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&■

PSYCHE

A Journal of Entomology

Volume 81
1974

Editorial Board

Frank M. Carpenter, Editor P. J. Darlington, Jr.

W. L. Brown, Jr.
E. O. Wilson
B. K. Holldobler

H. W. Levi
J. M. Burns

R. E. SlLBERGLIED

Published Quarterly by the Cambridge Entomological Club
Editorial Office: Biological Laboratories
1 6 Divinity Avenue
Cambridge, Massachusetts, U.S.A.

The numbers of Psyche issued during the past calendar year (1974)
were mailed on the following dates:

Vol. 80, no. 4, December, 1973: February 20, 1974

Vol. 81, no. 1, March, 1974: May 21, 1974

Vol. 81, no. 2, June, 1974: September 26, 1974

CjLL-

-f’&l

Psyche

A JOURNAL OF ENTOMOLOGY

Vol. 81 March, 1974 No. 1

CENTENNIAL ISSUE

CONTENTS

Centennial Issue 1

History of the Cambridge Entomological Club. J. R. Matthews 3

Supplementary Studies of Ant Larvae: T etramyrmex.

G. C. Wheeler and J. Wheeler 38

Modification of the Intersegmental Region in the Pterothorax of Cry-
phocricos (Heteroptera : Naucoridae). M. C. Parsons 42

The Polytypic Genus Celotes (Lepidoptera : Hesperiidae: Pyrginae)
from the Southwestern United States and Northern Mexico. J. Burns 51

A Remarkable New Island Isolate in the Ant Genus Proceratium
(Hymenoptera : Formicidae). W. L. Brown , Jr. 70

Sexual Biology, Chromosomes, Development, Life Histories and Para-
sites of Boreus, Especially of B. notoperates. A Southern California
Boreus. II. (Mecoptera: Boreidae). K. W. Cooper 84

Emergence Pattern of the Subalpine Dragonfly, Somatochlora semi -
circularis (Odonata: Corduliidae) . R. L. Willey 121

The Milliped Genus Bollmanella (Diplopoda, Choreumida, Conotyli-
dae). W. A. Shear 134

A New Genus of Primitive Meloidae from West Texas (Coleoptera)

F. G. Werner 147

Polymorphism in Stelopolybia areata (Hymenoptera, Vespidae)

R. L. Jeanne and R. Fagen 155

Behavior of the North American Termite, T enuirostritermes tenuiros -
tris, with Special Reference to the Soldier Frontal Gland Secretion,

Its Chemical Composition, and Use in Defense.

W. L. Nutting , M. S. Blum, and H. M . Fates 167

The Castianeirinae of Mexico. I. Castianeira dugesi (Becker) (Ara-
neae: Clubionidae) . J. Reiskind 178

The Soldier of the Ant, Camponotus ( Colobopsis ) fraxinicola, as a
Trophic Caste. E. (). Wilson 182

Chemical Defense and Sound Production in Australian Tenebrionid
Beetles ( Adelium spp.). T. Eisner, D. Aneshansley, M. Eisner,

R. Rutowski, B. Chong, and J. Meinwald 189

Synchronous Visualization of Video-taped Sounds and Motions of
Insects. P. A. Wussow, R. B. Willey, and J. Steinberg

209

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1973-1974

President B. K. Holldobler

Vice-President W. D. Winter, Jr.

Secretary H. E. Nipson

Treasurer F. M. Carpenter

Executive Committee R. E. SiLBERGLlED

L. P. Lounibos

EDITORIAL BOARD OF PSYCHE
F. M. Carpenter (Editor), Fisher Professor of Natural History,
Emeritus, Harvard University

J. M. Burns, Associate Professor of Biology, Harvard University
W. L. Brown, Jr., Professor of Entomology , Cornell University ,
and Associate in Entomology , Museum of Comparative Zoology
P. J. Darlington, Jr., Professor of Zoology , Emeritus, Harvard
University

B. K. HoLLDOBLER, Professor of Biology , Harvard University
H. W. Levi, Alexander Agassiz Professor of Zoology, Harvard
University

R. E. Silberlied, Asssitant Professor of Biology , Harvard University
E. O. Wilson, Professor of Zoology , Harvard University

PSYCHE is published quarterly by the Cambridge Entomological Club, the
issues appearing in March, June, September and December. Subscription
price, per year, payable in advance: $4.50 to Club members, $6.00 to all other
subscribers. Single copies, $2.00.

Checks and remittances should be addressed to Treasurer, Cambridge
Entomological Club, 16 Divinity Avenue, Cambridge, Mass. 02138.

Orders for missing numbers, notices of change of address, etc., should be
sent to the Editorial Office of Psyche, 16 Divinity Ave., Cambridge, Mass.
02138. For previous volumes, see notice on inside back cover.

IMPORTANT NOTICE TO CONTRIBUTORS

Manuscripts intended for publication should be addressed to Professor
F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge,
Mass. 02138.

Authors are expected to bear part of the printing costs, at the rate of
$15.50 per printed page. The actual cost of preparing cuts for all illustra-
tions must be borne by contributors: the cost for full page plates from line
drawings is ordinarily $12.00 each, and for full page half-tones, $20.00 each;
smaller sizes in proportion.

The December, 1973 Psyche (Vol. 80, no. 4) was mailed February 20, 1974

The Lexington Press. Inc.. Lexington. Massachusetts

PSYCHE

Vol. 8 1

March, 1974

No. 1

CENTENNIAL ISSUE

The Cambridge Entomological Club, the second oldest of the
entomological societies existing in the United States, was founded
on January 8, 1874. A few months later, on May 8, the first issue
of its journal, Psyche , was published. In recognition of this anni-
versary, the Editorial Board of Psyche has designated this number
of the journal, March, 1974, as the Centennial Issue. The leading
article is a history of the Club, prepared by Janice R. Matthews
and based on her careful study of the historical documents in the
Club’s archives. The members of the Cambridge Entomological
Club are indebted to Mrs. Matthews for making her manuscript
available to the editorial board and for her permission to publish
it in Psyche.

The other articles in this issue are not directly related to the
Centennial but it is worthy of note that all of them are authored,
at least jointly, by members of the Cambridge Entomological Club.

We are indebted to Ward’s Natural Science Establishment,
Rochester, New York, for a helpful contribution towards the cost
of printing this issue of Psyche.

— Frank M. Carpenter, editor.

I

PSYCHE.

ORGAN OF THE CAMBRIDGE ENTOMOLOGICAL CLUB.

EDITED BY B. PICKMAN MANN.

Vol. I.] Cambridge, Mass., May, 1874. [No. 1.

Introductory.

The Cambridge Entomological Club was formed
January 9, 1874, by the following persons, who met at Dr.
Hagen’s house, No. 7 Putnam Street, Cambridge, Massachu-
setts, namely : Messrs. E. P. Austin, Edward Burgess, G. R.
Crotch, of Cambridge, England, George Dimmock, J. H.
Emerton, Dr. IT. A. Hagen, Messrs. Samuel Henshaw, B, P.
Mann, H. K. Morrison, J. C. Munro, of Lexington; Dr. A. S.
Packard, of Salem, Messrs. Eugene Schwarz, and S. H. Scud-
der. It has since added to its number Messrs. J. A. Allen,
Walter Faxon, A. W. Gould, Prof. C. E. Hamlin, Messrs.
Holmes Hinkley, H. G. Hubbard, Baron C. R. Osten Sacken,
Messrs. F. G. Sanborn, G. D. Smith, P. S. Sprague, Roland
Thaxter, of Newtonville, and C. P. Whitney, of Milford, N. H.

At the fourth monthly meeting, held April 10, 1874, the
Club decided to undertake the publication of a monthly
organ, to be called Psyche. This organ will contain such
parts of the proceedings of the Club as seem to be of general
interest, biological contributions upon Arthropoda from any
competent person, lists of captures, with time and locality,
miscellaneous entomological information, and especially a
Bibliographical Record, in which last a list will be given
of all writings upon Entomology published- in North America,
and of all foreign writings upon North American Entomology,
from the beginning of the year 1874, with a brief note of the'
contents of each. For the greater perfection of this list,
authors and societies are requested to forward their works to
the editor at the earliest date possible.

Each number will contain at least four pages, and as soon
as the returns are sufficient to make it possible, a greater
number of pages will be given. The subscription price in

FRONTISPIECE: Facsimile of page one of the first issue of Psyche, May,
1874.

HISTORY OF THE

CAMBRIDGE ENTOMOLOGICAL CLUB*

By Janice R. Matthews
Department of Entomology
University of Georgia
Athens, Ga. 30601

The Beginnings of the Cambridge Entomological Club

On a Friday evening, January 9, 1874, Dr. Hermann A. Hagen,
Professor of Entomology at Harvard College and Curator at the
Museum of Comparative Zoology, invited a group of twelve men
to his home at 7 Putnam Street in Cambridge, to consider the
question of forming an entomological society. Most of them had
been meeting informally for several years as a section of the Boston
Society of Natural History, but some had more ambitious plans.
Wanting to publish a journal, to meet outside of Boston, and to
have members from all over the country, they desired to form
“an organization independent of any other” — which was to be the
Cambridge Entomological Club.

Among the twelve present, probably the two most influential that
first evening were Dr. Hagen and Samuel Scudder [1,2]. Dr.
Hagen was the first professor of entomology in the United States;
he had left Germany in 1867 at the invitation of Louis Agassiz to
take charge of the entomological department of the Museum of
Comparative Zoology in Cambridge, and had been appointed to his
professorship at Harvard in 1870, at the age of 53. f But although

*This article is based on a term paper submitted in partial fulfillment
of the requirements for the degree of Master in Arts in Teaching from
Harvard University, 1967. The secretaries’ records, minutes of the Club
meetings, and other pertinent documents were placed at my disposal by the
officers of the Club. In the present account, quoted passages without specific
references are taken directly from the minutes of the meetings.

The original manuscript has been placed on permanent file with other
Cambridge Entomological Club historical documents in the Museum of
Comparative Zoology; it was revised and updated for publication here by
the editor, F. M. Carpenter.

fEntomology had been recognized in America as a serious branch of
science since the latter part of the eighteenth century, however. William
Dandridge Peck [3], the first native born American entomologist, initiated
the scientific study of insects at Harvard as that institution’s first professor
of natural history; as early as 1837 his student, Thaddeus W. Harris, while
acting as librarian of Harvard College, gave a course in entomology that
included brief field excursions [3,4]. Following Hagen’s arrival in Cam-
bridge, Harvard became a center of entomological activity, involving
undergraduate and graduate students as well as more mature investigators.

3

Psyche

[March

Dr. Hagen had held his professorial position for four years now, his
first course of lectures had been given only the previous summer,
and its enrollment had been but one student, J. H. Comstock; when
he did formally teach, Dr. Hagen’s courses consisted of “lectures,
given at rare intervals to advanced students.” As this might indicate,
Dr. Hagen’s principal work and real devotion were centered about
the Museum of Comparative Zoology, and to him other interests
were secondary. So although most influential in the formation of the
Entomological Club, and an enthusiastic supporter of its activities,
Dr. Hagen did not wish any responsibility toward running it. Thus,
when at this first meeting Dr. Hagen declined to take the chair (as
he declined, or resigned from every office for which he was ever pro-
posed in the Club), Samuel Scudder was chosen as chairman.

A graduate of Williams College and Harvard’s Lawrence Scien-
tific School, Samuel Scudder had been an assistant to Agassiz, and
at the time of the Club’s founding was nearly 38 years old. Once
considered “the greatest Orthopterist America has produced,” he also
worked on the diurnal Lepidoptera. and, as the foremost American
student of fossil insects in his time, served as paleontologist to the
U.S. Geological Survey from 1886 to 1892 [2]. Scudder was also
a competent editor and a bibliophile; he served as assistant librarian
of Harvard College and librarian of the American Academy of Arts
and Sciences. Yet despite these and many other time consuming
activities and before illness finally forced his withdrawal from active
participation in 1903, he held various formal offices in the Cambridge
Entomological Club for a total of eighteen years.

Samuel Scudder having been appointed to the chair, the meeting
moved on to the first order of business — the establishment of some
guidelines for the new organization. Voting to keep it as informal
as possible, “no more rules being made than are necessary,” the
members decided that the new Cambridge Entomological Club should
have only one permanent officer, a secretary; to fill this position,
they wisely chose 26 year old Benjamin Pickman Mann [5]. The
son of Horace Mann, well known as a teacher and advocate of public
schools, Benjamin had graduated from Harvard College only four
years previously. He was a conscientious researcher, a specialist in
entomological literature and bibliography, who for many years to
come would not only keep careful record of all Club proceedings, but
serve as treasurer, librarian, and editor of the Club’s publication.

After Mann’s appointment and the decision to hold the next meet-
ing at Scudder’s home, the Scientific Communications of the evening

1974]

Matthews — Cambridge Entomological Club

5

began. Dr. Hagen commented on the discovery of fossil galls, ap-
parently caused by insects, preserved on a twig in amber from Mary-
land; this was of special interest to Hagen, since he had published
extensively on Baltic amber insects while he was still in Germany.
There then ensued a general discussion of “the senses by which in-
sects are caused to assembly for sexual or other purposes.” This
must have been a particularly interesting discussion because of the
varied backgrounds represented. For example, there was Dr. A. S.
Packard, who had been one of Agassiz’s students and who had just
finished his third year as State Entomologist of Massachusetts [6].
He and Scudder were nearly the same age and they had been close
friends since their undergraduate days, but Packard’s experiences had
been more varied: he had been a surgeon in the Civil War, a Cus-
todian of the Boston Society of Natural History, a lecturer on ento-
mology at Massachusetts Agricultural College and Bowdoin College;
and he had studied marine life along the southeastern coast, and
had published his well-known “Guide to the Study of Insects.”
However, his active association with the Entomological Club was
to be very brief, for he was appointed to a professorship at Brown
University in 1878, a position which he held until his death in 1905.

Then, in contrast, there was Edward Burgess, at the age of 26, a
recent graduate of Harvard College and a former assistant in the
Museum of Comparative Zoology; he was currently Instructor in
Entomology at the College, giving the “course of elementary instruc-
tion in the study of insects.” Although he became known in ento-
mological circles for his published accounts of insect morphology,
Burgess later won renown for his contributions to naval architecture
[7]. At about the same age, there was James H. Emerton, who had
already foretold his life interest by collecting spiders at over a hun-
dred localities in New England [8]. A skilled artist, he had recently
finished the first of innumerable illustrations he would make for
A. S. Packard, S. H. Scudder, and many other zoologists. A trip
to Europe and a position as curator in the museum of the Peabody
Academy of Science at Salem soon removed him from the Cambridge
scene for a few years but he continued to publish extensively on
spiders and to take an active part in the Entomological Club until
his death in 1931. Another member was Samuel Henshaw; at the
age of 22 and without college training, he was at the time beginning
to work on the insect collection at the Boston Society of Natural
History; he subsequently became an assistant in entomology at the
Museum of Comparative Zoology and later (1912- 1927) its director

6

Psyche

[March

[9]. Of nearly identical age was George Dimmock, a Harvard fresh-
man who had a strong interest in insects, especially Coleoptera and
Lepidoptera. Although on graduating from college he spent several
years at the University of Leipzig in Germany, from which he re-
ceived his doctorate, he later returned to Cambridge and for many
years continued to be active in the Entomological Club [10]. Young-
est of all the founders of the Club was Herbert K. Morrison, only
19 years old, an energetic and serious student of noctuid moths. His
experience on the first of the Club’s excursions to the White Moun-
tains in New Hampshire, a few months later, induced him to become
a professional insect collector. During the next decade, he collected
extensively in the United States, especially in such little-known re-
gions as Washington Territory, Arizona, New Mexico and Nevada,
and he 'furnished countless specimens of many orders to specialists in
this country and Europe. His death at the early age of 31 terminated
a brilliant entomological career [ 1 1 ] .

Also at this first meeting there was a European coleopterist, Eugene
A. Schwarz. Born in Germany, he received his training at the Uni-
versities of Breslau and Leipzig. In 1872, at the age of 28, he came
to the Museum of Comparative Zoology as an assistant to Hagen.
He was to stay in Cambridge only a few years, however, leaving in
1875 on several collecting trips and finally joining other entomolo-
gists in the U.S. Department of Agriculture in Washington, where
he remained until his death in 1928 [12]. Very little can be said
about the two remaining members present at the meeting. E. P.
Austin, who was in the mining business, was an amateur coleopterist
and published several papers on beetles in the course of the next few
years, but he was not active in the Club after 1882. Even less is
known of J. C. Munro, who lived in Lexington; he appears not to
have attended any other meetings of the Club.

One individual, George R. Crotch, although not present at the
first meeting, or in fact any other meeting of the Club, was regarded
by all as one of the founders. He had become interested in insects,
especially Coleoptera, while an undergraduate at Cambridge Uni-
versity in England. He had collected extensively in Europe and in
late 1872 he had come to this country to collect insects in the western
states. He was a very energetic and enthusiastic entomologist and a
prolific writer [13]. In late 1873, at the age of 31, he accepted a
position as assistant at the Museum with Hagen. By the end of that

Matthews — Cambridge Entomological Club

1974]

drawing by J. H. Emerton in the C.E.C. archives.]

8

Psyche

[March

year, however, he had developed tuberculosis and was unable to
attend the first meeting of the Club. He died six months later.*

At the second meeting, held at Scudder’s house (156 Brattle
Street), there were discussions of such topics as the identity of a
borer destroying an elm tree at Henry W. Longfellow’s house
(Hagen), of the metamorphosis of the Saturniidae (Morrison) and
of the preparation of lepidopterous larvae for preservation (Hagen,
Scudder, Morrison). Seven new members were elected: J. A. Allen,
C. E. Hamlin, and C. R. Osten Sacken, all assistants at the Mu-
seum; Dr. Walter Faxon, curator at the Museum; H. G. Hubbard
and Roland Thaxter, both Harvard undergraduates; and C. P.
Whitney, of Milford, New Hampshire, the first non-resident mem-
ber. Osten Sacken began collecting insects, especially Diptera, when
he was a boy in Russia; he was on the staff of the Russian Legation
in this country for 27 years but at the age of 45, in 1873, he resigned
to become an assistant to Hagen. He was an active participant in
the Entomological Club for the entire period during which he was
working at the Museum, but after experiencing two winters in
Cambridge he moved to Rhode Island (a choice “influenced by the
temperate winter-climate”) ; and in 1877, his work on the Diptera
of North America finished, he returned to Europe [14]. Hubbard
became acquainted with E. A. Schwarz at the meetings of the Club
and shortly after they formed a collecting team, ultimately resulting
in the famous “Hubbard and Schwarz” collection of Coleoptera [15]-
The other undergraduate, Thaxter, started as an entomologist and
was active in the Club for many years, his first ten papers being
published in Psyche. However, his interest was directed by Pro-
fessor Farlow towards fungi parasitic on insects, and he subsequently
became Professor of Cryptogamic Botany at Harvard, with most of
his research being on these parasitic fungi, especially the Laboul-
beniales.

After the third meeting, the Cambridge Entomological Club
gathered at a little building nicknamed the “eritomologicon,” situated
in the backyard of B. P. Mann’s residence at 19 Follen Street [16].
These early meetings had no planned program; at each meeting, a
different member was chosen chairman and the minutes and acquisi-
tions to the Club’s library were read. The remainder of the meeting
was then opened to general discussion and the exhibition of new

*The list of signatures of the founding members in the minutes of the
first meeting includes a line that reads: “This was to have been the place
for the name of George Robert Crotch, Cambridge, England.”

1974]

Matthews — Cambridge Entomological Club

9

1ST e w - England.

JOURNAL OF EDUCATION.

Boston, Mass., August 21, 1875.
The Cambridge Entomological Club.

