidium long,

apically truncate.

Variation. — Only subtle differences exist in the markings, punctation and struc¬

ture of the paratypes.

Systematics. — I have not assigned this peculiar Perdita to any of the existing

species groups or subgenera. In some characters it appears to be related to species

in both of the subgenera Cockerellia and Procockerellia , but major differences are

evident. For example, P. bohartorum has a wide malar space, long galea, flat

clypeal area, and prominent swellings on the sterna. Species of Cockerellia lack

232

PAN-PACIFIC ENTOMOLOGIST

a malar space, the galea is shorter, the clypeus of the males is protuberant in

profile and the abdomen lacks sternal projections. However, species of Prococke-

rellia have similar pubescence and long mouthparts. One ponders the function of

the distinctive sternal swellings since no other described species in this genus has

such abdominal modifications.

Types. — Holotype S and 11 6 paratypes: UTAH: Emery Co., sandy ridge E. of

Little Gilson Butte, 5100', May 29, 1981 (F. D. Parker, S. F. Parker). Holotype

deposited in the U.S. National Museum #100071. Metatypes, 21 6 and 7 9 (3

mating pairs), same data except: June 3, 1982 (F. D., J. H., S. F., and A. D.

Parker, T. L. Griswold).

Floral records. — All specimens were collected on flower heads of Wyethia scabra

Hook. This composite is a multiflowered perennial found on hill tops and ridges

in southern Utah, adjacent New Mexico and Arizona. The bees were collected

during the afternoon (1-3:30 p.m.). Males were resting or flying around the flower

heads and females were observed collecting pollen. This species was found only

at the type location; large patches of the host plant, even within 0.5 km of the

type locality, had no bees at the flowers. In both 1981 and 1982 we examined

many Wyethia plants at several other locations without finding additional pop¬

ulations of this Perdita.

Predator.—A female of Philanthus pulcher D.T. was netted after she had cap¬

tured a male of P. bohartorum on a Wyethia flower head.

Etymology.— It is a pleasure to name this distinctive bee after the brothers

Bohart who have contributed so much to our present day knowledge of the sys-

tematics and biology of Hymenoptera. Richard was my major advisor during my

graduate years at Davis, and George (Ned) was my “boss” and now is a colleague

in the bee lab at Logan, Utah.

Literature Cited

Cronquist, A., A. H. Holmgren, N. H. Holmgren, and J. L. Reveal. 1972. Intermountain flora. Vol.

1. Hafner Publ. Co., New York, 270 pp.

Hunt, C. B. 1974. Natural regions of the United States and Canada. W. H. Freeman and Co., San

Francisco, 725 pp.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 233-239

Expanding Geographical and Ecological Range of Platynota stultana

in California (Lepidoptera: Tortricidae)

Jerry A. Powell

Department of Entomological Sciences, University of California, Berkeley, Cal¬

ifornia 94720.

The moth that later became known to American economic entomologists as

the “omnivorous leaf-roller,” Platynota stultana (Walsingham), was described

from specimens collected by H. K. Morrison in the “Province of Sonora,” Mexico

(Walsingham, 1884). Probably the locality was near the international border be¬

cause Morrison’s biographers state that he worked in Arizona during 1881 and

New Mexico in 1882 (Mann, 1885), or 1882 and 1883 in Arizona (Smith, 1885),

but they do not mention Mexico among his expeditions. The species was collected

in what is now Cochise County in southeastern Arizona in the 1890’s by F. H.

Snow (AMNH, CU, KU) 1 and in Sinaloa, Mexico, as early as 1893 (USNM), so

the native range probably included semiarid parts of northwestern mainland Mex¬

ico and adjacent southwestern United States.

As the common name implies, a broad array of larval hostplants is known.

Atkins et al. (1957) summarized records representing about 20 families of An-

giospermae, most of which arose from urban and agricultural situations, whence

the insect has expanded its geographical range during the past 80 years. Platynota

stultana evidently was not native in cismontane southern California, because it

was not encountered in the early years of citrus investigations, during the 1880’s

and 1890’s, yet it became a conspicuous pest by 1913-1915 (Woglum, 1920).

History of Occurrence in California

Data associated with specimens examined (see acknowledgments) and in the

files of the California State Department of Food and Agriculture indicate the

following history of introduction and spread within the state.

The earliest records I have seen are in Los Angeles County, 1898 at La Mirada

and 1904 at Los Angeles (USNM). Had the species been native in the vicinity of

southern California towns, it certainly would have been encountered by Coquillett,

who reared and collected many microlepidoptera during 1883-1892 in the Los

Angeles area. As a comparison, there are a number of 1880’s records for the

orange tortrix, Argyrotaenia citrana (Fernald), a moth with similar larval and

adult habits and life history (Powell, 1964). By 1913-1915, however, P. stultana

had become an economic problem in several citrus and cut flower growing areas

of Los Angeles and Orange counties (Woglum, 1920; Bohart, 1942). Other early

collections of P. stultana include occurrences at San Diego, 1913 (SDNH), Needles

on the Colorado River, 1918 (CU), and San Bernardino by 1925. The species

also expanded southward, into Baja California, having been taken at Ensenada,

in 1941 (USNM).

1 See acknowledgments for list of institutional abbreviations.

234

PAN-PACIFIC ENTOMOLOGIST

Northward, P. stultana was taken at Saticoy, Ventura County, in 1925 (USNM),

and it reached coastal Santa Barbara County by 1940, although the date of initial

establishment and continuous residency there is unknown. There is a record for

Carpinteria in 1940 (CDFA), and P. stultana was reared from larvae during the

Channel Islands Biological Survey, on Anacapa Island, the same year (LACM).

The species evidently did not continue its northward expansion along the coast,

as there are no records north of Santa Barbara in field situations during the

subsequent 20 years.

Once established in southern California, a diversity of ornamental and agri¬

cultural plants was adopted. Records include tomato ( Lysopersicon ) (1898); Por-

tulaca (1904); Citrus and carnation {Diant hus) (1913); avocado {Persea) and wal¬

nut husks {Juglans) (1925); pepper pods ( Capsicum) (1926); pigweed ( Chenopodium )

and Malva (1929); youngberries ( Riibus) (1932); sugar beet (Beta) and Cyclamen

(1934) (USNM records). There were many interceptions of larvae in bell peppers

from Sinaloa at Nogales and Los Angeles during the late 1920’s and 1930’s,

resulting in a federal quarantine (Busck, 1933).

By contrast, there are few records of P. stultana on native plants. Goeden and

Ricker (1976a,b) encountered the species on native ragweeds {Ambrosia) in desert

areas and in roadside, abandoned cropland, and other artificial expansions of

Ambrosia habitat in cismontane southern California. Also in the 1970’s, the

species was reared by Gorelick from Eriogonum grande Greene, an insular en¬

demic plant, on Santa Catalina Island (CIS).

There are specimens from Sacramento reared in 1932 from alfalfa, and in

nursery situations during 1943-1945 (CDFA), but it appears that populations

have not been established continuously in the Sacramento Valley, because there

are no records for field situations between 1932 and 1955. Similarly, the omniv¬

orous leaf-roller was recorded in greenhouses in the San Francisco Bay area in

1940-1941 (Bohart, 1942); at Salmas, Monterey County, in 1955 (CDFA); and

in Alameda County at San Leandro (1957) (CDFA). However, prior to 1955,

there are no confirmed records of P. stultana established in the field anywhere

north of the Transverse Range, which separates cismontane southern California

from the rest of the state.

In the late 1950’s and during the following several years, populations of this

insect greatly expanded their geographic and ecological ranges in California. The

data suggest that changes in physiological tolerance developed in relation to phys¬

ical factors.

In 1956 adults of P. stultana were taken at lights in urban Davis, Yolo County

(20 km west of Sacramento) (CIS). Although I did not encounter the species during

three months’ sampling in Kern County in 1955, it appeared in agricultural field

situations there in 1959, in a light trap at Bakersfield in July (CIS), and feeding

on cotton flowers and fruit at Edison (10 km ESE of Bakersfield) in August (CDFA).

The following year P. stultana was taken on cotton at McFarland (40 km NNW

of Bakersfield), and during 1961-1963, the omnivorous leaf-roller quickly spread

throughout the San Joaquin and southern Sacramento valleys (Map 1). It acquired

considerable notoriety as a new pest of vineyards, beginning in 1963 (Lynn, 1969;

All Niazee and Stafford, 1972).

Platynota stultana reached the Monterey Bay area and inner Contra Costa

County (Antioch) by 1967 and Glenn County in the northern Sacramento Valley

VOLUME 59, NUMBERS 1-4

235

by 1968 (CIS, CDFA). Although there had been sporadic nursery and greenhouse

records in the San Francisco Bay area for more than 25 years, P. stultana was not

recorded out of doors until 1967, when it was reared from potted Senecio at

Albany, Alameda County (Frick and Hawkes, 1970; CIS). Adults first appeared

at lights in 1979, when they were collected in Berkeley (CIS). Its late establishment

on Santa Cruz Island, in the mid 1970’s (Powell, 1981) is postulated to be the

result of introduction by man following broader ecological adaptation by popu¬

lations of adjacent mainland areas, where P. stultana apparently was localized

and was rarely encountered during the 1940’s and 1950’s.

Increased Diversity of Larval Hostplants

In the process of the 1960’s expansion, P. stultana adapted to a wider diversity

of plants, including conifers and both dicotyledons and monocotyledons, a much

greater taxonomic array featuring more diverse chemical and physical character¬

istics, than the herbaceous angiosperms adopted in southern California during

the early part of the century. Thus, the physiological tolerance by P. stultana

expanded to encompass gymnosperms, including Pinaceae ( Pinus ) (Yolo Co.,

1966), Cupressaceae ( Juniperus ) (Fresno Co., 1966), and Taxaceae ( Taxus ) (Mon¬

terey Co., 1967); monocotyledons, Poaceae, including Zea (Tulare Co., 1963 and

Kern Co., 1969), Sorghum seed (Glenn Co., 1969); various additional legumes,

including Phaseolus (Stanislaus Co., 1961), Albizia (Kings Co., 1964), and Arachis

(Fresno Co., 1966); as well as Vitaceae and exotic representatives of many plant

families, such as Begoniaceae, Celastraceae, Aquifoliaceae, Theaceae, Aizoaceae,

Ginkgoaceae, and Ebenaceae (CDFA records), additional to host records sum¬

marized by Atkins et al. (1957).

Conifers, monocotyledons and legumes are kinds of plants generally used by

microlepidoptera that are specialized in terms of larval foods (Powell, 1980), and

none of these hosts had been recorded as a foodplant of P. stultana in California

prior to 1950. There is some indication that expanded host selection has taken

place in southern California populations as well, for example use of Ginkgo (1961),

Juniperus (1970), and Trifolium (1971) in Santa Barbara County (CDFA).

Whether physiological tolerance to physical factors such as high rainfall, low

temperature winters also broadened is problematic. There are early records for

desert areas, e.g., Needles (1918); Yuma (1925, 1928); and in San Diego County

at San Felipe Wash (1935) and Borrego (1941) (AMNH, USNM), indicating that

survival in arid climatic situations of extreme temperature ranges occurred. None¬

theless, range expansion of field populations into northern California seemed to

take place contemporaneously with broader hostplant use.

Discussion

Inconsistencies of sampling notwithstanding, it seems apparent that after col¬

onization around the turn of the century, Platynota stultana became widely es¬

tablished and adapted in southern California during the following 20 years, re¬

mained stable for another 30 years, then expanded its range in a relatively short

time to encompass low elevation areas in most of the rest of the state. The

maximum area (greatest right-angle, straight line distances) occupied in California

during 1915-1950, some 80,000 square km, more than tripled, to ca. 270,000

square km, during 1956-1968, with an increase in latitudinal range from 1°45 /

I968r •

Fig. 1. Geographical distribution of Platynota stultana in California: by 1925 (dark shading);

additional areas known to be inhabited by 1940 (cross hatched); later dated localities refer to first

records in counties north of the Transverse Ranges.

to 6°45 / (Fig. 1). At the same time there was adaptation to a considerably greater

spectrum of environmental conditions, particularly larval foods, shorter frost-free

season, and increased total precipitation.

A similar kind of range expansion, establishment followed by gradual encroach¬

ment in a restricted area for a long time, then rapid enlargement of the area

occupied, has been observed for other introduced insects. For example, the Eu¬

ropean skipper butterfly, Thymelicus lineola (Ochsenheimer), was discovered in

southern Ontario in 1910. Its early progress is poorly documented, but because

236

PAN-PACIFIC ENTOMOLOGIST

i96i km

CALIFORNIA INSECT SURVEY

Department of Entomology and Parasitology

UNIVERSITY OF CALIFORNIA

DRAFT I9SS 27/Uum

VOLUME 59, NUMBERS 1-4

237

butterflies are the best surveyed group of insects and because this species is a pest

of Timothy grass, the progress of its distribution in later years could be recon¬

structed with reasonable confidence (Burns, 1966). From 1910 to 1950 the range

did not enlarge much, south westward as far as Findlay in northern Ohio (1927)

and the vicinity of Ann Arbor, Michigan (1949), and northeastward to Toronto

(1945) and Niagara Falls (1948), a maximum range of ca. 46,000 square km.

Suddenly, during 1953-1965, records from widely scattered areas appeared, in¬

dicating that populations had expanded in all directions, to encompass most of

the northeastern U.S. and adjacent parts of Canada, from New Brunswick west

to Lake Superior and south to Maryland, a maximum area of ca. 1.5 million

square km. In addition, a colony was discovered in British Columbia, which may

have resulted from an independent introduction from Eurasia or via railroad from

the east (Bums, 1966).

It is probable that saltatorial range expansions following a period of slow prog¬

ress by colonist species often are masked by inadequate documentation of the

history of spread, or even that such bursts might appear to have occurred because

of gaps in the record. Thus, the Old World earwig, Euborellia annulipes (Lucas)

seems to have undergone a history in California similar to that of Platynota

stultana\ i.e., widespread establishment in southern California and at coastal

stations northward, from 1880 (and probably much earlier) to the 1920’s, then a

rapid expansion during 1932-1941, in which the Central Valley, Coachella Valley,

and Colorado River areas were occupied (Langston and Powell, 1975). By contrast,

an apparent accelerated expansion of the European tortricid moth, Croesia for-

skaleana (L.) in the northeastern U.S., was interpreted to be an artifact of lack of

effort by resident collectors (Powell and Burns, 1971). In both examples early

collection records are sporadic, however, and precise details of the range extensions

cannot be retraced.

The best documentation of insect introductions and subsequent spread have

come from examples of classical biological control. DeBach (1965) summarized

criteria of colonization and pointed out that only ca. 20-25% of attempted intro¬

ductions have resulted in successful establishment and then complete preadap¬

tation to the new environment usually seems to have been the rule. DeBach

discussed two possible cases of latent increase in adaptive fitness in entomopha-

gous insects. In one, Compenella bifasciata Howard (Aphelinidae), large releases

of progeny from a small founder colony (5 mated females) were made in various

parts of southern California in 1942. By 1946 it was apparent that establishment

was successful only at Riverside. During 1948-1957, however, the colonist pop¬

ulations spread westward and southward to inland districts of Los Angeles and

San Diego counties and continued to increase in abundance during the next several

years.

On theoretical evolutionary grounds, the most plausible explanation of delayed

ecogeographical expansions by introduced insects is one of increased genetic fitness

to environmental conditions in areas peripheral to the new home. It is generally

believed that introductions often consist of few founders. When their genetic

preadaptation sufficiently meets physical and biotic demands of the environment

at the place of introduction, colonization may be successful, often with establish¬

ment aided by release from biotic population controls in source populations. This

sets the stage for possible acceleration of evolutionary change. Selection pressure

238

PAN-PACIFIC ENTOMOLOGIST

at the margins of expanding founder populations may promote increased phys¬

iological adaptedness to environmental situations to which the founders, or even

the source populations, were not preadapted.

DeBach (1965) warned that observed long-term changes in distribution and

abundance of introduced species may be due to ecological factors rather than

genetic change. In particular, as demonstrated by population trends in several

aphelinid parasitoids of scale insects in California, competitive displacement either

by later arriving introduced species or by competitors already established, may

comprise an important, though cryptic influence affecting changes in populations.

Thus, for example, before assuming that increase in the distribution of Thymelicus

lineola was due to genetic change, we should consider whether ecological factors,

especially competition with native species of grass-eating Hesperiinae, might have

been involved. It would be interesting to search for changes in relative abundance

or distribution of native hesperiine species in the Great Lakes-Northeastern U.S.

region since 1950, and in changes in host grass selection by T. lineola.

By contrast, competitive displacement seems less likely in phytophagous, ho-

lometabolous species that are polyphagous as larvae, or in general feeders and

scavengers such as earwigs. Certainly it would be much more difficult to document

than in examples of closely related, specific feeders occupying similar niches to

one another. If altered genetic fitness is responsible for latent bursts of expansion,

the humbers of generations required presumably would vary widely with differing

selective regimes. On the average, for a univoltine insect such as Thymelicus, a

considerably longer period might be expected (e.g., 40-50 years), than for a homo¬

dynamic species such as Comperiella or Platynota with several annual generations

(e.g., 5-20 years).

ACKNO WLEDG MENTS

Collection data for early records were compiled by the late R. L. Lambert,

Ottawa, Canada, although I have reexamined most of the specimens. T. D. Eichlin,

California State Department of Food and Agriculture, Sacramento, provided cop¬

ies of all collection records for California counties north of Santa Barbara, many

of which are not represented by extant specimens. G. A. Gorelick provided spec¬

imens from Santa Catalina Island. Cooperation by authorities of institutional

collections enabled study of specimens in their care: ANMH—American Museum

of Natural History, New York; CDFA—California Department of Food and Ag¬

riculture, Sacramento; CIS—California Insect Survey, University of California,

Berkeley; CU—Cornell University, Ithaca, New York; KU—Snow Entomological

Museum, University of Kansas, Lawrence; LACM—Los Angeles County Museum

of Natural History, Los Angeles; SDNH—San Diego Natural History Museum,

San Diego, California; USNM—U.S. National Museum of Natural History, Wash¬

ington, D.C.

Dedication

This paper is dedicated to Prof. R. M. Bohart in commemoration of his long

and remarkably productive career, which included early work with Platynota

stultana and other California microlepidoptera. Twenty-nine years ago Bohart

and I participated in the University of California summer field course in Ento¬

mology, his initiation as an instructor and mine as a student. No doubt he re-

VOLUME 59, NUMBERS 1-4

239

covered from the experience in a few years, but I never did, for my attention had

been irrevocably turned from large to small insects, the really “innersting” ones.

Literature Cited

Ali Niazee, M. T., and E. M. Stafford. 1972. Notes on the biology, ecology, and damage of Platynota

stultana on grapes. J. Econ. Ent. 65:1042-1044.

Atkins, E. L. Jr., M. H. Frost, Jr., L. D. Anderson, and A. S. Deal. 1957. The “omnivorous leaf

roller”, Platynota stultana Wlshm., on cotton in California: Nomenclature, life history, and

bionomics (Lepidoptera, Tortricidae). Ann. Ent. Soc. Amer. 50:251-259.

Bohart, R. M. 1942. Platynota stultana as a pest of field-grown carnations. J. Econ. Ent. 35:399-

403.

Bums, J. M. 1966. Expanding distribution and evolutionary potential of Thymelicus lineola (Lep¬

idoptera: Hesperiidae), an introduced skipper, with special reference to its appearance in British

Columbia. Canad. Ent. 98:859-866.

Busck, A. 1933. In: Minutes of the 449lh regular meeting of the Entomological Society of Washington,

Oct. 5, 1933. Proc. Ent. Soc. Wash, 35:191-192.

DeBach, P. 1965. Some biological and ecological phenomena associated with colonizing ento-

mophagous insects, pp. 287-303. In: H. G. Baker and G. L. Stebbins, eds.. The genetics of

colonizing species. Academic Press, New York and London.

Frick, K. E., and R. B. Hawkes. 1970. Additional insects that feed upon tansy ragwort, Senecio

jacobaea, an introduced, weedy plant, in the western United States. Ann. Ent. Soc. Amer. 63:

1085-1090.

Goeden, R. D., and D. W. Ricker. 1976a. The phytophagous insect fauna of the ragweed Ambrosia

dumosa in southern California. Environ. Ent. 5:45-50.

-. 1976b. The phytophagous insect fauna of the ragweed Ambrosia psilostachva in southern

California, U.S.A. Environ. Ent. 5:1169-1177.

Langston, R. L., and J. A. Powell. 1975. The earwigs of California (Order Dermaptera). Bull. Calif.

Insect Survey 20:1-25.

Lynn, C. D. 1969. Omnivorous leaf roller, an important new grape pest in the San Joaquin Valley.

Calif. Agric. 23(4): 16-17.

Mann, B. P. 1885. Herbert Knowles Morrison. Psyche 4:287.

Powell, J. A. 1964. Biological and taxonomic studies on tortricine moths, with reference to the

species in California (Lepidoptera: Tortricidae). Univ. Calif. Publ. Ent. 32:1-317.

-. 1980. Evolution of larval food preferences in microlepidoptera. Ann. Rev. Ent. 25:133-159.

-. 1981. Five insects believed to be newly established or recolonized on Santa Cruz Island,

California (Dermaptera, Lepidoptera). Bull. So. Calif. Acad. Sci. 79:97-108.

Powell, J. A., and J. M. Bums. 1971. Colonization of the northeastern United States by two Palearctic

moths (Lepidoptera: Tortricidae). Psyche 78:38-48.

[Smith, J. B., ed.] 1885. Obituary note. Ent. Amer. 1:100.

Walsingham, Lord T. 1884. North American Tortricidae. Trans. Ent. Soc. London 1884:121-147.

Woglum, R. S. 1920. A recently discovered citrus pest, Platynota tinctana (Walk.) in California.

Calif. Dept. Agric. Mo. Bull. 9:341-343 [based on P. stultana misidentified by A. Busck].

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 240-245

Identification and Synonymies of Two Western Palearctic

Cerceris : maculata Radoszkowski and hathor n. sp.

(Hymenoptera: Sphecidae)

WOJCIECH J. PULAWSKI

California Academy of Sciences, Golden Gate Park, San Francisco, California

94118.

The identities of Cerceris orientalis Mocsary and haematina Kohl have been a

problem for hymenopterists because their original descriptions were inadequate

and the types could not be found. This paper presents the results of my study of

these problems and is dedicated to the anniversary of Dr. R. M. Bohart. I sincerely

thank Arnold S. Menke for discussing various aspects of the study, and Mary

Ann Tenorio for drawing the figures.

Cerceris maculata Radoszkowski

Cerceris maculata Radoszkowski, 1877:57, 2. Holotype 2: Uzbek SSR: Kyzil Kum

Desert (Zool. Mus. Moscow Univ.). —Shestakov, 1918:141.

Cerceris orientalis Mocsary, 1883:47, 2. Holotype 2: “Russia mer.-orient.” = Ka¬

zakh SSR, not Caucasus (lost, see below). Neotype: holotype of Cerceris mac¬

ulata Radoszkowski, present designation. Nec F. Smith, 1856. New synonym.

Cerceris eugenia Schletterer, 1887:390, new name for Cerceris orientalis Mocsary,

1883. —Shestakov, 1918:131; Kazenas, 1978:58.

Cerceris Moscaryi Kohl, 1888:139 (incorrect original spelling), new name for

Cerceris orientalis Mocsary, 1883. —Schletterer, 1889:890; ? Shestakov, 1918:

146 (as mokzaryi).

Cerceris haematina Kohl, 1916:111, 2. Holotype 2: origin unknown (lost, see

below). Neotype: holotype of Cerceris maculata Radoszkowski, present desig¬

nation. New synonym.

Diagnosis. — The female of maculata can be distinguished from other members

of the bupresticida species group by the roundly triangular lamella of sternum V.

The male can be recognized by the absence of micropunctation on the frons

adjacent to the frontoclypeal suture between the subantennal sclerite and tentorial

pit. See the description of hathor for further details.