Camt of the Cambridge Entomological Club, )

Mr. Washington, July, 1875. I

Mr. Editor: — This is the first uncomfortable clay we have had.
The clouds rise from the valley and descend from the summit by
turns, driven by the shifting currents of air, and we get more of
the fog and drizzly showers here on the middle of the mountain
than they have either above or below us. Once in a while we
catch a glimpse of Ml. Carter, with half a dozen little clouds play-
ing a stately game of tag among his green ravines, or of the Glen
House in the sunny valley, but it is only for an instant, and
though we can hear the stage rattling along half a mile overhead,
we have not seen the road above the trees since sunrise.

Alter passing the five-mile post an extensive view is opened to-
ward Conway, taking in several very picturesque mountains and
lakes, the summit of Mt. Washington tower on the right, and in
the middle distance the rugged sides of the south wall of Tucker-
man’s ravine. Here is the beginning of the habitat of the Mount-
ain UutterHy, a species peculiar to this locality and eagerly sought
by nearly all our party. They have the curious habit of flattening
their wings down upon the ground or rock when they alight tc
avoid the wind, but such is the force of habit that they do so when
it is a calm also, raising them slowly afterwards as if it were a sec-
ond thought. The caterpillars live on a coarse kind of sedge
which grows here.

Proceeding to the summit we arrived in season to witness the
ascent of the singular looking little engine and car. The engine
being built for up-hill work seems, as one of the party aptly ex-
pressed it, to “ tip down” as soon as it comes on the piece of level
track in part of the platform. Most of us, after a rapid glance at
three States and a hundred lakes and rivers, devoted our time to
hunting Alpine beetles which abound under the rocks. Nearly all
these are species peculiar to high mountains, but on fair days many
butterflies, flies, and wasps wander up from the valleys. Return-
ing by moonlight, we did ample justice to the fried hominy and
syrup which the stay-at-homes had provided, and at ten o’clock
when the rain came splashing down in torrents, most of us were
too drowsy to think that the morrow’s projected trip to Tucker-
man’s ravine must be abandoned.

Yours truly, Walter IIoxie.

Portion of a letter written by Club member Walter Hoxie while at the
C.E.C. camp, Mt. Washington, July, 1875, and published in the New-England
Journal of Education. [Copy in C.E.C. archives.]

10

Psyche

[March

materials and curiosities. For its first three years, the Club continued
in this strictly informal manner, and included not only regular meet-
ings but excursions to areas df entomological interest.

At the 7th meeting, July, 1874, athe Chairman had to be con-
tented with sitting on a rock instead of a chair, a. feat which he
performed with sufficient grace and dignity, wrapped in a blanket.”
This peculiar situation occurred because, during the summer of 1874,
an “Entomologists’ Camp” was held on Mt. Washington in New
Hampshire, a “quarter of a mile below the Halfway House and far
enough into the woods to be out of sight of the road.” The party,
including members and non-members alike, left Boston by Portland
steamer, and remained in New Hampshire for almost a month; the
expenses, including round-trip fare from Boston, were about twenty
dollars apiece, and provisions, tents, etc., were provided by the Club
Excursion Committee — Dimmock, Austin and Mann. A regular
Club meeting was held, although it was “several times disturbed by
Mr. Morrison’s frantic attempts to capture the moths attracted by
the sole luminary of the occasion, his own lantern.” But in the main,
these summer excursions were light-hearted affairs, and when the
next Mt. Washington announcement, for July, 1875, stated that
“members may invite the attendance of ladies,” the ten men who
appeared at the meeting had fourteen women with them.

In the early years, there was no intention to limit place of meetings,
which were often held outside the borders of Massachusetts. Nor
was there distinction made between resident and non-resident mem-
bers. Both of these policies speedily changed the Club from a local
organization to one including members from many parts of the coun-
try. By January, 1879, the secretary reported 47 members residing
outside of New England, and only 19 within the area, most of them
in the vicinity of Boston and Cambridge.

The Beginnings of Psyche

At the fourth meeting, on April 10, 1874, Samuel Scudder pro-
posed that the Cambridge Entomological Club should begin publica-
tion of a monthly journal. A lively and lengthy discussion followed
this proposal, ending in the decision to undertake such a project.
The title of this new “Organ of the Cambridge Entomological Club,”
proposed by Scudder, was to be Psyche , derived from the Greek word
for butterfly. B. P. Mann was elected as the editor for this new
publication and “charged with the execution of all but the scientific

1974]

Matthews — Cambridge Entomological Club

1 1

Camp of the Cambridge Entomological Club,

HALF-WAY HOUSE,

Mt. Washington, N. H.

All matters relating to tents, their location, etc. will be atten-
ded to by B. PICKMAN MANN, Camp Master, C. E. C.

NO SUGARING OF TREES
ALLOWED WITHIN 500 FEET
OF THE CAMP!!!

Memorabilia of the Mt. Washington camp of the Entomological Club,
1874 and 1875. Top: letterhead of camp stationery; middle and bottom:
reproductions of signs posted at the camp.

work, which latter the members were engaged to supply.” The first
number, to consist of four pages, was to be ready by the next monthly
meeting of the Club, and the subscription price was set at one dollar
per year. The Club thus began Volume i of Psyche (which, as
completed, covers the years 1874-1876), as a place for publishing
“biological contributions upon Arthropoda from any competent per-
son,” and miscellaneous entomological information, while assiduously
avoiding “all discussion of vexed questions.” However, economic
entomology and taxonomic descriptions were less ‘favored than con-
tributions to general anatomy and biological entomology. But the
most important part of Psyche , in the opinion of the founders, was
to be the Bibliographic Record. Through this the Club set out
ambitiously to record all writings upon entomology published in
North America, and all foreign writings upon North American
entomology, from the beginning of the year 1874, with a brief note
on the contents of each. The original model for the Bibliographical

12

Psyche

[March

Record was clearly Hagen’s Bibliotheca entomologica, which ap-
peared in two volumes in 1862 and 1863.

The position of Psyche in the history of the Cambridge Entomo-
logical Club was to be a paradoxical one, for while it brought the
Club into a position of national renown, at the same time it led to
financial problems. Accordingly, early in 1876, the Club voted to
establish annual dues of $2 for New England members and also
designated a committee to raise a publication fund for Psyche , “the
principal of the fund to be invested in trust securities” and the
income to be used for the publications of the Club. A year later,
January 12, 1877, there being no financial improvement and nothing
in the publication fund, it was decided that additional measures
should be “taken to increase the effectiveness of the work of the
Club and to obtain money to defray the expenses of the Club and of
the publication of Psyche ” Scudder’s proposal, which was adopted,
was that “an act of incorporation should be performed” and he ad-
vised the adoption by the Club of a Constitution and By-laws, “which
must be in force as a preliminary to the act of incorporation.” The
Constitution and By-laws were promptly approved. At the following
meeting, February 9, 1877, with a Justice of the Peace for Middle-
sex County in attendance, Scudder was elected President and Mann
was elected Secretary and Treasurer, and these officers and the mem-
bers of the executive committee signed the agreement of association.
The Secretary of the Commonwealth, Henry B. Pierce, formally
signed the Certificate of Incorporation on March 9, 1877.*

At the time of incorporation of the Club there were 48 members,
half of the number being resident in the Boston-Cambridge area.
The meetings were well attended, with an average of 1 1 members in
addition to a few guests, and they were active affairs, having lively
discussions. Most meetings were held at various members’ residences,
though some were held at an office that Scudder used for editing his
journal, Science . Many additional non-resident entomologists were

*The Corporation was established according to the provisions of Chapter
375 of the Acts of the General Court of Massachusetts, passed in the same
year as the founding of the Club, 1874. The incorporation apparently had
no effect on the Club. No annual reports of the financial holdings of the
Club were ever submitted to the Secretary of the Commonwealth, as re-
quired by law, and on March 24, 1964, eighty-seven years later, the Secre-
tary of the Commonwealth dissolved the Corporation, in accordance with
Chapter 180, section 26A, of the General Laws. Revival of the Corporation
is, however, provided for in the legislative actions — Editor

1974]

Matthews - — Cambridge Entomological Club

13

Samuel Scudder’s study in the yard of his house, 156 Brattle St., Cam-
bridge. Consisting of a single large room, it included an extensive cabinet
of insect drawers on one wall, a fireplace, and book shelves on two walls.
The Cambridge Entomological Club held virtually all of its meetings here
from 1888-1901. [Photograph, probably taken about 1890, in M.C.Z. archives.]

elected to membership, even some as officers of the Club, presumably
as a means of increasing subscriptions to Psyche. One of the major
interests of the members at this time, apart from Psyche , was the
Club Library, the goal being to have in one place as nearly complete
a collection of entomological publications as possible for the use of
the members. This is not surprising, since both Scudder and Mann
were bibliophiles. At first the secretary of the Club had the respon-
sibility of recording all these accessions but in 1880 a librarian was
elected. By 1886 the Club library included 1652 volumes and sepa-
rates, which were at first housed in Mann’s office but later transferred
to Scudder’s study.

The Lean Years

By 1890 the membership of the Club had changed greatly. Mann
had left Cambridge permanently in 1887 to do bibliographic work
for the Federal Department of Agriculture; and several other of
the original members, including Austin, Dimmock, Morrison, Pack-
ard and Schwarz, had moved away from the Cambridge area, most

14

Psyche

[March

of them beyond New England. Financial problems at the Museum
of Comparative Zoology, following Agassiz’s death, reduced the
funds available for assistants. Hagen, though he lived for another
three years, was stricken with paralysis in 1890 and the February
meeting of that year was the last he attended. During the period
from 1890 to 1900, when the meetings were held in Scudder’s study,
only five resident members were elected to the Club, but four of
these were to play a most important part in the history of the or-
ganization. One of them, A. P. Morse, then 28 years of age, was
an assistant in the zoology department of Wellesley College; later
he became associated with the Boston Society of Natural History,
and still later with the Carnegie Institution of Washington, as a
specialist in Orthoptera. He continued to be active in the Ento-
mological Club for a total of 43 years, until his health failed in
19 35 [17]. Another of the new members was F. C. Bowditch, an
amateur coleopterist with special interests in the Chrysomelidae ; in
the course of his life he built up an extensive and important collection
of the Chrysomelidae of the world, now at the Museum of Compara-
tive Zoology [18]. The third of the members was J. W. Folsom,
who was active in the Club while he was a graduate student at
Harvard University. His early interests were in morphology and
physiology of insects and he later taught entomology at the Univer-
sity of Illinois before becoming associated with the U.S. Bureau of
Entomology [19]. The fourth of this group was W. L. W. Field,
who joined the Club at the age of 19, while he was a first year
student in Harvard College. He was an enthusiastic lepidopterist
and published several papers in Psyche on inheritance in butterflies.'*
He attended meetings of the Club regularly and, as editor of Psyche
from 1904-1909, was responsible for making significant improvements
in its nature and content. Field did not continue in entomology,
professionally, but taught biology at Milton Academy until 1917,
when he became headmaster, a position that he held until 1942.
These four members, in addition to Scudder, Henshaw and Roland
Hayward (an amateur coleopterist who joined the Club in 1879
[20] ) , were the only individuals that attended the Club meetings

*At the Club’s meeting in September, 1907, Professor William Bateson of
Cambridge University, England, was scheduled to be the speaker; last
minute changes prevented his coming, so W. L. W. Field “gave an inter-
esting talk on the breeding experiments” that were being conducted by
Bateson, who has “thus brought to the attention of the world again the
long-neglected or forgotten Hereditary Laws discovered by Mendel.”

1974]

Matthews — Cambridge Entomological Club

15

from 1900 to 1903. The average attendance at these meetings, also
held in Scudder’s study, was between three and four.

The Harris Club

Just prior to 1900, another entomological club was formed, this
time in the city of Boston. The moving force for this was Mr. H. H.
Newcomb, an amateur lepidopterist and general insect collector. An
organizational meeting was held on November 24, 1899, in his office
on Court Street, Boston. The ten who were present were enthusiastic
amateurs; most were in business or law, although a few were college
students, not yet established professionally. W. L. W. Field, already
a member of the Cambridge Club, was one of the ten and served as
secretary of the new club for the three years of its existence. At
their second meeting, the members decided on the name Harris Club,
“in honor otf Thaddeus William Harris, eminent among early Ameri-
can Entomologists, whose entire life was spent in the neighborhood
of Boston.”*

In many ways the Harris Club paralleled the earlier days of the
Cambridge Club. From the beginning, the members agreed “that
the organization should be as informal as possible.” The constitution
generally expressed the same goals as that of the Cambridge Club.
And as did its counterpart, the Harris Club had a library, but of a
much less formal and extensive nature. They were an extremely
enthusiastic group and held several field excursions — only in their
case to Mt. Katahdin in Maine. By 1903 the Harris Club included
41 local members, although the average attendance at the meetings
for the previous year had been only 12. As Field noted in one of
his annual reports, there had been a progressive decrease in the per-
centage of members attending as the total membership increased.
Field undoubtedly provided a liaison with the Cambridge Club and
was almost certainly responsible for the suggestion that the Harris
Club merge with the olddr organization. On January 13, 1903, Field
and Newcomb proposed:

“That the Harris Club be merged in the Cambridge Entomologi-
cal Club, all members of the Harris Club of record January 13,

1903, to be nominated on one ballot for membership in the Cam-
bridge Entomological Club. The latter is an incorporated Club
with a long and distinguished list of members, past and present.

*This was not the same organization as the Harris Entomological Club,
which was founded in 1864 as a section of the Boston Society of Natural
History and which was discontinued in 1886.

Psyche

[March

1 6

It has a Publication Fund, and maintains a monthly journal, Psyche.

It is believed by many local entomologists that one large club will
be better than two small ones and that the members of the Harris
Club will find many advantages in an alliance with the older or-
ganization. If the plan is carried out, the Cambridge Entomological
Club will hereafter hold regular meetings in Boston.”

This was unanimously voted by the Harris Club and at the February
13 meeting of the Cambridge Club the 38 active members of the
Harris Club were nominated by Field and Hayward for membership.
At the next meeting, March 13, all were elected, with only Field,
Hayward and Bowditch (all amateurs) representing the Cambridge
Entomological Club, Scudder being unable to take part because of his
paralysis.

The change in the number and type of members rapidly led to a
change in the nature of the Cambridge Entomological Club. The
concept of a Club library very shortly came under critical examina-
tion, in part because of Scudder’s illness and of the necessity of mov-
ing the library from his study. In three months the library com-
mittee, with Field as chairman, recommended that the Club library
turn over whatever volumes the Boston Society of Natural History
desired for its library, and to dispose of the rest among its members
by auction. Thus, by a series of auctions presided over by H. H.
Newcomb, most of the library was gradually disposed of at ridicu-
lously low prices, until in 1909 the remaining works were sold for
twenty dollars to a new member, W. M. Wheeler.

At the October meeting, 1903, the members considered the future
of Psyche, which had been edited since 1891 by Samuel Henshaw.*
A committee chaired by A. P. Morse, and including Bolster, Field,
and Henshaw, was appointed to consider whether or not to continue
its publication. In December, Field reported that “the committee was
thoroughly in favor of maintaining Psyche as the journal of the Club
and was ready to assume charge of the journal if that was the pleas-
ure of the Club.” It was and the editorship was assumed by W. L.
W. Field; in November, 1906, the Club members discussed the Bib-
liographic Record, which had been the main part of the journal under
Mann and Scudder, and decided to discontinue it and to devote the
space to contributors’ articles.

Other changes also began to manifest themselves. After 1903 the
Club began to possess a very local outlook; the attempt to maintain
a wide non-resident membership appears to have been abandoned.
The attention of the Cambridge Entomological Club from 1904 to

1974]

Matthews — Cambridge Entomological Club

17

CAMBRIDGE ENTOMOLOGICAL CLUB.

Regular meeting of the Club Friday, 14 Dec. 1883, at 7.45 p. m.

Mrs. A. K. Dimmock will show a collection of insects from Betula alba (the
white birch).

Mr. G. Dimmock will describe a new mode of making preparations to show
the double nature of the wing-membrane of insects, and will show some micro-
scopical preparations of other parts of insects.

The meeting will be held at the Secretary’s house, 54 Sacramento
street, Cambridge. [The Porters station horse cars of the Charles
river road, or the North avenue or Arlington cars of the Cambridge
road, pass Sacramento street.]

| "gp* Annual election of officers at the meeting of 11 Jan. 1881.

George Dimmock, Secretary.

Notice of the Club meeting for December 14, 1883, to be held at Dimmock’s
house on Sacramento St., with information about horse car transportation.

1909 seemed to focus almost totally upon its biggest project, the
annual exhibitions of insect collections held in the late fall at the
Appalachian Mountain Club, 1050 Tremont Building, Boston, and
open to the general public. Several times, the announced topics for
regular meetings were totally ignored, the entire program being de-
voted to the arrangements of exhibits and informal conversation.
Often, nearly every other member had a box of specimens with him,
mainly Lepidoptera, and during this period, members regularly ex-
hibited entomological curiosities at the meetings as well.1**

The Bussey Institution

In 1907 another event occurred which had great significance for
the Cambridge Entomological Club. As far back as 1871, a few
years before the Club was founded, Harvard College had established
the Bussey Institution, located in Boston, as a place for courses in
“practical agriculture,” and for some years after that time offered

*Henshaw not only edited these volumes with great care but he personally
covered all deficits resulting from their publication. At the meeting of
March 8, 1901, the Club elected Henshaw a Life-Member, “as a token of
the Club’s appreciation of his generosity.”

**This is the origin of the tradition, still maintained, that notices of the
meetings should include the statement: “Members are invited to bring speci-
mens to the meeting for demonstration and discussion.”

i8

Psyche

[March

The Bussey Institution, Forest Hills (Boston). The Club meetings were held in the library of this building
from 1910-1929. This was the editorial office of Psyche and the official address of the Club. [Photograph in the
Archives of the Harvard College Library.]

1974]

Matthews — Cambridge Entomological Club

19

entomological instruction to undergraduates. The program was not
very successful, however, and in 1907 the Bussey Institution was
reorganized for graduate study and research in biology, including
entomology. In 1908 William Morton Wheeler, a distinguished
zoologist and entomologist, then curator of invertebrate zoology at
the American Museum of Natural History, was appointed Professor
of Economic Entomology at the Bussey Institution [21]. On his
arrival he was elected to the Club and personally welcomed by
President C. W. Johnson.*

In the following year Charles T. Brues was added to the Bussey
Staff as Instructor in Entomology; and in November the Club gained
by his election another member who would have an immediate and
lasting influence on its future [23]. At the very next meeting, Presi-
dent Bolster reported the resignation of Field as editor of Psyche.
“Mr. Brues, being called upon by the chair, said he would try to
arrange things so that he could take up the work,” thus beginning his
distinguished term as editor, which would span the next 37 years.

The membership of Wheeler, Brues, and their graduate students
served to draw the Cambridge Entomological Club closer to the
Bussey Institution, not only intellectually but physically as well.
Meetings during the post- Harris Club period had been held at various
addresses around Boston: in the council room at the Boston Society
of Natural History; at Emerton’s room, Clarendon Street; at the
Appalachian Mountain Club, Tremont Building; and at Newcomb’s
office, Court Street. At the February meeting, 1910, newly elected
president Wheeler reported that certain rooms at the Bussey were
being remodeled and he expressed his hope that future meetings of the
Club might be held there. The suggestion was enthusiastically ac-
cepted and the members voted to hold the next meeting at the Bussey
Institution, in Boston. That was the place of meeting for the follow-
ing 19 years.

At this time a task of primary importance facing the Club and
especially the new editor, C. T. Brues, was that of putting Psyche
on a satisfactory financial basis; despite change in format, expenses
continued to exceed subscription income and the contributions from

*C. W. Johnson became a member of the Club in 1903, shortly after the
election of the Harris Club members, when he arrived in Boston to be
curator at the Boston Society of Natural History, as a specialist in Diptera.
A genial personality, with youthful and vigorous enthusiasm, he was a
true naturalist and helpful to all who consulted him. He was active in the
Club until his death in 1932 [22].