Nomenclature.—The correct interpretation of eugenia and haematina is par¬

ticularly difficult because the original descriptions are inadequate, and the type

material is apparently lost. As Dr. M. Fischer kindly informed me in his letter of

16 November 1981, no specimen named eugenia, haematina, or orientalis can

be found in the Naturhistorisches Museum, Vienna, a situation already partly

reported by de Beaumont (1951b:340). A specimen thought to be the holotype

of orientalis Mocsary, housed at the Hungarian National Museum, Budapest, was

kindly submitted to me by Dr. J. Papp. It bears a handwritten label “ Cerceris

mocsaryi Kohl, orientalis Mocsary” in Mocsary’s handwriting, but no locality

label. It is a female of fischeri (see below) and is apparently the specimen studied

VOLUME 59, NUMBERS 1-4

241

Table 1. Comparison of presumed holotype of orientalis Mocsary (Hungarian National Museum,

Budapest) with original description (1883).

Body part

Specimen from Budapest

Mocsary, 1883 description

1. Scape

all yellow

black basodorsally

2. Clypeal apex

brownish, truncate

black, crenulate

3. Tergum II

yellow basally

yellow laterally

4. Tergum IV

black

yellow posteriorly and laterally

5. Tibiae

all yellow

yellow, darkened mesoventrally

by Schletterer (1887, 1889) and by Kohl in 1888 (Kohl, 1888:139, 1916:111) and

regarded by them as Mocsary’s holotype. Actually, it cannot be the true holotype

of orientalis, because it differs significantly from the original description, as shown

in Table 1.

The discrepancy in the color of terga II and IV is particularly striking and I

doubt that Mocsary would have made such an error. Schletterer redescribed

orientalis twice: in 1887 under the replacement name eugenia Schletterer, and in

1889 under the replacement name mocsaryi Kohl. As Kohl (1888) pointed out,

Schletterer’s 1887 redescription was based on a specimen not conspecific with the

holotype. The same must now be said about his 1889 redescription: it clearly

indicates a specimen different from the material Mocsary (1883) had before him,

and it well agrees with the specimen labelled orientalis received for study from

the Hungarian National Museum. This suggests that the holotype of orientalis

was confused by Schletterer with two other specimens, so that a total of three

must have been involved : 1. the holotype of orientalis (its possible fate is discussed

below), 2. an Egyptian female of hathor on which Schletterer (1887) based his

description of eugenia (now lost, see also under hathor below), and 3. a female

of fischeri which Kohl (1888) and Schletterer (1889) regarded as the holotype of

orientalis, on which they based their redescriptions of mocsaryi, which was then

returned to Budapest as the holotype of orientalis, and which finally was submitted

to me.

It is also very probable that the holotype of haematina Kohl was the misplaced

holotype of orientalis Mocsary. Descriptions of these two species show an almost

identical set of characters, including the characteristic lateral spots on tergum II.

The original locality or area of haematina is unknown, and the holotype of ori¬

entalis had no origin label, a coincidence which also suggests that the same spec¬

imen may have been involved. Most probably, the true holotype of orientalis was

kept unrecognized in the Vienna Museum while the supposed false holotype was

returned to Budapest after publication of Schletterer’s monograph (1887, 1889).

When casually discovered by Kohl (1916) many years later, it was then described

by him as haematina.

According to the original description, the holotype of orientalis originated from

“Russia meridionalis vel Caucasus.” However, this origin is corrected to “Russia

mer.-orient,” in Mocsary’s characteristic handwriting in a reprint of the original

paper (a xerox copy kindly sent by Dr. J. Papp). “Russia mer.-orient.” of Mocsary

is almost certainly today’s Kazakh SSR.

Several characters given in Mocsary’s original description help in recognition

242

PAN-PACIFIC ENTOMOLOGIST

of orientalis : clypeus convex (without discal process), with crenulate free margin;

propodeal enclosure unsculptured; propodeum and gastral tergum I reddish; ter-

gum II with lateral spots; terga III-V fasciate (fasciae broadened laterally). These

characters suggest that orientalis is identical to maculata. I have seen the holotype

and four additional females of maculata (three of them kindly sent by Dr. V. L.

Kazenas, Alma Ata). None is identical in coloration to the holotype of orientalis,

but the observed variation of the latter strongly suggests that they are conspecific

with it. For example, the propodeum in the specimens studied is all black, or with

small red spots, or largely red. Gastral tergum II has an oblong, yellow, lateral

spot or the yellow covers the whole tergum except a semilunar apicomedian area

(as in the holotype of maculata). There is little doubt that many specimens of

maculata have exactly the same coloration as Mocsary’s holotype. Because of the

observed variation, I designate the holotype of maculata as the neotype of both

orientalis and haematina, even if the yellow area of tergum II is larger in the first

than indicated in the descriptions of the latter two. This decision will end a

complex nomenclatorial problem.

Cerceris fischeri Spinola

Cercerisfischeri Spinola, 1839:493, 9, S. Syntypes: Egypt (Inst. Zool. Univ. Turin),

see de Beaumont, 195la: 175.

Cerceris mocsaryi: Kohl, 1888:139; Schletterer, 1889:890.

The specimen which Kohl (1888) and Schletterer (1889) erroneously regarded

as the holotype of orientalis Mocsary and which they redescribed under the name

mocsaryi Kohl (see above) is a female of fischeri, a member of the rybyensis

species group of de Beaumont (1951b). The female of fischeri can be recognized

by the following combination of characters: propodeum around enclosure im-

punctate or with a few scattered, punctures; and sternum V prominent postero-

laterally. The male can be recognized by the presence of a well defined, basal

platform on sternum II combined with acutely angulate posterolaterally sternum

VI. Subsidiary recognition features of both sexes are: hypoepimeral area carinate

below, propodeal enclosure unsculptured.

Cerceris hathor Pulawski, New Species

Cerceris eugenia: Schletterer, 1887:390; Kohl, 1888:139, 1916:111; de Beaumont,

1951a:180, 1951b:339, 1953:122, 1956:186, 1958:59; de Beaumont and By-

tinski-Salz, 1959:122; Pulawski, 1964:73.

Cerceris tricolorata: Mochi, 1938:190; Giner Marl, 1941:174.

Cerceris vidua: Honore, 1941:150, 1942:69.

Etymology.— Hathor, a goddess of ancient Egyptians, in apposition.

Nomenclature.—As Kohl (1888) pointed out, Schletterer (1887) examined the

holotype of Cerceris orientalis Mocsary and renamed the species eugenia, but his

description was based on a specimen belonging to a different species. Kohl (1888)

and also de Beaumont (1951b) applied the name eugenia to this latter species,

but Article 72(d) of the Code is clear: the replaced name ( orientalis ) and the

replacement name {eugenia) must have the same holotype. This means that the

species described by Schletterer under the name eugenia must bear a different

name.

VOLUME 59, NUMBERS 1-4

243

Schletterer (1877) mentioned two geographic areas in which eugenia supposedly

occurs: southeastern Russia (= present Kazakh SSR) and Egypt. Since the holotype

of eugenia originated from the first area, his description must have been based

on an Egyptian specimen. Even though the latter was subsequently lost, de Beau¬

mont (1951b) was able to recognize and to characterize (as eugenia) the species

to which it belonged. The species has no available name and is here described as

hathor.

Systematics. — Cerceris hathor is a member of the bupresticida species group of

de Beaumont (1951b). The group, redefined by Krombein (1981), includes the

following species:

bidentula Maidl, 1926; southern India, Sri Lanka, Thailand, Malaya, Celebes

(= langkasukae Pagden, 1934)

ssp. bidentula spiniventris Tsuneki, 1963; Thailand

bupresticida Dufour, 1841; Mediterranean basin, Transcaspia, Afghanistan

hathor Pulawski, n. sp.; North Africa, Israel, Azerbaidzhan

kokuevi Shestakov, 1912; Transcaspia

(= egregia Kazenas, 1977, according to Y. L. Kazenas’s identification labels)

maculata Radoszkowski, 1877; Transcaspia

(= eugenia Schletterer, 1887, haernatina Kohl, 1916)

mastogaster Smith, 1856; India, Sri Lanka

odontophora Schletterer, 1887; Balkans, Turkey, Iran, Transcaspia

palmetorum de Beaumont, 1951; North Africa

supposita Kohl, 1916, as interpreted by Kazenas, 1978 (perhaps not conspecific

with the holotype of supposita); Transcaspia

tricolorata Spinola, 1839; North Africa

This list may be incomplete, and other described species, especially Oriental and

Ethiopian, may belong here. Some of the species listed above are insufficiently

known, and some were unavailable for comparison during this study. My inter¬

pretation of supposita and kokuevi is based on specimens so labelled and kindly

sent to me by Dr. V. L. Kazenas.

Diagnosis.— The female of hathor can be recognized by the combination of:

propodeum sparsely punctate, sternum V nondentate posterolaterally, and lamella

of sternum V evenly arcuate. The shape of gastral segment VI is the same in males

of hathor, maculata and odontophora: its sternum is dentate posterolaterally, and

its tergum is not. Unlike maculata, the face in males of hathor is densely micro-

punctate including the area above the frontoclypeal suture, and unlike odonto¬

phora, the propodeum is sparsely punctate outside the enclosure.

Comparative description.— At least some punctures of the propodeal dorsum

around enclosure are about 1 diameter apart (punctures compressed against each

other in bidentula spiniventris, bupresticida, and odontophora, and in some males

of maculata). Hindcoxa with medioventral carina (carina absent in kokuevi and

palmetorum). Gastral sternum II with indistinctly delimited basal platform (sim¬

ilar platform present in kokuevi, maculata, mastogaster, supposita, and female of

bidentula spiniventris ; platform absent in bupresticida, odontophora, tricolorata,

and male of bidentula spiniventris). Thorax black or partly red, but tegula and in

most populations also pronotum and metanotum pale yellow (metanotum black

244

PAN-PACIFIC ENTOMOLOGIST

maculata

Fig. 1. Species of Cerceris. Ventral view of female sternum V showing apical outline. Small drawing

to right of each illustration shows cross-section profile of sternal apex.

in odontophora, mesoscutum and propodeum partly to all yellow in kokuevi and

palmetorum).

Female. — Clypeal middle section with two admedian tubercles near free margin

(tubercles absent in tricolorata). Sterna III-V without sublateral process (process

present in mastogaster : Krombein, 1981). Gastral sternum V (Fig. 1) tuberculate

posterolaterally (similarly in odontophora and tricolorata, angulate in maculata

and supposita, dentate in bidentula spiniventris). Apicomedian lamella of sternum

V evenly rounded, its midlength less than its width (lamella almost identical in

bidentula spiniventris', with semicircular notch in bupresticida', roundly triangular

in maculata', strong, erect, triangular in mastogaster: Krombein, 1981; semicir¬

cular, shallowly notched apically in odontophora ; trapezoidal, with width about

twice length in supposita', with widely concave free margin m tricolorata). Lateral

carina of tergum VI not expanded (expanded at midlength in supposita).

Male.—Face with large punctures and numerous micropunctures between ten¬

torial pit, antennal socket and orbit (micropunctures indistinct in bidentula spi¬

niventris, absent adjacent to frontoclypeal suture in maculata and supposita).

Gastral terga I and II not elongate: spiracles of tergum I closer to tergal hindmargin

than to each other (terga I and II elongate in bidentula spiniventris and tricolorata:

spiracles of tergum I about equidistant from each other and tergal hindmargin).

A spine-like, posterolateral projection present on sternum VI, but absent from

tergum VI (similar in maculata and odontophora ; projection absent on tergum

and sternum in kokuevi and palmetorum:, and present on tergum and sternum in

bidentula spiniventris, bupresticida, mastogaster, and tricolorata).

Geographic distribution.— Desert habitats of North Africa and Israel, probably

also Azerbaidzhan SSR.

Material examined. — Holotype 2: EGYPT, Ghiza near Cairo, 20 Apr. 1958,

W. J. Pulawski collector (W. J. Pulawski collection). Paratypes: EGYPT, Kom

VOLUME 59, NUMBERS 1-4

245

Oshim on Ghiza-Fayum road, 19 April to 13 May 1958, same collector and

depository (1 $, 4 6).

Literature records. — MOROCCO: Midelt (de Beaumont, 1951b, 1953). AL¬

GERIA (de Beaumont, 1951, 1958): Biskra, Laghouat, Tadjemout, Tassili des

Ajjer (Oued Teneouene, Oued Ti’Harat). CHAD: Tibesti: Zouarke (de Beaumont,

1956). EGYPT: Ghiza near Cairo, Kom Oshim on Cairo-Fayum road (Pulawski,

1964), Gebel Asfar near Cairo (Honore, 1941). ISRAEL (Negev Desert): Beer-

sheba, Gvulot, Kfar Yeroham, Revivim (de Beaumont and Bytinski-Salz, 1959).

AZERBAIDZHAN SSR: Adzhikend (Kohl, 1916).

Literature Cited

de Beaumont, J. 1951a. Synonymie de quelques Cerceris. 2. (Hym., Sphecid.). Mitt. Schweiz. Ent.

Ges. 24:175-180.

-. 1951b. Contribution a l’etude des Cerceris nord-africains. EOS Rev. Espanola Ent. 27:299-

408.

-. 1953 (1952). Hymenopteres recoltes par une mission suisse au Maroc (1947). Sphecidae 2.

Bull. Soc. Sci. Nat. Phys. Maroc 32:107-131.

-. 1956. Sphecidae (Hym.) recoltes en Libye et au Tibesti par M. Kenneth M. Guichard. Bull.

Brit. Mus. (Nat. Hist.) Ent. 4:165-215.

-. 1958. Hymenopteres Sphecides de la mission du Tassili des Ajjer (1949). Trav. Inst. Rech.

Sahar. 3:55-71.

de Beaumont, J., and H. Bytinski-Salz. 1959. The Sphecidae (Hymen.) of Eretz Israel. II. Subfam.:

Nyssoninae (tribes: Gorytini, Nyssonini, Alyssonini) and Philanthinae. Bull. Res. Counc. Israel

B8:99—151.

Giner Mari, J. 1941. Los Cerceris de Africa paleartica. EOS Rev. Espanola Ent. 17:135-291.

Honore, A. 1941. Notes sur quelques Cerceris de la faune egyptienne (Hymenoptera: Sphegidae).

Bull. Soc. Fouad I er Ent. 25:145-162.

-. 1942. Introduction a l’etude des Sphegides en Egypte (Hymenoptera: Aculeata). Bull. Soc.

Fouad I er Ent. 26:25-56.

Kazenas, V. L. 1978. Royushchiye osy Kazakhstana i Sredney Azii (Hymenoptera, Sphecidae).

Opredelitel. Izdatelstvo Nauka Kazakhskoy SSR, Alma Ata, 172 pp.

Kohl, F. F. 1888. Neue Hymenopteren in den Sammlungen des k. k. naturhistorischen Hofmuseums.

Verh. Zool. Bot. Ges. Wien 38:133-156.

-. 1916 (1915). Beitrag zur Kenntnis der Gattung Cerceris auf Grundlage der O. Radoszkov-

skyschen Sammlung. Arch. Naturgesch., Abt. A, 88 (7): 107-125.

Krombein, K. V. 1981. Biosystematic studies of Ceylonese wasps, VIII, A monograph of Philanthidae

(Hymenoptera: Sphecoidea). Smithsonian Contrib. Zool. No. 343:i—iii, 1-75.

Mochi, A. 1938. Revisione delle specie egiziane del genere Cerceris Latr. (Hymenoptera: Sphegidae-

Philanthinae). Bull. Soc. Fouad l er Ent. 22:136-229.

Mocsary, A. 1883. Hymenoptera nova Europaea et exotica. Ertek Termeszettud. Korebol 13(11):

1-72.

Pulawski, W. J. 1974. Etudes sur les Sphecidae (Hym.) d’Egypte. Polskie Pismo Ent. 34:63-155.

Radoszkowski, O. 1877. Sphegidae. In: Voyage au Turkestan d’A. P. Fedchenko, fasc. 14, tome 2,

partie 5. Bull. Soc. Imper. Amis. Sci. Nat. 26:1-87.

Schletterer, A. 1887. Die Hymenopteren-Gattung Cerceris Latr. mit vorzugsweiser Beriicksichtigung

der palaarktischer Arten. Zool. Jahr., Abt. Syst. 2:349-510.

-. 1889. Nachtragliches iiber die Hymenopteren-Gattung Cerceris Latr. Zool. Jahr., Abt. Syst.

4:879-904.

Shestakov, A. V. 1918 (1917). Materiaux pour servir a une faune des guepes du genre Cerceris Latr.

(Hymenoptera, Crabronidae). Ann. Mus. Zool. Acad. Sci. Russie 22:1 18-166.

Spinola, M. 1839 (1838). Compte-rendu des Hymenopteres recueillis par M. Fischer pendant son

voyage en Egypte, et communiques par M. le Docteur Waltl a Maximilien Spinola. Ann. Soc.

Ent. France 7:437-546.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 246-248

A Fluorescing Insect, Cysteodemus armatus LeConte

(Coleoptera: Meloidae)

R. W. Rust and R. C. Bechtel

(RWR) Biology Department, University of Nevada; (RCB) Nevada State De¬

partment of Agriculture, Reno, Nevada.

This paper is not about Sphecidae, Vespidae or Chrysididae. It is not even

about Hymenoptera, but concerns a very unusual, perhaps unique beetle. Because

the uniqueness of the beetle reflects many of the same qualities possessed by Dick

Bohart, apparent to both his students and peers, we honor him on his seventieth

birthday by dedicating this paper to him.

In the animal world, very few species possess structures or structural modifi¬

cations that fluoresce. Some deep sea fishes (Bond, 1979) and marine chaetop-

terians (Maglitsch, 1972) and cephalopods (Barns, 1980) fluoresce, many with the

aid of fluorescing bacteria. Of the terrestrial animals, scorpions (Williams, 1968;

Williams and Hadley, 1967) and some agelenid spiders (T. Lugaski, pers. comm.)

fluoresce. Here we report on the fluorescence observed for the first time in an

insect, the meloid beetle, Cysteodemus armatus LeConte.

While collecting scorpions with ultraviolet (UV) lamps near Lava Dune, 10 km

WNW Lathrop Wells, Nye County, Nevada (116 o 30 , W-36°41"N) on May 15,

1982, a roundish yellow-green fluorescing “rock” was found. The “rock” got up

and walked away from the radius of the UV light. It proved to be a living specimen

of Cysteodemus armatus (Meloidae: Meloinae: Eupomphini). The specimen was

placed in a glass specimen vial and returned to camp for further observation.

Continued collecting with UV lamps that evening produced no more specimens

of C. armatus. The fluorescence had been observed with light from a 120 volt 6

watt black light tube (F6T5/BLB). In white light or sun light the beetle had a light

yellow dusting on the dorsal surface of the exoskeleton which was responsible for

the fluorescence observed with the UV light (Fig. 1). The material, a cuticular

secretion most likely wax or lac epicuticle, was easily picked off with a dissecting

needle or fine scalpel. The material was not soluble in ethanol, chloroform or

water and retained its fluorescing property in those liquids. Eleven additional

specimens of C. armatus in the collection of the Nevada State Department of

Agriculture also fluoresced under UV fight. These specimens were from several

different locations in Nevada. Fourteen genera of meloid beetles were checked for

UV fluorescence, but none fluoresced. One or more species in the following genera

were checked: Meloe, Cordylospasta, Eupompha, Spastonyx, Phodaga, Pleuros-

pasta, Tegrodera, Pyrota, Lytta, Epicauta, Li ns ley a, Nemognatha, Tricrania and

Pseudozonitis.

Other Coleoptera with “dusty” exoskeletons were examined under UV fight for

fluorescence, but none fluoresced. Species examined included: (Elateridae) Alaus

melanops LeConte, A. myops (Fabricius), A. oculatus (L.) and Chaleolepidius

asperata discreta Blaisdell, C. variolosa Horn, Phloeodes diabolicus LeConte, P.

VOLUME 59, NUMBERS 1-4

247

Fig. 1. Cysteodemus armatus LeConte photographed with UV light and Kodak® Plus X film with

30 second exposure. The waxy epicuticle has been removed from the right elytron to show the

contrasting pattern of fluorescence.

pustulosus LeConte, Zopherus elegans Horn, Z. laevicollis Solier, Z. nodulosus

haldemani Horn and Z. nodulosus nodulosus Solier.

Two questions arise from this observation: how and why fluroesce? The “how”

question will be answered by biochemists when the fluorescing compound is

known. Many organic molecules are known to fluoresce (anthracene, perylene,

pyrene, quinine) and the fluorescence produced is from radiative transitions be¬

tween states of like multiplicity and visible most often in LTV light (Cowan and

Drisko, 1976). The “why” question is more difficult to answer. However, a simple

explanation may be that certain organic compounds fluoresce and one of these

compounds is found in the waxy layers of the C. armatus exoskeleton. Thus, the

fluorescence would have nothing to do with evolution or selection in the past

history of the beetle. On the other hand, the compound that fluoresces could be

involved with antipredator behavior, waterproofing, light or heat reflectance, stor¬

age of a waste product, or be more basic in mate selection or location. We do not

have enough information to know.

248

PAN-PACIFIC ENTOMOLOGIST

We thank Drs. Gary Bloomquist and Glenn Miller for their thoughts and com¬

ments on the chemistry of fluorescing compounds.

Literature Cited

Baras, R. D. 1980. Invertebrate zoology. Saunders Publ., Philadelphia.

Bond, C. E. 1979. Biology of fishes. Saunders Publ., Philadelphia.

Cowan, D. O., and R. L. Deisko. 1976. Elements of organic photochemistry. Plenum Press, New

York.

Meglitsch, P. A. 1972. Invertebrate zoology. Oxford University Press, New York.

Williams, S. C. 1968. Methods of sampling scorpion populations. Proc. Calif. Acad. Sci. (4)36:221-

230.

Williams, S. C., and N. F. Hadley. 1967. Scorpions of the Puerto Penasco area (Cholla Bay), Sonora,

Mexico, with description of Vejovis baergi, new species. Proc. Calif. Acad. Sci. (4)35:103-116.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 249-253

A New Spider Parasitoid, Ocnaea boharti, from Arizona and

New Mexico (Diptera: Acroceridae)

Evert I. Schlinger

Department of Entomological Sciences, University of California, Berkeley, Cal¬

ifornia 94720.

The genus Ocnaea Erichson (1840) has 19 included species, ranging from Cal¬

ifornia and Georgia, south to Brazil and Ecuador. Nine species occur in the United

States, five in Central America and the Caribbean, and three in Brazil and Ecuador.

In addition to the new species described here, I have on hand four additional

species from Brazil, Venezuela and Costa Rica. Species from Chile described

earlier in Ocnaea were discussed by Schlinger (1968) and were referred to Ar-

rhynchus Philippi. All species described in Pialeoidea Westwood were placed in

Ocnaea by Schlinger (1961), and other Ocnaea species have been removed and

placed in Exetasis Walker (see Schlinger, 1968 and 1972).

Biological notes on species of Ocnaea are rare, and the only recorded host is

the California trapdoor spider, Bothriocyrtum californicum, from which Jenks

(1940) reared Ocnaea smithi Sabrosky (1948). Known host records and immature

stages of Ocnaea and related panopine genera were published by Schlinger (1972).

Octiaea. species are rather large flies (5-20 mm), yet published records on known

specimens, including the type specimens, have numbered less than 25 since the

genus was described 142 years ago. During the past 30 years that I have been

identifying acrocerids from collections from around the world, I have examined

only an additional 15 specimens. These data attest to the peculiar rare status of

these “large” flies in that only 40 specimens are known to have been collected or

reared in the past 150 years.

A further complication to the study of this genus is the high degree of sexual

dimorphism known in the few species where both sexes have been associated.

Other than the new species described below, only O. smithi has both sexes de¬

scribed. This indicates that some new synonomy may be expected as further

specimens become available.

I wish to thank Dr. Mont Cazier, Arizona State University at Tempe, for the

loan of the holotype specimen and Dr. James Sublette, New Mexico State Uni¬

versity at Las Cruces for the loan of the paratype specimen. These type specimens

will be deposited in the California Academy of Sciences, San Francisco, California.

Special acknowledgment is given to Mrs. Celeste Green, my friend and former

Scientific Illustrator in the Department of Entomology, University of California

at Berkeley, for her excellent color rendering of this new species.

Ocnaea boharti Schlinger, New Species

(Plate I, Figs. 1, 2)

Male. — Length of entire specimen 10.00 mm, wing length 7.00 mm.

Color.— Yellow, white, brown, and black, with faint metallic bluish-brown re¬

flexions on mesonotum, scutellum and on medial tergal spots; black are eyes and

250

PAN-PACIFIC ENTOMOLOGIST

occiput; dark brown are outer and dorsal margins of antennal segment III, antennal

segments I—II, posterior Vi of antennal tubercle, mesonotum, scutellum, coxae,

most of pleura, sternal area (almost black), most of tergite I, large medial spot on

tergites II and V, smaller medial spot on tergites III-IY, and tarsal claws; yellow

are remainder of tergal areas, most of sternites I-IV, most of legs, squamal rim

and wing veins; light brown are inner and ventral margins of antennal segment

III, anterior V2 of antennal tubercle, area around and beneath wing base, post-alar

callus, inner sides of femora I and II, tiny spot at knees, tibial spurs, tip of basitarsi,

pulvilli, halter stem and knob, small lateral spot on sternite IV and most of sternite

V; dull white are humerus, squama (nearly opaque), wing membrane (hyaline),

narrow posterior fascia on tergites I and II and sternite IV, and somewhat wider

fascia on sternites II and III; genitalia are yellowish-brown, and ocelli are orange-

brown.

Pile. — Light brown and dense on eyes and occiput, reaching out to base of

antennal segment II; longer below near mouth and above near ocelli; that on

thorax dense, white, long, nearly obscuring ground color, somewhat longer than

the longest pile on eyes; that on abdomen mostly white, long and dense on tergite

II dorsally and tergites V and VI laterally; that on medial tergal spots light brown

and shorter; remainder white, shorter, and less dense than on mesonotum.

Head. — V 3 higher than long (PI. I), antenna V 3 longer than head height; antenna

asetate except, for small group of dorsal setae on segment II; proboscis not evident,

but a group of stout hairs project from underneath tiny “proboscial” covering.

Thorax .—With shining ground color, difficult to detect under dense pile; hu¬

merus strongly developed and directed anteriorly, reaching in front of eyes, be¬

coming nearly acuminate (PI. 1); humerus length subequal to that of head height;

femur III hardly longer than I and II; tibia III swollen and about as long as femur

III; (PI. I), venation with cell R 5 open at wing margin and membrane without

macro trichiae.

Abdomen.— Shiny, rather narrow, not much wider than thorax; tergites II-IV

subequal in width; genital capsule large, aedeagus formed as in Fig. 1.

Female.— Length of entire specimen 10.00 mm, wing length 10.00 mm. Similar

to that described for male above except for obvious differences as follows:

Color. — Black also are antennal segment III (except extreme base dark brown),

most of coxae and tergite I; dark brown with bluish-brown to black reflections

on large dorso-medial spot, and extreme lateral margins of tergites II and III, all

of tergites IV-VI, and most of sternites I-VI (except for narrow whitish-light

brown fascia on sternites III and IV); dark brown are squamal rim, halter knob,

femora, inner side of tibia and tarsi (most legs are broken at end of tibia or absent

altogether); whitish-brown are outer basal V3 of tibiae and humerus; light brown

are wing veins.

Pile. — Not dense throughout, more dark brown on eyes, that on mesonotum

lightly mixed with light brown; that on mesonotum lightly mixed with light brown;

that on abdomen more brown than white.

Head. — Much smaller than male (Fig. 2), more round, about as high as long;

antenna IV 2 times longer than head height; antennal segment III with small but

distinct group of dorsal, subapical setae.

Thorax. — Shiny, ground color not obscured by pile; humerus Vs longer than

head height, slightly more curved and more acuminate than in male (Fig. 2); tibia

VOLUME 59, NUMBERS 1-4

251

Plate I. Ocnaea boharti, holotype male.

Ill not swollen, but tibia II shortened and sinuated; squamal membrane nearly

clear; wing membrane lightly infuscated throughout; sparse but distinct macro-

trichiae present in membrane throughout length of costal cell; venation similar

to male but some cells (especially R 5 ) are longer and therefore somewhat differently

shaped.

Abdomen. — Little wider than thorax, widest at tergite III (broken off and re¬

glued).

Types .—Holotype <5, ARIZONA: 2 mi. NE Portal, November 3, 1960 (M. A.

Cazier), from the Arizona State University at Tempe Collection. Paratype 2, NEW

MEXICO: Roosevelt Co., October 31, 1964 (A. Berg) from the New Mexico

University Collection in Las Cruces.

I should note that while I have little doubt that the paratype is the female of

this species, certain secondary sexual characters which are evident in this specimen

are known to occur in both sexes of certain other species of Ocnaea. One such

character is the presence of a distinct, but sparse series of macrotrichiae in the

costal cell. Another is the darker brown wing veins and lightly infuscated wing

membrane. Finally, the small, more-rounded head of this female is obviously

different (compare PI. I with Fig. 2), and while this trait occurs in some species

of certain related genera, i.e. Pialea Erichson, it is not known in Ocnaea species.

252

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Ocnaea boharti, aedeagus in lateral view, holotype.

Fig. 2. Ocnaea boharti, head and humerus in lateral view, para type female.

VOLUME 59, NUMBERS 1-4

253

This new species, with its stiletto-like humeri, shows no close relationship to

any known Ocnaea, and seems to form its own unique species group. Its color

combination of yellow, brown and shiny bluish-brown reflections are very rem¬

iniscent of the Chilean species, Arrhynchus vittatus Philippi. The relationship of

Arrhynchus to Ocnaea was discussed in detail by Schlinger (1968), and while these

genera are closely related, no species within either genus has this distinctly mod¬

ified humerus (in either sex) while O. boharti n. sp., has this character in both

sexes.

It is with the greatest pleasure that I name this new species for my colleague

and former major professor, Dr. Richard M. Bohart, for whom I have the greatest

respect both as a person and as a professional entomologist.

Literature Cited

Erichson, W. F. 1840. Die Henopier, eine Familie aus der Ordnung der Dipteren, pp. 135-175, pi.

I, figs. 7-10. Entomographien, Untersuchungenin dem Gebiete der Entomologie mit besonderer

Benutzung der Konigl. Sammlung zu Berlin, vol. 1, Berlin.

Jenks, G. E. 1940. The spider’s “uninvited” fly brings doom. Natural History (Magazine) 45:157-

161.

Sabrosky, C. W. 1948. A further contribution to the classification of the North American spider

parasites of the family Acroceratidae (Diptera). Amer. Mid. Nat. 39:382-430.

Schlinger, E. 1. 1961. New species of Acrocera from Arizona and Ocnaea from California, with

synonymical notes on the Genus Ocnaea. Ent. News 72:7-12.

-. 1968. Studies in neotropical Acroceridae, Part 1. A revision of Arrhynchus Philippi and a

key to the genera of the Ocnaea branch of the Panopinae. Rev. Chilena Ent. 6:47-54.

-. 1972. A new Brazilian panopine species, Exetasis eickstedtae, reared from the theraphosid

spider, Lasiodora klugi (Koch), with a description of its immature larval stages (Diptera: Ac¬

roceridae). Papeis Avulsos Zool. S. Paulo 26:73-82.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 254-255

A New Species of Macrobiotus from Tierra del Fuego

(Tardigrada: Macrobiotidae)

Robert O. Schuster 1

Department of Entomology, University of California, Davis, California 95616.

A new tardigrade from Tierra del Fuego is described in this paper. This species

has claws which are sufficiently different from typical Macrobiotus claws to strong¬

ly suggest that it represents an undescribed genus. However, the difference is only

an extreme modification of a single character which is normally somewhat variable

in other species of Macrobiotus, and this species is tentatively placed there.

Macrobiotus tridigitus Schuster, New Species

(Figs. 1, 2)

Holotype. — Total length, from extended buccal opening to end of claws IV, 246

gm. Cuticle smooth, pores not evident by phase microscopy. Without eye pig¬

mentation. Buccal apparatus (Fig. 1) 47 ^m long; tube including buccal ring 34

g m; placoid strands 13 gm; tube with long ventral lamina. Pharynx contains

apophyses, 3 macroplacoids with first and third of similar size and perceptibly

longer than second, and microplacoid. Claw sequence 2, 1, 1,2 (Fig. 2), accessory

points of primary claw branches very long, not attached along claw margin; lunulae

large, without teeth.

Type series. — Holotype and 1 paratype from Sierra Martial, Tierra del Fuego,

ARGENTINA, in cryptogams, tree line (2000-2500 feet), 1-19-1979, Arthur M.

Shapiro. Deposited with the Department of Entomology, University of California,

Davis.

Etymology.— Latin; masculine, tres, three; digitus, finger, descriptive of the

claws.

Discussion.— The buccal ring of this species is either longitudinally grooved or

completely divided into buccal lamellae but the exact structure cannot be deter¬

mined from the available specimens. If ten buccal lamellae do exist, the only

departure from typical Macrobiotus species (see Ramazzotti, 1972, and Schuster

et al., 1980) is the extreme elongation of the cuticle forming the accessory points

of the primary claw branches. Normally the cuticle is attached along the edge of

each primary claw branch and terminates in two small, subapical spines. The

accessory points of this species terminate as thin cuticular processes that arise at

the base of the primary branch, are much longer than the branch, and are not

attached to it. These processes seem to be quite inflexible and probably provide

some mechanical advantage to locomotion. They do distinguish M. tridigitus from

all other Macrobiotus.

1 I appreciate having the opportunity to participate in this tribute to the systematic contributions

by Dr. Richard M. Bohart.

VOLUME 59, NUMBERS 1-4

255

Figs. 1, 2. Macrobiotus tridigitus. 1, detail of buccal tube and placoids. 2, claws of leg II.

Literature Cited

Ramazzotti, G. 1972. II Phylum Tardigrada. Seconda edizioni aggiomata. Mem. 1st. Ital. Idrobiol.

28:1-732.

Schuster, R. O., D. R. Nelson, A. A. Grigarick, and D. Christenberry. 1980. Systematic Criteria of

the Eutardigrada. Trans. Amer. Micros. Soc. 99:284-303.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 256-266

New North American Pulverro Pate with a Key to the

Species (Hymenoptera: Sphecidae)

Norman J. Smith

Fresno County Department of Agriculture, 1730 South Maple Avenue, Fresno,

California 93702.

Pulverro Pate is a member of the sphecid subfamily Pemphredoninae and its

distribution is limited to western North America. These wasps are relatively small,

mostly black and average 3 mm in length. They are predaceous on thrips and

usually nest in sandy ground or soil that has been recently disturbed (Bohart and

Grissell, 1972). Most species occur in arid areas of the west where they visit native

flowers but show no particular specificity.

Pulverro contains 11 species, 6 previously described ( monticola Eighme, costano

Pate, mescalero Pate, constrictus (Provancher), laevis (Provancher), and colum-

bianus (Kohl)) and 5 described in this paper (boharti, patei, eighmei, coconino

and mexicanus). In Pate’s second publication on the subtribe Ammoplanma (1939)

he suggested the presence of two species groups in Pulverro but did not set them

apart with characters. With the collection of additional specimens and new species,

it appears that there are two fairly well defined species groups, the mescalero group

and the laevis group. The mescalero group contains 5 species and is distinguished

by the following: males; mandible with ventral edge simple, without tooth or

carina, narrowing apically (Fig. 5); scutum with apicolateral corners squared (Fig.

13), extending forward over pronotum (Fig. 2); females; scutum same as males;

frons with tumid areas basad of antennal sockets, median area usually depressed.

The laevis group contains 6 species and is distinguished by the following: males;

mandible with ventral edge having a prominent tooth medially or carina apically

(Figs. 3, 4); scutum with apicolateral corners rounded (Fig. 12), not extending

forward over pronotum (Fig. 1); females; scutum same as males; frons without

tumid areas basad of antennal sockets; median area forming a smooth carina,

usually terminating approximately halfway between median ocellus and basal edge

of clypeus.

In this paper I describe 5 new species and provide a key to the valid species.

A later paper will present a revision of Pulverro and discuss relationships between

the species.

Key to Male Pulverro

1. Mandible with ventral edge simple, without tooth or carina, narrowed

apically (Fig. 5); scutum with apicolateral corners squared (Fig. 13),

extended forward over pronotum (Fig. 2) ( mescalero group) . 2

- Mandible with ventral edge having prominent tooth medially (Figs. 3,

4) or carina apically, mandible enlarged apically; scutum with apicolater¬

al corners rounded (Fig. 12), not extended forward over pronotum (Fig.

1 ) (laevis group). 6

VOLUME 59, NUMBERS 1-4

257

2. Mandible with medio ventral translucent tooth on inner margin (Fig. 11)

. 3

- Mandible without medioventral translucent tooth on inner margin ... 4

3. Clypeus mediobasally with prominent angular process; abdominal ster¬

num 8 occasionally with apicomedian tooth larger than lateral teeth (as

in Fig. 8). columbianus (Kohl)

- Clypeus mediobasally with small, nipple-like projection, abdominal

sternum 8 with teeth approximately equal size (as in Fig. 7).

. monticola Eighme

4. Antennal flagellomeres 1-5 (at least) nodulate ventrally (Fig. 17) . 5

- Antennal flagellomeres not nodulate ventrally . mescalero Pate

5. Clypeus broadly triangular, lacking basomedial nipple-like projection;

flagellomeres ventrally mostly brown to black . mexicanns N. J. Smith

- Clypeus narrowly triangular, with basomedial nipple-like projection;

flagellomeres ventrally entirely yellow . boharti N. J. Smith

6 . Clypeus basally with highly polished area. 7

- Clypeus basally without polished area .. 8

7. Sterna 4 and 5 with erect tufts of white hair when viewed laterally;

sternum 5 with median, tooth-like carina projecting distally; sternum 8

rounded apically (Fig. 8) . coconino N. J. Smith

- Sterna 4 and 5 without erect tufts of hair; sternum 5 without median

tooth-like carina; sternum 8 pointed apically (Fig. 7) .

. constrictus (Provancher)

8 . Mandibles with prominent tooth medioventrally (Figs. 3, 4); clypeus

apicolaterally with sharp points. 9

- Mandibles with only a low carina apicoventrally; clypeus apicolaterally

without sharp points . 10

9. Mandible with ventral tooth large and rounded apically (Fig. 3); clypeus

polished, without silver hair. laevis (Provancher)

- Mandible with ventral tooth moderate in size and pointed apically (Fig.

4); clypeus not polished, with silver hair. costano Pate

10. Clypeus apically with 3 serrations (Fig. 16), mediobasally without a

nipple-like projection; notauli present but obscure .. . eighmei N. J. Smith

- Clypeus apically truncate, mediobasally with a nipple-like projection;

notauli deeply impressed. patei N. J. Smith

Key to Female Pulverro

1. Scutum with apicolateral corners squared (Fig. 13), extended over prono-

tum (Fig. 2); frons with tumid areas above antennal sockets, median

area usually depressed ( mescalero group) .

- Scutum with apicolateral corners rounded (Fig. 12), not extended over

pronotum (Fig. 1); frons without tumid areas above antennal sockets,

median area with low carina, usually terminating about halfway between

median ocellus and clypeus ( laevis group).

2. Upper frons punctures 1-3 puncture diameters apart, interspaces usually

polished .

- Upper frons punctures no more than 1 diameter apart, usually less,

interspaces dull .

258

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-11. 1, 2, lateral view of scutum and pronotum; 1, P. costano, female; 2, P. monticola,

male, G—gena, SC—scutum, T—tegula, PN—pronotum. 3-5, male, lateral view of mandible and

lower part of head; 3, P. laevis; 4, P. costano; 5, P. monticola. 6, P. mexicanus, male, front view of

head. 7-9, male sternum 8, ventral view; 7, P. constrictus ; 8, P. coconino, AES—apical end of sternum,

SH —shaft; 9, P. costano. 10, P. patei, female, face. 11, P. monticola, male, dorsal view of mandible.

VOLUME 59, NUMBERS 1-4

259

Figs. 12-17. 12, 13, dorsal view of scutal shoulders, male; 12, P. costano\ 13, P. monticola. 14,

15, front view of clypeus and antennal sockets, female; 14, P. constrictus ; 15, P. coconino. 16, P.

eighmei, male, front view of clypeus. 17, P. mexicanus, male, lateral view of basal flagellomeres; DE —

distal end, PE—proximal end.

3. Clypeus evenly covered with sparse silvery pubescence, head obviously

wider than long (0.88 as long as wide, n = 3). mescalero Pate

- Clypeus medially highly polished, sparse silver pubescence laterally,

head about as long as wide (1.00 as long as wide, n = 3) .

. boharti N. J. Smith

4. Frons with tumid areas strongly delineated; head much wider than long

(0.82 as long as wide, n = 3); clypeal disc wider than long, slightly

polished . monticola Eighme

- Frons with tumid areas slightly delineated; head slightly wider than long

(0.96 as long as wide, n = 3); clypeal disc about as long as wide, highly

polished . columbianus (Kohl)

5. Clypeus globose, without silvery pubescence, polished; occipital carina

present ventrally . 6

- Clypeus slightly convex, somewhat flattened, covered with silvery pu¬

bescence; occipital carina absent or evanescent ventrally . 8

260

PAN-PACIFIC ENTOMOLOGIST

6 . Upper frons sparsely punctate, polished, concave; scutum flattened, pol¬

ished, notauli deeply impressed. patei N. J. Smith

- Upper frons densely punctate, not polished, somewhat convex; scutum

convex, not polished, notauli moderately impressed . 7

7. Frontal carina sharp; species collected in interior areas of northern and

central California ... costano Pate

- Frontal carina evanescent; species collected along southern California

coast . laevis (Provancher)

8 . Stigma light brown to yellow; frontal carina evanescent, scapal basins

not well differentiated; occipital carina evanescent ventrally.

. eighmei N. J. Smith

- Stigma dark brown to black; frontal carina and scapal basins well dif¬

ferentiated; occipital carina absent ventrally. 9

9. Clypeus extended dorsad past midline of antennal sockets (Fig. 15), with

sparse silvery pubescence, apical half often yellow-brown; northern Ar¬

izona and southern Utah. coconino N. J. Smith

- Clypeus not extended dorsad past midline of antennal sockets (Fig. 14),

with moderate silvery pubescence, only tip yellow-brown; mostly south¬

ern California deserts. constrictus (Provancher)

Pulverro boharti N. J. Smith, New Species

Holotype male .—Length 3 mm. Black; mouthparts, antennae ventrally, fore¬

tibia, mid and hindtibia at knees, foretarsi and mid and hindtarsi at base yellow.

Wing veins brown, stigma dark brown. Body with sparse hair, abdominal segments

with sparse hair, clypeus and lower frons with sparse silver pubescence. Mandibles

bidentate, ventrally with apical half not having low carina, narrowing apically;

inside ventral margin without hyaline tooth. Clypeus medially subtriangular, api¬

cal edge truncate, mediobasal edge with nipple-like projection. Lower frons with

low, broad frontal carina medially; scapal basins slightly differentiated, shallow;

upper frons rugose. Antennae with basal flagellomeres nodulate ventrally (as in

Fig. 17); pedicel strongly nodulate. Midventral line approximately l h length of

forecoxa.

Scutum with apicolateral corners moderately squared and slightly extended over

pronotum (as in Figs. 2, 13), dorsal surface convex and moderately hairy, surface

slightly rugose; notauli moderately impressed. Propodeum dorsally with two par¬

allel carinae; reticulate between. Mesopleuron polished, setigerous punctures sparse.

Sternum glabrous, highly polished.

Sternum 8 apically serrate and rounded, moderately hairy (as in Fig. 9).

Female. —Length 3.5 mm. Markings as in male. Head approximately as long

as wide (1.00, n = 3). Mandibles as in male. Clypeus broadly and slightly concave

apicomedially; clypeal disc with sparse silver pubescence laterally, highly polished

medially. Lower frons with tumid areas basad of antennal sockets moderately

developed, converging basomedially, medially with shallow depression; scapal

basins strongly defined, lightly silvered apically. Frons with small, shallow, central,

circular depression. Upper frons with setigerous punctures moderately spaced (2-

VOLUME 59, NUMBERS 1-4

261

3 puncture diameters apart), polished between in Wyoming specimens, faintly

rugose in Arizona specimens.

Scutum with apicolateral corners moderately squared and extended over prono-

tum (as in Figs. 2, 13); notauli obscure; scutum dorsally convex and rugose.

Pygidium trigonal and margined by sharp carina with few punctures laterally.

Diagnosis.—P. boharti is a member of the group mescalero . Major distinguish¬

ing characters of the male are the nipple-like projection on the basomedial edge

of the clypeus, clypeus with sparse silver pubescence; basoflagellomeres nodulate

ventrally (as in Fig. 17) and sternum 8 rounded (as in Fig. 9). Males of boharti

most closely resemble those of mescalero and mexicanus. All three lack the trans¬

lucent tooth on the inner medioventral margin of the mandible but mescalero

lacks the nodulate basoflagellomeres common to boharti and mexicanus. The

nipple-like projection of the clypeus of boharti males separates the species from

mexicanus, and the latter has a rectangular occipital area behind the compound

eyes.

Major female distinguishing characters are: clypeal disc polished medially and

with silver pubescence laterally; head approximately as long as wide and upper

frons (at least in the Wyoming specimens) sparsely punctate and polished between

punctures. Females of mescalero are most closely related to those of boharti. They

are related by their upper frons punctures being only 1-2 puncture diameters apart

and polished between punctures. P. boharti differs from mescalero in having a

medially polished clypeus which is sparsely silvered laterally and the head is about

as long as wide. P. mescalero lacks the polished area on the clypeus and the head

is wider than long.

Type material. — Holotype 8 (UCD): WYOMING, Sweetwater Co., 12 mi. s.

Green River, 28 July 1969, R. M. Bohart. Paratypes, 15 <5, 1 2, same data as

holotype. Other localities (all specimens collected by P. H. Timberlake, UCR,

unless otherwise stated): ARIZONA: Navajo Co.: Hotevilla, 27 Sept. 1977, 4 <3,

2 2 (R. M. Bohart, UCD); Hotevilla, 30 Aug. 1976, 1 5, 1 2 (R. M. Bohart, UCD);

3 mi. w. Old Oraibi, 24 Sept. 1964, 1 6, 1 2; 23 mi. n. Indian Wells, 14 Sept.

1960, 1 2. Coconino Co.: Tuba City, 24 Sept. 1964, 10 <3, 6 2. Apache Co.: 16.6-

51.5 mi. se. Round Rock, 28 Aug. 1961, 9 8, 1 1 2; 18 mi. sw. Ganado, 29 Aug.

1967, on Gutierrezia microcephila, 1 8. NEW MEXICO: McKinley Co.: 29 mi.

s. Gallup, 17 Sept. 1968, 1 8 ; 8 mi. sw. Ramah, 17 Sept. 1968, 1 8. UTAH: Uintah

Co.: Bonanza, 8-30 Aug. 1975, 2 <3, 1 2 (G. E. Bohart, USU). Duchesne Co.:

Fruitland, 30 Aug. 1949, 1 2 (G. F. Knowlton, USU). COLORADO: Rio Grande

Co.: 5 mi. nw. South Fork, 14 Aug. 1978, 9 8 (T. Griswold, personal).

Etymology.—P. boharti is named after R. M. Bohart in recognition of his sys¬

tematic work on the ammoplanines and the many specimens and new species

that he has collected over the years.

Pulverro coconino N. J. Smith, New Species

(Figs. 8, 15)

Holotype male .—Length 3.25 mm. Black; mouthparts yellow; foretibia, mid

and hindtibia at knees and all tarsi burnished yellow. Wing veins light brown,

stigma dark brown. Body mostly glabrous but tiny hairs can be found over most

262

PAN-PACIFIC ENTOMOLOGIST

parts. Clypeal disc with sparse silver pubescence. Mandibles bidentate apically;

ventrally with carina developed into 2 (one larger than the other), small, blunt

teeth medially. Clypeus with median triangular platform, truncate apically, sur¬

rounded laterobasally by highly polished vertically rising ridge, smoothly pro¬

duced mediobasally to extend well basad and above midline of antennal sockets

(as in Fig. 15). Lower frons flat medially, scapal basins not differentiated; upper

frons rugose, convex. Antennae entirely dark brown; basal flagellomeres slightly

nodulate ventrally. Midventral line of head approximately one-fourth length of

forecoxa.

Scutum with apicolateral comers rounded and not extending over pronotum,

dorsal surface convex with setigerous punctures moderate (1-2 puncture diameters

apart) and polished between. Propodeum dorsal surface with two subparallel

carinae, reticulate between. Abdominal sterna 4 and 5 with median erect tufts of

hair when viewed laterally; sternum 7 with erect tuft of hair easily visible laterally,

pointing back toward tufts on sterna 4 and 5; sternum 5 with median carina

produced into tooth pointing distally.