20

Psyche

[March

members were never great enough to give the Club working capital
to use. Early in 1916 Brues recommended that the length of the
articles be normally limited to eight pages and that articles of greater
length should be paid for by the authors, who would also be required
to bear the cost of the engravings for their illustrations and to pay
the cost price for their reprints. Although this was approved, for
many years the financial status of Psyche continued to fluctuate,
primarily because of the lamentable absence of any substantial en-
dowment to fall back upon in critical periods. Deficits did occur,
sometimes amounting to several hundreds of dollars, but various
members, notably Wheeler, Brues, Johnson, and Thomas Barbour,*'
made sufficient personal contributions to balance the accounts. Never-
theless, this was clearly the turning point in the history of Psyche ;
never again did the journal pass through such a protracted period of
financial struggle as between 1895 and 1915. The Club members
had every reason to be pleased at the Annual Meeting of 1919 as
they heard the report of the acting treasurer, A. C. Kinsey: “At the
close of a year of increased cost o'f everything it would not have
been surprising if a financial report for the Club had presented a
deficit .... Contrary to this, however, the cash balance to date is
$118.35. This improvement in conditions should be credited in part
to the increased efforts in collecting money due, but in a larger part
to the efforts of the editor of Psyche. An adoption of this report
should include an expression of thanks to the editor for his careful
management of the cost of publishing the magazine.”

During the first decade (1908-1918) of meetings at the Bussey,
the Club was very busy giving a cordial welcome to the many new
members elected during the period — a total of 92. Many of these
were graduate students at Harvard and they enlivened the programs
with accounts of their research and especially of their collecting trips.
Among these, for example, was W. M. Mann, who, as one of
Wheeler’s students, collected extensively during his graduate years,
even in such remote regions as the East Indies and South America.
He published occasionally on ants after leaving the Bussey but for

*Dr. Barbour was elected a member of the Club in November, 1909,
along with Brues, while he was a graduate student at Harvard. A verte-
brate zoologist, he was soon appointed curator of reptiles and amphibians
in the Museum of Comparative Zoology, and later served as Director of
the Museum (1927-1946). However, he continued his membership in the
Club and was very generous in his contributions towards the cost of pub-
lishing Psyche .

1974]

Matthews — Cambridge Entomological Club

21

most of his life he was Director of the National Zoological Park and
had little time for entomological studies [24]. Another student,
James W. Chapman, was also a myrmecologist ; in 1916, after grad-
uating from the Bussey, he joined the staff of Silliman University,
on the Island of Negros, Philippines, where he and Mrs. Chapman
remained until the Japanese occupation in World War II [25].
A few of the Harvard faculty joined the Club and were active
throughout this period, notably Nathan Banks, Curator of Insects
at the Museum of Comparative Zoology, and G. H. Parker, Pro-
fessor of Zoology at Harvard and for many years chairman of that
Department. During this time also the Club included among its
members a substantial number of enthusiastic amateurs, who took an
active and important part in the meetings. P. G. Bolster, a prac-
ticing attorney, was a very effective collector, mostly of Coleoptera;
he was president of the Club twice during this period, in 1909 and
1913 [26].'* Another amateur active at this time was L. W. Swett,
a proprietor of a store in Lexington; he built up a very large and
useful collection of geometrid moths, now housed in the Museum
of Comparative Zoology, and he was president of the Club in 19 11
[27]. One of the most active of all Club members at this time was
C. A. Frost, a civil engineer in Framingham. He was almost “the
last of the old-time general students of Coleoptera and was most
helpful to other coleopterists” [28] ; he published about eighty papers,
many of them in Psyche, and at various times served the Club as
secretary and treasurer, and twice as president.

Among the newest elements in the Club membership in this decade
were eight of the entomologists employed at the Gypsy Moth Parasite
Laboratory of the Federal Bureau of Entomology, in Melrose High-
lands. Included was the head of the control program for the Gipsy
moth, A. F. Burgess, president of the Club in 1916.

As might be expected, the meetings were well attended, with an
average attendance of about twenty. The programs were varied but
usually included a main speaker and two or three shorter presenta-
tions. The meeting of June 17, 1919, might well serve as an example
of one of this period, as well as of the decade to follow. The main
feature of the evening was an address by A. C. Kinsey, a graduate
student, on the origin of some biological characteristics of gall wasps.

*In 1909 Bolster prepared a manuscript on the history of the Club, the
substance of which was read to the members on his retirement from the
presidency in 1909. This was never published but is included in the ar-
chives of the Club.

22

Psyche

[March

(Qanthriiiyc Eutnmuliujiral (5luli

LECTURES ON INSECTS

These lectures will treat in a popular way
of the habits, growth and structure of Insects
and their adaptation to their surroundings.
They will he abundantly illustrated by the
stereopticon and by motion pictures as far as
they can be obtained.

Saturday afternoons at 2:30, March 12
and 19, April 16 and 23.

The lectures will be by the following well-
known members of the Club:

March 12. The Butterflies — by W. L. W.
Field, Headmaster of Milton Academy.

March 19. The Dragonflies — by R. Heber
Howe, Jr., Master of Natural Science at the
Middlesex School.

April 16. Ants and the Social Life of Insects —
by W. M. Wheeler, Professor of Entomology
at Harvard University.

April 23. Plant Galls and the Insects that Pro-
duce them — by A. C. Kinsey, of Indiana
University.

Tickets for the course, $2.00, for sale at
the Bussey Institution, Forest Hills, Boston,
Mass. Sent by mail on receipt of price.
Make checks payable to Cambridge Entomo-
logical Club. Course and single tickets for
sale at Tfemont Temple on days of lectures.

Announcement of the public lectures given in Boston (1921) by members
of the Cambridge Entomological Club.

1974}

Alatthews — Cambridge Entomological Club

23

Mr. Emerton followed, speaking of his recent spider-collecting trip
to Cape Cod; Dr. Wheeler exhibited specimens of the Japanese
beetle, recently introduced into New Jersey, and a drawing of a
“peculiar, elongate ant” from South America. Mr. Frost gave a
number of collecting notes on beetles and spoke of his success in
rearing sumac borers from dead twigs. The meeting concluded with
a discussion of insects as food by a visiting entomologist from Bel-
gium, Dr. J. C. Bequaert, his account being based on experiences in
the Congo region of Africa; and Dr. Wheeler remarked on his
observations of similar foods used by the Australian natives.

One of the important projects of the Cambridge Entomological
Club in the early twenties was a series of lectures given in Boston
for the public at large. The first formal note of this idea was in
October, 1920, when a committee of four was appointed to propose
plans. In the course which these affairs often follow, three of the
four ended up as lecturers as well — W. L. W. Field, R. H. Howe,
and W. M. Wheeler; the fourth member of the committee, J. H.
Emerton, having reached 72 years, preferred not to undertake a lec-
ture, and A. C. Kinsey, then at Indiana University, was selected to
take his place. All were to speak on their “well known specialities”
at Tremont Temple, on four successive afternoons during the spring
of 1921. The printed announcement of the lectures stated: “These
lectures will treat in a popular way of the habits, growth and struc-
ture of Insects and their adaption to their surroundings. They will
be abundantly illustrated by the stereopticon and by motion pictures
as far as they can be obtained.”

The lectures were quite a success, with an average attendance of
about 150 persons, and income from ticket sales ($2 for the entire
course, or fifty cents per lecture at the door) nearly equalled the
expenses. Inspired by this, the Club voted the following year to
expand the series to six lectures. Post card notices were sent to the
Audubon Society’s mailing list of 4,000 names, advertisements were
placed in the Boston Transcript , and printed posters sent to libraries,
schools and clubs. The lecturers included L. O. Howard, W. T.
M. Forbes, J. C. Bradley, C. T. Brues, Miss Edith M. Patch, and
even J. H. Emerton. However, the series of six lectures appears to
have been too long, for only 41 course tickets were sold, more than
half of each audience paying at the door, and attendance varied
greatly from lecture to lecture. The average attendance was smaller
than in 1921 as well, only between 50 and 100 persons. Because of
its enthusiastic advertising campaign, the club went $230 in debt,

24

Psyche

[March

Seal of the Cambridge Entomological Club, designed by A. P. Morse and
adopted by the Club in January, 1922. The White Mountain Butterfly
( Oeneis melissa semidea) is shown resting on rock-fragments, with Mt.
Washington and the rest of the presidential range (New Hampshire) in
the background.

and, having no funds set aside for this, had to solicit for the deficit
from its members. For this reason, the lecture series was discontinued
and never revived.

At the very same meeting at which the Club made the decision to
hold their first public lecture series, they also (upon the motion of
Dr. Wheeler) appointed a committee to look for a design for a Club
seal. Emerton, Wheeler, and Morse were appointed, and for the next
two years they examined and exhibited at meetings the numerous
designs submitted by members. Finally, at the Annual Meeting,
January, 1922, the committee recommended a design by A. P. Morse,
showing the White Mountain Butterfly (then known as Oeneis semi-
dea) , “perched characteristically on the dark grey, deeply weather-
bitten rock-fragments of its mountain home, whose tints and texture
its own so closely resemble, that when lying on its side with wings
closed to escape the wind it becomes almost invisible. Beyond it at
the right is suggested the sedgy slope of ‘Semidea plateau’ (so chris-
tened by Scudder) with its rock-rivulets in whose crannies the but-

1974]

Matthews — Cambridge Entomological Club

25

terfly often seeks shelter from the furious blasts which sweep over
the summits even in midsummer. Beyond, from the depths of the
Great Gulf, rise the slopes of the northern peaks, Mts. Jefferson,
Adams, and Madison, with Mt. Washington suggested at the left.
Over all float the summer clouds which often shroud the summit of
Washington for days at a time even when the other peaks are free”
[29]. The seal was used on the cover of Psyche for the next 37
years, until 1959, and is reproduced on page 24 of the present issue.

During the second decade o'f the Bussey meetings 62 members were
elected — less than in the previous period but a clear indication of
the continuing vitality of the Club. Some of these new members were
to have an active part in the future of the Club. For example,
George C. Wheeler, one of W. M. Wheeler’s students, was elected
in 1920; still an active member of the Club, he is a frequent con-
tributor to Psyche. F. M. Carpenter, a high school senior who had
been attending some of the meetings since 1920 as a guest of F. W.
Dodge, was elected in April, 1922; he became associate editor of
Psyche eight years later and then editor after B rues’ retirement in
1947. Dr. J. C. Bequaert, who had been a guest at several earlier
meetings, was elected to membership in 1923, having been appointed
in medical entomology in the Harvard Medical School ; later, he
succeeded Banks as Curator of Insects at the Museum of Compara-
tive Zoology and participated in the Club meetings until his retire-
ment in 1956. At the meeting of January 8, 1924, P. J. Darlington,
Jr., an undergraduate in the College, was elected to membership; he
also took an active part in the Club’s affairs and followed Dr.
Bequaert as Curator of Insects at the Museum. That particular
meeting, incidentally, was the 50th anniversary of the founding of
the Club, although the minutes refer to it only as the 50th annual
meeting. J. H. Emerton, still regularly attending the meetings, gave
a brief account of the history of the Club from 1874 to 1910 [30]
and W. L. W. Field spoke about the Harris Club and read letters
from two of the original members of the Cambridge Club — B.
Pickman Mann and E. A. Schwarz.

Return to Cambridge

Towards the end of the twenties, plans were made by the Harvard
Administration to terminate the Bussey Institution as a graduate
school and to centralize biological instruction and research in a com-
plex of buildings near the Museum of Comparative Zoology. The

26

Psyche

[March

new building, first named the Biological Institute but later termed
the Biological Laboratories, was to be ready for occupancy in 1931.
Impatient to become established in Cambridge, Professor Wheeler
moved to temporary quarters in the Museum of Comparative Zoology
in 1929, bringing with him the rest of the entomological staff and
their graduate students. At the October meeting, 1929, the members
of the Club voted to hold future meetings in Cambridge and left it
to the executive commitee to find appropriate quarters for the gather-
ing. Since the privilege of smoking was apparently a requirement,
satisfactory housing was found in the basement of the Peabody Mu-
seum, adjoining the Museum of Comparative Zoology. Accordingly,
on November 12, 1929, with President Carpenter in the chair, the
Club met again in Cambridge. As the secretary of the Club, R. P.
Dow, recorded in the minutes: “This was the first meeting of the
Club in Cambridge since March, 1903, when the entire Harris Club
was elected to membership.” For the next two years the Club met
in the Peabody Museum, in an exotic-looking room, decorated with
examples of ancient Indian cultures, until the Biological Institute
was completed. On October 13, 1931, the 498th meeting of the
Club was held in room B-455 of the Biological Institute, with Presi-
dent C. A. Frost in the chair. Since that time, except for a short
interval during World War II, the Club has met in that room,
over a period of 43 years, and the formal address of the building, 16
Divinity Avenue, has been the official address of the Club.

Two months later, December, 1931, a most unusual meeting was
held — the 500th meeting, which apparently holds the record for

Explanation of Photograph on Opposite Page

Some members of the Cambridge Entomological Club in 1929. Front row
(left to right), Charles W. Johnson, Curator at Boston Society of Natural
History; Nathan Banks, Curator of Insects, Museum of Comparative Zool-
ogy; Elizabeth Bryant, Assistant Curator of Insects, M.C.Z. ; J. C. Bequaert,
then Associate Curator, later Curator of Insects, M.C.Z. Back row (left
to right) : Albert P. Morse, Curator of Natural History, at Peabody Museum,
Salem; Arthur Loveridge, then Assistant Curator of Herpetology, M.C.Z.,
and collector of African insects; Charles T. Brues, Associate Curator of
Insects, M.C.Z., and later Professor of Entomology at Harvard; E. T.
Learned, Boston physician and lepidopterist ; Samuel E. Cassino, Salem
publisher (Naturalists’ Directory), engraver, and lepidopterist; Frank M.
Carpenter, then postdoctoral fellow at Harvard and president of the
Cambridge Entomological Club. The photograph was taken in one of the
entomological rooms in the M.C.Z. [Original in archives of Cambridge
Entomological Club.]

1974]

Matthews — Cambridge Entomological Club

27

Some members of the Cambridge Entomological Club, 1929. For explanation, see opposite page.

28

Psyche

[March

length and variety. In recognition of the occasion, a special program
and dinner at the Faculty Club of Harvard University were held for
the 29 members and one guest present. W. L. W. Field was toast-
master of the evening; he read the minutes from the first Club
meeting in 1874, and a letter from Henshaw expressing regret at
being unable to attend. The first after-dinner speaker was Nathan
Banks, who “emphasized the need for specialization and the further-
ance of systematic entomology.” Next, C. W. Collins, of the Parasite
Laboratory in Melrose Highlands, brought the felicitations of the
Bureau of Entomology and traced the. development of economic
entomology during the Club’s existence. C. T. Brues gave a humor-
ous account of “some of the peculiar and amusing incidents” con-
nected with his editorship of Psyche. C. W. Johnson and A. P.
Morse spoke on the history of the Club (with no extant record of
their remarks) . Following these speeches, a motion picture on the
life history of the yellow fever mosquito was shown. And then the
regular scientific program of the evening began, a talk by Dr. Joseph
Bequaert on “Some Problems of Medical Entomology in Guatemala.”
When the 500th meeting finally adjourned, it was 1 1 :00 p.m.

Between 1920 and 1940, there occurred a striking transition in
the composition of the Cambridge Entomological Club. Through the
early twenties, the Club consisted of “not-all-terribly-old amateurs,
more or less old professionals, and a few young people” [31]. But
it was the custom in those days to pass the presidency and vice-presi-
dency of the club around among the senior people, and so rotated that
every few years the same man would be president again. In 1929,
however, the tradition was broken by the election of Frank M.
Carpenter, then a research fellow, to the presidency. His address as
retiring president at the end of the year was published in the next
issue of Psyche [32]. Dr. Carpenter was reelected as president for
the following year, thus once again breaking tradition, for he was
only the third person in the history of the Club to hold two successive
terms of this office.* From this time the Club began to have an
increasing number of younger members, including graduate students,
as its officers.

*The others were Edward Burgess, 1879 and 1880; and J. H. Emerton,
1904 and 1905. In later years there were two other reelections: C. T.
Brues, 1944 and 1945; and E. O. Wilson, 1954 and 1955.

1974]

Matthews — Cambridge Entomological Club

29

World War II

During the thirties, the Club meetings were enthusiastically at-
tended, the average being about 22. But this situation was shortly
to change with entrance of the country into World War II. The
December, 1941, meeting was held only two days after Pearl Harbor.
No recorded mention of the war was made, however, until the fol-
lowing February, when a symposium on Insects and the War was
held at the regular meeting. Discussions were presented by Dr.
Bequaert of the Medical School, Dr. J. E. Gordon of the Harvard
Medical Unit in England, Dr. A. Getty of the Massachusetts
Bureau of Public Health, and Dr. C. B. Philip of the U. S. Public
Health Service. These speakers considered the expected increase in
insect-borne diseases among the public in general, as a result of the
rapid transfer and movements of armed forces.

The Second World War had many more direct effects on the
Club than the First did. By 1943, attendance at meetings was down
to 7 or 8 members and it was apparent that the situation was going to
become worse in the days immediately ahead, as one after another of
the Club’s members left to serve, directly or indirectly, in the war.
In December, 1943, with only Carpenter, George Erickson (the
Club’s secretary), and one other member present, the meeting was
held at the Harvard Faculty Club library, “where they discussed
possible means whereby an active nucleus of the Club might be main-
tained during the War.” For all of 1944 this small group of three
or four met at the library on the second Tuesday of each month to
carry on the Club’s tradition. Some type of program was always
included. At the meeting of February 15, for example, Private Floyd
Werner (elected a member in 1939), on leave from the army, pre-
sented a paper on the “Distribution of Certain Species of the Beetle
Genus EpicautaC In another year, the war was coming to a close
and members began returning; and by the end of 1945 the meetings
reverted to their pre-war time, place and vigor. The “active nucleus’*
had indeed been maintained, and the continuity of the Club insured.

Recent Years

Since 1950 the Club has continued with notable consistency, though
with some obvious and distinctive trends. The attendance at meet-
ings, for example, has been on the increase. The average attendance
for the meetings of the decade 1953-54 to 1962-63 was 29, with the
election of 62 new members; the average for the period from 1963-64
to 1972-73 was 38, with election of 133 new members. Although

30

Psyche

[March

many of the members elected were graduate students living in the
vicinity of Cambridge, there has been a noticeable increase in the
number of amateurs that have joined the Club and that have regu-
larly attended the meetings. From 1903, when the Harris Club
members were elected, through the Bussey meetings in the twenties,
non-professionals made up about half of the membership in attend-
ance. This may have been due in part to W. M. Wheeler’s en-
couragement of amateur naturalists and his open expression of high
regard for their contributions both to research and to the training of
biologists.* However, during the thirties and forties, perhaps as a
consequence of the depression and of World War II, the number of
non-professional members in the Club dropped significantly. In the
course of the past two decades, this trend has been reversed.

Other trends in the nature of the membership during the past two
decades are apparent. For example, more women have joined the
Club and taken part in the programs than previously. In the late
fifties the Club experienced the greatest infusion of feminine talent
in the entire history of its administration, with the election of Ruth
Lippet Willey as secretary (1955) and Margaret C. Parson as secre-
tary (1957), vice-president (1958) and, in 1950, as the first lady
president of the Entomological Club.

Another striking change has been the increase in the number of
arachnologists in the Club. The study of spiders was well repre-
sented in the Club 'from the very beginning by J. H. Emerton, one
of the founders; as shown by the minutes of the meetings, he often
spoke on habits of spiders — flying spiders were his favorite topic —
and he published many papers in Psyche. After his death in 1932,
spiders were almost never mentioned at the meetings until 1956, when
Dr. Herbert Levi was appointed at the Museum of Comparative
Zoology and became a member of the Club. He was promptly joined
by a series of enthusiastic students of spiders (and even of milli-
pedes). Such was their productivity that in 1964 Dr. Carpenter,
while exhibiting the December issue of Psyche , felt compelled to note
“what he interpreted as a hopeful trend in the fact that the number

*In his address, “The Dry-Rot of Our Academic Biology,” at the Boston
meeting of the American Society of Naturalists in 1922 Wheeler advocated
the “utilization by the instructor of competent amateur naturalists as oc-
casional assistants” and he continued: “We have all known amateurs who
could make an enthusiastic naturalist out of an indifferent lad in the course
of an afternoon’s ramble, and, alas, professors who could destroy a dozen
budding naturalists in the course of an hour’s lecture.” [33].

1974]

Matthews — Cambridge Entojnological Club

3i

of pages devoted to insects was greater than the number devoted to
spiders.”

In March of 1965 the meeting commemorated the hundredth an-
niversary of Professor W. M. Wheeler’s birth. Miss Adeline
Wheeler, his daughter, attended the meeting and made available an
exhibit of some of his original drawings. A complete set of his
publications was also shown and several members discussed his con-
tributions to the study of social insects. Three of the members
present, Dr. F. M. Carpenter, Dr. E. O. Wilson and Robert W.
Taylor (president of the Club), represented three successive genera-
tions of ant students that began with Professor Wheeler.

The story of Psyche since 1950 has been one of slow but steady
growth with respect to both its size and the number of subscribers.
With the retirement of Professor Brues as editor in 1947, the Club
elected as the new editor Frank M. Carpenter, who had been asso-
ciate editor of Psyche since 1928. The journal now has the largest
number of subscribers in the Club’s history; 115 of these are Club
members, the rest being libraries or other institutions. The policy
approved by the Club in 1916, i.e., requiring payment by authors for
at least partial cost of printing of their articles, has prevented the
occurrence of the deficits that plagued the journal in its earlier years.
The more general acceptance o'f this policy by educational and other
institutions, including the federal government, has allowed the publi-
cation in Psyche of more articles than would otherwise have been
possible. In the past twenty years the average number of pages per
volume has increased from 130 to 370.

The back issues of Psyche , now comprising 80 volumes, have
created a storage problem. In 1930, when the editorial office moved
to Cambridge, they were stored in the attic of the Museum of Com-
parative Zoology. In 1964 space needs of the Museum staff neces-
sitated the removal of the volumes and on April 14, 1964, the Club
voted approval of a contract for the storage of the back issues with
the Johnson Reprint Corporation in New York, which also was
authorized to serve as the Club’s agent in selling the back issues. In
1972, however, as a result o'f a generally depressed economic situa-
tion, the Reprint Corporation asked that the contract be terminated
and that the volumes be disposed of in some way. Fortunately, very
satisfactory space for them was found in the basement of the Mu-
seum, along with its own publications, and early in 1973 the back
volumes, weighing some 6,000 pounds, were returned to Cambridge
and deposited in the Museum.

32

Psyche

[March

The Centennial of the Founding of the Club

At the May meeting, 1973, the Club voted that the officers for
I973_74 (including the elected members of the Executive Committee
and the members of the Editorial Board of Psyche) constitute a
Centennial Committee, with authority to make arrangements for
appropriate recognition of the 100th anniversary of the Club. In
October President Holldobler reported for the Committee that the
centennial celebration was planned for April 8 and 9, 1974. Pro-
fessor Thomas Eisner of Cornell University, a member of the Club
since 1950, would present a public lecture on the first day, on a
topic of general interest. The entomological and arachnological sec-
tions of the Museum of Comparative Zoology and the Museum
Laboratories would hold open-house on the afternoon of the second
day, followed by a dinner and the Centennial Meeting at the
Harvard Faculty Club. The March issue of Psyche was to be the
Centennial Issue and would include an article on the history of
the Club.

These plans were effectively carried out. Dr. Eisner’s lecture,
entitled On Insects and How They Live as Chemists , was given to
a capacity audience in the main lecture room of the Biological
Laboratories. At the open-house on Tuesday, the new entomological
facilities and equipment in the Museum Laboratories were demon-
strated by Professors Holldobler, Wilson and Levi, and the insect
collections at the Museum were shown by Dr. John Lawrence.
Photographs and documents from the Club’s archives were on display.

The dinner at the Faculty Club was attended by fifty members
and guests; among the guests were Dr. Miriam Rothschild of
England and Dr. Clark A. Elliott of the Harvard University
Archives. Following the dinner, Professor F. M. Carpenter spoke
on the subject, Aspects of the History of the Cambridge Entomologi-
cal Club — Somewhat Anecdotal. He discussed the general status
of entomology and related fields of biology when the Club was begun,
with examples from the Club’s records, and reviewed the achieve-
ments of the thirteen men who founded the Club. In the course of
the past century about 800 individuals have been members of the
society. With a current membership of slightly more than a hundred
members, mostly local, the Club continues to be vigorous and active.
Psyche J with an accumulated pagination of about 20,000 in its eighty
volumes, now has some five hundred individuals and institutions on
its subscription list. In his final remarks, Professor Carpenter em-

1974]

Matthews — Cambridge Entomological Club

33

phasized the close association that has existed between the Cambridge
Entomological Club and the Museum of Comparative Zoology,
beginning with the founding of the society by Professor Hagen and
his associates at the Museum. He stressed the need for continuing
this relationship.

With this Centennial Meeting, the Club began its second century.

References

1. Henshaw, S. 1894. Hermann August Hagen. Proc. Amer. Arts Sci.
29: 419-423.

2. Mayor, A. G. 1924. Samuel Hubbard Scudder. Mem. Nat. Acad. Sci.
17: 81-103.

3. Wheeler, W. M. 1936. Entomology at Harvard University. [In]
Notes Concerning the History and Contents of the Museum of Com-
parative Zoology, pp. 22-32 [Tercentennial of the Founding of Harvard
College].

4. Grote, A. R. 1889. The Rise of Practical Entomology in America.
20th Ann. Report Ent. Soc. Ontario, pp. 75-82.

5. [Anon.] 1926. B. Pickman Mann. Psyche 33 : 172.

6. Cockerell, T. D. A. 1920. Biographical Memoir of Alpheus Spring
Packard, 1839-1905. Nat. Acad. Sci. Biog. Mem. 9: 181-236.

7. Scudder, S. H. 1891. The Services of Edward Burgess to Natural
Science. Proc. Boston Soc. Nat. Hist. 25 : 358-365; also: Anon., 1891.
Edward Burgess. Psyche 6: 131.

8. Banks, N. 1932. J. H. Emerton. Psyche 39: 1-8.

9. Jackson, R. T. 1944. Samuel Henshaw. Proc. Amer. Acad. Arts Sci.
75: 167-169.

10. Emerton, J. H. 1930. George Dimmock. Psyche 37: 299.

11. Mann, B. P. 1885. Herbert Knowles Morrison. Psyche 4: 287.

12. Howard, L. O., H. S. Barber, and A. Busck. 1928. Dr. E. A. Schwarz.
Proc. Ent. Soc. Wash. 30: 154-183.

13. Edwards, H. 1874. A Tribute to the Memory of George Robert Crotch.
Proc. Calif. Acad. Sci. 5: 332-334.

14. Aldrich, J. M. 1906. Baron Osten Sacken. Ent. News 17: 269-272.
Also, C. W. Johnson. Ent. News 17: 273-275.

15. Schwarz, E. A., L. O. Howard and T. N. Gill. 1901. Henry Guernsey
Hubbard. Proc. Ent. Soc. Wash. 4: 350-360.

16. Cambridge Entomological Club, secretary’s report for 1881.

17. Dow, R. 1937. The Scientific Work of Albert Pitts Morse. Psyche 44:
1-11.

18. Banks, N. 1927. The Bowditch Collection of Coleoptera. Ent. News
38: 79.

19. Davis, J. J. 1936. Justus Watson Folsom. Journ. Econ. Ent. 29:
1178-1179.

20. Henshaw, S. 1906. Roland Hayward. Psyche 13: 101-103.

34

Psyche

[March

21. Brues, C. T. 1937. William Morton Wheeler. Psyche 44: 61-96.
Also, M. A. and H. E. Evans, 1970. W. M. Wheeler, Biologist.
Harvard Univ. Press, pp. 1-363.

22. Brues, C. T. 1933. Charles Willison Johnson, 1863-1932. Ent. News
44: 113-116.

23. Melander, A. L. 1955. Charles Thomas Brues. Ann. Ent. Soc. Amer.
48 : 422-423.

24. Wilson, E. O. 1959. William M. Mann. Psyche 66: 55-59.

25. Chapman, J. and E. Chapman. 1947. Escape to the Hills. Cattell
Press, pp. 1-247.

26. Darlington, P. J., Jr. 1933. Percy Gardner Bolster. Psyche 40: 87-88.

27. Johnson, C. W. 1930. Louis William Swett. Psyche 37: 300.

28. Darlington, P. J., Jr. 1963. Charles Albert Frost. Psyche 70: 3-6.

29. Morse, A. P. 1922. The Seal of the Cambridge Entomological Club.
Psyche 29 : 42.

30. Emerton, J. H. 1924. Early History of the Cambridge Entomological
Club. Psyche 31: 1-6.

31. From a transcription of an interview with Dr. P. J. Darlington, Jr.,
Dec. 13, 1968.

32. Carpenter, F. M. 1930. A Review of our Present Knowledge of the
Geological History of the Insects. Psyche 37: 15-34.

33. Wheeler, W. M. 1922. The Dry-Rot of our Academic Biology. Ad-
dress of the Retiring President of the American Society of Naturalists,
December 29, 1922. Published in Science, 1923, 57: 61-71.

1974]

Matthews — Cambridge Entomological Club

35

OFFICERS OF THE CAMBRIDGE ENTOMOLOGICAL CLUB

Date of

Election

President

Vice-President

Secretary

Treasurer

Jan.

1874

B.P. Mann

Jan.

1875

B.P. Mann

Jan.

1876

B.P. Mann

Feb.

1877

S.H. Scudder

B.P. Mann

B.P. Mann

Jan.

1878

E.P. Austin

B.P. Mann

B.P. Mann

Jan.

1879

E. Burgess

B.P. Mann

B.P. Mann

Jan.

1880

E. Burgess

B.P. Mann

B.P. Mann

Jan.

1881

E.L. Mark

B.P. Mann

B.P. Mann

Jan.

1882

S.H. Scudder

W. Trelease

W. Trelease

Jan.

1883

B.P. Mann

G. Dimmock

S. Henshaw

Jan.

1884

S.H. Scudder

G. Dimmock

B.P. Mann

Jan.

1885*

S.H. Scudder

G. Dimmock

B.P. Mann

Jan.

1886

S.A. Forbes

R. Hayward

B.P. Mann

Jan.

1887

J.H. Emerton

R. Hayward

B.P. Mann

Jan.

1888

W. Trelease

R. Hayward

S. Henshaw

Tan.

1889

S.H. Scudder

R. Hayward

S. Henshaw

May 1890

C.W. Woodworth

R. Hayward

S. Henshaw

Jan.

1891

G.H. Snow

R. Hayward

S. Henshaw

Jan.

1892

W.J. Holland

R. Hayward

S. Henshaw

Jan.

1893

W.H. Ashmead

R. Hayward

S. Henshaw

Jan.

1894

T.E. Beau

R. Hayward

S. Henshaw

Jan.

1895

C.M. Weed

R. Hayward

S. Henshaw

Jan.

1896

H.S. Pratt

R. Hayward

S. Henshaw

Jan.

1897

H.G. Dyar

R. Hayward

S. Henshaw

Jan.

1898

T.E. Beau

R. Hayward

S. Henshaw

Jan.

1899

A.G. Mayer

R. Hayward

S. Henshaw

Jan.

1900

J.W. Folsom

R. Hayward

S. Henshaw

Jan.

1901*

J.W. Folsom

R. Hayward

R. Hayward

Jan.

1902*

J.W. Folsom

R. Hayward

R. Hayward

Apr.

1903

A.P. Morse

W.L.W. Field

R. Hayward

Jan.

1904

J.H. Emerton

W.L.W. Field

R. Hayward

Jan.

1905

J.H. Emerton

J.W. Dow

R. Hayward

Jan.

1906

W.L.W. Field

A.H. Clark

R. Hayward

Jan.

1907

H.H. Newcomb

C.A. Frost

F.C. Bowditch

Jan.

1908

C.W. Johnson

C.A. Frost

F.A. Sherriff

Jan.

1909

P.G. Bolster

W.M. Wheeler

C.A. Frost

F.A. Sherriff

Jan.

1910

W.M. Wheeler

W.L. Fiske

C.A. Frost

F.A. Sherriff

Jan.

1911

L.W. Swett

W. Reiff

C.A. Frost

F.A. Sherriff

Jan.

1912

C.T. Brues

C.A. Frost

W.M. Mann

F.A. Sherriff

Jan.

1913

P.G. Bolster

J.W. Chapman

W.M. Mann

F.W. Dodge

Jan.

1914

A.P. Morse

J.W. Chapman

W.M. Mann

F.W. Dodge

Jan.

1915

C.W. Johnson

T. Barbour

H.M. Parshley

F.W. Dodge

Jan.

1916

A.F. Burgess

F.G. Carnochan

H.M. Parshley

F.W. Dodge

Jan.

1917

F.G. Carnochan

S.W. Denton

H.M. Parshley

C.A. Frost

*No elections at the annual meeting; officers held over.

36

Psyche

[March

Date of

Election

President

Vice-President

Secretary

Treasurer

Jan. 1918

W.M. Wheeler

S.W. Denton

A.C. Kinsey

H.A. Preston

Jan. 1919

S.W. Denton

C.A. Frost

A.C. Kinsey

L.R. Reynolds

Jan. 1920

C.A. Frost

W.L.W. Field

J.H. Emerton

F.H. Walker

Jan. 1921

N. Banks

L.R. Reynolds

J.H. Emerton

F.H. Walker

Jan. 1922

W.M. Wheeler

L.R. Reynolds

J.H. Emerton

F.H. Walker

Jan. 1923

A.P. Morse

R.H. Howe

J.H. Emerton

F.H. Walker

Jan. 1924

C.T. Brues

R.H. Howe

J.H. Emerton

F.H. Walker

Jan. 1925

J.H. Emerton

C.W. Johnson

J.C. Bequaert

F.H. Walker

Jan. 1926

W.L.W. Field

O.E. Plath

J.C. Bequaert

F.H. Walker

Jan. 1927

O.E. Plath

S.M. Dohanian

F.M. Carpenter

F.H. Walker

Jan. 1928

J.C. Bequaert

F.M. Carpenter

J.W. Wilson

F.H. Walker

Feb. 1929

F.M. Carpenter

C.W. Collins

J.W. Wilson

F.H. Walker

Feb. 1930

F.M. Carpenter

C.W. Collins

R.P. Dow

F.H. Walker

Jan. 1931

C.A. Frost

C.W. Collins

P.J. Darlington, J

r. F.H. Walker

May 1932

C.W. Collins

C.W. Johnson

J.W. Johnston

F.H. Walker

May 1933

A.P. Morse

P.J. Darlington, J

r. M. Bates

F.H. Walker

May 1934

P.J. Darlington, Jr.

O.E. Plath

R.P. Dow

F.H. Walker

May 1935

J.C. Bequaert

O.E. Plath

D. Davenport

F.H. Walker

May 1936

O.E. Plath

F.M. Carpenter

D. Davenport

F.H. Walker

May 1937

C.A. Frost

J.C. Bequaert

V.G. Dethier

R.P. Dow

Oct. 1938

W.S. Creighton

C.H. Blake

V.G. Dethier

R.P. Dow

May 1939

C.H. Blake

F.M. Carpenter

C.T. Parsons

R.P. Dow

May 1940

F.M. Carpenter

P.J. Darlington, J

r. R.P. Holdsworth

R.T. Holway

May 1941

P.J. Darlington, Jr.

J.C. Bequaert

C.M. Williams

R.T. Holway

May 1942

J.C. Bequaert

L.G. Wesson

G. Erikson

T. Rhyder

May 1943

N. Banks

C.T. Brues

G. Erikson

J.C. Bequaert

May 1944

C.T. Brues

K. Arbuthnot

G. Erikson

F.M. Carpenter

May 1945

C.T. Brues

K. Arbuthnot

G. Erikson

F.M. Carpenter

May 1946

P.J. Darlington, Jr.

J.C. Bequaert

N. Bailey

F.M. Carpenter

May 1947

G.A. Edwards

C.L. Remington

N. Bailey

F.M. Carpenter

May 1948

N. Bailey

W.L. Brown, Jr.

F.G. Werner

F.M. Carpenter

May 1949

F.G. Werner

W.L. Nutting

J. Woodland

F.M. Carpenter

May 1950

K. Christensen

A.G. Humes

R.J. Goss

F.M. Carpenter

May 1951

A.G. Humes

F.Y. Cheng

T. Eisner

F.M. Carpenter

May 1952

T. Eisner

W.L. Brown, Jr.

P.A. Adams

F.M. Carpenter

May 1953

W.L. Brown, Jr.

S.K. Harris

E.O. Wilson

F.M. Carpenter

May 1954

E.O. Wilson

T. Eisner

N.W. Gillham

F.M. Carpenter

May 1955

E.O. Wilson

P.A. Adams

R.H. Lippitt

F.M. Carpenter

May 1956

P.A. Adams

B.R. Headstrom

R.B. Willey

F.M. Carpenter

May 1957

R.B. Willey

S. Duncan

M.C. Parsons

F.M. Carpenter

May 1958

N. Gilham

M.C. Parsons

A.M. Stuart

F.M. Carpenter

May 1959

M.C. Parsons

A.L. Bull

A.M. Stuart

F.M. Carpenter

May I960

H.W. Levi

N. Bailey

G.L. Bush

F.M. Carpenter

May 1961

J.J.T. Evans

C. Walcott

A.R. Brady

F.M. Carpenter

1974] Matthews — Cambridge Entomological Club 37

Date of
Election

President

Vice-President

Secretary

Treasurer

May 1962

L.M. Roth

A.R. Brady

E.G. MacLeod

F.M. Carpenter

May 1963

E.G. MacLeod

J. Beatty

J. Reiskind

F.M. Carpenter

May 1964

R.W. Taylor

J. Reiskind

H. Reichardt

F.M. Carpenter

May 1965

H. Reichardt

S. Vogel

C.C. Porter

F.M. Carpenter

May 1966

J. Reiskind

C.C. Porter

F.C. Coyle

F.M. Carpenter

May 1967

F.C. Coyle

R.W. Matthews

L. Pinter

F.M. Carpenter

May 1968

W.G. Eberhard

C.F. Moxey

S.B. Peck

F.M. Carpenter

May 1969

S.B. Peck

R.E. Silberglied

W. Shear

F.M. Carpenter

May 1970

R.E. Silberglied

C.F. Moxey

C.S. Henry

F.M. Carpenter

May 1971

C.S. Henry

T.P. Webster III

H.F. Nijhout

F.M. Carpenter

May 1972

F. Nijhout

T.H. Hlavac

R. Swain

F.M. Carpenter

May 1973

B. Holldobler

W.D. Winter

H.E. Nipson

F.M. Carpenter

EDITORS OF PSYCHE

Volume

Years

1

1874-76

B. Pickman Mann

2

1877-79

George Dimmock and B. P. Mann

3

1880-82

George Dimmock and B. P. Mann

4

1883-87

George Dimmock and B. P. Mann

5

1888-90

George Dimmock and Samuel Henshaw7

6

1891-93

Samuel Henshaw

7

1894-96

Samuel Henshaw

8

1897-99

Samuel Henshaw

9

1900-02

Samuel Henshaw

10

1903

Samuel Henshaw

11

1904

W. L. W. Field

12

1905

W. L. W. Field

13

1906

W. L. W. Field

14

1907

W. L. W. Field

15

1908

W. L. W. Field

16

1909

W. L. W. Field

17

1910

Charles T. Brues

i

i

53

1946

Charles T. Brues

54

1947

Frank M. Carpenter

i

81

Present

Frank M. Carpenter

SUPPLEMENTARY STUDIES ON ANT
LARVAE: TERATOMYRMEX1

By George C. Wheeler and Jeanette Wheeler
Laboratory of Desert Biology
Desert Research Institute
University of Nevada System
Reno 89507

The first collector of this Australian genus, T. Greaves, Esq. (in
1942), must have been surprised when he first viewed his specimens
under magnification ; he probably thought he had collected diseased
ants. The describer (J. J. McAreavey 1957) was evidently some-
what astonished for he gave it the name T eratomyrmex , from the
Greek teras ( teratos ) monster, marvel, wonder + myrinex ant.
We have in English teratism the worship of monsters, teratology
the study of monstrosities and teratoma a tumor derived from more
than one embryonic layer and made up of a mixture of tissues.
T eratomyrmex is certainly an appropriate name, for the worker
looks as if it has a huge tumor on the top of its thorax (see Fig. 7
and 8). Aside from this unique peculiarity the genus is quite ordi-
nary and belongs in the Formicini, an ordinary tribe of Formicinae,
which includes such anatomically commonplace genera as Acantho-
myops, Lasius and Formica.