Sternum 8 serrate, sub truncate apically with a large median tooth, not obviously

spatulate, moderately hairy (Fig. 8).

Female.— Length 3.25 mm. Markings as in male except antennae almost entirely

yellow, clypeus burnished yellow on apical half. Clypeus sparsely silvered. Man¬

dibles bidentate apically and without ventral carina or teeth. Clypeus medially

with slightly raised trigonal platform, apically evenly rounded, convex, medio¬

basally produced basad of midline of antennal sockets (Fig. 15). Lower frons

medially with low, sharp carina, ending approximately halfway between median

ocellus and clypeus; scapal basin shallow, moderately defined; upper frons with

setigerous punctures moderate (1-2 puncture diameters apart) and polished be¬

tween. Midventral line of head long due to absence of occipital carina.

Scutum as in male. Propodeum dorsally with subparallel carinae weakly de¬

veloped. Abdomen ventrally without distinguishing characters. Pygidium trigonal,

margined with sharp carina, punctate evenly over entire surface.

Diagnosis. —P. coconino is a member of the laevis group. Distinguishing char¬

acters for male coconino are the mediobasal polished area on the clypeus, the

unique erect tufts of hair on sterna 4, 5 and 7, and the tooth-like median carina

on sternum 5. The closest relative is P. constrictus, based on the shared mediobasal

polished area of the clypeus found in coconino but constrictus lacks the abdominal

characters.

Distinguishing characters for female coconino are the apical half of clypeus

burnished yellow and sparsely silvered and clypeus extending upward between

antennal sockets (Fig. 15). P. coconino and constrictus share a flattened, slightly

convex, sparsely silvery pubescent clypeus and the absence of the occipital carina

ventrally. The clypeus of constrictus is entirely black and the mediobasal area is

somewhat truncate, not extended past the midline of the antennal sockets (Fig.

14).

Type material.— Holotype 8 (CAS): ARIZONA, Coconino Co., Tuba City, 24

Sept. 1964, on Chrysothamnus, P. H. Timberlake. Paratypes; 6 8, 5 2 (7 with same

data as holotype): ARIZONA: Coconino Co.: 26 mi. e. Tuba City, 24 Sept. 1964,

on Chrysothamnus, 3 8, 1 2 (P. H. Timberlake, UCR). Numerous other specimens

VOLUME 59, NUMBERS 1-4

263

were studied from Emery and Wayne Counties in southern Utah collected at

approximately the same time of year by F. D. Parker, D. Viers and T. Griswold.

Pulverro eighmei N. J. Smith, New Species

(Fig. 16)

Male holotype. — Length 3 mm. Black; mouthparts, ventral side of antennae

yellow; foretibia, mid and hindtibia at knees and all tarsi burnished yellow. Wing

veins hyaline to light yellow; stigma brown to yellow (variable in paratypes).

Clypeus with sparse to moderate silver pubescence; scapal basins with sparse

silver pubescence apically. Mandible bidentate apically; apicoventrally with low

carina, mandible widened apically; inside edge without tooth. Clypeus and median

area triserrate apically (Fig. 16), with sparse silver pubescence, moderately convex

discally. Lower frons without frontal carina, but with flat, narrow, smooth line

from clypeus to median ocellus, scapal basins shallow, not well defined; upper

frons convex, rugose. Antennal flagellomeres mostly terrete, appearing slightly

nodulate ventrally in some specimens, light brown dorsally. Midventral line ap¬

proximately Vi length of forecoxa. Occipital carina ventrally evanescent, actually

appearing recessed.

Scutum with apicolateral corners rounded (as in Fig. 12), not extended forward

over pronotum (as in Fig. 1); dorsal surface convex, rugose; notauli obscure.

Mesopleuron shiny but not smooth, somewhat rugose. Propodeum dorsally with

numerous, low, longitudinal carinae.

Sternum 8 slightly spatulate, apically serrate and sparsely hairy.

Female. — Length 3.5 mm. Markings as in male except stigma only rarely light

brown. Mandibles as in male except apicoventrally without carina. Clypeus convex

discally with sparse silver pubescence, slightly convex and burnished yellow api¬

cally. Lower frons with low, broad carina medially, ending approximately %

distance between clypeus and median ocellus, changing to smooth line at that

point; upper frons polished but with setigerous punctures dense (equal to or less

than one puncture diameter apart); scapal basins shallow, rather broad, tumidity

basad of antennal sockets absent. Midventral line as in male; ventral, occipital

carina as in male. Scutum, propodeum and mesopleuron as in male. Pygidium

margined by sharp carina, medially with longitudinal tumid area, laterally im-

punctate.

Diagnosis.—P. eighmei belongs to the laevis group. Major distinguishing char¬

acters for male eighmei are the tri-serrate clypeus and the evanescent, recessed

ventral area of the occipital carina. The tri-serrate clypeus alone will distinguish

eighmei from all other species in the genus.

Distinguishing female characters are the yellow stigma and the evanescent re¬

cessed occipital carina. The yellow stigma is unique and will distinguish this sex

from all other Pulverro females.

P. eighmei does not appear to have close relatives, however the evanescent,

recessed occipital carina in the female suggests a relationship with coconino and

constrictus which have lost their occipital carina.

Type material— Holotype <5 (UCD): CALIFORNIA, San Bernardino Co., Kra¬

mer Hills, 8 May 1978, on Machaeranther a tortifolia, R. W. Brooks. Paratypes:

86 3, 63 2, same data as holotype. Numerous other specimens have been collected

264

PAN-PACIFIC ENTOMOLOGIST

throughout the desert regions of southern California, eastern Nevada, western

Arizona and northern Baja, California.

Etymology.—P. eighmei (pronounced a-me-I) is named after Dr. Lloyd E.

Eighme, who has contributed significantly to work on Pulverro. Dr. Eighme also

supplied valuable taxonomic information and specimens as well as criticism on

the manuscript.

Pulverro mexicanus N. J. Smith, New Species

(Figs. 6, 17)

Male holotype. — Length 3 mm. Black; mouthparts yellow; foretibia, mid and

hindtibia at knees, fore and midtarsi burnished yellow. Wing veins dark brown,

stigma black. Clypeus and apical edge of scapal basins with dense silver pubes¬

cence. Mandibles bidentate apically; apicoventrally lacking the low carina; inner

median edge lacking hyaline tooth. Clypeus medially with pentagonal-shaped

platform, concave discally, very slightly concave apically. Lower frons medially

with broad carina, ending approximately halfway between clypeus and median

ocellus; scapal basins sharply defined; upper frons convex, rugose. Occipital region

boxlike, extending straight back from compound eye before turning to middle

(Fig. 6). Antennal flagellomeres ventrally with basal 6 nodulate (Fig. 17). Mid-

ventral line approximately half length of forecoxa.

Scutum with apicolateral corners squared and extending forward over pronotum

(Figs. 2, 13), convex dorsally, rugose; notauli moderately excised. Mesopleuron

shiny but not polished, slightly rugose. Propodeum dorsally with numerous, lon¬

gitudinal, low carinae. Female unknown.

Diagnosis.— P. mexicanus is a member of the mescalero group. Major distin¬

guishing characters are: flagellomeres 1-6 ventrally nodulate (Fig. 17); clypeus

medially with a pentagonal platform; and occipital area in back of the compound

eyes is rectangular (Fig. 6). P. boharti and mescalero are the nearest relatives,

their relationship being discussed under the boharti diagnosis. The last two major

distinguishing characters will separate mexicanus from all other species of the

genus. Females of mexicanus are not known but would probably resemble those

of mescalero and boharti.

Type material. — Holotype S (USU): MEXICO, Oaxaca, 3 mi. se. Yahuitlan,

17 Sept. 1974, G. E. Bohart & W. Hanson. Paratypes: 1 <3, same data as holotype

(UCD). MEXICO: Coahuila: Monclova, 21 Sept. 1974, 1 <3 (G. E. Bohart & W.

Hanson, USU).

Etymology.—P. mexicanus, known only from the 3 males listed above, was

collected from two widely separated localities in Mexico, and is named for the

country of origin.

Pulverro patei N. J. Smith, New Species

(Fig. 10)

Holotype male. — Length 3 mm. Black; mouthparts, foretibia, mid and hindtibia

at knees and all tarsi yellow. Wing veins light brown, stigma dark brown. Body

mostly glabrous; abdominal segments sparsely hairy, clypeus with moderate silver

pubescence; lower frons and especially scapal basins with dense silver pubescence.

Mandibles bidentate, ventrally with apical half having low carina; inner edge with

small, hyaline tooth. Clypeus medially subtriangular, concave discally, apex lat-

VOLUME 59, NUMBERS 1-4

265

erally with tiny, blunt teeth, mediobasally with nipple-like projection. Frons ridged

medially, approximately halfway from clypeus to midocellus; scapal basins well

defined, shallow; upper frons sparsely punctate but faintly rugose. Antennae en¬

tirely black, all segments terete. Midventral line between occipital carina and

hypostomal carina approximately % length of forecoxa.

Scutum with apicolateral corners rounded and not extending over pronotum

(as in Figs. 1, 12); notauli deeply impressed; setigerous punctures sparse, minutely

reticulate between, Propodeal dorsum reticulate with two, parallel longitudinal

carinae. Mesopleuron with setigerous punctures sparse, highly polished between.

Sternum 6 glabrous, highly polished.

Sternum 8 spatulate, serrate and with long hair.

Female.— Length 3 mm. Markings as in male except antennae are almost en¬

tirely yellow. Head appearing narrowed, longer than wide (Fig. 10). Mandibles

lack low carina and hyaline tooth. Central clypeal disc roundly tumid and highly

polished, basal edge extending between antennal sockets, apical edge truncate.

Clypeus and frons without silver pubescence. Frons ridged medially, approxi¬

mately Vs distance from clypeus to midocellus; upper frons with setigerous punc¬

tures sparse, highly polished between; upper median area concave. Propodeum

tricarinate. Pygidial plate trigonal and margined by sharp carina, with few punc¬

tures laterally.

Diagnosis. — P. patei belongs in the laevis species group. Unique in this species

is the fact that males and females appear to be more closely related to different

species in their respective sexes. Males, except for the species group characters,

would appear to be close to monticola and columbianus because of the presence

of a small hyaline tooth medioventrally on the inside of the mandible (as in Fig.

11) and a small, nipple-like projection mediobasally on the clypeus. Females on

the other hand show a close affinity to costano and laevis, primarily due to the

strongly tumid, polished disc on the clypeus much as in costano and laevis.

Characters that definitely relate the male and female are the sparse setigerous

punctures on the upper frons and scutum and the smooth, polished appearance

of both areas, as well as deeply impressed notauli. Locality records also bear out

their relationship.

Good distinguishing characters for the male are the antennae entirely black,

terete; the basomedial nipple-like projection on the clypeus, and the highly pol¬

ished scutum with its sparse, setigerous punctures and deeply impressed notauli.

Distinguishing characters for the female: the tumid, polished clypeal disc, the

highly polished upper frons with its sparse setigerous punctures and central con¬

cavity; the overall narrow appearance of the head (Fig. 10); the dorsally flattened

scutum with its sparse setigerous punctures and polished appearance.

Type material. — Holotype S (LTCD): ARIZONA, Navajo Co., Hotevilla, 27 Sept.

1977, on Chrysothamnus, R. M. Bohart. Paratypes, 4 S, 1 9, same data as holotype.

Other paratypes (all collected by P. H. Timberlalce, UCR, unless stated otherwise):

ARIZONA: Navajo Co.: Dinnebeta Wash, 30 Aug. 1976, 2 <5 (R. M. Bohart,

UCD); 3 mi. w. Old Oraibi, 24 Sept. 1964, on Chrysothamnus, 12 <5, 17 9; 11 mi.

n. Holbook, 22 Sept. 1964, on Gutierrezia lucida, 1 6. Coconino Co.: Tuba City

and 26 mi. e. Tuba City, 24 Sept. 1964, on Chrysothamnus, 5 6, 6 $. Apache Co.:

24 mi. nw. Concho, 25 Sept. 1964, on Chrysothamnus, 1 $, 1 2; 18 mi. sw. Ganado,

25 Sept. 1964, on Baileya, 1 <5; Steamboat Canyon, 14 Sept. 1961, on Chryso-

266

PAN-PACIFIC ENTOMOLOGIST

thamnus, 1 3. UTAH: Kane Co.: 23 mi. e. Kanab, 23 Sept. 1964, on Chryso-

thamnus, 1 3; Glenn Canyon, 25 Sept. 1964, on Gutierrezia lucida, 1 3. Other

specimens studied: NEW MEXICO: Socorro Co.: 3 mi. w. Bingham, 12 Sept.

1961, on Baileya pleniradiata, 29 3, 1 2 (2 3, coll, by P. D. Hurd, UCB). Lincoln

Co.: 24 mi. w. Carrizozo, 12 Sept. 1961, on Gutierrezia, 5 3. De Baca Co.: Fort

Sumner, 2 Oct. 1965, on Gutierrezia , 1 3 (G. E. Bohart, USU).

Etymology.—P. patei is named after V. S. L. Pate, a prolific worker on the

Ammoplanina.

Acknowledgments

I would like to thank the following museums and individuals for their generous

loan of specimens: University of California at Riverside (UCR, P. H. Timberlake,

S. Frommer), at Berkeley (CIS, J. Powell), and at Davis (UCD, R. Schuster); Utah

State University (USU, G. E. Bohart, W. Hanson); Pacific Union College (PUC,

L. Eighme); Terry Griswold (personal); U.S. National Museum (USNM, A. Menke);

California Department of Food and Agriculture (CDFA, M. Wasbauer); Phila¬

delphia Academy of Natural Sciences (ANSP, D. Otte); American Museum of

Natural History (AMNH, M. Favreau); Canadian National Collection (CNC, L.

Masner); and Cornell University (COR, L. Pechuman). Holotypes selected from

U.C. Davis specimens are deposited at Davis. Holotypes from loaned specimens

are deposited as stated for each species.

I would like to thank Professor Richard M. Bohart for all his advice and patience

during this study. It could not have been accomplished without his previous work

on the genus, his contribution of specimens and his generous sharing of personal

research money for invaluable collection trips. Professors Robbin W. Thorp and

Albert A. Grigarick also read the manuscript and made helpful comments. Others

who were helpful in collecting specimens or making suggestions were Lloyd E.

Eighme, Robert W. Brooks, Lynn S. Kimsey, Robert L. Kimsey and Larry D.

French.

Literature Cited

Bohart, R. M., and E. E. Grissell. 1972. Nesting habits and larva of Pulverro monticola. Pan-Pac.

Ent. 48:145-149.

Pate, V. S. L. 1939. Studies in the pemphredonine wasps. II. Records and descriptions of new forms

in the ammoplanoid complex from southwestern United States. Trans. Amer. Ent. Soc. 64:

373-420.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 267-280

Taxonomic and Nomenclatural Studies on American Polistine

Wasps (Hymenoptera: Vespidae)

Roy R. Snelling

Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los

Angeles, California 90007.

The paper is dedicated to my colleague Dr. Richard M. Bohart, with thanks

for his encouragement and patient assistance over the past thirty years.

The recent monograph by Richards (1978) of the social wasps (except Vespinae)

of the New World has done much to improve our understanding of the taxonomy

of these wasps. Such a monumental work cannot be without errors, nor can all

the taxonomic decisions made therein be final or universally accepted. Therefore,

in the spirit of continuing and partially correcting the study so admirably initiated

by Richards, I offer the following in the hope of further clarifying the taxonomy

and nomenclature of some of these wasps.

Genus Polistes Latreille

Polistes is a large, cosmopolitan genus of social wasps. The increasing interest

in the social behavior of wasps has focused on Polistes because species occur

almost everywhere, the wasps themselves are relatively mild tempered, and nest

activity takes place on a single exposed comb.

The taxonomy of Polistes is, however, less amenable to easy study: often the

subtle differences between species have not been appreciated by earlier workers,

whose species concepts were often broad enough to include several closely related

species by modern criteria. Many species are apparently widespread, sometimes

exhibiting a bewildering range of color variants. These are considered localized,

or geographical, races or varieties and often have been formally named; some

species have acquired a formidable clutter of supposed geographic subspecies. In

some cases, these forms are truly nothing more than color variants. In others,

they appear to be sympatric or allopatric sibling species. Often, morphological

evidence is inconclusive and some decisions as to the status of a given form may

be nothing more than intuitive.

Subgenus Aphanilopterus Meunier

Polistes exclam ans of Authors

Richards (1978) treated seven forms as geographical segregates of P. exclamans.

This was in line with my own views on this wasp (Snelling, 1954, 1955, 1970).

Previously the following have been considered subspecies of P. exclamans : ari-

zonensis Snelling, bahamensis Bequaert and Salt (= louisianus Bequaert), bili-

neo/aftt-S Bequaert and Salt, durangoens /^Snelling, lineonotus R. Bohart, picturatus

Bequaert and Salt, as well as exclamans Viereck (s. s.)- I no longer believe this

interpretation to be correct. Rather, I am of the opinion that there are several

species, morphologically almost identical, largely allopatric and, possibly, behav-

268

PAN-PACIFIC ENTOMOLOGIST

iorally different. Most of these forms are wholly allopatric and internally homo¬

geneous. I now interpret the group as follows:

P. arizonensis New status

P. bahamensis New status

= P. exclamans louisianus (syn. by Snelling, 1955)

= P. bahamensis bilineolatus New synonymy

= P. bahamensis picturatus New synonymy

P. exclamans New status

= P. exclamans durangoensis New synonymy

= P. instabihs coahuilae New synonymy

P. lineonotus New status

The form that occurs in southern Arizona and adjacent Mexico was originally

described as P. e. arizonensis . Throughout its range, this is a remarkably consistent

wasp and its color pattern exhibits no tendency to vary toward the richer markings

of P. exclamans to the east nor toward P. lineonotus in Lower California. The

head and body are reddish-brown, without mesonotal stripes and with very narrow

apical fasciae on the first to third terga. Both P. exclamans and P. lineonotus have

abundant yellow marks on the thorax, including longitudinal mesonotal stripes

in P. lineonotus, and broad fasciae on all the abdominal terga and most sterna.

In addition to the characteristic color pattern, P. arizonensis differs from P.

exclamans in the more sharply lobate humeral angle of the pronotal keel (as in

Fig. 2, versus Fig. 4) and the narrower first tergum (about 0.85-1.00 times as

wide as long) which is distinctly depressed beyond the middle when viewed in

profile (much as in Fig. 1, versus Fig. 3). Very short (0.02-0.03 mm long) hairs

are present on the eyes and are sufficiently abundant to be easily visible at 100X

magnification. A very few hairs may be present on the eyes of P. exclamans, but

these are consistently less than 0.01 mm long and easily overlooked at 100X

magnification.

In the more southern parts of its range in Mexico, P. exclamans is represented

by a color phase which is strikingly similar to P. arizonensis. The body is wholly

ferruginous, except for the following yellow marks: narrow inner orbital stripe;

narrow postorbital line; line or pronotal keel and on scutellum and metanotum;

very narrow submedian pair of propodeal stripes; narrow distal fasciae on first

three terga and second and third sterna. The wings are yellowish brown. I have

seen specimens of these from the States of Jalisco (Guadalajara and Jocotopec),

Hidalgo (14 mi NW Ixmiliquilpan) and Guanajuato (Roque). I have also examined

specimens intermediate between this form and more typical P. exclamans from

Zacatecas (Ojo Caliente, Nochistlan) and Guanajuato (64 km SE La Sauceda).

In 1955 I described P. e. durangoensis from a few specimens from the States

of Durango and Chihuahua, Mexico. They differed from the typical form in

possessing greater amounts of black on the thorax and in the more extensively

yellow abdomen. This variant is a trivial one and is synonymized here. Richards

(1978) described P. instabilis coahuilae from two specimens from Las Delicias,

Coahuila, Mexico. I have examined these specimens; in my opinion they are of

the same color phase as P. e. durangoensis and, therefore, also a synonym of P.

exclamans (New synonymy).

Bohart (1949) described P. e. lineonotus from Lower California, where it is

VOLUME 59, NUMBERS 1-4

269

common. Geographically isolated from all its more closely related congeners, this

wasp represents a self-consistent population. Although variable in the expression

of the markings, they do not tend toward those of either P. exclamans or P.

arizonensis and I believe this wasp should be accorded specific status.

The wasp here called P. bahamensis occurs in tfie Bahamas and Florida; it

ranges sporadically along the Gulf Coast to New Orleans, Louisiana. Parts of its

range in Florida and Louisiana are sympatric with that of P. exclamans. In these

areas there is no evidence among the several hundred specimens that I have seen

that the two forms hybridize. It is largely on this basis that I feel compelled to

regard P. bahamensis as a separate entity.

The two remaining forms, P. bahamensis bilineolatus and P. b. picturatus, are

localized, insular variants of P. bahamensis. Although each does differ slightly

from P. bahamensis in the extent of yellow markings, neither is consistently

marked on any of the islands and some specimens, which, on the basis of locality

should be one or the other subspecies, are marked like typical P. bahamensis.

Under these circumstances, I can see neither need nor justification for treating

these variants as subspecies.

Polistes dominicus (Yallot)

Vespa dominica Vallot, 1802:173.

Polistes cincta Lepeletier, 1836:522. 2. Preoccupied, New synonymy.

Polistes cinctus cinctus: Richards, 1978:478. 2, <$.

Polistes cinctus subsp. barbadensis Richards, 1978:479. 2. New synonymy.

This is the wasp previously treated as Polistes cinctus by various authors. Long

unnoticed is the fact, pointed out to me by A. S. Menke (pers. comm.), that P.

cinctus is preoccupied by Vespa cincta Drury (1770), a synonym of P. annularis

(Linne). The older name of Vallot is based on specimens figured by Reamur (1742

6 : pi. 14, figs. 9 & 10). The Reamur specimens were presumably from Santo

Domingo and apparently are no longer in existence. Among the Lesser Antillean

wasps, only the present species matches the figures at all well.

In the absence of verifiable original material, and in order to stabilize the

nomenclature, a female Neotype is here designated which agrees with the char¬

acteristics cited by Richards (1978) for P. cinctus cinctus. The neotype is from St.

Pierre, Martinique, 23 April 1965 (D. R. Davis) and is in the USNM. Additional

specimens, all from Martinique, are neoparatypes and deposited in AMNH, LACM

and USNM.

The form from Barbados, described as P. cinctus subsp. barbadensis, is a minor

color variant, here placed in synonymy. In addition to Martinique and Barbados,

I have seen material of P. dominicus from St. Kitts, Montserrat, St. Vincent, Sta.

Lucia and Trinidad. Richards (1978) recorded P. c. barbadensis from Mustique

and Carriacou Islands.

Polistes bequaertellus Snelling, New Species

(Figs. 1, 2)

Diagnosis. — Known from females only. Morphologically most similar to P.

exclamans, but differs in the laterally depressed pronotal keel (Fig. 2) and the

Figs. 1-7. Polistes spp. 1, 2, P. bequaertellus, profile of first tergum and frontal view of pronotal

collar. 3,4 , P. exclamans, same. 5-7, P. boharti, profile of first tergum, frontal view of head of male

and female. Scale line = 1.0 mm.

VOLUME 59, NUMBERS 1-4

271

more slender first tergum, which is distinctly depressed beyond mid-length when

viewed in profile (Fig. 1); eyes without erect hairs.

Description female holotype. — Measurements (mm): Head width 4.00; head

length 3.54; wing length 17.0; total length 18.0.

Structurally almost identical to P. exclamans, to which it will run in Richards’

(1978) key; pronotal keel distinctly depressed on either side of the humeral angle

(Fig. 2); first tergum (Fig. 1) slender (1.2X longer than wide) and, in profile, is

distinctly less convex than in P. exclamans and depressed at about the distal one-

third; eyes without erect hairs.