The first specimens of T eratomyrmex we have seen were a gift
from Rev. B. B. Lowery (of Norwood, South Australia) in 1967.
In the accompanying letter he wrote: “I have also taken the liberty
of filling up the vacant spaces in the box with a few specially chosen
mountings for your collection. Make sure you have a look at the
T eratomyrmex and Epopostruma frosti under the microscope. Both
these species, by the way, are very rare.” In a later note he wrote:
“The species [T. greavesi] appears to be rare in the ranges near
Murwillumbah NE. NSW. Forages on low shrubs and in leaf litter.
Nests in very moist places, usually in white-rotten timber. It is
very easily mistaken for a small black Iridomyrmex.” He also in-
cluded a quotation from a personal communication he had received
from Tom Greaves: “ ‘T. greavesi is a dominant ant in residual
forest,’ i.e., in the Jackall Ranges, about 75 miles north of Brisbane.”

^ymenoptera : Formicidae: Formicinae.

Manuscript received by the editor December 10, 1973

38

1974]

Wheeler & Wheeler — Teratomyrmex

39

Recently Dr. W. L. Brown sent us a worker of T eratomyrmex
(see Fig. 7 and 8) together with several larvae. Given such a weird
worker could one reasonably expect its larva to be somewhat bizarre
also? We hoped it would. And we were disappointed: the larva is
quite ordinary. Its profile is pogonomyrmecoid, a character shared
with 19 other genera in the Formicinae; its mandibles are campono-
toid in common with 23 other formicine genera. In fact, in our key
to the larvae of the Formicidae (1974), in which most ant larvae
were differentiated to genera or at least to tribes, we could only add
T eratomyrmex to the residual lump of 22 undifferentiated genera:
“Tribes Formicini, Gesomyrmecini, Gigantiopini, Melophorini and
Plagiolepidini.”

The index of specialization (see our 1974) for T eratomyrmex is
14; that of the tribe Formicini is 14. [The most specialized ant
larvae — the Leptanillinae — have an index of 35, while the Formi-
cinae as a whole are less specialized with 17. The index for the
family as a whole is 22.]

Genus Teratomyrmex McAreavey

Body pogonomyrmecoid. Entire integument spinulose. Body hairs
long and moderately numerous; of 3 types: denticulate; long, smooth
and flexuous; 2- or 3-branched, smooth with the branches long and
flexuous. Head hairs moderately numerous; unbranched, denticulate
throughout most of length. Labrum deeply bilobed. Mandibles
camponotoid. Maxillae with paraboloidal apex; palp a paxilla; galeae
digitiform. Labrum subrectangular in anterior view; palp a short
paxilla ; opening of sericteries wide and salient.

Teratomyrmex sp. (near greavesi McAreavey) Fig. 1-6. Length
(through spiracles) about 3.7 mm. Shape pogonomyrmecoid (i.e.,
diameter greatest near the middle of the abdomen, decreasing gradu-
ally toward head and more rapidly toward posterior end, which is
rounded ; thorax more slender and forming a rather stout neck which
is curved ventrally) but abdomen rather slender. Anus posteroventral
and with 2 small lips. Wing and gonopod vestiges present. Ten pairs
of spiracles. Entire integument spinulose, the spinules most promi-
nent on the venter of T1-3 and AI-AIII, sparse and scattered else-
where. Body hairs moderately numerous and long. Of 3 types:
(1) 0.075-0.15 mm long, unbranched and with numerous short
denticles, longest and most numerous on the thorax; (2) 0.2-0.33 mm
long, unbranched, smooth, very long, slender and flexuous, none on

Psyche

[March

40

Textfigure 1-6, larva. 1, head in anterior view, X71; 2, left mandible in
anterior view, X145; 3-5, three types of body hairs X267 ; 6, larva in side
view, X 14.

Textfigure 7 and 8, worker. 7, head and dorsum of pronotum in anterior
view, X34; 8, head, thorax and petiole in dorsal view, X34.

1974] Wheeler & Wheeler — Teratomyrmex 41

Ti, more numerous on abdomen; (3) 0.09-0.15 mm long, 2- or 3-
branched, with long slender tips, on all somites. Cranium suboctag-
onal; broader than long. Antennae small, each a slight elevation
with 3 sensilla, each of which bears a small spinule. Head hairs
moderately numerous, rather long (0.05-0.09 mm) and rather stout,
unbranched and with numerous fine denticles throughout length.
Mouth parts large. Labrum bilobed; each lobe with 2 hairs about
0.05 mm long with small denticles and with 5 sensilla on and near
the ventral border; posterior surface spinulose, the spinules in short
to long rows, the rows radiating from each dorsolateral angle, and
with 5 sensilla on each half near the midline. Mandibles camponotoid
(i.e., subtriangular ; base broad, width 2/3 length; apex forming a
round-pointed tooth; one small subapical tooth); anterior and pos-
terior surfaces with numerous short longitudinal rows of minute
spinules. Maxillae with the apex paraboloidal ; palp a paxilla with
5 (3 apical and bearing a spinule each, 1 subapical and encapsulated,
1 lateral and bearing a spinule) sensilla; galea digitiform, with 2
apical sensilla. Labium subrectangular in anterior view; anterior
surface with minute spinules in short rows; palp a short peg with 5
(3 apical and bearing a spinule each and 2 lateral — 1 encapsulated
and 1 bearing a spinule) sensilla; opening of sericteries wide and
salient, with 2 ventral projections. Hypopharynx densely spinulose,
the spinules minute and arranged in numerous subparallel rows; the
rows grouped in 2 subtriangles which have their bases near the
middle.

Material studied: 14 larvae from Queensland: Cedar Creek Falls,
Tamborine Mts., Ross & Cavagnaro, 1962; courtesy of Dr. W. L.
Brown.

Literature Cited

McAreavey, J. J.

1957. [Teratomyrmex gen. nov.] Mem. Nat. Mus. Viet. No. 21, p. 54-56.
Wheeler, G. C., and Jeanette Wheeler.

1974. Ant larvae: review and synthesis. Mem. Entom. Soc. Washing-
ton [in press].

MODIFICATION OF THE
INTERSEGMENTAL REGION IN THE
PTEROTHORAX OF CRYPHOCRICOS
(HETEROPTERA: NAUCORIDAE ) *

By Margaret C. Parsons
Department of Zoology, University of Toronto

Toronto, Ontario M5S 1A1

Introduction

In most Hydrocorisae (aquatic Heteroptera) the hindlegs are used
for swimming, and their extrinsic muscles, which originate in the
metathorax, are well developed. The tergal and pleural depressors
of the metathoracic trochanter (Muscles 70 and 71, respectively, of
Larsen, 1945) are especially large in many Hydrocorisae, including
the Naucoridae (Larsen, 1945) and the closely related family
Aphelocheiridae (Parsons, 1969). These muscles slant anterolaterally
in the metathorax, their anteriormost fibers originating near its
boundary with the mesothorax.

In at least three genera of Naucoridae ( Limnocoris , Ambrysus,
and Cryphocricos) the anterior part of the metathoracic episternum,
on which Muscle 71 originates, usually projects into the cavity of
the preceding segment. This makes it appear, at first glance, that
the anteriormost fibers of the muscle attach on the mesothorax. The
metapleural projection, which will be described in a later publication,
appears to be produced by post-ecdysial growth of the skeleton, since
it is absent in newly-moulted adults and is present only in older ones.

Another example of post-ecdysial skeletal change occurs in the
tergum of adult Cryphocricos barozzii and makes the anteriormost
fibers of Muscle 70 appear to originate on the mesothorax rather
than on the metathorax. This tergal modification seems to occur only
in micropterous specimens, with reduced forewings. I have not ob-
served it in macropterous Cryphocricos or in Ambrysus , Limnocoris ,
or Pelocoris, all of which possess forewings of normal length.

Materials and Methods

Cryphocricos barozzii Signoret, collected in Nova Teutonia, Brazil,
and preserved in 70% alcohol, were dissected in 80% alcohol under

* Manuscript received by the editor January 20, 1974.

42

1974]

Parsons — Pterothorax of Cryphocricos

43

a stereoscopic microscope. The attachment of Muscle 70 was studied
by cutting the tergum sagittally, with a razor blade; the cut Was
made half way between the ventral phragmal process and the base of
the mesothoracic wing. A few Pelocoris femoratus Palisot-Beauvois,
preserved in Bouin’s fluid, were also examined for comparison.

Most of the Cryphocricos were micropterous, lacking hindwings
and indirect flight muscles and possessing forewings which reached
only as far as the third abdominal segment. Only two of the rare
macropterous forms, with forewings and hindwings extending nearly
to the tip of the abdomen, were available for the investigation. Al-
though they were not extensively dissected, their indirect flight mus-
cles appeared to be either degenerate or absent.

Newly-moulted Cryphocricos could be distinguished from older
individuals by the thinness of their exoskeletons and by the distinct-
ness and relative thickness of the underlying epithelial layer. In
older specimens the epithelium was thinner, less distinct, and easily
torn, and the thickened exoskeleton had a layered appearance, prob-
ably owing to the deposition of successive internal layers of endo-
cuticle after ecdysis, as Neville (1970) has observed in several in-
sects. These layers could be peeled away from each other in an older
specimen of Cryphocricos which had been immersed for 24 hours in
a concentrated solution of potassium hydroxide.

Observations
Typical Naucoridae
(figs. 1, 5)

In most macropterous naucorids, such as Pelocoris, the posterior
margin of the mesothoracic notum (fig. 1 ; N II) is evaginated, form-
ing a double-walled scutellar lobe (SL). The opening into the lobe
(OSL) extends anterolaterally to the base of the forewing (WB).
Immediately ventral to this opening lies the well developed meso-
thoracic postnotum (PN), which bears the second phragma (PH).
The postnotum separates the metathoracic notum (N III) from the
scutellar lobe and is mostly concealed by the latter externally. Only
its most lateral part, which joins the mesothoracic epimeron (EM),
forming a postalar bridge, is externally visible. The postalar bridge
bears a large sensory membrane (SO), part of the mesothoracic
scolopophorous organ (Larsen, 1957).

The concealed medial portion of the postnotum forms a two-
walled, vertical invagination, the second phragma (PH). The

44

[March

Psyche

Figures 1-4, Diagrammatic internal views of dorsolateral intersegmental
region of pterothorax in various adult Naucoridae. Left side of body has
been cut parasagittally through terga (cut edge), between ventral phragmal
process (not shown) and wingbase (WB). Fig. 1. Typical macropterous
Naucoridae (based on Pelocoris femoratus) . Fig. 2. Macropterous Crypho-
cricos barozzii. Fig. 3. Newly-moulted micropterous C. barozzii. Fig. 4.
Older micropterous C. barozzii (dotted line indicates approximate position
of opening into scutellar lobe, which is no longer visible).

1974]

Parsons — Pterothorax of Cryphocricos

45

phragma is especially pronounced on either side of the midline, where
it bears a pair of large ventral processes (fig. 5; VP) upon which
two of the mesothoracic indirect flight muscles attach (Muscles 30
and 31 of Larsen, 1945; present but degenerate in most Pelocoris) .

According to Snodgrass (1935) the phragma represents the pri-
mary boundary (antecosta) between the metathorax and mesothorax,
and the mesothoracic postnotum is thus an intersegmental plate which
is only secondarily associated with the mesothorax. This interpreta-
tion, which has been challenged by Matsuda (1970), is supported by
the fact that the posterior wall of the phragma is continuous with
the metathoracic episternum (fig. 1 ; ES).

Muscle 70 (fig. 1 ; M. 70) originates posterior to the phragma
and postnotum, its fibers attaching only on the metathoracic notum.
The latter forms a sharp angle with the posterior wall of the
phragma. The angle is sclerotized medially (fig. 1) but contains a
narrow membrane laterally (fig. 5; MEM). The membrane lies
immediately anterior to the origin of Muscle 70.

Macropterous Cryphocricos

(fig- 2)

Most of the typical structural relationships described above are
also present in macropterous Cryphocricos. In the latter, however,
the postnotum forms a definite, double-walled second phragma only
near the midline, at and between the ventral processes (position same
as in micropterous Cryphocricos , figs. 6, 7). Lateral to the ventral
processes the postnotum does not seem to be invaginated, and appears
merely as a transverse thickening (fig. 2, PN) just posteroventral to
the opening into the scutellar lobe (OSL). Neither of the two
available macropterous specimens were newly-moulted. It is thus
possible that a low, double-walled phragmal invagination is present
in this region immediately after ecdysis. As new layers of endocuticle
are subsequently laid down, the structure of the phragma could
become obscured, giving it the appearance of an uninvaginated thick-
ening in older specimens.

Laterally the anterior portion of the postnotal thickening forms a
well-defined postalar bridge with the mesothoracic epimeron (EM).
The bridge contains a large sensory membrane (SO) similar to that
of typical naucorids. The posterior portion of the postnotal thicken-
ing is continuous laterally with the metathoracic episternum (ES)
and medially with the metanotum (N III). Its boundary with the

1974]

Parsons — Pterothorax of Cryphocricos

47

latter is marked dorsolaterally by a narrow, unpigmented band of
sclerotization, the homologue of the membrane (fig. 5; MEM)
which is present in typical Naucoridae. Muscle 70 originates only on
the metanotum, posterior to this band, as in other macropterous
nau corids, and its fibers do not extend onto the postnotum (fig. 2).

Micropterous Cryphocricos
(figs. 3, 4, 6, 7)

In short-winged Cryphocricos as in macropterous ones, a well-
defined second phragma (figs. 6, 7; PH) is present only at and
between the ventral phragmal processes (VP). In addition, the
mesothoracic postnotum appears to be incomplete in micropterous
Cryphocricos. These skeletal modifications are more easily observed
in newly-moulted adults (figs. 3, 6) than in older ones in which parts
of the thickened exoskeleton have coalesced (figs. 4, 7).

In newly-moulted specimens the portion of the postnotum which
usually lies immediately lateral to the ventral phragmal processes
appears to have disappeared. The scutellar lobe (figs. 3, 6; SL) is
shorter than in macropterous Cryphocricos , and its ventral wall is
directly continuous medially with the metanotum (N III), upon
which Muscle 70 originates. The junction between these two regions
forms a fold (F) which projects anteriorly and is continuous with
the phragmal process (fig. 6). Short, sclerotized struts connect its
edge secondarily with the mesonotum (N II). The anteromedial
fibers of Muscle 70 originate on the edge of this fold, immediately
beneath the opening into the scutellar lobe (fig. 3; OSL). They thus
attach farther anteriorly than in macropterous Cryphocricos (fig. 2),
in which they originate well posterior to the opening.

The more anterolateral fibers of Muscle 70 do not attach to the
edge of the fold but are separated from it by a roughly triangular
area which represents the lateral portion of the incomplete postnotum
(fig. 3; PN). There is no membrane or unpigmented band marking
its posterior boundary with the metanotum. It is ventrally continu-

Figures 5-7, Diagrammatic internal views of intersegmental part of
pterothoracic terga in various adult Naucoridae. Terga have been cut
parasagittally, medial to wingbases, and removed from body; only left
side, between ventral phragmal process (VP) and cut edge, is shown.
Arrow indicates level at which anteriormost fibers of Muscle 70 originate.
Fig. 5. Pelocoris femoratus (ventral phragmal process cut off at base).
Fig. 6. Newly-moulted micropterous Cryphocricos barozzii. Fig. 7. Older
micropterous C. barozzii.

48

Psyche

[March

ous with the mesothoracic epimeron (EM), and the postalar bridge
contains a very reduced scolopophorous organ (SO).

In older micropterous Cryphocricos skeletal boundaries are much
more difficult to interpret, owing to the thickening of the cuticle, the
differential growth of some regions, and the obliteration of folds,
making them appear as solid thickenings rather than hollow invagina-
tions or evaginations. Comparison of older specimens (fig. 4) with
newly-moulted ones (fig. 3) indicates that during post-ecdysial de-
velopment several changes have occurred in the lateral and dorso-
lateral intersegmental region of the pterothorax. The scolopophorous
sense organ is no longer visible on the internal surface of the postalar
bridge. The cavity of the scutellar lobe (fig. 4; SL) has been
reduced, and the opening into the lobe (fig. 4; dotted line) is no
longer visible. The ventral wall of the lobe merges indistinguishably
with the fold which borders on it ventrally. In newly-moulted
Cryphocricos this well-defined fold (fig. 3; F) marks the boundary
of the scutellar lobe with the metanotum medially (fig. 3; cut edge)
and with the postnotum laterally (PN). In older adults these
boundaries have become obscured. The fold and scutellar lobe appear
as three separate layers only posteriorly; more anteriorly they appear
as a single, somewhat thickened layer (fig. 4).

In addition, the anteromedial fibers of Muscle 70, which origi-
nate on the metanotum, attach considerably more anteriorly in older
Cryphocricos than in younger ones (figs. 3, 4; M. 70; figs. 6, 7;
arrows). This suggests that the fold (figs. 3, 6; F) between the
scutellar lobe and the metanotum grows anteriorly as new layers of
endocuticle are added to the internal surface of the skeleton. One
older specimen, in which the muscle attached at the level indicated
by the arrow in Figure 7, was treated in potassium hydroxide so that
the more internal layers of endocuticle could be peeled away from
the outermost ones. Although the inner layers lacked a fold, the
more external ones showed a definite one which lay posterior to the
level at which the muscle fibers had originated.

Discussion

In these descriptions I have taken the view that Muscle 70 origi-
nates on the metanotum in all four types of Naucoridae. In the two
macropterous forms (figs. 1,2) its origin is separated from the meso-
scutellar lobe by the postnotum. In micropterous Cryphocricos (figs.
3, 4), however, a portion of the postnotum, immediately lateral to the

1974]

Parsons — Pterothorax of Cryphocricos

49

ventral phragmal process, is absent. In this region, consequently, the
part of the metanotum on which Muscle 70 originates borders
directly on the scutellar lobe. This atypical intersegmental boundary
appears to grow anteriorly during post-ecdysial development.

Figures 3 and 4 could be interpreted in tWo other ways, either
( 1 ) that there are no post-ecdysial changes in skeletal boundaries
and that in Figure 4 the anteriormost fibers of Muscle 70 have
simply shifted anteriorly, onto the mesonotum, or (2) that a. post-
notum is present lateral to the phragmal process and is represented
by the fold (fig. 3; F) upon which the anteromedial fibers of Mus-
cle 70 attach. According to this second view the origin of the muscle
has shifted from the metanotum onto the homologue of the thickened
postnotum of macropterous Cryphocricos (fig. 2; PN). Conse-
quently, it is the boundary between the mesothoracic scutellum and
postnotum (secondary intersegmental boundary of Snodgrass, 1935)
which appears to grow anteriorly after ecdysis, rather than a primary
intersegmental boundary. Neither of these two interpretations has
been adopted here because in Cryphocricos and at least two other
genera of Naucoridae post-ecdysial skeletal growth takes place in
the pterothoracic pleuron (see p. 46) and almost certainly occurs
along the primary intersegmental boundary between the metathoracic
episternum and the mesothoracic epimeron.

Degeneration or absence of the three mesothoracic indirect flight
muscles, and consequent loss of the ability to fly, is a common oc-
currence among Hydrocorisae (Larsen, 1950). Extreme alary di-
morphism such as that of Cryphocricos is much less common. Larsen
(1950) studied Aphelocheirus ( Aphelocheiridae) which has both
micropterous and macropterous forms, and found that the size and
shape of the ventral phragmal processes, upon which two of the
indirect flight muscles attach, differ in the two types. Similar dif-
ferences in the phragmal processes occur in flying and flightless
Ilyocoris (Naucoridae; Larsen, 1970). Both forms of Ilyocoris
possess forewings and hindwings of normal length, but most in-
dividuals lose their indirect flight muscles soon after the final moult.
Larsen did not, however, observe post-ecdysial skeletal changes, ab-
sence of the lateral part of the second phragma, or loss of a portion
of the mesothoracic postnotum in either Aphelocheirus or Ilyocoris.
All these features appear to occur in micropterous Cryphocricos.

Acknowledgements

I wish to thank Fritz Plaumann (Nova Teutonia, Brazil) for

50

Psyche

[March

collecting the Cryphocricos , John Polhemus (Englewood, Colorado)
for identifying them, my husband, Thomas Parsons, for examining
the manuscript, and Donald Chant and the Department of Zoology,
University of Toronto, for making available the laboratory facilities
for this research. The investigation was made possible by a grant
from the National Research Council of Canada.

Literature Cited

Larsen, O.

1945. Der Thorax der Heteropteren. Skelett und Muskulatur. Lunds
Univ. Arsskr. 41(3): 1-96.

1950. Die Veranderungen im Bau der Heteropteren bei der Reduktion
des Flugapparates. Opusc. Ent. 15: 17-51.

1957. Truncale Scolopalorgane in den pterothorakalen und den beiden
ersten abidominalen Segmenten der aquatilen Heteropteren. Lunds
Univ. Arsskr. 53 (1): 1-66.

1970. The flight organs of Ilyocoris cimicoides L. (Hem., Naucoridae).
Ent. Scand. 1 : 227-235.

Matsuda, R.

1970. Morphology and evolution of the insect thorax. Mem. Amer.
Ent. Inst. 76: 1-431.

Neville, A. C.

1970. Cuticle ultrastructure in relation to the whole insect. In: Insect
Ultrastructure, A. C. Neville (ad.), Sympos. Roy. Ent. Soc.
Lond. 5: 17-40.

Parsons, M. C.

1969. Skeletomusculature of the pterothorax and first abdominal seg-
ment in micropterous Aphelocheirus aestivalis F. (Heteroptera :
Naucoridae). Trans. Roy. Ent. Soc. Lond. 121: 1-39.

Snodgrass, R. E.

1935. Principles of insect morphology. New York: McGraw-Hill.

THE POLYTYPIC GENUS CELOTES
(LEPIDOPTERA: HESPERIIDAE: PYRGINAE)
FROM THE SOUTHWESTERN UNITED STATES
AND NORTHERN MEXICO*

By John M. Burns

Museum of Comparative Zoology, Harvard University

Although most species of American hesperiids are hard to deter-
mine, some are not; and none is more immediately distinct than the
“streaky skipper,” Celotes nessus (Edwards). Ever since it was
described (in genus Pholisora) in 1877, nothing else like it has been
known. Indeed, since the turn of the century — despite abortive
attempts of various workers to jam it into polytypic genera (such as
Pyrgus [then commonly called Hesperia ], Systaseaj and Antigonus)
— it has properly stood alone in the monotypic genus Celotes (God-
man and Salvin 1899). I feel, therefore, a measure of remorse in
now describing a second species of Celotes that closely resembles the
first.

Although I have accumulated data on Celotes since New Year’s
Eve of 1961, when I first recognized it as polytypic, I still see no
clear clues to the evolutionary differentiation of C. nessus and the
species to be described: they overlap fully in space and in time, and
at least partly in choice of larval foodplants. The new species is a
multivoltine mallow-eater occurring in a montane strip that runs
northwest-southeast through the middle of the range of C. nessus.

Because C. nessus is both peculiar and familiar, the following
description is comparative. And because original descriptions can be
too tedious for words, this one is largely visual.

In preparing it, I have examined the genitalia o'f all specimens of
Celotes available to me and possessed of an abdomen — a total of
529 individuals, of which 97 represent the new species. Wherever
possible, I have also examined the metathoracic pouch of males.
Each pinned specimen studied has received a sex-and-determination
label.

The new specific name is a noun in apposition.

Celotes limpia new species

Holotype. — cf, Limpia Canyon, 5000 feet, Davis Mountains,
4 miles WNW of Fort Davis, Jeff Davis County, Texas, May 2,

* Manuscript received by the editor March 1, 1974.

51

52

Psyche

[March

1959 (J- M. and S. N. Burns) [Museum of Comparative Zoology
no. 31888].

Size. — On average, limpia is larger than nessus ; and, in both
species, females are larger than males, although size varies consid-
erably (table 1). In nessus , the wingspread of Texas specimens
exceeds that of Arizona specimens by at least 1 mm.

Table 1. Maximum wingspread in well-mounted specimens of Celotes
from Texas.

Species

Sex

Number

measured

Wingspread

Mean

(mm)

Range

nessus

$

35

23.7

22-26

$

22

25.1

23-27

limpia

$

39

25.9

23-29

9

24

27.8

25-31

Facies. — Fig. iE-H. Very like nessus (fig. iA-D), but with
slightly larger and more conspicuous hyaline spots and a generally
paler aspect.

Noteworthy in the latter connection is a set of pale areas between
dark marks on the proximal 60% of the ventral secondary (i.e. from
the three hyaline spots of the secondary inward) : these pale areas
are enlarged and whitened in limpia and hence usually more pro-
nounced than in nessus. In dorsal view, unworn (particularly
reared) specimens of limpia may show considerable grayish overscaling
on the body and adjacent wing bases. The usually paler shades of
brown and tan in limpia tend to make it reflect more and appear
colder and harder and sometimes more contrasty. All of these
average differences involving color are perceptible chiefly when
limpia is compared with nessus from Texas; nessus from Arizona
(where limpia does not occur) often essentially duplicates the facies
of limpia — except insofar as the remarkably small average size of
Arizona nessus gives its pattern a special sharpness. Wear and fading

Data for specimens in Fig. 1:

A, B — Sitting Bull Falls, 4650 feet, Guadalupe Mountains, Eddy County,
New Mexico, V-26-1959 (J. M. and S. N. Burns).

C, D — Topotype. San Antonio, Bexar County, Texas, VI-30-1963 (J. M.
Burns).

E, F — Holotype. Limpia Canyon, 5000 feet, Davis Mountains, 4 mi. WNW
of Fort Davis, Jeff Davis County, Texas, V-2-1959 (J. M. and S. N.
Burns) .

G, H — Allotype. As for holotype, except V-l-1959.

1974]

Burns — Polytypic Genus Celotes

53

Fig.

1.

Facies of Celotes.

C. nessus

C. limpia

l

dorsal

J A

/ c

f E
( G

B -» $

D -4 $

F -» $

H — > $

i

ventral

54

Psyche

[March

raise additional hell with these subtle color characters, but no matter:
the genitalia (q.v.) last and last.

Male scent-spreading sex characters . — Males of limpia , like
those of nessus, have a narrow costal fold on the primary and a
metathoracic system of ventral pouch and tibial tufts. The meta-
thoracic pouch of limpia (fig. 2B) bears short linear scales over most
or all of its dorsal surface (except at the lateral margin). In nessus
(fig. 2A), linear scales are altogether absent (particularly in Arizona
males) or, if present (as in Texas males), are much less dense and/or
proximal in distribution; even in extreme cases, they never attain
the distal apex of the pouch, and the pouch is therefore clad — en-
tirely or for its greater part — with much broader, flat-lying shingle-
like scales (cf. figs. 2A and 2B). Because the linear scales of limpia
are creamy, whereas some to all of the broad shinglelike scales of
nessus are gray to brown, the general color of the dorsal surface of
the metathoracic pouch differs in the two species. (Interior morphol-
ogy of costal folds has not been studied.)

Fig. 2. Metathoracic pouch of the male; abdomen removed, dorsopos-
terior view. A. Celotes nessus from Austin, Travis County, Texas, V-26-
1966 (J. M. Burns). B. Celotes limpia, a paratype reared out IX-10-1966
from a larva collected on Abutilon malacum, 15 mi. SE Redford, 2500 ft.,
Presidio County, Texas, VIII-18-1966 (R. O. Kendall).

Male genitalia. — Fig. 4. Although basically similar to nessus
(fig. 3), there are abundant differences, of which the most salient
include the following. ( 1 ) In limpia , a long thin flattened projection,
like a curved spatula, arises from the dorsoposterior rim of the body
of the valva and starts dorsad but at once curves caudad, extending
far back to become the caudalmost valval element and to end in a

1974]

Burns — Polytypic Genus Celotes

55

slightly flared tip with a peripheral array of fine sharp teeth (fig.
4A-C). Its homolog in nessus is a robust spike arising in a similar
position, except that the distinctive high dorsal curvature of the body
of the valva in nessus places the origin of the spike well down on the
medial surface of the valva (fig. 3A-C) ; this tapering spike extends
chiefly dorsad, but usually bends slightly caudad at its apex, which
is pointed and often entire or bifurcate (fig. 3A,B), but sometimes
trifurcate, or very rarely quadrifurcate ; although occasionally broad-
ened or lengthened and bent strongly caudad apically, the spike never
approaches the form of the spatula in limpid. (2) Anterior end of
tegumen, in limpid , large and well-developed, projecting far cephalad
(fig. 4E,F) ; but in nessus, extraordinarily reduced (fig. 3E,F).
(3) In limpid, uncus broad, and the paired terminal prongs of the
uncus heavy and stubby (fig. 4F) ; but in nessus, uncus relatively
narrow, and its terminal prongs more delicate (fig. 3F).

Femule genitdlid. — Fig. 6. Sterigma of limpid altogether more
massive (fig. 6A,B) and, viewed ventrally, more nearly square in
outline (fig. 6A) ; heavy central sclerotization in lamella postvagi-
nalis like a wide triangle tapering quickly toward ostium bursae
(fig. 6A). Sterigma of nessus (fig. 5) less massive (fig. 5A,B) and,
in ventral view, narrower posteriorly, suggesting in outline a cau-
dally truncated triangle (fig. 5A) more than a square; heavy central
sclerotization in lamella postvaginalis limited to a comparatively
narrow midventral strip that tapers but little toward ostium bursae
(fig. 5A). Ostium bursae of limpid a relatively narrow curved slit,
like a crescent bowed dorsad (fig. 6C), but that of nessus large and
more or less round, like a manhole (fig. 5C).

Spdtidl distribution. — A widespread species, nessus ranges from
about the 97th meridian in southcentral Oklahoma and central Texas
west to northwestern and southeastern Arizona and south in Mexico
to at least southern Sonora, southern Chihuahua and Coahuila
(fig. 7). By contrast, so far as known, limpid occurs only along the
southern Rocky Mountain axis in Trans-Pecos Texas and along the
Mexican counterpart of this axis, the northern segment of the Sierra
Madre Oriental, as far south at least as southern Coahuila (fig. 8).
Having been taken in numbers in the Guadalupe, Davis, and Chisos
mountains of Trans-Pecos Texas, limpid may confidently be expected
(1) in many other west Texan ranges — such as the Delawares,
Chinatis, and Santiagos — that are more or less associated with the
scattered southern Rocky Mountain system, (2) northward in at
least the New Mexican extension of the Guadalupes, if not in the
Sacramentos and more northern chains, and (3) southward in Chi-

56

Psyche

[March

Fig. 3. Male genitalia of Celotes nessus. A. Lateral view of right valva.
B. Medial view of right valva. C. Dorsal view of both valvae. D. Dorsal
view of aedeagus. E. Left lateral view of uncus, tegumen, gnathos, vin-
culum, and saccus. F. Dorsal view of uncus, tegumen, vinculum, and saccus.
[A and B, drawn from male from Palo Duro Canyon, 2800 feet, Randall
County, Texas, V-10-1959 (J. M. and S. N. Burns) ; C to F, from male from
Austin, Travis County, Texas, V-26-1966 (J. M. Burns).]

1974] Burns — Polytypic Genus Celotes 57

Fig. 4. Male genitalia of Celotes limpia. A to F as in fig. 3. [A and B,
drawn from holotype; C to F, from paratype from type locality, V-4-1959
(J. M. and S. N. Burns).]

58

Psyche

[March

Fig. 5. Female genitalia of Celotes nessus. A. Ventral view of bursa
copulatrix, sterigma, and ovipositor lobes. B. Right lateral view. C. Ven-
troposterior view. [Drawn from female from Austin, Travis County, Texas,
IV-18-1967 (J. M. Burns).]

1974]

Burns — Polytypic Genus Celotes

59

Fig. 6. Female genitalia of Celotes limpia. A. Ventral view of bursa
copulatrix, sterigma, and ovipositor lobes. B. Left lateral view. C. Ventro-
posterior view. [Drawn from paratype from Limpia Canyon, 4700 feet,
Davis Mountains, 5 mi. NE of Fort Davis, Jeff Davis County, Texas, V-4-
1959 (J. M. and S. N. Burns).]

6o

Psyche

[March

Fig. 7. Spatial distribution (based on material examined) of Celotes
nessus — southwestern United States and northern Mexico. Rivers and the
borders of states are shown. Type locality indicated by arrow.

huahua, as well as Coahuila, and, in all likelihood, beyond these
states. Indeed, either limpia or nessus (see N omenclatural epilogue
below) was long ago reported from Durango city (Godman and
Salvin 1899), and both species probably range well down the Central
Plateau of Mexico. The distribution of limpia appears to lie within
that of nessus.

Over its much broader distribution, nessus occurs from sea level
to somewhere between 5000 and 6000 feet; limpia , with its strictly
interior distribution, is now known from an elevational range of
1900-5700 feet. In montane regions of the southwestern United
States, both skippers are species of alluvial fans, foothills, lower
canyon reaches, and lower elevations generally; they avoid higher
elevations.

The distribution of nessus in the United States is not perfectly
known. The skipper may conceivably occur in the New York and
Providence mountain region of southeastern California, in southern
Nevada, in southern Utah, possibly in extreme southwestern or south-
eastern Colorado, and certainly in considerably more of New Mexico

1974]

Burns — Polytypic Genus Celotes

61

Fig. 8. Spatial distribution (based on material examined) of Celotes
limpia — Trans-Pecos Texas. Mountains and the borders of counties are
shown. Type locality indicated by arrow. The range as plotted spans a
north-south distance of about 220 miles; but, since preparing this map, I
have examined one male from southern Coahuila, Mexico, about 250 miles
southeast of the Big Bend.

than fig. 7 indicates. On the other hand, this map probably gives a
close approximation of the eastern distributional limit of nessus.
This meridional eastern limit coincides neatly with that of many other
organisms — including other pyrgine skippers of the southwestern
United States and northern Mexico such as Pyrgus philetas Edwards
(Burns and Kendall 1969) and Erynnis meridianus Bell (Burns
1964, and unpublished) — and reflects an important biogeographic
barrier.

There are two freaky records of streaky nessus east of the 97th
meridian. The first [Burton, Georgia, V-21-1911 (J. C. Bradley)
(Cornell University collection)] was published by Harris (1950)
and repeated [as “Georgia (once!)”] by Klots (1951), but was not
even mentioned later by Harris (1972) and should be discounted.
The second [Tallulah, Louisiana, between VIII-1926 and X-1931
(collected in the day at an altitude of 20 feet by an airplane)] was
published by Glick (1939) ; but though it has been repeated (Lam-
bremont 1954; Mather and Mather 1958), the feat itself has not.

62

Psyche

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Temporal distribution. — Both limpia and nessus are multivoltine,
and both fly at the same times. Nearly all of the dated wild-caught
adults of Celotes that I examined were collected from March to
September: in Texas, every half-month interval from the beginning of
March to mid-September includes records of both limpia (N = 68)
and nessus (N = 1 58 ) ; however, for Arizona nessus (N = 183),
the months of May and June together yield but 3 records, which
leaves a major gap between one large cluster of records in March
and April and another in July, August, and the first half of Septem-
ber. The only other dated wild-caught specimens examined are 5
nessus from far southern Arizona collected in mid-January and
5 nessus from the last half of September and the first half of October
from central and, as might be expected, extreme southern Texas (the
lower Rio Grande Valley). In the experience of Kendall (1965),
nessus flies from early March to mid-November in Texas.

The fact that temporal distribution is essentially continuous
through the warm season for both limpia and nessus in Texas but
distinctly bimodal for nessus in Arizona probably relates to the dif-
ferent patterns of rainfall in the two states. For the most part,
Texas has much more evenly distributed precipitation than does
Arizona, where it occurs in two widely disjunct winter and summer
periods. The second flight period of nessus in Arizona coincides with
the summer rainy season, to which it is presumably geared. In the
course of his field studies and laboratory rearings, Kendall (1965,
and unpublished) has observed that full-grown larvae of Celotes are
able to enter diapause (facultatively) to beat heat and drought, as
well as to get through the cold of a winter season. Kendall has
shown in the laboratory that such diapause can be broken by pro-
viding moisture, whereupon larvae pupate and produce adults
promptly. Given this capacity to be flexible, variable larval growth
rates, and the usual vicissitudes of weather, individuals of a single
generation must often get out of synchrony with the result that gen-
erations overlap broadly and irregularly, forming no definite number
per year in either species.

There are no indications that limpia and nessus are temporally
displaced with respect to each other where they spatially coexist in
Trans-Pecos Texas. Rather, my records show that adults of both
species have repeatedly been collected in association — unwittingly,
of course, since the collectors were unaware of the existence of more
than one species of Celotes. On nine occasions altogether, five dif-
ferent collectors (or pairs of collectors) have caught a total of 37

1974]

Burns — Polytypic Genus Celotes

63

limpia and nessus at the same place on exactly the same date — the
places occurring in Jeff Davis and Brewster counties and the dates
in March, May, June, July, and August. Many other records of
both species at a single locality differ by only one to a few days.

Larval foodplants. — Numerous studies in Texas by R. O. and
C. A. Kendall show that both limpia and nessus feed on various
species of Malvaceae. For both skippers, the Kendalls have witnessed
oviposition — • directly on larval foodplants — in the wild, and have
collected numerous eggs and larvae from which they have reared
adults.

In the field, limpia eats Abutilon malacum Wats., A . incanum
(Link) Sweet, Sphaeralcea angustifolia var. lobata (Woot.) Kern.,
and Wissadula holosericea (Scheele) Gke., and, in the laboratory,
also Malvastrum americanum (L.) Torr. and Althaea rosea Cav.
In the field, nessus eats Abutilon incanum, Sphaeralcea angustifolia
var. lobata, Wissadula holosericea , W. amplissima (L.) R. E. Fries,
and Sida filipes Gray, and, in the laboratory, also Althaea rosea.
Both skippers very likely. eat other mallows, as well.