Integument generally reddish brown, the following yellow: head, except supra-

clypeal area and ocellar area; broad stripe confluent with pronotal keel, extending

to lower corner; stripe on dorsal margin of pronotum and along posterior margin;

pair of long, submedian stripes on mesoscutum and spot, or very short stripe,

next to tegula; transverse anterior stripe on scutellum; anterior half of metanotum;

large spot on upper mesepisternum; short longitudinal bar at side of propodeum

and narrow submedian stripes; narrow stripe along distal margins of terga 1-5,

that of second extended forward along lateral margin nearly to base; most of sixth

tergum; broad stripes along distal margins of sterna 2-5, slightly constricted sub-

laterally; entire sixth sternum; apices of all femora; broad outer stripe on pro-

and mesotibiae; basal one-third of metatibia; tarsi. Wings yellowish, veins and

stigma light reddish brown.

Type material. — Holotype and 11 paratype 9: Cockburn Town, San Salvador,

EL SALVADOR, no date (P. Bartsch). Holotype and 8 paratypes in USNM, 3

paratypes in LACM.

Etymology. — This species is named in memory of the late Joseph C. Bequaert,

in recognition of his many outstanding contributions to the systematics of the

Vespoidea.

Discussion. — Variation among the paratypes is not very great. Head width, as

an indicator of size, varies from 3.64 to 4.10 mm. The mesoscutal stripe adjacent

to the tegula, is always present and is usually broad and complete, but in two

specimens it is narrow and briefly interrupted. The lateral stripes ofthe propodeum

may be short and separated from the submedian stripes by about their own width,

or long and broadly connected to the submedian stripes along their distal one-

half (approximately).

The morphological distinctions between this species and P. exclamans are slight,

but consistent. In P. exclamans the pronotal ridge is very slightly, or not at all,

depressed mesad of the humeral angle, and often not depressed below it (Fig. 4).

In that species, too, the first tergum is a little stouter, about 1.0-1.1 times longer

than wide, and in profile is more convex and weakly, if at all, depressed in the

apical one-third (Fig. 3). The first abdominal tergum of P. bequaertellus is a little

more slender (1.15-1.28 times longer than wide, x = 1.20). In P. exclamans the

thorax is more abundantly marked with yellow and the distal stripes of the ab¬

dominal terga are broader and sublaterally constricted.

Superficially, P. bequaertellus resembles P. cubensis Lepeletier, but that species

has conspicuous short hairs on the eyes, lacking in P. bequaertellus. The common

Central American species, P. instabilis Saussure, is also similar but is extensively

black on the thorax and abdominal venter, lacks mesoscutal stripes and is generally

less richly marked (though with four propodeal stripes).

272

PAN-PACIFIC ENTOMOLOGIST

Subgenus Epicnemius Richards

Polistes pacificus Fabricius

Polistes pacificus Fabricius, 1804:274.

Polistes liliaciosus Saussure, 1854:97, pi. 11. fig. 7. 2. New synonymy.

Polistes liliaceusculus Saussure, 1854:98. 2. New synonymy.

Polistes modestus F. Smith, 1862:38. 2. New synonymy.

Polistes pacificus var . flavopictus Ducke, 1918:360, 374. 2. New synonymy.

Polistes pacificus var. trinitatis Bequaert, 1937:195, 197. 2. New synonymy.

Richards (1978) treated all of the above synonymous forms as subspecies of P.

pacificus, a species which ranges from southern Texas to Argentina and Paraguay.

These are all based on varying color combinations of black, ferruginous and

yellowish markings. The darkest form with reduced yellow and ferruginous mark¬

ings is P. p. modestus in the north. The southern form, in Brazil, Paraguay and

Bolivia, is P. p. liliaciosus, with abundant yellow markings, including broad me-

sonotal stripes.

Each of these forms appears distinctive when isolated examples are seen. It is

also true that when several hundred specimens from throughout the range of the

species are available, there is a continuous cline of color variation. Richards noted

the difficulty of dealing with this complex of overlapping forms, but opined that

“several . . . will ultimately turn out to be species . ...” I do not agree with this

prediction; too much obvious intergradation is present in the samples I have seen

for this prediction to hold up. Nor do I see any utility in recognizing a series of

subspecies based on selected populations in a highly variable species.

Polistes boharti Snelling, New Species

(Figs. 5-7)

Diagnosis. — Related to P. pacificus, to which it will run in Richards’ key. Male

differs in having clypeus yellow, with apex acute, and digitus of genitalia with

only a few straight setae. Female differs in having apex of clypeus acute, genal

ridge complete to base of mandible; greatest width of gena equal to greatest width

of eye, in profile.

Description male holotype.— Measurements (mm): Head width 3.74; head length

3.59; wing length 13.5; body length (head + thorax + extended abdomen) 19.5.

Head (Fig. 6): Slightly broader than long in frontal view, posterior ocelli anterior

to slightly convex occipital margin; least length of malar area subequal to mini¬

mum antennal socket diameter. Clypeal breadth (between inner orbits) and length

subequal; clypeus contiguous with eye margin for distance equal to maximum

antennal socket diameter; apex of middle lobe acute, margins nearly straight;

lateral sinus deep. Interocellar distance about 1.3 times diameter of anterior ocel¬

lus; ocellocular distance about 2.1 times diameter of anterior ocellus; ocelloccipital

distance about 3.0 times diameter of anterior ocellus. Antennal segment 13, in

profile, hardly curved in ventral view, about 1.4 times longer than broad. Gena,

in profile, widest at about midlength, about equal to profile width of eye; genal

ridge distinct, somewhat sinuate, continuous to base of mandible, though weak

below sinuation.

Thorax: Pronotal keel well developed, narrowly lamellate, abruptly rounded

onto side, blunt below on side; fovea well developed. Epicnemial groove strong;

VOLUME 59, NUMBERS 1-4

273

dorsal groove of mesepisternum absent; entrance to spiracular chamber slightly

raised, about half as wide as subalar plate. Groove between metepisternum and

propodeum very weak, median furrow of propodeum broad, with fine, well spaced,

transverse striae in middle which do not extend onto side except below; muscle-

slit with sides converging above, apex narrowly rounded.

Abdomen: First tergum, in dorsal view, about as broad as long, spiracle prom¬

inent; in profile, sloping up steeply behind insertion of muscle (Fig. 5). Last ventral

segment broad, apex rounded, disc depressed.

Terminalia: Digitus with only a few straight setae.

Sculpture: Clypeus dull, distal one-fourth closely micropunctate, remainder of

disc contiguously micropunctate; head and thorax generally dull, contiguously

minutely granulopunctate.

Color: Black. The following ferruginous: mandible; lateral lobes of clypeus;

antenna (last 6 segments darker above): most of legs (except coxae, trochanters

and dorsal portions of femora); irregular blotch on each side of second tergite.

The following yellow: minute mandibular spot; clypeus mostly; inner orbital stripe,

well into ocular sinus; outer orbital stripe (somewhat reddened); pronotal collar

and dorsal margin and stripe down side almost to lower corner and spot at lower

corner; margin of tegula; very narrow, evanescent stripe along anterior margin of

scutellum; distinct stripe along anterior margin of metanotum; small spot on upper

mesepisternum; broad submedian propodeal stripes; propodeal valve; antero-

dorsal stripe and ventral blotch on mesocoxa; anterodorsal stripe on metacoxa;

apical fasciae on first three terga, somewhat broadened laterally on first and second;

narrow apical fascia on second sternum, weak across middle. Wings yellowish,

slightly clouded; veins and stigma yellowish brown.

Female allotype. — Measurements (mm): Head width 3.64; head length 3.33;

wing length 11.5; total length 15.0 (paratypes: head width 3.69-4.10; head length

3.33-3.79; wing length 11.5-13.5; total length 15.0-17.0).

Head (Fig. 7): Slightly broader than long in frontal view; ocelli, in frontal view,

well anterior to nearly straight occipital margin; minimum length of malar area

distinctly greater than minimum antennal socket diameter. Clypeal breadth slight¬

ly greater than length; clypeus contiguous with eye for distance about equal to

minimum antennal socket diameter; apex of middle lobe acute, margins nearly

straight, lateral sinus deep. Interocellar distance about 1.6 times diameter of

anterior ocellus; ocellocular distance about 2.5 times diameter of anterior ocellus;

ocelloccipital distance about 3.4 times diameter of anterior ocellus. Genal profile

widest at about midlength and equal to greatest width of eye in profile; genal ridge

distinct to base of mandible, weaker below, slightly sinuate.

Remainder about as in male, except usual sexual differences. Lower gena and

malar area tessellate and moderately shiny, with sparse, conspicuous, moderate

punctures (about 0.04 mm diameter).

Color: Black. The following ferruginous: mandible; most of malar area, trans¬

verse clypeal band; antenna (last six segments infuscated above); pale stripe on

upper two-thirds of outer orbit; femoral apices; tibiae; tarsi. The following yellow:

most of clypeus; irregular blotch on malar area adjacent to eye; inner orbital stripe,

extending into ocular sinus; pronotal collar and dorsal margin and stripe down

side to below level of fovea; side of axilla; anterior transverse stripe on scutellum

and metanotum; spot on subalar plate; tegula; spot on upper mesepisternum; pair

274

PAN-PACIFIC ENTOMOLOGIST

of submedian propodeal stripes; propodeal valve; anterodorsal stripe on meso-

and metacoxae; broad apical stripes on first three terga, sublaterally constricted

on second and third segments, moderately expanded at side; medial one-half of

margin of fourth tergum; very narrow margin of second sternum, greatly expanded

at side. Wings as in male.

Type material —All from MEXICO: Holotype male: Compostela, Nayarit, 26

Aug. 1959 (A. S. Menke and L. A. Stange). Allotype female: 3.9 km NE Taxco,

elev. 1707 m, Guerrero, 16 Sept. 1976 (C. D. George and R. R. Snelling). Para-

types: 1 9, 64 mi W. Tehuantepec, Oaxaca, 21 July 1952 (E. E. Gilbert and C. D.

MacNeil); 1 9, Terra Blanca, Vera Cruz, 15 Aug. 1962 (H. E. Milliron); 1 9, Santa

Lucia, elev. 4000 ft, Sinaloa, 4 Aug. 1964 (W. R. M. Mason). Holotype and

allotype in LACM; paratypes in CNC and UCB.

Etymology ’.—This species is dedicated to R. M. Bohart, colleague and early

inspiration to my own work in vespoids.

Discussion. — Richards (1978) tentatively recorded under P. pacificus a female

from Vulkan, Colima, MEXICO, which he thought might represent a different

species; evidently this specimen is P. boharti.

The paratype females are somewhat variable in the extent of the ferruginous

markings. In one (Tehuantepec), the clypeal stripe is broad and the outer orbital

stripe is connected to the expanded blotch of the malar area; the yellow malar

blotch is reduced to a narrow stripe along the lower orbit of the eye. There is a

very obscure, reddish blotch on each side of the second tergum. The two remaining

paratypes have only minute, ferruginous spots on the clypeal disc; both have

distinct, ferruginous blotches on the second tergum.

Although this species is obviously close to P. pacificus the differences noted

above should easily separate the two. The characteristic setation of the male digitus

of P. boharti is especially distinctive. Although the male genitalic structures of

the two species are very similar, the digitus of P. pacificus males bears many dark,

stout, distally curved or hooked setae.

Females of the two species are similar. In females of P. pacificus the genal ridge

is usually weak below, often not reaching much below the level of the lower one-

fifth to one-fourth of the eye; if it extends below this level, it is very weak and

rarely attains the base of the mandible. In P. boharti the ridge is as strong at the

mandibular base as elsewhere. The gena, in profile, is narrower than the greatest

eye width in P. pacificus, greater in P. boharti. The lower (lateral) extension of

the pronotal ridge is not depressed in P. pacificus and a lobate humeral angle is

not defined; in P. boharti the pronotal ridge is definitely depressed below the

humeral angle and the latter is thus lobate.

Polistes palmarum Bequaert

Polistes major var. palmarum Bequaert, 1936:11. 9.

Polistes major subsp. slevini R. Bohart, 1949:103. 9. New synonymy.

This wasp is completely allopatric to P. major Beauvois, of which it has been

considered a subspecies. The color pattern is remarkably stable and different from

that of P. major and P. palmarum is therefore considered to be a separate species.

The range of P. palmarum extends from canyons along the western margin of the

Colorado Desert in California to the Cape Region of Lower California. Specimens

VOLUME 59, NUMBERS 1-4

275

from the Cape Region were described as a separate subspecies, P. major slevini,

by Bohart (1949). This form, with somewhat darker red integument than in more

northern samples, is a minor variant of P. palmarum.

Genus Clypearia Saussure

Clypearia naumanni Richards

Clypearia naumanni Richards, 1978:197-198. 2.

Richards (1978) described this species from a single female from Barro Colorado

Island, Canal Zone, PANAMA. The following specimens extend the range to the

north: 1 2, Hacienda Comelco, 24 km NW Canas, Guanacaste Province, COSTA

RICA, 13 Jan. 1972 (E. R. Heithaus; LACM); 1 2, Los Mangos, Los Tuxtlas, Vera

Cruz, MEXICO, 11 Sept. 1973 (M. Sousa; LACM).

Genus Mischocyttarus Saussure

Subgenus Kappa Bequaert

Mischocyttarus atrocyaneus Zikan

Mischocyttarus atrocyaneus Zikan, 1949:204. 2.

Mischocyttarus {Kappa) atrocyaneus: Richards, 1978:297. 2.

Previously known only from PANAMA (Boquitos, Volcan de Chiriqui), COS¬

TA RICA (San Jose) and MEXICO (Yucatan, no further data). New distribution

data: Fortin de las Flores Vera Cruz, MEXICO; 2.5 mi W Quezaltepeque, EL

SALVADOR; 9 km NW Gamboa, Canal Zone, PANAMA. The short series from

near Gamboa is associated with two nests. One of these is small (12 cells), with

a centrally placed pedicel; a single wasp was on the nest. The second nest, oblong,

with the pedicel at one end, consists of 74 cells (many barely begun) and had

seven or eight wasps when collected. Both nests were suspended under leaves.

Mischocyttarus immaginatus Richards

Mischocyttarus immarginatus Richards, 1940:182. 2.

Mischocyttarus oaxacanus Zikan, 1949:168-169. 2, 8. New synonymy.

Mischocyttarus immarginatoid.es Zikan, 1949:236. 2 {nomen nudum).

Zikan (1949) described M. oaxacanus from a few specimens from several lo¬

calities in MEXICO (Nuevo Leon and Oaxaca). Richards (1978) omitted mention

of M. oaxacanus in his treatment of this genus. Dr. Richards informed me {in

litt.) that he failed to find specimens that agreed with Zikan’s description either

in Paris or in Zikan’s collection. The description and figures of M. oaxacanus

agree, however, with specimens of M. immarginatus and I think it safe to assume

them conspecific; M. immarginatus, described in 1940, has priority over M.

oaxacanus (New synonymy).

Richards (1978) recorded M. immarginatus from MEXICO (Guerrero, Nayarit,

San Luis Potosi, Vera Cruz and Yucatan), NICARAGUA and COSTA RICA. I

have seen specimens from MEXICO (Guerrero, Jalisco, Nayarit, Oaxaca, Sinaloa

and Yucatan), GUATEMALA (El Rancho), HONDURAS (Tegucigalpa), NIC¬

ARAGUA (Managua; San Juan del Sur) and COSTA RICA (Hacienda Comelco

and Nicoya, Guanacaste Province).

276

PAN-PACIFIC ENTOMOLOGIST

Mischocyttarus tolensis Richards

Mischocyttarus tolensis Richards, 1941:128. 2.

This species was known from a few specimens from PANAMA, (Tole; Barro

Colorado Island; Alajuela). I have seen the following additional material: 1 8, 9

km NW Gamboa, Canal Zone, PANAMA, 4-8 Aug. 1975 (E. M. and J. L. Fisher;

LACM); 4 2, Puerto Viejo, Heredia Province, COSTA RICA, 6 Aug. 1965 (R. J.

Hamton; LACM).

The male of M. tolensis is unknown and the male collected near Gamboa is

only provisionally assigned to M. tolensis. In the key to males of Kappa by

Richards (1978), it runs to couplet 7 where it fails to agree with either alternative.

The last antennal segment is about 2.5 times longer than wide and is distinctly

shorter than the third flagellar segment; the occiput is sharply margined; the

propodeal furrow bears abundant fine punctures and sparse coarser punctures;

the propodeal valves are black; the clypeus and mandibles are largely ferruginous.

Except for the flagellar characters and usual sexual differences, the male is very

similar to females of M. tolensis, but lacks erect hairs on the eyes.

Subgenus Monocyttarus Richards

Mischocyttarus flavitarsis (Saussure)

Polybia flavitarsis Saussure, 1854:199. 2.

Mischocyttarus flavitarsis var. centralis Bequaert, 1933.129. 2. New synonymy.

Mischocyttarus flavitarsis var. idahoensis Bequaert, 1933:133. 2. New synonymy.

Mischocyttarus flavitarsis \ ar. kaibabensis Bequaert, 1933:133. 2. New' synonymy.

Mischocyttarus flavitarsoides Zikan, 1949:236 ( nomen nudum).

The range of M. flavitarsis extends from the Pacific Coast to the Rocky Moun¬

tains, from British Columbia and Alberta to northern Mexico. Throughout this

range there exist a number of color phases which have been treated as subspecies:

M. f. centralis Bequaert, M. f idahoensis Bequaert, M. f. kaibabensis Bequaert,

and M. f navajo Bequaert, in addition to the nominate form. These various forms

are based on differing combinations of the basic black and yellow color pattern,

including replacement of black by ferruginous.

The northern, melanic, form is M. f. idahoensis, which is black, with greatly

reduced yellow markings. The xanthic, southern, form is M. f. kaibabensis, in

which the black color is completely replaced by ferruginous. Complete replacement

of black by ferruginous also occurs in some populations of M. f. centralis from

northern Mexico, but the wings are yellowish rather than brown as in M. f.

kaibabensis.

While the various color phases, in their ideal phenotypes, are distinct from one

another, it is also true that considerable intergradation takes place in areas between

such populations. In general, the variation is clinal and one form gradually replaces

another. This is especially true of the series M. f idahoensis -► M. f. flavitarsis

M. f. centralis -* M. f kaibabensis. These forms do not, in my opinion, merit

formal recognition. Each represents a selected point in a long clinal series. There¬

fore, M. f. centralis = M. f. idahoensis = M. f kaibabensis = M. flavitarsis.

One form, M. f. navajo, seems to be much more stable, perhaps due to its

relatively isolated distribution in central and southern Arizona. I have not yet

VOLUME 59, NUMBERS 1-4

277

seen any material which I can consider to be intermediate between M. f navajo

and any other of the color phases attributed to M. flavitarsis. Therefore, I am

inclined to believe that M. navajo should be elevated to species level until evidence

is advanced to the contrary.

Mischocyttams navajo Bequaert

Mischocyttarus flavitarsis var. navajo Bequaert, 1933:133. 9.

This wasp is found in Arizona, southwestern New Mexico and northern Sonora.

It is common and quite stable in its color pattern. The wings are dark fuscous

and the thorax is principally blackish, with limited yellow markings on the prono-

tum, spot below tegula and a pair of submedian propodeal stripes; the first tergum

is black, with a broad apical yellow band. Richards (1978) remarked that this is

“rather larger than subsp. flavitarsis .” I have measured head width of 50 randomly

selected females of M. flavitarsis and 50 of M. navajo, no two individuals with

the same collection data.

The results confirm Richard’s assumption. For M. flavitarsis the head width

range is 3.08-3.95 mm, with the mean at 3.52 mm; only 24% of the specimens

measured had a head width in excess of 3.75 mm. Head width, in the measured

specimens of M. navajo ranged between 3.18 and 4.21 mm, with a mean width

of 3.75 mm; 62% of the measured specimens have a head width of 3.75 mm or

more.

Subgenus Monacanthocnemis Ducke

Mischocyttarus chalucas Snelling, New Species

(Figs. 8-11)

Diagnosis. — First tergum slender, anterior width less than distal width of meta¬

femur; dorsal mesepistemal groove distinct; mesoscutum closely to densely coarsely

punctate, interspaces with abundant very fine punctures; last antennal segment

of male weakly curved, about 2.5 times longer than broad at base; and fifth to

seventh flagellar segments a little broader than long.

Description female holotype. — Measurements (mm): Head width 2.32; head

length 1.65; wing length 7.5; total length 11.5.

Head (Fig. 10): About 1.4 times broader than long; ocelli at top of flat vertex.

Malar space very narrow, eye nearly touching base of mandible. Clypeus narrow,

about 1.2 times longer than wide, contiguous with eye for distance about twice

minimum antennal socket diameter; apical margin obtuse. Interantennal distance

about twice minimum antennal socket diameter; minimum antennal socket di¬

ameter about 1.5 times antennocular distance and slightly greater than anten-

noclypeal distance. Flagellum stout, first segment about 0.8 times length of scape

and longer than combined second and third segments. Frons very slightly bulging

and very weakly depressed in middle. Interocellar distance about 1.6 times anterior

ocellus diameter; ocellocular distance subequal to anterior ocellus diameter. Gena

about one-half as wide as eye profile at sinus.

Thorax: Slender, about 1.5 times longer than wide. Pronotal keel, in dorsal

view, concave, strongly lamelliform laterad, weak in middle, strongly lobate and

somewhat reflected at humerus. Proepisternum bi-reflexed in front. Propodeal

furrow broad, weak, without median ridge; valves narrow. Mesotibia with two

PAN-PACIFIC ENTOMOLOGIST

278

Figs. 8-11. Mischocyttarus chalucas. 8, 6 head, frontal view; 9, 3 head, lateral view; 10, $ head,

frontal view; 11, petiole dorsal view. Scale line = 1.0 mm.

apical spurs. Inner, distal margin of last two metatarsal segments not produced,

outer margin strongly produced.

Gaster: Petiole slender, about 3.5 times longer than greatest width, when mea¬

sured from propodeal insertion (Fig. 11); spiracle at about midlength and short

of apex of metacoxa.

Sculpture; Clypeus and supraclypeal area dull, subcontiguously to densely mi-

cropunctate and with dense to close, moderate punctures; gena dull, closely mi-

cropunctate and with sparse, somewhat coarser punctures; occiput shiny, with

irregularly spaced moderate punctures and a few micropunctures; remainder of

head and thorax shiny, with dense to close, moderate punctures, interspaces closely

micropunctate.

VOLUME 59, NUMBERS 1-4

279

Color: Black, but irregularly replaced with ferruginous on head (especially clyp-

eus and gula) and thorax (especially below and on side). Following pale yellow:

broad stripe along dorsal mandibular margin; transverse bar at apex of clypeus;

inner orbital line to top of sinus; curved bar above each antennal socket; outer

orbital line nearly to top of eye; keel, most of dorsal margin and lower corner of

pronotum; subalar and lower posterior spot on mesepisternum; broad anterior

stripe on scutellum; anterior stripe, narrowed in middle, and lateral extensions

of metanotum; large, submedian, posterior spots on propodeum; valves; narrow

distal fascia on first five terga and second to fifth sterna, that of second tergum

extended along side nearly to base; short lateral stripe on sixth tergum and sternum;

short apicoventral stripe on procoxa; outer stripe on mesocoxa; two dorsal stripes

on metacoxa; small apicodorsal spot on each femur; outer stripe on protibia; basal

spot, long median stripe and small apical spot on mesotibia; short stripe on outer

face of probasitarsus and on outer face of prodistitarsus. Tibial spurs dirty white.

Wings clear, slightly infuscated; veins and stigma brown.

Male allotype. — Measurements (mm): Head width 2.10; head length 1.77; wing

length 7.0; total length 11.0.

Similar to female, except usual sexual differences, and the following.

Head (Fig. 8): About 1.2 times wider than long. Mandible tridentate. Anten-

noclypeal distance a little greater than minimum antennal socket diameter; min¬

imum antennal socket diameter about twice antennocular distance. Frons bifaced,

i.e., there is a more or less vertical anterior portion and a definite dorsal face

forming a flat plane in front of anterior ocellus (Fig. 9). Scape stout-pyriform,

about 1.6 times longer than broad, apical width about 1.6 times basal width; first

flagellar segment flattened, about 1.8 times longer than broad, about twice longer

than second; fifth to seventh segments broader than long; eight as broad as long

and narrower than seventh; ninth and tenth a little longer than broad; apical

segment about 2.5 times longer than wide at base, slightly curved in profile.

Color: Very similar to that of female, but underside of scape, ventral surface

of thorax and coxae yellow.

Type material. — Holotype female and allotype: Tuxtla Gutierrez, Chiapas,

MEXICO, 7 June 1964 (J. D. and D. Pallister) in AMNH.

Etymology. — Chalucas, for whom this species is named, is a trickster god and

folk hero of the Zoque Indians indigenous to tlie Tuxtla Gutierrez region.

Discussion.— This species appears to be most similar to the South American

M. punctatus (Ducke). The petiolar spiracle, however, is anterior to the end of

the metacoxa (slightly beyond in M. punctatus ), the dorsal mesepisternal groove

is stronger, and the mesoscutal punctures are clearly a mixture of moderate (0.04-

0.06 mm diameter) and very fine (less than 0.01 mm diameter) punctures. In M.

punctatus, the finest punctures are about 0.02 mm diameter and grade up to the

punctures of moderate size (about 0.05 mm diameter) and the punctures, irre¬

spective of size, are very evenly spaced, while the moderate punctures of M.

chalucas are irregularly spaced from subcontiguous to interspaces of more than a

puncture diameter.

Acknowledgments

For making available material utilized in this study, I wish to thank: P. H.

Arnaud, Jr., California Academy of Sciences (CAS); M. Favreau and J. G. Rozen,

280

PAN-PACIFIC ENTOMOLOGIST

Jr., American Museum of Natural History (AMNH); L. Masner, Canadian Na¬

tional Collections (CNC); A. S. Menke, National Museum of Natural History

(USNM); J. A. Powell, University of California, Berkeley (UCB). Other specimens

were from the collections of the Natural History Museum of Los Angeles County

(LACM).

Literature Cited

Bequaert, J. C. 1933. The nearctic social wasps of the subfamily Polybiinae (Hymenoptera, Vespidae).

Ent. Amer. (n.s.) 13:87—150.

-. 1936. Two new color forms of Polistes major Palisot de Beauvois from California and

Arizona (Hymenoptera, Vespidae). Ent. News 47:7-13.

-. 1937. The American Polistes with prepectal suture. Their structural characters, distribution

and variation. Archiv. Inst. Biol. Vegetal. 3:171-205.

Bohart, R. M. 1949. Notes on North American Polistes with descriptions of new species and sub¬

species (Hymenoptera: Vespidae). Pan-Pac. Ent. 15:97-103.

Drury, D. 1770. Illustrations of natural history. London. Vol. 1, xxviii + 130 pp.

Ducke, A. 1918. Catalogo das Vespas do Brasil. Rev. Mus. Paulista 10:314-374.

Fabricius, J. C. F. 1804. Systema Piezatorum. Hafniae, 572 pp.

Lepeletier de St. Fargeau, A. L. M. 1836. Histoire naturelle des insectes. Hymenopteres. I. Roret’s

Suites a Buffon, Paris,

Reamur, R. A. F. de. 1742. Memoires pour servir a l’histoire des insectes, 6. Paris, 608 pp.

Richards, O. W. 1940. Preliminary diagnoses of some new species of Mischocyttarus Saussure

(Hymenopt., Vespidae). Ann. Mag. Nat. Hist. (1 1) 5:176-183.

-. 1978. The social wasps of the Americas, excluding the Vespinae. Publ. 786, Brit. Mus. (Nat.

Hist.), London, 580 pp.

Saussure, H. de. 1854. Etudes sur la Famille des Vespides 2. Monographie des guepes sociales on

de la tribu des Vespiens. Paris and Geneva, pp. 97-256.

Smith, F. 1862. Description of new species of Hymenoptera collected at Panama by R. W. Sketch,

with a list of described species. Trans. Ent. Soc. London (3) 1:29-44.

Snelling, R. R. 1954. Wasps of the genus Polistes in California and Arizona (Hymenoptera: Vespidae).

J. Kansas Ent. Soc. 27:151-155.

-. 1955. Notes on some Polistes in the American Museum of Natural History, with descriptions

of New North American subspecies (Hymenoptera, Vespidae). Amer. Mus. Novit. 1701, 9 pp.

-. 1970. The social wasps of Lower California, Mexico (Hymenoptera: Vespidae). L. A. Co.

Mus. Contr. Sci. 197, 20 pp.

Vallot, J. N. 1802. Concordance systematique, servant de table des matieres a l’ouvrage de Reamur

intitule Memoires, Paris, 198 pp.

Zikan, J. F. 1949. O Genero Mischocyttarus Saussure (Hymenoptera, Vespidae), con a descrifao de

82 especies novas. Bol. Parq. Nac. Itatiaia 1:1-251.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 281-297

A Synopsis of the Genus Epanthidium Moure with the

Description of a New Species from Northeastern

Mexico (Hymenoptera: Megachilidae ) 1

Lionel A. Stange

Bureau of Entomology, Florida Department of Agriculture and Consumer Ser¬

vices, P.O. Box 1269, Gainesville, Florida 32602.

The genus Epanthidium consists of 12 described species from southern South

America. Therefore, it was surprising to discover a 13th species from northeastern

Mexico. Although amphitropical distribution is relatively uncommon in bees, it

is not unknown even in the Anthidiini. For example, Michener and Ordway (1964)

described a species of the anthidiine bee genus Nananthidium Moure from north¬

eastern Mexico. This genus was previously known only from southern South

America.

I have not seen all of the primary types (those seen are identified by an excla¬

mation point after the depository) and for some species interpretations I have

relied on the original descriptions and specimens identified by Schrottky, Friese

and Moure. However, it must be pointed out that the original descriptions are

based largely on color, which is often variable within species and also similar in

different species, so that a few nomenclatorial corrections may be anticipated. I

hope to treat the generic relationships of the New World genera of Anthiidini at

a future date since data gathered after Michener’s 1948 revision of the tribe

suggest some different groupings.

I present here a key to the known species which should aid in future type and

revisionary studies, and also to suggest the relationship of the new species de¬

scribed here. New records and floral associations gathered to date are also pro¬

vided.

Acknowledgments

I would like to acknowledge the valuable assistance of Padre Jesus Moure in

species identification. Jack Neff has provided many new floral records. Special

thanks are due also to Charles Porter, Manfredo Fritz, and George and Richard

Bohart for many valuable specimens. Other important collaborators are acknowl¬

edged in the list of collections consulted. Field work in South America was sup¬

ported by the National Geographic Society (1977) and the Consejo Nacional de

Investigaciones Cientificas y Tecnicas (1976), Buenos Aires, Argentina.

Collections Studied and Type Depositories

AMNH—American Museum of Natural History, New York 10024. J. G. Ro-

zen, Jr.

1 Contribution No. 549. Bureau of Entomology, Division of Plant Industry, Florida Department of

Agriculture and Consumer Services, Gainesville, Florida.

282

PAN-PACIFIC ENTOMOLOGIST

Berlin—Zoologisches Museum der Humboldt—Uni versitat, East Berlin, Ger¬

many.

BM —British Museum (Natural History), London, England. I. H. H. Yarrow.

Curitiba—Department of Zoology, University of Parana, Curitiba, Brasil. J. Moure.

FSCA —Florida State Collection of Arthropods (Hymenoptera Collection),

Gainesville, Florida 32602. H. V. Weems, Jr.

Fritz Collection—Private collection. Instituto Entomologico de Salta, P.O. Box

3, Rosario de Lerma, Salta, Argentina. Manfredo Fritz.

IML—Instituto Miguel Lillo, San Miguel de Tucuman, Argentina. A. Willink.

KU—Snow Entomological Collection, University of Kansas, Lawrence, Kansas

66045. C. D. Michener.

Logan—Bee Biology & Systematics Laboratory, Utah State University, Logan,

Utah 94321. G. E. Bohart.

LACM—Natural History Museum of Los Angeles County, Los Angeles, California

90007. R. R. Snelling.

MLP—Museo de Ciencias Naturales, La Plata, Argentina. L. de Santis.

Paris—Musee National d’Histoire Naturelle, Paris, France. S. Kellner-Pillault.

Porter Collection—Private collection. 301 N. 39th Street, McAllen, Texas 78501.

Sao Paulo—Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo,

Brasil.

UCD—Entomology Museum, University of California, Davis, California 95616.

R. M. Bohart.

USNM—U.S. National Museum of Natural History, Washington, D.C. 20560.

A. S. Menke.

Vienna—Naturhistorisches Museum, Vienna, Austria. M. Fischer.

Epanthidium Moure, 1947

Type species: Hypanthidium tigrinum Schrottky, by original designation.

This genus was reduced to subgeneric status (under Dianthidium ) by Michener

(1948), but its structural characters warrant generic status in my opinion. These

include the absence of the preoccipital carina, male hindcoxa not spined, presence

of upper propodeal pits at least laterally, modified metasomal sternum VI of the

female and sternum II of the male, and the elevated lamella of the posterior lobe

of the pronotum. Although little information on nesting habits is available, one

nest I examined was made from adobe clay (5 cells), and no resin was used, which

contrasts with the better known nests of Dianthidium (Grigarick and Stange, 1968)

which have plant resins as the common bonding material. Anthidium inerme

Friese, 1908, was included in the genus by Michener (1948) but my studies of the

type material have convinced me that it is not congeneric and probably belongs

to a new genus.

Diagnosis. — Body robust, mesoscutum broader than long; interalveolar carina

absent; female mandible 4-toothed; maxillary palpus 2-segmented; preoccipital

carina absent; arolium present; pronotal lobe lamellate, elevated, sometimes low

and opaque; anterior and lateral faces of mesepisternum separated by a carina;

propodeum with sulcus behind spiracle; row of pits along upper margin of pro-

podeum strongly developed at least laterally; posterior coxa of male without spine

(although a small tooth present in a few species); metasomal sternum VII of male

VOLUME 59, NUMBERS 1-4

283

trilobed or bilobed; sternum II with a transverse elevation usually ending sub-

laterally in a spine; female sternum VI variously modified with processes or ridges.

Epanthidium boharti Stange, New Species

(Figs. 1, 8, 11)

Diagnosis.— The bilobate metasomal tergum VII of the male (Fig. 1) is dis¬

tinctive in the genus (other species are trilobate) except for E. paraguayense

(Schrottky) from South America. These 2 species can be separated in the male

by the deeper emargination of tergum VII in E. boharti. Female E. boharti can

be distinguished from all other known species except E. paraguayense and E.

erythrocephalum by the lack of a median carina on metasomal tergum VI. From

these 2 species, female E. boharti can be distinguished by the sublateral tooth on

sternum VI (from erythrocephalum) and the distance between the lateral ocellus

and the posterior margin (from paraguayense). The black and ivory coloration in

both sexes of E. boharti is distinctive from the South American species which

usually have red or orange markings.

Holotype male.— Length 14 mm, forewing 10 mm; width of metasoma (at

middle) 5 mm. Base color black with ivory markings as follows: clypeus except

apex and dorsal margin to tentorial pit; paraocular stripe from clypeus to about

level of antenna; small spot near dorsal margin of eye; anterior margin of meso-

scutum inteiTupted widely at middle where it points posteriorly, and extending

along lateral margin to near middle of tegula; axillae nearly completely; meta-

notum with large spot laterally interrupted medially by a triangular black area;

metasomal tergum I with large sublateral spot extending nearly to middle, broadly

emarginate posteriorly; following 3 terga with sublateral spots decreasing in size

posteriorly; terga III-VI with transverse stripe at middle decreasing in length

posteriorly; pubescence mostly whitish except for closing faces of tarsi which are

golden color. Forewing uniformly dark infuscate with violaceous reflection, except

apical l A.

Pubescence—Basitarsus of foreleg and midleg with many elongate setae on

posterior margin, about 3 times as long as greatest width of tarsus.

Puncturation—Vertex posterior to ocelli shiny with setiferous punctures about

one puncture apart. Tegula shagreened but nearly impunctate at middle; upper

row of propodeal pits absent at middle; area anterad of sternum II ridge sparsely

punctate.

Structure—Clypeal apex with 4 fairly evenly spaced teeth; scape shorter than

pedicel and first 3 flagellomeres together; distance between lateral ocellus and

hind margin greater than interocellar distance; height of posterior lobe of prono-

tum less than midocellus diameter, opaque; mesepisternum not produced ven-

trally into small tooth; midventral carina of mesosoma low throughout; tegula

with posterior margin narrow, thin (as in Fig. 20); midcoxa without tooth at mesal

posterior margin; curved ridge of sternum II strongly raised with strongly sloping

posterior face, sublateral tooth moderately developed; tergum VII bilobate (Fig.

1); lateral margins of terga V-VI without teeth or tubercles; genitalia as in Fig. 8.

Female. — As described for male except as follows: clypeus black, vertex with

posterior margin orange-colored; scopa whitish; foreleg and midleg without elon¬

gate setae on basitarsi; tegula densely punctured at middle; mandible quadri-

284

PAN-PACIFIC ENTOMOLOGIST

punctate; tergum VI without longitudinal carina and with subapical margin strong¬

ly angled laterally followed by undulating outline to sting emargination, without

teeth; sternum VI with sublateral ridge produced into tooth sublaterally, medial

process narrowly rounded (Fig. 11).

Types .—Holotype <5: MEXICO, Tamaulipas: 55 km S. Ciudad Victoria, Ruta

85 ca. Llera, 22, 25-VI-1981, B. Miller, C. Porter, L. Stange (FSCA). Paratypes

as follows: 2 9, same data as holotype (FSCA, Porter Collection). MEXICO: Nuevo

Leon: Cola de Caballo, ca. El Cercado, 15-23-VI-1976, 2 9, C. Porter (FSCA,

Porter Collection); Tamaulipas: Ruta 180, ca. Soto la Marina, 75 km E. Ciudad

Victoria, 1-VII-1981, 1 9, B. Miller, C. Porter, L. Stange (FSCA).

Etymology. — This species is named for Richard M. Bohart in celebration of his

seventieth year.

Epanthidium anisitsi (Schrottky)

(Fig. 6)

Dianthidium anisitsi Schrottky, 1908:231. Syntypes, 3, 9, Asuncion, Paraguay,

Anisits Collection (not located).

Anthidium olympinum Strand, 1910:546 (synonymy after Schrottky, 1920:211).

Three syntypes, 6, Asuncion, Paraguay (Berlin).

Taxonomy: Moure, 1947:35 (in Epanthidium).

This is one of the largest species in the genus, with the spine of the curved ridge

of the male sternum II unusually large and somewhat erect. Male tergum VII is

distinctive in that the middle process is below the plane of the lateral process (Fig.

6). The female is unknown to me. I have not seen the types, but I have seen a

specimen identified by Schrottky in 1910.

New record.—PARAGUAY: Cuaguazu, XI-79, M. Fritz coll. (1 <3—FSCA).

Epanthidium autumnale (Schrottky), New Combination

(Figs. 2, 9, 12)

Dianthidium autumnale Schrottky, 1909a:218. Holotype 3, Paraguay, Puerto Ber-

toni, 2-III-1909 (not located).

This species is, according to my records, commonest in arid northwestern

Argentina. A few specimens seen from Brazil and Argentina (Misiones) have more

extensive yellow bands on the metasomal terga. The male has a distinctive tergum

VII with the emargination between the middle and lateral process very shallow

(Fig. 2). The yellow spot on the male hindcoxa is distinctive except for E. ni-

grescens and E. bicoloratum.

New records (37 3, 24 9).—ARGENTINA: Catamarca: El Rodeo, IV, Porter (1

3, 1 9—FSCA); 20 km E. Belen, flowers of Zuccagnia, X, Stange (1 9—FSCA).

Salta: Yacochuya, III-IV, Porter, Stange, Willink collectors (6 3, 17 9—IML,

FSCA); Tacuil, I, Stange, Willink collectors (1 9—FSCA); 5 km W. Agua Blanca,

V, Stange (2 3—FSCA); Rio Pescado, IX, Porter (1 9—FSCA). Tucuman: San

Pedro de Colalao, XI, Stange (2 3—FSCA); Tacanas, XI, XII, Stange (2 3—FSCA);

Raco, I, Stange (1 3, 1 9—FSCA); Amaicha del Valle, I, Stange (1 9—FSCA).

Jujuy: 7 mi. S. Volcan, III, Stange (1 3—FSCA); Humahuaca, I, Stange (1 9—

FSCA). La Rioja: Angulos, XII, Porter & Stange (1 3—FSCA). Cordoba: Los

Cocos, I, Stange (1 3, 2 9—FSCA); Saltos Blancos, I, Stange (1 3—FSCA). Misiones:

VOLUME 59, NUMBERS 1-4

285

Cataratas del Iguazu, XI, Porter & Stange (1 <3, 2 2—FSCA). BOLIVIA: Santa

Cruz: General Saavedra, VII, Porter & Stange (1 2—FSCA). BRAZIL: Santa

Catalina: Nova Teutonia, XI, Plaumann (1 2—KU).

Epanthidium bertonii (Schrottky)

(Figs. 5, 17)

Dianthidium bertonii Schrottky, 1905:3. Holotype 2, Paraguay, Puerto Bertoni,

Bertoni (not located).

Dianthidium brethesi Schrottky, 1909a:217. Holotype <5, Catamarca, III-08, C.

Bruch (not located). New synonymy.

Anthidium argentinum Friese, 1908:72. Syntypes, <3, 2, Tucuman and Mendoza,

XI (not located). New synonymy.

Taxonomy: Schrottky, 1908:230 (description <3); Moure, 1947:35 (in Epanthi¬

dium).

My identifications are based on studies at Curitiba in 1974 and on the literature.

There are 2 color forms; dark populations ( brethesi ) exist in the Subandean desert

of Argentina, and more reddish populations occur in more humid areas. This

species has no unusual features which would provide easy diagnosis.

New records (11 <3, 2). —ARGENTINA: Catamarca: 3 km N. Belen, XII, Porter

& Stange (3 <3—FSCA); Andalgala, J. Neff (3 6, 2 2—FSCA). Cordoba: Sierras,

XII, Williner (1 2—FSCA); Formosa: Riacho Negro, XI, Willink (3 2—IML,

FSCA). La Rioja: Carrizal, G. Bohart, XI (1 <3—FSCA); Famatina, XII, Porter &

Stange (1 2—FSCA). Misiones: San Ignacio, XII, Willink (1 3— FSCA). Salta: San

Carlos, I, Monroe & Willink (1 3— IML). San Luis: Villa Dolores, II, Porter (1

2—FSCA). Tucuman: San Pedro de Colalao, I, Stange (1 2—FSCA); Amaicha,

XI, Stange (2 2-FSCA, IML). BOLIVIA; Santa Cruz: Rio Piray, VII, Porter &

Stange (1 <3-FSCA). PARAGUAY; Villarrica, I, Schade (1 <3-FSCA).

Epanthidium bicoloratum (Smith)

(Figs. 12, 19)

Anthidium bicoloratum Smith, 1879:88. Holotype 2, Mendoza (BM!).

Anthidium bicoloratum tucumanum Vachal, 1904:14. Holotype 2, Tucuman, coll.

Vachal (Paris). New synonymy.

Taxonomy: Schrottky, 1908:230 (in Dianthidium, description of <3); Friese, 1908:

74 (description of <3); Moure, 1947:35 (in Epanthidium).

This species is easily identified in both sexes by the tegula (Fig. 19) which is

drawn out posteriorly, thickened and compressed laterally.

New records (29 3 , 32 2). —ARGENTINA: Catamarca: Andalgala, flowers Pro-

sopis alba , X, Stange Sc Willink (1 2—IML); Belen, X, Stange (2 3— FSCA). Chaco:

San Bernardo, VI, Sorio (1 2—Fritz Coll.). Cordoba: Quebrada Honda, Unquillo,

I, Stange (l 2—FSCA); Los Cocos, I, Stange (1 2—FSCA). Corrientes: Paso de la

Patria, XI, Porter (1 3, 3 2—FSCA); Santa Ana, V, Porter & Stange (1 2—FSCA).

La Rioja: Famatina, XII, Porter Sc Stange (1 2 —FSCA); 7 mi. S. Villa Casana,

XII, Porter & Stange (1 2—FSCA); Santa Cruz, X, Stange (1 3— FSCA); Villa

Union, XII, Porter Sc Stange (1 2—FSCA). Misiones: Posadas, XI, Porter Sc Stange

(1 <3—FSCA). San Juan: Albardon, XII, Porter Sc Stange (1 2 —FSCA). San Luis:

San Jeronimo, II, Fritz (1 2 —Fritz Coll.). Salta: Yacochuya, 9 km N.W. Cafayate,

286

PAN-PACIFIC ENTOMOLOGIST

IV, Porter & Stange (2 6, 2 9—FSCA). Santiago del Estero: Los Tigres, I, Golbach

(1 6—FSCA); Las Termas del Rio Hondo, IV, Porter & Stange (1 9—FSCA).

Tucuman: Amaicha, XI, Stange (1 6—FSCA); 11 km W. Las Cejas, X, I, Stange

(1 6, 1 9—FSCA); Villa Padre Moure, IV, Stange (1 6—FSCA).

Epanthidium confusum (Smith)

(Fig. 16)

Anthidium confusum Smith, 1879:80. Holotype <3, Mendoza (BM!).

Anthidium dentiventre Friese, 1906:100. Holotype 9, Mendoza (Friese Coll., Ber¬

lin). Preoccupied by Anthidium dentiventris Friese, 1904.

Anthidium jenseni Friese, 1908:69. New name for A. dentiventre Friese, 1906.

Taxonomy: Joergensen, 1912:137 (jenseni = confusum)', Moure, 1947:35 (in

Epanthidium ).

This very distinctive species occurs in the Larrea Subandean desert of Argentina

but also in N.E. Argentina. The abruptly raised midventral carina of the thorax

is distinctive in both sexes, and the abruptly raised longitudinal carina of tergum

VI is distinctive in the female. The tricolored abdomen, consisting of red anteriorly

and dark posteriorly, marked with large transverse pale spots is also distinctive.

New records (8 6, 15 9).—ARGENTINA: Catamarca: Ampajango, XII, Stange

(1 6, 1 9—FSCA); El Pucara, Neff (1 9—FSCA); Cuesta de la Chilca, XII, Neff (2

6—FSCA). Cordoba: Bialet Masse, II, Willink (1 9—IML); Alta Gracia, XI, Stange

(1 9 —FSCA); Quilpo, II, Fritz (1 9—FSCA); Capilla del Monte, II, Fritz (2 9—

Fritz Coll.). Corrientes: Ituzaingo, XII, Fritz (2 9—Fritz Coll., FSCA). Entre Rios:

Liebig, II, Fritz (1 6—Fritz Coll.). Mendoza: Potrerillos, II, Stange (1 6—FSCA).

Salta: Yacochuya, 9 mi. N.W. Cafayate, X, XII, Porter & Stange (1 6, 2 9 —FSCA).

Tucuman: Amaicha, XI, Stange (1 6—FSCA).

Epanthidium erythrocephalum (Schrottky)

(Figs. 13, 21)

Anthidium erythrocephala. Schrottky, 1903:453, PI. XIII, Fig. 8. Syntypes, 6, 9,

Campinas, Sao Paulo (Sao Paulo).

Taxonomy: Schrottky, 1910:271 (in Dianthidium); Moure, 1947:35 (in Epanthi¬

dium).

Both sexes are distinguished by the angled axilla (Fig. 21). The lack of a lon¬

gitudinal carina on tergum VI of the female is shared only by E. bo hart i and E.

paraguayense. Schrottky (1903) gave a color figure of this species which has

distinctive coloration among the known Brazilian species.

New records (14 6, 9 9).—ARGENTINA: Misiones: Bompland, III, Joergensen

(1 6—La Plata); Cataratas del Iguazu, XI, Porter & Stange (1 9—FSCA). Salta:

Rio Pescado, YPF Estacion, XI, Porter (1 6—FSCA); Yacochuya, 9 km N.W.

Cafayate, XII, IV, Porter & Stange (2 3, 1 9—FSCA). San Juan: Pocitos, 1(16—

FSCA). Tucuman: Horco Molle, 700 m, II, Stange (1 6—FSCA); Tacanas, XII,

Stange (1 9—FSCA). BOLIVIA: Santa Cruz: Saavedra, Est. Experimental, I, Stange

(1 9 —FSCA); 38 km N. Santa Cruz, I, Stange (1 9—FSCA); Buena Vista, VII,

Stange (2 6—FSCA). BRAZIL: Goias: Aragarcas, I, Oliviera (1 9—FSCA). PAR¬

AGUAY: Rio Ypone, Cororo, II, Fritz (5 6, 4 9—FSCA, Fritz Coll.).