Documentation: Eggs and larvae of Celotes were found on
Sphaeralcea angustifolia var. lobata at Davis Mountains State Park,
Jeff Davis County, V-1-61, by R. O. and C. A. Kendall. At the
laboratory in San Antonio, those larvae that were not preserved were
reared perforce on Althaea rosea. One adult was obtained from a
larva on V-28-61 and two from eggs on VI-6 and 9-61. Although
all were reported as nessus (Kendall 1965), I find that only the
adult that emerged VI-6 is nessus whereas those that emerged V-28
and VI-9 are limpia (paratypes).

From 3 larvae found by the Kendalls on Wissadula holosericea in
Musquiz Canyon on state highway 118, Jeff Davis County, VIII-
14-66, 1 adult (a paratype of limpia) was obtained on IV-14-67.

From 2 larvae found by the Kendalls on Abutilon incanum near
Rio Grande Village in Big Bend National Park, Brewster County,
X- 1 2-67, 2 adults of limpia (paratypes) were obtained (after a
lengthy diapause) on VI-19 and 22-68. At the same locality, III-
27-68, the Kendalls saw a female of Celotes oviposit on A. incanum
but got only the egg and not the female; from this egg, an adult of
limpia (paratype) was obtained on V-29-68.

The Kendalls collected 23 larvae on Abutilon malacum, VIII-
18-66, at a point on ranch road 170 overlooking the Rio Grande,
about 15 miles southeast of Redford, Presidio County; and, using
A. incanum, Malvastrum americanum, and especially Wissadula

64

Psyche

[March

holosericea as larval food in the laboratory, they reared 22 adults of
limpia (all paratypes) (one seemingly parasitized larva was pre-
served). At exactly the same spot, X- 11-67, the Kendalls collected
about 40 more Celotes larvae on A. malacum and reared 23 adults
(apparently limpia , but none of them examined by me).

Foodplant records for nessus are legion (Kendall 1959, 1965, and
unpublished). Most of them are of Abutilon incanum; and these
come from many counties in central Texas: Bexar (innumerable
records), Blanco, Bosque, Comal, Crockett, Kerr, Kimble, Llano,
Maverick, McCulloch, Medina, Nueces, San Patricio, Travis,
Uvalde, and Val Verde. I have examined much of the nessus ma-
terial reared on A. incanum.

The one valid record of nessus found as an egg on Sphaeralcea
angustifolia var. lobata in Jeff Davis County, Trans-Pecos Texas,
and reared to an adult in the laboratory on Althaea rosea was dis-
cussed above; the other two adults reared from this lot of eggs and
larvae on Sphaeralcea are actually limpia , though they, too, were
reported (Kendall 1965) as nessus.

Three larvae found on Sida filipes in San Antonio, Bexar County,

V- 9-63, and kept on this plant, produced adults of nessus (not seen
by me) on VI-8, 14, and VII-11-63 (Kendall 1965).

A larva found by the Kendalls on Wissadula holosericea in San
Antonio, Bexar County, V-24-65, yielded an adult of nessus (seen
by me) on VI-i 1-65.

Three larvae found on W . amplissima in extreme southern Texas
at the Laguna Atascosa National Wildlife Refuge, Cameron County,
IV-21-62, produced adults of nessus (all seen by me) on V-15, VI-4,
and VIII-20-62 (Kendall 1965).

Hypodigm. — Of limpia : the holotype cf specified earlier, an allo-
type 9 with identical data (except V-1-59), and paratypes comprising
64 cf 30? from Trans-Pecos Texas and 1 cf from Mexico. Paratypic
data arranged in the system described by Burns (1964: 19-20), with
names of recurring collectors abbreviated: J. M. and S. N. Burns
to B, H. A. Freeman to F, and R. O. and C. A. Kendall to K.

MEXICO. Coahuila. 25 mi. N Saltillo, IX-19-69, 1 cf (J- A.
Scott) .

UNITED STATES. Texas. Brewster County: Alpine, V-20-
26, 1 cf (O. C. Poling) (MCZ) ; V-21-26, 1 cf (O. C. Poling)
(MCZ) ; VI-4-42, 1 c? (F) ; VI-5-42, 1 <? (F) ; VI-6-42, 3c? (F) ;

VI- 7-49, icf (F); VII-10-49, 1? (F) ; VIII-9-52, 2tf (F) ;

VII- 28-53, icf (F); VI-16-60, 19 (F); III-14-61, 1 (W. S.

1974]

Burns — Polytypic Genus Celotes

65

McAlpine) ; III-27-61, 7cf (W. S. McAlpine) (2 In PMY) ;

VI- 5-61, 1 cf (F). 8 mi. N Study Butte, 3500 ft., VII-15-63, 1 cf
(H. Clench) (CM). Big Bend National Park: The Basin, 5500 ft.,
IV-30-65, 1$ (K) ; IX-8-65, icf (K). Oak Spring, 5700 ft.,
IX-9-65, 1$ (K). Rio Grande Village, 1900 ft., III-27-68, 3cf
(K) ; V-29-68, icf reared from egg (K) ; VI-19, 22-68, 2cf reared
from larvae (K). Window, 5500 ft., VI 1 1- 16-66, 3cf (K).
Culberson County: 2 mi. N Pine Springs, 5700 ft., Guadalupe Mts.,

VII- 19-63, 6(f (H. Clench) (CM). Jeff Davis County : 11 mi. N
Alpine, VI-20-68, 5 cf (J. A. Scott). 13 mi. NW Alpine on state
highway 118, IV-17-60, 1 cf 1$ (K. Roever). Big Aguja Canyon,
IV-2-68, 5cf (K). Davis Mts. State Park, V-28-61, 1 cf reared
from larva (K) ; VI-9-61, i? reared from egg (K). Ft. Davis,

VIII- 27-54, icf (R. M. Bohart). Limpia Canyon, 4800 ft., Davis
Mts., 1 mi. N Ft. Davis, IV-28-59, icf (B) (MCZ). Limpia
Canyon, 5000 ft., Davis Mts., 4 mi. WNW Ft. Davis, IV-28-59, 2?
(B)' (MCZ); V-1-59, 2 cf 1? (B) (MCZ); V-2-59, 2$ (B)
(MCZ); V-4-59, icf (B) (MCZ). Limpia Canyon, 4700 ft.,
Davis Mts., 5 mi. NE Ft. Davis, V-4-59, icf 1? (B) (MCZ).
Musquiz Canyon on state highway 118, VIII-14-66, 2cf (K) ;
IV- 1 4-67, 1$ reared from larva (K). Presidio County: 30 mi. SE
Presidio along Rio Grande, III-26-66, 2$ (D. J. Lennox). 15 mi.
SE Redford on ranch road 170, 2500 ft., VIII-29, 30, 31-66 and

IX- 1, 4, 5, 6, 7, 8, 9, 10, 11, 17, 29-66, 7 cf 15? reared from
larvae (K).

Hypodigm of nessus summarized in table 2.

Table 2. Specimens of Celotes nessus examined.

COUNTRY

State

$

$

N

MEXICO

Chihuahua

3

1

4

Coahuila

2

2

Sonora

7

7

UNITED STATES

Arizona

169

24

193

New Mexico

8

2

10

Oklahoma

1

1

2

Texas

147

67

214

N

II

04

04

Cn

97

432

66

Psyche

[March

Nomenclatural epilogue. — A species of what we now call Celotes
was named three times within less than a decade, 1877 to 1884 (see
e.g. Lindsey, Bell and Williams 1931; Evans 1953). In order of
publication, the names are Pholisora nessus Edwards, Spilothyrus
notabilis Strecker, and Carcharodus radiatus Pldtz. The fact that I
have made no effort to examine types is not so cavalier as it might
seem. On the one hand, genus Celotes is such a distinctive element
of the American fauna that one may safely assume that all three
names have long since been assigned to it correctly. On the other
hand, one may also assume that all three refer to nessus rather than
to limpia for the following reasons. The type locality for both
nessus and notabilis is San Antonio (or perhaps the nearby town of
New Braunfels for the latter), at the eastern edge of the main range
of nessus and therefore a few hundred miles removed from the near-
est populations of limpia in Trans-Pecos Texas. Because intervening
areas have now been well-collected, the total lack of limpia from
eastcentral Texas is real. The type locality for radiatus is simply
Texas; but, inasmuch as the name was published in 1884, the ma-
terial on which it was based almost certainly came from central
Texas, too. The west Texas areas in which limpia occurs are rela-
tively remote. Indeed, in all of the material assembled from museums
for this study, there were only two specimens of limpia that had been
collected prior to 1961 — and they were taken in 1926.

Godman and Salvin (1899) proposed Celotes as a monotypic genus
with Pholisora nessus Edwards as its type-species. There is some
historical and distributional interest in noting that their hypodigm
of “nessus” undoubtedly included nessus but may have been mixed.
As they themselves said, it comprised 3 specimens from Texas pro-
vided by Strecker and 3 from Mexico (Northern Sonora and Du-
rango city). Strecker’s notabilis ( — nessus) was one of a series of
lepidopterans that he described from material collected “'by Mr. J.
Boll, mostly in the vicinity of New Braunfels and San Antonia [sic]”
(Strecker 1878). Owing to its geographic origin, the Strecker ma-
terial certainly — and that from Northern Sonora almost certainly
— would have been nessus. Furthermore, the male genitalia that
Godman and Salvin clearly figured (pi. 91, fig. 29) belong without
question to nessus. The Durango material is problematic, however.
To judge from the locality, it might almost as well be limpia ‘as
nessus. Godman and Salvin’s color illustrations of dorsal and Vteiitral
aspects of a female of Celotes (pi. 91, figs. 27, 28) could be taken
for either species, the dorsal view (fig. 27), in particular, being

1974]

Burns — Polytypic Genus Celotes

67

rather more reminiscent of limpia with respect to both facies and size.
Unfortunately, the geographic source (s) of the female (s) on which
these figures are based was not indicated, but probability favors
Mexico: Evans (1953) records that the holdings of the British
Museum by then included 10 specimens of nessus from Texas, of
which just 2 are females, but still only 3 from Mexico, to wit, “ic?
2?” from “N. Sonora. Durango.” I would not be surprised to find
that Godman and Salvin depicted a fetnale of limpia from Durango.

Summary

Celotes limpia , a new species of skipper butterfly in a hitherto
monotypic genus, is described and extensively compared with C.
nessus, which it closely resembles in superficial appearance and biol-
ogy. Of 529 specimens examined, 97 represent the new species.
Whereas C. nessus ranges widely (from central Texas and Oklahoma
west to northwestern Arizona and south to at least the northern tier
of Mexican states), C. limpia is relatively restricted, occurring within
the range of C. nessus in Trans-Pecos Texas and interior northern
Mexico. In Trans-Pecos Texas, where the only direct comparisons
could be made, the two species are not only sympatric but also syn-
chronic, flying in an indefinite number of generations from March to
September. Larvae of both skippers are known, in the field, to eat
various plants in the Malvaceae: Abutilon incanum, A. malacum
( limpia only), Wissadula holosericea, W. amplissima {nessus only),
Sphaeralcea angustifolia var. lobata, and Sida filipes {nessus only).
In the laboratory, additional malvaceous foodplants are Althaea rosea
and Malvastrum americanum {limpia only). Celotes limpia and
C. nessus differ slightly in size and strikingly in genitalic morphology
(both male and female) and in the morphology of certain scales on a
male secondary sex character having a presumed communicative
function.

Acknowledgements

Thank you : for lending specimens, J. A. G. Rehn and D. C.
Rentz and Academy of Natural Sciences of Philadelphia, F. H.
Rindge and American Museum of Natural History, C. D. MacNeill
and P. H. Arnaud, Jr. and California Academy of Sciences, J. A.
Powell and California Insect Survey, H. K. Clench and Carnegie
Museum, L. M. Martin and J. P. Donahue and Los Angeles County
Museum of Natural History, J. F. G. Clarke and W. D. Field and

68

Psyche

[March

National Museum of Natural History, C. L. Remington and Pea-
body Museum of Yale University, F. H. Chermock, H. A. Freeman,
R. O. Kendall, D. J. Lennox, C. D. MacNeill, B. Mather, D. Pat-
terson, K. Roever, and J. A. Scott; for discovering foodplants , R. O.
and C. A. Kendall; for drawing genitalia , R. G. Gillmor; for photo-
graphing adults , T. P. Webster; for photographing metathoracic
pouches, F. White; and for providing financial support, National
Science Foundation grants GB-5935 and GB-37832.

Literature Cited

Burns, J. M.

1964. Evolution in skipper butterflies of the genus Erynnis. Univ.
California Publ. Entomol. 37: 216 pp., 1 pi.

Burns, J. M. and R. O. Kendall

1969. Ecologic and spatial distribution of Pyrgus oileus and Pyrgus
philetas (Lepidoptera : Hesperiidae) at their northern distribu-
tional limits. Psyche 76: 41-53.

Edwards, W. H.

1877. Descriptions of new species of butterflies belonging to the N.
American fauna. Canad. Entomol. 9: 189-192.

Evans, W. H.

1953. A catalogue of the American Hesperiidae indicating the classi-
fication and nomenclature adopted* in the British Museum (Natu-
ral History). Part III. Pyrginae. Sec. 2. London: British Mu-
seum. 246 pp., pis. 26-53.

Glick, P. A.

1939. The distribution of insects, spiders, and mites in the air. U. S.
Dept. Agr., Tech. Bull. no. 673. 150 pp.

Godman, F. D. and O. Salvin

1879-1901. Biologia Centrali-Americana. Insecta. Lepidoptera — Rho-
palocera. Vol. 2, 782 pp.; vol. 3, 113 pis.

Harris, L., Jr.

[1950]. The butterflies of Georgia (revised). Georgia Soc. Nat., Bull,
no. 5. vii + 33 pp.

1972. Butterflies of Georgia. Norman: Univ. Oklahoma Press, xxii +
326 pp.

Kendall, R. O.

1959. More larval foodplants from Texas. J. Lepidopterists’ Soc. 13:
221-228.

1965. Larval food plants and distribution notes for twenty-four Texas
Hesperiidae. J. Lepidopterists’ Soc. 19: 1-33.

Klots, A. B.

1951. A field guide to the butterflies of North America, east of the
Great Plains. Boston: Houghton Mifflin Co. xvi + 349 pp., 40 pis.

Lambremont, E. N.

1954. The butterflies and skippers of Louisiana. Tulane Stud. Zool. 1 :
125-164.

197+]

Burns — Polytypic Genus Celotes

69

Lindsey, A. W., E. L. Bell and R. C. Williams, Jr.

1931. The Hesperioidea of North America. Denison Univ. Bull., J.
Sci. Lab. 26: 1-142.

Mather, B. and K. Mather

1958. The butterflies of Mississippi. Tulane Stud. Zool. 6: 63-109.
Strecker, H.

[1878] “1877.” Lepidoptera, Rhopaloceres and Heteroceres, indigenous
and exotic; with descriptions and colored illustrations. No. 14.
Pp. 125-143.

A REMARKABLE NEW ISLAND ISOLATE
IN THE ANT GENUS PR0CERAT1UM
(HYMENOPTERA: FORMICIDAE)

By William L. Brown, Jr.*

Department of Entomology
Cornell University
Ithaca, New York 14850

In 1969, I collected in southern Africa and Madagascar, and then
moved on for work in southern India, stopping en route for four
days (29 March- 1 April) on Mauritius in order to sample the small
but interesting endemic ant fauna known from this island in the
Indian Ocean. My experience there was greatly enriched by aid and
information from Mr. Leo Edgerly, retired chief forester of Mauri-
tius, whose knowledge of the island and its plants, animals and people
is probably unmatched.

On the second day of my stay, on Mr. Edgerly’s advice, I climbed
the small mountain known as Le Pouce (“The Thumb”) in the
Moka Range in the northern part of the island. This mountain,
formerly collected by the Mauritian entomologists R. Mamet and
J. Vinson, is described in Donisthorpe’s paper (1946) on the ants
of Mauritius. The Moka Range rises from a sea of sugar cane, and
Le Pouce is separated from neighboring mountains in the narrow
volcanic chain by deep saddles. It is an elongate massif, with tall,
steeply dropping cliffs on all sides and a topside “plateau” sloping up
gently eastward to a sudden sharp peak that gives the mountain its
name. The plateau averages about 700 m in elevation; it is still
largely covered with a low, gnarled native forest much invaded by
guava and grassland. A mostly gentle trail hugs the cliffs and crosses
the plateau.

The first day on Le Pouce was a bright, sunny Sunday, and the
few well-marked trails through the pleateau scrub were invaded by
bands of teenagers from the lowlands, equipped with portable radios.
Accordingly, I stuck to the heavy vegetation. Ants were very scarce,
and nearly all that I found were colonies of the small dark Solenop-

*This work was supported by United States National Science Foundation
Grant GB-31662 and an earlier grant from the same agency; the support is
gratefully acknowledged.

Manuscript received by the editor February 15, 1974

70

1974]

Brown — Ant Genus Proceratium

71

sis mameti in rotten wood. Near the end of the day, when the sun
was getting low, I was at the eastern end of the plateau where it
begins to rise suddenly to the peak. Here a 2-centimeter-thick hollow
rotting stick lying on the leaf litter unexpectedly yielded a nest of
sluggish red ponerines, obviously Ectatommini in habitus. I got the
nest alive into a plastic “live tube” and kept it for a week until it
succumbed to mold. I was much excited by the find, although I as-
sumed at the time that I had a species of Gnamptogenys belonging
to the Indo-Australian “Stictoponera” group, because of the rather
large size and more or less shining integument of the workers, and
the distinct (though small to a hand lens) eyes. Even this would
have been a remarkable range extension.

On the next day I collected in other parts of the island, including
the scrubby “peat forest” on high ground in the south, then being
destroyed and replanted (disastrously!) with hurricane-vulnerable
exotic pines. The bag was discouraging, consisting almost entirely of
introduced ant species. A rapid traverse with Mr. Edgerly of the
southern Cocotte Mountain, badly stripped by a recent hurricane,
turned up nothing but introduced ants.

On the first of April, though I was scheduled to leave on an eve-
ing flight to Bombay, I tried Le Pouce again. A telephone call to
Mr. J. Vinson, who had collected some of Donisthorpe’s material,
convinced me that the main path on the LePouce plateau should not
be avoided. I arrived there early in the afternoon ; the day was heavily
overcast, threatening rain on the peaks, and it took me about an hour
to walk up to the plateau. Whereas the sunny Sunday in the scrub
shade had yielded almost no ants foraging, I now found foragers on
foliage and on the hard-packed earth of the trail every few meters
of the way.

These were mostly Camponotus aurosus and Pristomyrmex spp.
(=Dodous) , native Mauritian species. Before long, on the trunk
of a small tree by the path, I found a sparse trail of bright red ants
climbing the bark. Closer examination revealed these to be pre-
dominantly the ectatommine I had collected on the previous Sunday,
but interspersed with these were workers of Pristomyrmex bispinosus
(Brown, 1971) which, with their gasters carried partly curled under,
looked remarkably like the ectatommines. It is hard to avoid the im-
pression that some kind of mimicry involves these two species in this
habitat. The ectatommines ascending the trunk nearly all carried in
their mandibles tiny whitish spherical objects that proved eventually
to be arthropod eggs — probably spider eggs.

72

Psyche

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I climbed the tree, which was only about 5 meters high, and soon
found the nest about 3 meters up. Where two of the gnarled branches
crossed, a thick pad of lichens surrounded the place where they
touched. Forcing the branches apart, I found a rotted-out pocket,
evidently caused by their rubbing together in high winds. The cavity
extended downward several centimeters into one of the branches, and
it was full of the ectatommine ants with brood and many of the
round white arthropod eggs; I estimate that I removed or saw at
least 200 workers, and there may have been more. The nest was very
inconveniently placed, and the branch was too thick to break or cut
with the small knife I had along, so I had to be satisfied with what
I could extract by means of a twig wetted with alcohol, and with my
fingers and forceps.