VOLUME 59, NUMBERS 1-4

287

Epanthidium joergenseni (Friese)

(Figs. 3, 15)

Anthidium joergenseni Friese, 1908:73. Syntypes, < 3 , 9 , Mendoza (Pedregal) and

Tucuman, Argentina (not located).

Dianthidium bruchi Schrottky, 1909a:217. Holotype <3, Catamarca, Argentina, C.

Bruch, III-08 (not located). New synonymy.

Taxonomy: Joergensen, 1912:137 (in Dianthidium ); Moure, 1947:35 (in Epan¬

thidium).

This is one of the largest species in the genus, apparently confined to the Larrea

Subandean desert in Argentina. The female has a characteristic subapical ridge

on sternum VI with a strongly developed lateral process much anterior to the

large apical process (Fig. 15), the 3 processes forming an almost equilateral triangle.

The male is less easily characterized but lacks reddish bristles on sternum 11 and

III, has tergum VII with the emargination between the middle and lateral processes

rather narrow (Fig. 3), and lacks specializations found in other species (axilla not

modified, midventral carina of thorax not raised, tegula neither drawn out nor

impunctate). I am synonymizing Dianthidium bruchi Schrottky based on the

original description.

New records (10 < 5 , 9 9 ).— ARGENTINA: Catamarca: Andalgala, X, I, flowers

of Verbecina, Neff, Stange & Willink (4 < 3 , 1 9 —FSCA, IML); Ampajango, XII,

Stange (1 9 — FSCA); Los Nacimientos de Abajo, I, III, Stange & Willink (1 S, 1

9 — FSCA, IML). Salta: Yacochuya, 9 km N.W. Cafayate, IX, Stange & Willink

(1 < 3 , 1 9 —FSCA, IML); Tacuil, XII, Stange & Willink (1 6— IML). Santiago del

Estero: Rio Salado, Wagner (1 9 — La Plata). Tucuman: Amaicha, XI, Stange (3

< 3 , 1 9 -FSCA).

Epanthidium nectarinioides (Schrottky)

(Fig. 22)

Anthidium nectariniaid.es Schrottky, 1903:451. Syntypes, < 3 , 9 , Campinas, Sao

Paulo, Brazil, 30-1 (Sao Paulo).

Taxonomy: Schrottky, 1910:271 (in Dianthidium ); Schwarz, 1933:21 (in Antho-

dioctes)\ Moure, 1947:35 (in Epanthidium).

This species is easily recognized by the color pattern. The black body with

metasomal terga III-VI yellowish orange is unique in the genus. However, this

pattern is common in Anthodioctes (Megachilidae) and Brachygastra (Vespidae)

and other Hymenoptera of the same region. In the original description, this name

is spelled “ nectariniaides” but this is obviously a typographical error since the

name is spelled “ nectarinioides ” in two other places (pp. 444, 445 in Keys). The

spelling “ nectarinioides ” has been used in all subsequent citations.

New records (5 < 3 , 4 9 ).— ARGENTINA: Jujuy: Camino St. Pedro, II, Monroe

& Willink (1 < 3 — IML). Misiones: I (1 S —La Plata). Salta: Pocitos, XI, IV, Porter

& Fritz (2 < 3 , 2 9 — FSCA, Fritz Coll.); Rio Pescado, YPF Estacion, IV, Porter (1

< 3 , 1 9 — FSCA).

Epanthidium nigrescens Friese, New Combination

(Fig. 18)

Anthidium nigrescens Friese, 1906:100. Syntypes, < 3 , 9 , Mendoza, Salta (AMNH!;

Berlin).

288

PAN-PACIFIC ENTOMOLOGIST

The male is distinguished by the combination of reddish bristles mesad of

sublateral tooth of transverse ridge of sternum II, densely punctate unmodified

tegula and raised ventral median longitudinal carina of mesothorax. The lack of

a definite sublateral spine-like process (Fig. 18) on female sternum VI is distinctive.

Also, the weak longitudinal carina of tergum VI is a helpful character to identify

the females. A nest was seen from Bolivia with 5 cells (no resin) in adobe clay

attached beneath a rock overhang.

New records (41 8, 38 2).—ARGENTINA: Jujuy: Palpala, I, Aczel (2 8— IML);

6 km S. Volcan, III, Stange & Willink (1 2—FSCA). Mendoza: Mendoza, I (1 8—

FSCA). Salta: Yacochuya, 9 km N.W. Cafayate, IV, Porter & Stange (32 8, 30

2—IML, FSCA, LACM); Tacuil, IV, Stange & Willink (2 <$, 2 2-IML, FSCA).

Tucuman: Amaicha, XI, I, Stange (3 8, 1 2—FSCA). BOLIVIA: Camargo, 2480

m, I, Weyrauch (2 <3-IML, FSCA).

Epartthidium paraguayense (Schrottky)

(Fig. 4)

Dianthidium paraguayense Schrottky, 1908:232. Holotype 2, Paraguay, Asuncion,

Anisits leg., 4-1 (not located).

Taxonomy: Moure, 1947:35 (in Epanthidium).

The male has a bilobate tergum VII which is distinctive except for E. boharti.

The female lacks a longitudinal carina on tergum VI which is also characteristic

for E. boharti and E. erythrocephalum. The tricolor pattern of E. paraguayense

distinguishes both sexes from E. boharti, whereas the non-angled axilla provides

an easy recognition character from E. erythrocephalum. All of the previous records

were from N.E. Argentina, Paraguay and S.E. Brazil so that the newly collected

material from N.W. Argentina is an important range extension.

New records (33 8, 30 2).—ARGENTINA: Salta: Yacochuya, 9 km N.W. Ca¬

fayate, IV, Porter & Stange (29 <3, 24 2—FSCA, LACM, Fritz Coll.); Pocitos, XI,

Fritz (1 2—FSCA). Catamarca: El Rodeo, IV, Porter (2 3—FSCA). Misiones:

Bompland, I, Joergensen (1 8— La Plata); Dos de Mayo, XI, Escobar & Claps (1

2—IML). Tucuman: 10 km W. Las Cejas, IV, Stange (1 8— FSCA). BRAZIL:

Xambore, XII, Azeveda (1 2—Curitiba); Matto Grosso, IV-V, Fairchild (1 2—

FSCA). PARAGUAY: Independencia, II, J. Foerster (1 2 —KU); Rio Ypane,

Cororo, II, Fritz (1 2—Fritz Coll.).

Epanthidium sanguineum (Friese)

(Fig. 10)

Anthidium sanguineum Friese, 1908:74. Syntypes, 8, 2, Mendoza & Tucuman

(Vienna!).

Taxonomy: Joergensen, 1912:138 (in Dianthidium ); Moure, 1947:35 (in Epan¬

thidium).

This species is distinguished easily from all other known Epanthidium by the

nearly impunctate tegula. The apically bispined subapical ridge of sternum VI of

the female is diagnostic for that sex. In the male the toothed midcoxa is char¬

acteristic. Coloration is highly variable with the metasoma reddish in terga I—II,

and III-VI much darker with or without pale areas on the terga. It is fairly

abundant in the Subandean desert of western Argentina.

VOLUME 59, NUMBERS 1-4

289

New records (25 3, 24 2). —ARGENTINA: Catamarca: 8 mi. N. Belen, flowers

of Zuccagnia, X, Stange (3 <3, 1 2—FSCA); Andalgala, flowers of Prosopis alba,

X, Willink & Stange (1 3—IML); 6 km S. Santa Maria, flowers of Larrea divaricata,

XII, Willink & Stange (1 2—IML); Ampajango, XII, Stange (1 2—FSCA). La

Parnpa: Sierra Lihuel Calel, II, Stange (1 2—FSCA). La Rioja: 7 km S. Villa

Casana, flowers of L. divaricata, XII, Porter & Stange (1 3, 5 2—FSCA); Villa

Union, XII, Porter & Stange (1 2—FSCA). Mendoza: La Pasarella, flowers of L.

divaricata, XI, Neff (1 3—FSCA). Rio Negro: Balneario Las Grutas, I, J. & L.

Stange (1 2—FSCA). Salta: Yacochuya, 9 km N.W. Cafayate, XII, Porter & Stange

(1 2—FSCA). Tucuman: Amaicha, XI, I, Stange (3 <3, 2 2—FSCA).

Epanthidium tigrinum (Schrottky)

(Fig. 14)

Hypanthidium tigrinum Schrottky, 1905:10. Holotype 2, Paraguay, Villa Encar-

nacion (Vienna!).

Anthidium multifasciatum Strand, 1910:547. Syntypes, 4 <3, Paraguay, Villa Morro

and Asuncion (Berlin). New synonymy.

Taxonomy: Schrottky, 1908:231 (in Dianthidium ; description of 3); Moure, 1947:

33 (type species of Epanthidium).

The male of this species is unique in the genus in having reddish bristles on

metasomal sternum III and by the small process of the mesepisternum. The female

is distinguished easily by the mostly yellow mandibles. The typical pale stripes

on the mesoscutum do not vary in my specimens, but occur in some other species.

New records (14 3, 13 2).—ARGENTINA: Misiones: Cataratas del Iguazu, XI,

C. Porter & L. Stange (13 <3, 9 2-FSCA, UCD, LACM). PARAGUAY: Villa

Encarnacion (2 2 —Vienna, LACM); Villarrica, II, F. H. Schade (1 2—KU). BRA¬

ZIL: Goias: Ihla do Bananal, Santa Isabel do Moro, VI, M. Alvarenga (KU);

Minas Gerais: Belo Horizonte (1 2—Curitiba). BOLIVIA: Santa Cruz: Buena

Vista, I, Porter & Stange (1 3—FSCA).

Floral Records

All floral records are from Argentina, mostly from Andalgala, Prov. Catamarca,

October 1973 to March 1974 by J. L. Neff, unless otherwise noted.

ACANTHACEAE

Justicia campestris Griseb.

BORAGINACEAE

Heliotropium sp.

CAPPARIDACEAE

Atamasquea emarginata Miers ex Hook.

& Arn.

COMPOSITAE

Hyalis argentea D. Don ex Hook. &

Arn.

Senecio sp.

EPANTHIDIUM spp.

confusum (1 3, 1 2, Cuesta de Chilca,

Catamarca)

bicoloratum (1 3, Fuerte Quemada, Tu¬

cuman, I, Neff)

sanguineum (4 3, 3 2, III)

joergenseni (13, XII)

bertonii (1 3, Belen, Catamarca, XII,

Neff)

bicoloratum (Joergensen, 1912)

290

PAN-PACIFIC ENTOMOLOGIST

Tessaria dodonaefolia Cabrera

T. absinthioides (Hook. & Arn.) DC

Wedelia glauca (Ortega) Blake

Verbesina octantha S. F. Blake

Verbesina sp.

EUPHORBIACEAE

Jatropha excisa Griseb.

LABI AT AE

Salvia gilliesii Benth.

LEGUMINOSAE

Hojfmanseggia sp.

Marrubium vulgare L.

Prosopis chilensis (Mol.) Stuntz

P. nigra (Griseb.) Hieron.

P. alba Griseb.

Adesmia muricata (Jacq.) A. DC

Adesmia sp.

Zuccagnia punctata Cav.

Cercidium praecox (R. & P. Harms.)

LOASACEAE

Caliphora sp.

bertonii (1 3, Andalgala, X, Bohart; 1 3,

Belen, Catamarca, XII, Neff)

bicoloratum (12)

joergenseni (1 <3, Chaquiago, Catamar¬

ca, XII, Neff)

bicoloratum (12)

bicoloratum (6 <3, 2 2, Andalgala, XI,

Bohart)

joergenseni (7 <3, 1 2, Andalgala, X,

Stange)

sanguineum (2 <3, XII)

bicoloratum (1 <3)

paraguayense (1 <3, Yacochuya, Salta, IV,

Stange)

confusum (after Joergensen, 1912)

joergenseni (after Joergensen, 1912)

bicoloratum (2 2, Yacochuya, XII,

Stange)

bertonii (2 2, Yacochuya, XII, Stange; 1

3, Salta, XII, Neff)

autumnale (12, Yacochuya, Salta, XII,

Neff)

erythrocephalum (13, Yacochuya, XII,

Stange)

joergenseni (2 3)

sanguineum (13)

bicoloratum (13 3, 4 2)

sanguineum (1 3, 1 2, Colpes, Catamar¬

ca)

bertonii (2 3, Carrizal, La Rioja, X, Bo¬

hart)

bicoloratum (5 3, 1 2, Andalgala, XI,

Bohart)

confusum (12, Yacochuya, XII, Stange)

autumnale (1 3, Yacochuya, Salta,

Stange)

bicoloratum (12)

bicoloratum (13)

confusum (Joergensen, 1912)

joergenseni (12, Yacochuya, Salta, XII,

Stange)

nigrescens (1 2, Yacochuya, Salta, XII,

Stange)

bertonii (12, Escoipe, Salta, II, Neff)

VOLUME 59, NUMBERS 1-4

291

OXALIDAE

Oxalis sp.

POLYGALACEAE

Monnina lorenziana Chod.

SOLANACEAE

Salpichroa origanifolia (Lam.) Thel-

lung

Lycium chilense Miers ex Bert.

VERBENACEAE

Lippia turbinata Griseb.

ZYGOPHYLLACEAE

Bulnesia foliosa Griseb.

Larrea cuneifolia Cav.

L. divaricata Cav.

confusum (1 2, Agua de las Palomas,

Catamarca, I, Neff)

sanguineum (1 2, III)

bicoloratum (1 <3)

bicoloratum (2 <3, 1 2)

bicolor atum (1 2, Los Cocos, Cordoba,

I, Stange)

autumnale (1 <3, 1 2, Los Cocos, Cor¬

doba, I, Stange)

bicoloratum (7 8, 1 2)

bertonii (1 <3, Tapia, Tucuman, II, Neff)

sanguineum (3 <3, 3 2, XII)

bicoloratum (3 <3)

sanguineum (2 8 , XII; 1 8, XI, La Pa-

sarella, Mendoza, Neff)

bicoloratum (2 8, Cebollar, La Rioja,

XI, Bohart)

joergenseni (1 8, Andalgala, XI, Bohart)

bertonii (1 8, Andalgala, X, Bohart)

Key to Species of Epanthidium Moure

Males

1. Seventh metasomal tergum bilobate (Figs. 1, 4); curved ridge of sternum

II ending laterally in blunt process, without associated reddish bristles;

height of opaque lamella of posterior lobe of pronotum less than mid¬

ocellus diameter . 2

- Seventh tergum trilobate (Figs. 2, 3, 5); curved ridge of sternum II usually

ending laterally in sharp tooth, sometimes with reddish bristles mesad

to tooth or on sternum III; height of translucent lamella of posterior

lobe of pronotum usually more than midocellus diameter (except E.

nectarinioides . 3

2. Black with ivory markings on head, pronotum, axillae, metanotum, and

some metasomal terga; emargination of tergum VII deeper than width

at apex (Fig. 1) (Mexico) . boharti Stange, n. sp.

- Black and yellow, with orange on vertex, metanotum, tegula and legs;

emargination of tergum VII about equal to width at apex (Fig. 14)

(Argentina, Brazil, Paraguay) . paraguayense (Schrottky)

3. Metasomal sternum II or III with prominent, usually dark reddish bris¬

tles mesad of tooth or lateral angle. 4

- Metasomal sternum II and III without bristles . 6

4. Metasomal sternum II with only 1 or 2 prominent bristles mesad of

292

PAN-PACIFIC ENTOMOLOGIST

1 2 3

boharti autumnale joergenseni

0 BOHARTI

Figs. 1-8. Epanthidum spp. 1-5, Male tergum VII, dorsal view. 6, 7, Side view of tergum VII. 8,

Male genitalia (a = ventral view, b = lateral view).

tooth but III with many bristles; mesepisternum with small tooth pos¬

terior to extreme ventral part of preepisternal carina; ventral median

longitudinal carina of mesosoma low throughout; tegula with many fine

punctures (Misiones, Argentina; Bolivia; Brazil; Paraguay) .

. tigrinum (Schrottky)

- Metasomal sternum II with more prominent, usually dark red bristles

mesad of tooth than on sternum III; mesepisternum without small tooth

VOLUME 59, NUMBERS 1-4

293

ventrally; ventral median longitudinal carina of mesosoma produced

toothlike near anterior margin, or midcoxa with tooth at mesal posterior

angle; tegula punctate or nearly impunctate . 5

5. Ventral median longitudinal carina of mesosoma low throughout its

course; reddish bristles on sternum II running nearly to midline; tegula

nearly impunctate; midcoxa produced toothlike at mesal posterior angle

(Subandean desert, Argentina) . sanguineum (Friese)

- Ventral median longitudinal carina of mesosoma produced toothlike

near anterior margin; reddish bristles end well before midline; tegula

with many fine punctures; midcoxa without tooth at mesal posterior

angle (Subandean desert, Argentina—dark form; Chaco, Argentina; Bo¬

livia-reddish form). nigrescens (Friese)

6. Tegula drawn out posteriorly and thickened dorso-ventrally (Fig. 19);

midcoxa with small tooth or angle at mesal posterior angle .

. bicolorutum (Smith)

- Tegula not drawn out and thickened posteriorly (Fig. 20); midcoxa with¬

out process at mesal posterior margin . 7

7. Mid ventral carina of mesosoma produced toothlike near anterior margin

(Subandean desert, Argentina) ..'. confusum (Smith)

- Midventral carina of mesosoma not produced toothlike . 8

8. Tergum VII with lateral lobe nearly continuous with median process

(Fig. 2) (Subandean desert, Chaco or Misiones, Argentina) .

... autumnale (Schrottky)

- Tergum VII with deep U-shaped emargination separating lateral lobe

from median process (Figs. 3, 5) . 9

9. Upper row of propodeal pits strongly developed for nearly complete

distance; axilla angled or metasoma with anterior terga dark and pos¬

terior terga orange . 10

- Upper row of propodeal pits weakly developed, fading toward middle;

axilla not angled; metasoma without contrasting dark and orange terga

10. Metasoma with terga I—II dark, III-VI orange; axilla not angled (Fig.

22) (N.W. Argentina, Bolivia, Brazil, Paraguay) . . nectarinioides (Schrottky)

- Metasoma with all terga dark with yellow spots or transverse stripes;

axilla angled (Fig. 21) (N. Argentina, Bolivia, S.E. Brazil, Paraguay) . .

. erythrocephalum (Schrottky)

11. Tergum VII with middle process projecting below plane of lateral process

(Fig. 6); tooth at lateral margin of transverse ridge of metasomal sternum

II very large (basal width about equal to basal width of hind basitar-

somere), elevated at about a 40° angle from surface (Paraguay) .

. anisitsi (Schrottky)

- Tergum VII with middle process on same plane as lateral process (Fig.

7); tooth of transverse ridge of sternum II small, only slightly raised

from surface. 12

12. Lateral margins of terga V-VI with toothlike projections; tergum VII

with width of lateral process less than 2 times that of emargination

between middle and lateral processes (Fig. 5) (N. Argentina, Bolivia)

. bertonii (Schrottky)

294

PAN-PACIFIC ENTOMOLOGIST

- Lateral margins of terga V-VI tuberculate; tergum VII with width of

lateral process more than 2 times that of emargination between middle

and lateral process (Fig. 3) (Subandean desert, Argentina).

. joergenseni (Friese)

Females

(Female of E. anisitsi unknown)

1. Tergum VI without longitudinal carina; mandible mostly dark; clypeus

without continuous impunctate raised band along midline, although

some impunctate areas may be present . 2

- Tergum VI with longitudinal carina for at least half the distance between

anterior margin and sting emargination, if weak or obscure (some ti-

grinum and nigrescens ), mandible mostly yellow or clypeus with strong

impunctate raised band on midline from apex to dorsal border . 4

2. Axilla strongly angled laterally (Fig. 21); sternum VI with subapical ridge

not produced into a prominent tooth sublaterally (Fig. 13); tergum VI

with sublateral band produced into spine laterally followed by small

series of teeth (best seen in ventral view); height of translucent lamella

of pronotal lobe more than midocellus diameter (N. Argentina, Bolivia,

S.E. Brazil) . erythrocephalum (Schrottky)

- Axilla rounded laterally (Fig. 22); sternum VI with subapical ridge pro¬

duced into a prominent tooth sublaterally (Fig. 11); tergum VI with

lateral angle of sublateral band rounded followed by a rounded lobe;

height of opaque pronotal lobe less than midocellus diameter. 3

3. Black with ivory markings; terga III-VI with transverse ivory band at

middle, black laterally; distance from lateral ocellus to posterior margin

much greater than interocellar distance, area below midocellus with

punctures about a puncture diameter apart (N.E. Mexico).

., . boharti Stange, n. sp.

- Black and yellow, with orange on vertex, metanotum, tegula and legs;

terga III-VI dark at middle with sublateral pale band; distance from

lateral ocellus to posterior margin about equal to or less than interocellar

distance, sculpture about the same as area below midocellus (Salta and

Misiones, Argentina; S.E. Brazil; Paraguay) . paraguayen.se (Schrottky)

4. Tegula drawn out posteriorly into a laterally compressed process (Fig.

19) (Argentina) . bicoloratum (Smith)

- Tegula not drawn out posteriorly (Fig. 20) . 5

5. Tegula nearly impunctate; subapical ridge of sternum VI with pair of

submedium spines at posterior middle (Fig. 10) (Argentina deserts) . .

... sanguineum (Friese)

- Tegula with many fine punctures; subapical ridge of sternum VI without

pair of spines . 6

6. Mandible predominately yellow; tergum VI with longitudinal carina

weak, sometimes obliterated (N.E. Argentina, Bolivia, S.E. Brazil, Par¬

aguay) . tigrinum (Schrottky)

- Mandible predominately dark (some bertonii have yellowish mandibles

but carina of tergum VI strongly developed)

VOLUME 59, NUMBERS 1-4

295

AUTUMNALE SANGUINEUM

BOHARTI

BICOLORATUM

ERYTHROCEPHALUM

T1GRINUM

JOERGENSENI

CONFUSUM

BERTONI1

NIGRESCENS

21 22

Figs. 9-22. Epanthidium spp. 9-18, Female sternum VI in ventral view. 19, 20, dorsal view of

tegula; 19, E. bicoloratum, 20, E. autumnale. 21, 22, Dorsal view of axilla; 21, E. erythrocephalum,

22, E. nectarinioides.

7. Body and terga I—II black, terga III-VII mostly orange (N. Argentina,

S.E. Brazil) . nectarinioides (Schrottky)

- Without this color combination, terga I—II with pale spots or transverse

stripes or mostly reddish. 8

8. Midventral carina of thorax raised abruptly at anterior margin; longi¬

tudinal carina of tergum VI abruptly raised near middle; terga I—II mostly

296

PAN-PACIFIC ENTOMOLOGIST

reddish and terga III-V dark with large pale-yellow transverse stripes

(Subandean desert, Argentina) . confusum (Smith)

- Midventral carina of thorax not raised; longitudinal carina of tergum

VI not abruptly raised near middle; different color combination on

abdominal ridge .. 9

9. Metasomal sternum VI with subapical ridge strongly produced laterally

and with a truncate (Fig. 17) or spine-like median process (Fig. 15);

longitudinal carina of tergum VI strong to posterior margin. 10

- Metasomal sternum VI with subapical ridge usually angled at most

laterally, when somewhat produced process not much larger than lateral

angle of tergum VI (Figs. 12, 18), without discrete median process;

longitudinal carina of tergum VI not reaching posterior margin or weaker

than impunctate raised area on clypeus. 11

10. Metasomal sternum VI with subapical ridge produced spine-like at mid¬

dle (Fig. 15); lateral angle of tergum V weakly developed, much smaller

than lateral angle of tergum VI (Subandean desert, Argentina).

. joergenseni (Friese)

- Metasomal sternum VI with subapical ridge truncate at middle (Fig.

17); lateral angle of tergum V strongly developed, often as large as lateral

angle of tergum VI . bertonii (Schrottky)

11. Median impunctate raised band on clypeus more strongly developed

than median longitudinal carina of tergum VI (which is usually strongest

near posterior margin) (N.W. Argentina). nigrescens (Friese)

- Median impunctate raised band on clypeus less developed than median

carina of tergum VI (which fades toward posterior margin) (N. Argen¬

tina, S.E. Brazil) . autumnale (Schrottky)

Literature Cited

Friese, H. 1906. Resultate einer Reise des Herrn A. C. Jensen-Haarup in die Gegend von Mendoza

(Argentina). Hymenoptera, Fam. Apidae. Flora og Fauna 3:89-102.

-. 1908. Die Apidae (Blumenwespen) von Argentina nach den Reisener gebnissen der Herren

A. C. Jensen-Haarup und P. Joergensen in den Jahren 1904-1907. Flora og Fauna (1908):

1-74.

-. 1911. Apidae. I. Megachilinae. In: F. E. Schulze, ed., Das Tierreich 28:1-440. R. Friedlander

und Sohn, Berlin.

Grigarick, A. A., and L. A. Stange. 1968. The pollen-collecting bees of the Anthidiini of California

(Hymenoptera: Megachilidae). Bull. Calif. Insect Survey 9:1-113.

Joergensen, P. 1912. Revision der Apiden der Provinz Mendoza, Republic Argentina (Hym.). Zool.

Jahrb. Syst. 32:89-162.

Michener, C. D. 1948. The generic classification of the anthidiine bees (Hymenoptera, Megachilidae).

Amer. Mus. Novitates (1381): 1-29.

Michener, C. D., and E. Ordway. 1964. Some anthidiine bees from Mexico. J. New York Ent. Soc.

72:70-78.

Moure, J. 1947. Novos agrupamentos genericos e algumas especies novas de abelhas sulamericanas.

Mus. Paranaense Publ. Avulsas (3): 1-37.

Schrottky, C. 1903. Ensaio sobre al abelhas solitarias do Brazil. Rev. Mus. Paulista 5:330-613.

-. 1905. Contribucion al conocimiento de los Himenopteros del Paraguay. Ann. Cient. Paraguay

(4): 1-14.

-. 1908. Nuevos Himenopteros. Ann. Soc. Cient. Argentina 65:225-239.

-. 1909a. Hymenoptera nova. Ann. Soc. Cient. Argentina 67:209-228.

-. 1909b. Himenopteros de Catamarca. Ann. Soc. Cient. Argentina 68:233-272.

VOLUME 59, NUMBERS 1-4

297

-. 1910. Neue sudamerikanischen Arten der Bienengattumg Anthidium Fabr. Ent. Ztg. Wien

29:267-271.

-. 1920. Himenopteros nuevos o poco conocidos sudamericanos. Rev. Mus. Paulista 12:179—

227.

Smith, F. 1879. Descriptions of new species of Hymenoptera in the collection of the British Museum.

Taylor and Francis, London, 240 pp.

Strand, E. 1910. Beitrage zur Kenntnis der Hymenopterenfauna von Paraguay auf Grund der

Sammlungen und Beobachtungen von Prof. J. D. Anisits. VII. Apidae. Zool. Jahrb. Syst. 29:

455-562.

Schwarz, H. 1933. Some Neotropical Anthidiinae: Anthidium, Heteranthidium, and Dianthidium.

Amer. Mus. Novitates (624): 1-27.

Vachal, J. 1904. Voyage de M. G. A. Baer au Tucuman (Argentine). Hymenoptera mellifera (Familia

unica: Apidae). Rev. d’Ent. (France) 23:9-26.

PAN-PACIFIC ENTOMOLOGIST

59(1-4), 1983, pp. 298-301

A Redefinition of the Ageniella partita Group with

Descriptions of Two New Species

(Hymenoptera: Pompilidae)

Marius S. Wasbauer

Analysis and Identification/Entomology, California Department of Food and

Agriculture, Sacramento, California 95814.

The partita group of the subgenus Ageniella was treated by Townes (1957). He

included five species, two known from both sexes, two from the female only and

one from the male only. The present paper redefines the group, excludes one

previously described species and provides descriptions of two new species based

on both sexes.

As far as is known, the Ageniella partita group is strictly Nearctic in distribution

and includes species of small size and fugitive habits which are not well represented

in most collections. Features which will separate this group from others in the

subgenus are: Anterior wing of both sexes hyaline, neither patterned nor infuscate

except often with a poorly defined dark band at apex; marginal cell more than its

own length from wing apex; third submarginal cell short, at least 0.65 as high as

long; second recurrent vein meets third submarginal cell usually considerably

beyond middle and joins subdiscoidal vein half or less distance from origin of

subdiscoidal to wing margin; brush on inner side of posterior tibia somewhat

narrowed but usually not interrupted toward apex; females always with mesosoma

or metasoma red or orange; males of some species with tibial spurs of anterior

and middle legs white.

An explanation of the abbreviations used in the following descriptions is given

by Evans (1966:4). L and W refer to length and width, respectively and A3 refers

to the third antennal segment.

For the loan of comparative material, I wish to thank A. S. Menke, Systematic

Entomology Laboratory, U.S.D.A., % United States National Museum, Wash¬

ington, D.C. (USNM), A. Newton, Museum of Comparative Zoology, Harvard

University, Cambridge (MCZ), W. J. Pulawski, California Academy of Sciences,

San Francisco (CAS) and R. O. Schuster, University of California, Davis (UCD).

Ageniella ( Ageniella ) joannae Wasbauer, New Species

(Figs. 1, 2)

Holotype female. — 5.4 mm. Anterior wing 4.10 mm. Integumental color of head

black, of lower face, clypeus, antennae, legs, mesosoma and metasoma medium

brown, the following areas suffused with black: mesonotum laterally and poste¬

riorly, scutellum laterally, postscutellum medially, base of petiole and tarsi. Ap-

pressed pubescence long, dense, pale golden, somewhat coarser at apices of T 2-

4, forming ill-defined apical bands, silvery on mesepisternum, propodeum and

posterior coxae dorsally. Erect hairs short, pale golden, scattered over entire body,

longer on vertex, clypeus, mandibles, prosternum laterally, pygidium and meta-

somal sterna. Head rounded, slightly broader than long, TFD/FD 1.01; MID/FD

VOLUME 59, NUMBERS 1-4

299

0.55; ocelli in a compact, somewhat acute triangle, laterals much nearer to com¬

pound eyes than to each other, POL/OOL 0.64. Length of clypeus 0.40 X width,

frontal surface gently convex, apical margin slightly convex with ill defined prom¬

inence medially, apical hairless portion shining, wider at middle, set olf from

remainder of clypeus by a distinct narrow groove. Antennae of moderate length,

WA3/LA3 0.28, LA3/UID 0.73. Length of anterior wing marginal cell 0.86 X

distance from its apex to apex of wing. Transverse median vein meeting media

at origin of basal. Second recurrent vein nearly straight posteriorly, but strongly

curved at anterior fourth, meeting third submarginal cell 0.75 X distance from

base to apex of cell.

Allotype male. — 5.07 mm. Anterior wing 3.13 mm. Integumental color of head

black, that of lower face, clypeus, mandibles, mesosoma and anterior segments

of metasoma, light orange-brown, the following areas suffused with black: anten¬

nae, palpi, mesonotum and scutellum laterally, postscutellum medially, mesepi-

sternum ventrally, tarsi, base of petiole and apical mctasomal segments. Tibial

spurs of anterior legs orange-brown, suffused with black at apices, spurs of middle

and posterior legs black. Appressed pubescence pale, coarse, silvery on lower face,

occiput, prosternum, mesepisternum, posterior face of propodeum, and posterior

coxae dorsally. Head rounded, no broader than long, TFD/FD 1.00. Compound

eyes slightly divergent dorsally, UID/LID 1.15. Ocelli in a slightly acute triangle,

nearer each other than to compound eyes, POL/OOL 0.67. Antennae moderately

long, WA3/LA3 0.42, LA3/UID 0.60, width of penultimate segment 0.41 X length,

Subgenital plate gradually tapering to a narrowly rounded apex, strongly convex

in cross section but without a basal, triangular, flat-topped area. Genitalia (Fig.

1) with ventral lobe of digitus nearly horizontal, exceeding dorsal lobe; dorsal

lobe nearly vertical, blade-like. Aedeagus parallel-sided, not expanded at apex.

Gonostylus in lateral view (Fig. 2) with dorsal margin strongly sinuate.

Types. — Holotype 9 (UCD): CALIFORNIA: San Diego Co.: Warner Springs,

Aqua Caliente Creek, elevation 3100', VIII-23/25, 1980 (J. Slansky). Allotype S

(UCD) and paratypes (9 9 , 11 6, CAS and author’s collection) with same data as

holotype except collectors P. Adams, M. Wasbauer. Some specimens were taken

in a Malaise Trap.

The collector informs me that the holotype was taken as it walked among

foraging ant workers and was difficult to distinguish from the ants because of

similar size, coloration and pilosity. The ant was subsequently determined as

Formica pilicornis Emery.

This species is affectionately dedicated to Joanne Slansky.

A discussion of the relationships of Ageniella joannae is given together with

that for A. boharti.

Ageniella ( Ageniella ) boharti Wasbauer, New Species

(Fig. 3)

Holotype female. —4.4 mm. Anterior wing 3.70 mm. Integumental color of head

and mesosoma, antennae, palpi and legs light orange-brown, posterior tarsi a little

darker; metasoma dark brown, nearly black. Appressed pubescence very sparse,

pale golden becoming silvery on mesepisternum anteriorly and posterior coxae

dorsally. Erect hairs pale golden, very short and scattered except for a few longer

hairs on vertex, clypeus, mandibles, pygidium and metasomal sterna. Head slightly

PAN-PACIFIC ENTOMOLOGIST

300

Figs. 1, 2. Ageniella joannae. 1, Male genitalia, ventral view. Vestiture omitted on right side. 2,

Left gonostylus, external view. Fig. 3. Ageniella boharti. Left gonostylus, external view.

longer than broad, TFD/FD 0.97, MID/FD 0.63. Ocellar bases darkened, ocelli

in a compact triangle, the laterals much nearer each other than to compound eyes,

POL/OOL 0.71. Length of clypeus 0.37 X width, frontal surface gently convex,

apical margin nearly straight, hairless portion of apex shining, set off from re¬

mainder of clypeus by an indistinct groove. Antennae relatively short, WA3/LA3

0.30, LA3/UID 0.56. Length of anterior wing marginal cell 0.60 X distance from

its apex to apex of wing. Transverse median vein meeting media at origin of basal.

Second recurrent vein strongly curved, meeting third submarginal cell 0.65 X

distance from base to apex of cell.

Allotype male.— 2.21 mm. Anterior wing 1.59 mm. Integumental color of head

and metasoma black; mesosoma, clypeus, lower face, scape and pedicel pale

orange-brown; antennal flagellum and tarsi dark brown, the following areas of

mesosoma suffused with black: mesonotum, scutellum, postscutellum and mes-

episternum anteriorly. Tibial spurs of anterior and middle legs pale yellowish

white, of posterior legs blackish. Appressed pubescence sparse, pale over entire

body, coarser, silvery on lower face, clypeus, occiput, and most of mesosoma.

Head rounded, slightly broader than long, TFD/FD 1.05. Compound eyes di¬

vergent dorsally, UID/LID 1.28. Ocelli in a compact, slightly acute triangle, nearer

each other than to compound eyes, POL/OOL 0.70; antennae moderately long,

WA3/LA3 0.40, LA3/UID 0.45, penultimate segment 0.55 as wide as long.

Subgenital plate narrow, tapering gradually from base to a narrowly rounded apex;

strongly convex but without a basal, triangular, flat-topped area. Genitalia as in

Ageniella joannae but gonostylus in lateral view (Fig. 3) with dorsal margin not

as strongly sinuate.

VOLUME 59, NUMBERS 1-4

301

Types.— Holotype $ (UCD): CALIFORNIA: San Diego Co.: Borrego Valley,

IV-11-1969, R. M. Bohart collector. Allotype 6 (UCD) and paratypes (5 2, 2 6,

CAS and author’s collection): CALIFORNIA: San Diego Co.: Borrego State Park,

Clark Dry Lake, III-31 and IV-1, 1977, M. Wasbauer, J. Slansky, collectors.

At Clark Dry Lake, both sexes frequented the margins of the lake where there

were numerous cracks in the dried mud surface. Females were found investigating

the cracks and possibly were using them for nesting.

This species is named for a long time friend and associate, Prof. R. M. Bohart,

in honor of his outstanding contributions to our knowledge of the biology and

taxonomy of aculeate wasps.

Ageniellapallida Banks, known from females only, was placed by Townes (1957)

in the A. partita group. It is a species with an orange mesosoma and in this regard

is similar to Ageniella joannae and A. boharti. I have examined the holotype of

pallida in the MCZ and other material in the USNM. The following features will

separate it. from A. joannae and A. boharti and exclude it from the partita group

as here defined: Anterior wing marginal cell long, 0.85-0.89 X its length from

wing apex; third submarginal cell long, height 0.54-0.55 length; second recurrent

vein meets third submarginal cell before middle and intersects subdiscoidal vein

more than half distance from origin of subdiscoidal to wing margin.

With A. pallida excluded, the only members of the partita group with orange

or brown mesosoma are Ageniella joannae and A. boharti. These species differ

as follows: for females, the body of Ageniella joannae is uniformly light brown

and exhibits a rather conspicuous pale golden appressed pubescence. The body

of A. boharti is noticeably bicolored with the mesosoma light brown and the

metasoma nearly black, with the appressed pubescence very dilute and much less

conspicuous. The ratio MID/TFD averages 0.61 for A. joannae and 0.58 for A.

boharti. LA3/UID averages 0.71 for A. joannae and 0.59 for A. boharti. There is

no overlap in the ranges of these two ratios for the type series of either species.

For males, A. joannae and A. boharti are similarly colored but the first two

metasomal segments of A. joannae are concolorous with the mesosoma and the

middle tibia! spurs are dark. In A. boharti, the metasoma is entirely dark and the

middle tibia! spurs are whitish. The genitalia are very similar but the dorsal margin

of the gonostylus is more strongly sinuate in A. joannae (Fig. 2) than in A. boharti

(Fig. 3).

Literature Cited

Evans, H. E. 1966. A revision of the Mexican and Central American spider wasps of the subfamily

Pompilinae (Hymenoptera: Pompilidae). Mem. Amer. Ent. Soc. 20:1-442.

Townes, H. 1957. Nearctic wasps of the subfamilies Pepsinae and Ceropalinae. U.S. Natl. Mus.

Bull. 209:1-286.

302

PAN-PACIFIC ENTOMOLOGIST

THE PAN-PACIFIC ENTOMOLOGIST

Index to new taxa in volume 59

aliquantillus Grigarick, Schuster and Nelson

( Echiniscus ), 73

apilosus Grissell ( Boharticus), 88

bequaertellus Snelling ( Polistes ), 269

bishopensis Irwin ( Pherocera ), 125

boharti Dailey and Sprenger ( Loxaulus ), 45

boharti Eigbme ( Diodontus ), 50

boharti Ferguson (. Philanthus ), 56

boharti Griswold ( Bembix ), 103

boharti Irwin ( Pherocera), 127

boharti Michener (Microthurge), 186

boharti Miller ( Hypogeococcus ), 194

boharti N. J. Smith ( Pulverro), 260

boharti Schlinger ( Ocnaea), 249

boharti Snelling (Polistes), 272

boharti Stange (Epanthidium), 283

boharti Wasbauer (Ageniella), 299

Boharticus Grissell, 80

Bohartiellus Marsh, 148

bohartorum Parker (Perdita), 229

boydi Irwin ( Pherocera), 128

breviclava Grigarick, Schuster and Nelson

( Oreella), 66

burnetti Dailey and Sprenger (Trichoteras), 43

chalucus Snelling (Polistes), 277

chewbacca Menke ( Polemistus ), 173

coconino N. J. Smith (Pulverro), 261

conversus Horning and Schuster (Pseudechi-

niscus), 108

cornutus Marsh (Bohartiellus), 150

crossotus Grigarick, Schuster and Nelson

(Bryodelphux) , 69

dickboharti Menke ( Polemistus), 168

eighmei N. J. Smith (Pulverro), 263

fragilis Dailey and Sprenger (Heteroecus), 46

gilli Miller (Hypogeococcus), 200

hamoni Miller (Hypogeococcus) , 202

hathor Pulawski (Cerceris) r 242

joannae Wasbauer (Ageniella), 298

latro Menke and Richards (Trypoxylon), 152

lyoni Dailey and Sprenger (Heteroecus), 47

margaretae Grissell (Boharticus), 86

margaretae Miller (Hypogeococcus), 205

marginoporus Grigarick, Schuster and Nelson

(Echiniscus), 75

mexicanus N. J. Smith (Pulverro), 264

Microthurge Michener, 181

mosaicus Grigarick, Schuster and Nelson

(Echiniscus), 76

nanellus Grissell (Boharticus), 87

nigragena Irwin (Pherocera), 132

parkeri Ferguson (Philanthus), 58

parvisentus Homing and Schuster ( Pseudechi-

niscus), 109

patei N. J. Smith (Pulverro), 264

perplexus Horning and Schuster ( Pseudechi-

niscus). 111

plaumanni Marsh (Bohartiellus), 151

richardi Grissell (Boharticus), 84

rufoabdominalis Irwin (Pherocera), 133

rupina Irwin ( Pherocera), 136

tom.entam.ala Irwin ( Pherocera ), 138

tridigitus Schuster (Macrobiotus), 254

vaderi Vincent (Polemistus), 172

yoda Vincent (Polemistus), 171

zolae Grissell (Rhopalicus), 90

NOTICE

This issue constitutes the whole volume for 1983.

Parts 3 and 4 of Volume 58 for the year 1982

will be published later this year. This unfortunate

circumstance could not be avoided.

Paul Arnaud, Editor

THE PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

Papers on the systematic and biological phases of entomology are favored, including short notes or articles up to ten printed pages,

on insect taxonomy, morphology, ecology, behavior, life history, and distribution. Excess pagination must be approved and will be

charged to the author. Papers arc published alter acceptance in approximately the order that they are received. Papers of less than a

printed page will be published as space is available, in Scientific Notes.

All manuscripts will be reviewed before acceptance.

Manuscripts lor publication, proofs, and all editorial matters should be addressed to the editor.

General. — The metric system is to be used exclusively in manuscripts, except when citing label data on type material, or in direct

quotations when cited as such. Equivalents in other systems may be placed in parentheses following the metric, i.e. “1370 m (4500

ft) elevation”. j

Typing. — Two copies of each manuscript must be submitted (original and one xerox copy or two xerox copies are suitable). All

manuscripts must be typewritten, double-spaced throughout, with ample margins, and be on bond paper or an equivalent weight

Carbon copies or copies on paper larger than 8'/r X 11 inches are not acceptable.

Underscore only where italics are intended in the body of the text. Number all pages consecutively and put authors name on each sheet.

References to footnotes in text should be numbered consecutively. Footnotes must be typed on a separate sheet.

Manuscripts with extensive corrections or revisions will be returned to the author for retyping.

First Page. — The page preceding the text of the manuscript must include (1) the complete title. (2) the order and family in parentheses,

(3) the author's name or names, (4) the institution with city and state or the author’s home city and state if not affiliated (5) the

complete name arid address to which proof is to be sent.

Names and descriptions of organisms. - The first mention of a plant or animal should include the full scientific name with the author

of a zoological name not abbreviated. Do not abbreviate generic names. Descriptions of taxa should be in telegraphic style. The

International Code of Zoological Nomenclature must be followed.

Tables. — Tables are expensive and should be kept to a minimum. Each table should be prepared as a line drawing or typed on a

separate page with heading at top and footnotes below. Number tables with Arabic numerals. Number footnotes consecutively for

each table. Use only horizontal rules. Extensive use of tabular material requiring typesetting may result in increased charges to the

author.

Illustrations. — No extra charge is made for line drawings or halftones. Submit only photographs on glossy paper and original drawings.

Authors must plan their illustrations for reduction to the dimension of the printed page (117 X 181 mm; 4V S x 7 ■/» inches). If possible,

allowance should be made for the legend to be placed beneath the illustration. Photographs should not be less than the width of the

printed page. Photographs should be mounted on stiff card slock, and bear the illustration number on the face.

Loose photographs or drawings which need mounting and/or numbering are nol acceptable. Photographs to be placed together should

be trimmed and abut when mounted. Drawings should be in India Ink, or equivalent, and at least twice as large as the pruned

illustration. Excessively large illustrations are awkward to handle and may be damaged in transit. It is recommended that a metric

scale be placed on the drawing or the magnification of the printed illustration be stated in the legend where applicable. Arrange figures

to use space efficiently. Lettering should reduce to no less than 1 mm. On the back of each illustration should be stated (1) the title

of the paper, (2) the author’s complete name and address, and (3) whether he wishes the illustration returned to him. Illustrations

not specifically requested will be destroyed. Improperly prepared illustrations will be returned to the author for correction prior to

acceptance of the manuscript.

Figure legends. — Legends should be typewritten double-spaced on separate pages headed EXPLANATION OF FIGURES and placed

following LITERATURE CITED. Do not attach legends to illustrations.

References. — All citations in text, e.g., Essig (1926) or (Essig 1958), must be listed alphabetically under LITERATURE CITED in the

following format;

Essig, E. O. 1926. A butterfly migration. Pan-Pac. Entomol.. 2:2 11-212.

Essig, E. O. 1958. Insects and mites of western North America. Rev. ed. The Macmillan Co., New York, 1050 pp.

Abbreviations for titles of journals should follow a recent volume of Serial Sources for the Biosis Data Base, BioSciences Information

Service. For Scientific Notes the citations to articles will appear within the text, i.e. . . . “Essig (1926. Pan-Pac. Entomol., 2:21 1-212)

noted ..

Proofs, reprints, and abstracts. — Proofs and forms for the abstract and reprint order will be sent to authors. Changes in proof will be

charged to the author.

Editorial and administrative charges. — All regular papers of one to ten printed pages are charged at the rate of S24.00 per page. Private

investigators or authors without institutional or grant funds may apply to the society for a grant to cover a portion of the editing and

administrative charges. In no case will society grants subsidize more than two thirds of these costs. Individuals receiving a society

subsidy thus will be billed a minimum of $8.00 per page. Pages additional to the first ten are charged at the rate of $30.00 per page,

without subsidy. Editorial and administrative charges arc in addition to the charge for reprints and do not include the possible charges

for extra pagination or the costs lor excessive changes after the manuscript has been sent to the printer.

Reprint costs. — Current charges for reprints are as listed below. These charges are subject to change, and authors will be billed at the

rate in effect at the time of publication.

Each Additional

1 pp

Printed Page

1st 100

23.75

5.00

Additional 100’s

16.60

2.85

PUBLICATIONS

OF THE

PACIFIC COAST ENTOMOLOGICAL SOCIETY

PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.

Vol. 1 (16 numbers, 179 pages) and Vol. 2 (9 numbers, 131 pages). 1901-

1930. Price $5.00 per volume.

THE PAN-PACIFIC ENTOMOLOGIST.

Vol. 1 (1924) to Vol. 51 (1975), price $10.00 per volume of 4 numbers, or

$2.50 per single issue. Vol. 52 (1976) to Vol. 57 (1981), price $15.00 per

volume, or $3.75 per single issue, except for Vol. 57, no. 1, $10.00. Vol. 58

(1982) and subsequent issues, $20.00 per volume or $5.00 per single issue.

MEMOIRS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.

Volume 1. The Sucking Lice by G. F. Ferris. 320 pages. Published October

1951. Price $10.00 (plus $1.00 postage and handling).*

Volume 2. The Spider Mite Family Tetranychidae by A. Earl Pritchard and

Edward W. Baker. 472 pages. Published July 1955. Price $10.00 (plus $1.25

postage and handling).*

Volume 3. Revisionary Studies in the Nearctic Decticinae by David C. Rentz

and James D. Birchim. 173 pages. Published July 1968. Price $4.00 (plus

$0.75 postage and handling).*

Volume 4. Autobiography of an Entomologist by Robert L. Usinger. 343

pages. Published August 1972. Price $15.00 (plus $1.25 postage and han¬

dling).*

Volume 5. Revision of the Millipede Family Andrognathidae in the Nearctic

Region by Michael R. Gardner. 61 pages. Published January 21, 1975. Price

$3.00 (plus $0.75 postage and handling).*

*(Add 6 % sales tax on all California orders (residents of Alameda, Contra Costa, San Francisco,

Santa Clara and Santa Cruz counties add 6 ‘/ 2 %). Members of the Society will receive a 20%

discount on the price of the memoirs.)

Send orders to:

Pacific Coast Entomological Society

% California Academy of Sciences

Golden Gate Park

San Francisco, California 94118

U.S.A.