A few minutes later along the trail, I found the entrance to
another nest in a small tree, also about 3 m above ground, and like
the first one of that day, also in a rot hole covered with lichens.
I took a few workers from near the entrance, and later found two
foraging workers on the hard-packed earth of the trail. Further
observations were made impossible by a sudden torrential rain, and it
was only with great difficulty that I got back down off the mountain,
partially dried out my clothes, and reached the airport in time for
the flight to India.

My hosts in Calicut, Drs. A. Benedict Soans and Joyce Soans, gave
me access to a stereomicroscope, and I quickly learned that the ecta-
tommines I had were not Gnamptogenys , but Proceratium! I offer
a formal description of the new Proceratium species, followed by a
discussion of its relationships and its evolutionary and zoogeographic
significance.

Proceratium avium new species
Figs. 1, 2

Holotype worker: TL 4.8, HL 0.96, HW 0.94 (Cl 98), ML
0.32, scape L 0.93, greatest diameter of eye 0.09, WL 1.43, L
petiole in side view 0.60, L petiolar node as seen from above 0.47,
W petiolar node 0.47, L hind tibia 0.97, L hind metatarsus 0.81 mm.
For head length (HL), measurement is taken from the anterior
lateral corners of the head (clypeus) ; head width (HW) excludes
the eyes, and is taken just behind them. The petiolar node length
excludes the brief anterior peduncle and is taken from the approxi-
mate base of the anterior nodal slope. The length of the gaster is

1974]

Brown — Ant Genus Proceratium

73

taken in side view from the dorsal side of the juncture with the post-
petiole straight to the most posterior part of the curve of the second
(downcurved) gastric segment (true abdominal segment IV).

Paratype workers (19 measured from 3 colonies at the type local-
ity) : TL 4. 7-5.0, HL 0.92-0.98, HW 0.91-0.98 (Cl 96-101), ML
0.31-0.34, scape L 0.90-0.99, greatest diameter of eye close to 0.09,
WL 1.38-1.49, L petiole in side view 0.59-0.61, L petiolar node
0.42-0.47, W petiolar node 0.41-0.47, L hind tibia 0.96-1.02, L hind
metatarsus 0.80-0.87 mm.

Composite description: Form of head and body more or less as
shown in Figs. 1 and 2. Variation occurs in the following traits:
Posterior border of head in full-face view varying from transverse,
nearly straight (holotype) to broadly rounded with or without a
narrow flattened or even feebly concave median portion, as in Fig. 2.
Nuchal carina (on cervical face of head) continuing as a ventro-
lateral margin halfway down each side of head. Sides of head vary-
ing from approximately straight and parallel (Fig. 2) to gently con-
vex and slightly converging anteriad. Eyes each composed of a single
clear, convex facet. Median lobe of clypeus with sides feebly sinuous,
as in Fig. 2, or merely weakly convex. Mandibles with 4 strong
teeth, but one of these is sometimes double; in addition a small offset
tooth is sometimes developed at the basal angle, normally hidden when
closure is complete. The inner margin of the peduncle of the mandi-
ble has a low, suboblong, sinuous or even bidentate ridge or lamella,
visible only when the mandible is open. The scapes vary in length
and slightly in apical thickness, so that when laid straight back they
surpass the posterior border of the head by amounts ranging from less
than half their apical thickness to more than their apical thickness.

Labral shield bilobate, the lobes separated by a broad V-shaped
notch; also, lateral to each lobe is a small thumb-shaped lobe or knob
extending dorsomesally (flexad) of the plane of the shield from its
basal ridge. Maxillary palpi 4-segmented, the basal segment short and
cylindrical, the second segment flat and attached to the basal segment
at nearly a right angle by a peduncle that arises from the side of II;
III and IV are short elliptical segments extending in line from II.
This is the characteristic form of maxillary palpi in Proceratium ,
Labial palpi 3-segmented.

Trunk convex from side to side, and from front to rear in side
view, but the portion of the outline from mesonotum to propodeal
declivity may be straight or may be interrupted by a feeble dip or
saddle in the region of the obsolete metanotal groove. The propodeal

74

Psyche

[March

angles may be smoothly rounded, as in Fig. 2, or may bear very low,
obtuse corners that also render the sides of the declivity vaguely sub-
marginate (30 March colony) as opposed to the rounded sides of the
declivity in the 1 April series (Fig. 2), taken less than half a kil-
ometer away. Seen from above, trunk broadly rounded in front,
sutureless, tapering gently caudad, with a feeble suggestion of con-
striction behind mesonotum.

Petiole briefly pedunculate in front, the peduncle with strong
anterolateral cornuae, the node varying from loaf-shaped with rela-
tively steep, rounded anterior face (Fig. 2) and rounded sides to a
lower version with more gradually sloping anterior face and sides
less bulging. Postpetiolar (first gastric) segment varying in dorso-
ventral depth, especially near the posterior quarter or third of its
length, and the individuals with deeper postpetiole tend to have the
tergum of this sgment more swollen (or humped as seen in side view)
than in the paratype depicted in Fig. 2.

Integument overall smooth and decidedly shining, but head, trunk,
node and postpetiolar (first gastric) segment thickly sown with cir-
cular, centrally-tuberculate and piligerous foveae or coarse punctures.
These foveae tend to be larger and more densely arranged, even sub-
contiguous in places (e.g., in the region behind the eyes), in the
30 March colony as compared with the series of 1 April, but the
latter series vary greatly in this respect, even within samples from the
same nest. The least strongly punctured individuals have the dorsa
of trunk, node and postpetiole almost completely smooth and shining,
the punctures here small and sparse. Antennae and legs much more
finely and densely punctate, becoming more opaque apicad. Median
clypeal lobe finely punctate-rugulose in a longitudinal direction ;
mandibles striate-punctate, but shining and coarsely punctate toward
apicolateral margins. Second gastric (IV true abdominal) segment
smooth, shining, with sparse piligerous punctulae. Gastric apex finely
and densely punctulate, subopaque.

Pilosity abundant, moderate and uneven in length, of fine, sub-
decumbent to suberect tapered hairs, a little longer but much less
abundant than usual in Proceratium, grading into a much shorter,
appressed to decumbent pubescence-like covering as one moves apicad
on antennae, legs and toward gastric apex; also a fine, appressed,
medially-directed pubescence in the space between eyes and frontal
carinae.

Color (of fully mature individuals) rich bright ferruginous red,
legs and scapes often lighter and more yellowish; region immediately

1974]

Brown — Ant Genus Proceratium

75

Fig. 1. Proceratium avium new species, paratype worker in side view; only the longer hairs seen in silhouette
are shown (from colony No. M-252, 1 April 1969).

Fig. 2. Same, full-face view of head. Drawings by Bente King.

76

Psyche

[March

around eyes often with darker pigment. Some specimens, presumably
nearer to being callow, are lighter, more yellowish-ferruginous in
general color.

Malpighian tubules numbered 6 each in 4 live workers dissected,
and 4 in a fifth worker, the last probably representing a mutilated
specimen (dissected under less than ideal conditions). Not crypto-
nephric. The count of 6 agrees with a worker of P. goliath dissected
in Costa Rica.

Queen, subergatoid (from 30 March colony) : TL 5.1, HL 0.90,
HW 0.92 (Cl 102), ML 0.34 (mandibles slightly opened), scape
L 0.83, greatest diameter of eye 0.14, WL 1.38, petiole L (from
above) 0.57, petiole W. 0.58, L hind tibia 0.87, L hind metatarsus
0.71 mm. Head as in worker, of the shape with transverse, only
feebly convex posterior border in full-face view; sides converging
slightly from behind eyes toward mandibles, and bulging again
slightly at lateral ends of clypeus. Mandibles with 5 moderately
strong teeth and a small offset basal tooth. Ocelli small but distinct;
compound eyes nearly circular in outline, moderately convex, with an
estimated 70 distinct ommatidia, relatively smaller than usual in
Proceratium queens. Scapes just barely overreaching posterior border
of head when laid straight back.

Trunk somewhat reduced, but the usual sclerites of the ptero-
thorax separated, except that the elements of the mesonotum are all
indistinguishably fused into one gently convex continuous shield ;
metanotum distinct but narrow, forming the usual acute median
process, but in short compressed form. Propodeum with slightly
raised, but bluntly rounded angles and slightly concave, submargined
declivity. Wing stumps present and blackened, but small.

Petiole shorter, broader and a little higher than in worker, the
node rounded above and with a rather steep anterior face. Gaster
much deeper and broader than in worker, extraordinarily volumi-
nous, but undercurved as in worker. Sting present, stout, protrusible.

Sculpture, color and pilosity much as in the worker over head,
trunk and petiole, but the punctures here shallower, less distinct and
sparser than in even the more weakly foveate workers; mesonotum
and node with obsolescent foveae, almost completely smooth, shining.
Gaster smooth and shining, without coarse punctures, but very finely
and densely punctulate, with associated dense, short, appressed pu-
bescence and a few short, delicate decumbent to erect hairs, mainly
at the posterior borders of the first segment and on the undersurfaces,
and becoming abundant near the gastric apex. Due to the great bulk

1974]

Brown — Ant Genus Proceratium

77

of the gaster, the fusion of the mesonotal sclerites, and the small
eyes, I doubt if this queen ever had flight-functional wings.

Male (from 30 March colony) : TL 4.7, HL 0.76, HW (in-
cluding eyes) 0.98, ML 0.29, scape L 0.64, greatest diameter of
compound eye 0.40, WL 1.55, L petiole 0.55, W petiole 0.35, L hind
tibia 0.93, L hind metatarsus 0.80, forewing 4.0 mm.

Head with high-domed vertex set with 3 large, clear ocelli; com-
pound eyes large and strongly bulging, with convex inner margins as
seen in full-face view of head. Median lobe of clypeus projecting,
broadly truncate. Frontal carinae parallel, low but sharp, vertically
raised. Mandibles triangular, each ending in a stout curved apical
tooth, masticatory borders curved and cultrate. Scapes overreaching
posterior border of vertex. Palpi segmented, as far as can be seen in
the undissected specimen, as in worker.

Alitrunk with well-developed flight sclerites and wings; notauli
lacking; scutellum semiglobose, protruding. Metanotum with the
usual stout, acute median tooth. Propodeal dorsum short, convex,
rounding obtusely into the declivity, which is indistinctly marginate
on the sides.

Petiole long and very low, with the highest part of the gently con-
vex node near midlength, but the greatest width at about the posterior
third. Postpetiole anteriorly wider than petiole, and widening still
more posteriad, but not quite as wide as the succeeding segment,
which is vaulted ventrad, but not as strongly as in worker and queen.

Genitalia largely retracted (not dissected), but it can be seen that
the parameres have thin, broadly rounded apices that are bent mesad
toward each other.

Legs long and slender, with the usual single pectinate spur on each
middle and hind tibial apex, and the tarsal claw slender and simple.

Wings of basic Proceratium pattern, with Rsf2-3 completely lack-
ing in the forewing; m-cu also gone; the enlarged “cubital” cell
receives only one free vein distally (Rsf5), because the apical free
abscissa of M is gone. In the hind wing, the large cell receives a
short remnant of Rs and the longer free apical abscissae of M and
CuA; 1 A is missing beyond cu-a. Wings hyaline, with light brown
veins. ( In the newly-discovered males of the related P. stictum —
from Brookvale, Queensland, and P. goliath — from Zent, Costa
Rica, the venation pattern is similar to that of avium , but darker in
goliath , still paler in stictum ; m-cu is preserved in stictum, but in
the hind wings of both, the free abscissa of Rs is reduced to obso-
lescence. )

78

Psyche

[March

Sculpture shining, with foveae much as in the worker, but also
with rugulae and more opaque around the eyes, posterior scutellum,
dorsum and sides of propodeum, metanotum and sides of petiole.
Pilosity fine, rather short, moderately abundant, decumbent to sub-
erect, generally distributed over body, scapes and legs. Color light
yellowish-brown, gaster more brownish; vertex infuscated near ocelli
and in back of compound eyes.

Holotype (from unnumbered colony, i April 1969) and paratypes
(colonies ICA-69, 30 March 1969; M-252 and an unnumbered
colony, plus strays, 1 April 1969) taken from Le Pouce (mountain),
Mauritius, in native forest between 700 and 800 meters elevation on
the plateau just below the peak (W. L. Brown, Jr.). Holotype and
paratypes deposited in Museum of Comparative Zoology, Harvard
University, Cambridge, Massachusetts, U.S.A. Paratypes in Cornell
University Collection, British Museum (Natural History), Aus-
tralian National Insect Collection, Canberra, Australia, and else-
where.

P. avium belongs to the stictum group of Proceratium , containing
two other species. P. stictum (Brown, 1958 ,*366) has been known
only from a single worker from North Queensland. (Now I have
seen males apparently belonging to stictum , taken at light by E. S.
Ross and D. Q. Cavagnaro at Brookvale, Queensland; and R. W.
Taylor showed me a series that I was only able to examine hurriedly,
but which looked to me like stictum , from his collections made in
North Borneo.) The second species of the stictum group is P. goliath
(Kempf and Brown, 1968:94 ff.) from Costa Rica. P. aviu?n is
distinct from both of these species in its much shinier and coarser
sculpture and in details of the shape of the trunk, particularly its
completely unarmed propodeum, in its longer antennae, and in its
less strongly undercurved gaster. P. avium is also much larger than
P. stictum , but smaller than P. goliath , and its eyes are relatively
larger than in either of these species. In fact, P. avium might well
go into a group of its own, but the structure of the clypeus, mandi-
bles, and petiole are so much like those of P. stictum and P. goliath
that the relationship of these three species is obvious.

The characters they share are also primitive for the genus Pro-
ceratium, and this with the widely discontinuous known distribution
of the group suggests that the stictum group represents the relicts of
an early dispersal wave of a primitive Proceratium stock that spread
widely over the earth and was overtaken (except on Mauritius) by
later waves of more advanced Proceratium groups. Possibly wide-

1974]

Brown — Ant Genus Proceratium

79

spread extinction of stictum- group stocks ensued ; although it is not
easy to visualize significant competition among such rarely-collected
species, I should point out that the adaptive zone (predation of spider
eggs) is a limited one. Furthermore, advanced methods of collecting,
at least in North America, have proven that some advanced Procera-
tium species are common in suitable microhabitats. These micro-
habitats are, so far as we can tell, all cryptic ones — above all, in
large masses of well-rotted wood. This may, of course, be an artifact
of biased collecting methods, but I doubt it. On the northern fringes
of the range of Proceratium — for example, in the environs of Bos-
ton — P. silaceum and P. pergandei are taken occasionally, but always
under rocks in the soil. Farther south in the U.S., almost all
collections are made deep in rotten wood or in humus and litter near
rotten wood. The type series of P. goliath came from a rotten log
in wet tropical forest.

The microhabitat of P. avium , to judge from the limited observa-
tions I have made, differs strikingly from that of other Proceratium.
Certainly, the nest site is different. The first nest of P. avium taken
on Mauritius came, it is true, from a hollow rotting stick lying on
the forest floor. This may be a common kind of site for the species,
yet very many such branchlets were examined on the day of collection
without success, and it is possible that the branchlet had fallen re-
cently from a tree above. In any case, I have never seen any of the
eight other Proceratium species collected alive by myself in various
countries nesting in such a large cavity in such an exposed site as was
the colony in this stick. The other two P. avium nests seen were of
course arboreal, with the foraging trails openly exposed for some
meters over open soil and tree trunk. The ground and arboreal nest-
ing sites for ants in wet tropical montane forest are rather academi-
cally distinguished in any case, but it seems to me that the exposure
of the nests and foraging trails of P. avium is what is significant here.
If other species of Proceratium forage so openly day or night, it has
not been noted as far as I know, and indications are against it.

Of greatest interest is the contrast in habitus, especially that part
owing to sculpture and pilosity, between P. avium and its congeners.
In P. avium , the looser, coarser, more shining integument and its
fairly long open pilosity may be compared with the finely rugulose-
punctate or reticulopunctulate sculpture and very short, fine, more or
less dense pilosity of the other Proceratium species. There is every
reason to believe that these fine, crowded punctures and their associ-
ated hairs are a specialized evolutionary development (pushed further

8o

Psyche

[March

in Discothyrea ) stemming from a condition in which the integument
was more coarsely sculptured, with larger foveae, each fovea bearing
a hair on a central tubercle. The Old World ( Indo-Melanesian)
Gnamptogenys stocks include species (e.g., G. menadensis) that meet
these specifications and are known to be epigaeic, even arboreal, for-
agers (personal observations), but Gnamptogenys has reduced palpal
segmentation and other characters that make it more likely a con-
vergent than an ancestral stock to Proceratium.

I believe that Acanthoponera is nearer to the ancestral line of
Proceratium because of the higher basic palpal segment number and
the traces in some Proceratium species of a median carina on the
frontal area and vertex of the head, characters of Acanthoponera
(Brown, 1958:188). But the main question we are led to consider
is: Has P. avium preserved some version of the archetypal Procera-
tium sculpture-pilosity pattern, or is its present condition a secondary
reversion from the fine-scale pattern characteristic of other Procera-
tium around the world ?

Possibly we shall never have a clearly definitive answer to this
question, but we do have one clue pointing toward the reversion
hypothesis. This clue is the rather unusual eyes of P . avium — a
rather large, glassy-looking orb on each side, backed by a perceptible
amount of dark pigment, and relatively larger than the character-
istically single-facetted “compound” eyes of other Proceratium species.
I think we have to suppose that the ancestors of P. avium, and also
of all other Proceratium, had already specialized for a cryptic ex-
istence to the point where minute single-facetted eyes, probably barely
enough to sense the difference between light and dark, served ade-
quately the lifeways of these animals. My guess is that such lifeways
also had forced selection for the fine sculpture-pilosity pattern of Pro-
ceratium in the ancestral lineage of this genus. If my reasoning is
correct, then P. avium is an interesting example of a “character-
released” species on a remote oceanic island with a depauperate
endemic ant fauna.

The known endemic ant fauna of Mauritius numbers only about
7 species, if we take Donisthorpe’s (1946, 1949) count as a base and
deduct obviously introduced species, including synonyms, and add my
collections. The endemics now appear to be restricted to the small
areas of upland native forest; the cane fields and other culture areas
are saturated with introduced ant species. In a sense, then, the moun-
tain forests represent the “real” island (s) of Mauritius as far as the
endemics are concerned.

1974]

Brown — Ant Genus Proceratium

8l

Among the ants of the present endemic fauna, it is difficult to pick
out any that might be serious competitors of Proceratium avium.
Our assumption here, of course, is that P. avium subsists primarily
on arthropod eggs, probably mainly the eggs of spiders. (But the
assumption rests on only a few observations, which need augmenta-
tion.). Perhaps Solenopsis mameti , a much smaller ant that nests
mainly in rotten wood in forest shade, would qualify as a competitor.
This judgement is based on the generalized feeding habits of similar-
sized Solenopsis elsewhere in the world, and we have absolutely no
direct information on the food of S. mameti. At least, the species
has not been seen foraging on open paths or tree trunks during
the day.

The bright red color and open-trail foraging of P. avium suggests
reduced predation pressure in the Mauritian native forest habitat,
but the possible mimicry with Pristomyrmex bispiwosus could on the
other hand indicate that predator pressure is appreciable, and in some
way answered by protective properties.

In summary, a reasonable hyothesis to explain the atypical “epi-
gaeic characters” of P. avium assumes that the ancestral stock reached
Mauritius a long time ago from Africa or Asia in a floating log or
rotting branch, and established itself in an ant-poor environment that
was perhaps also weak in the kinds of predators that attack open-
foraging ants. Evolution in such an environment, it is argued, led to
the reacquisition of characters that had been lost by the parent Pro-
ceratium stock during continental specialization to cryptic environ-
ments in which arthropod (spider?) eggs had become its main food.

Another hypothesis is that the Mauritian Proceratium retains a
sculptural-pilosity pattern primitive for the genus, and that its arthro-
pod egg-feeding proclivities were acquired in an open-foraging situa-
tion that elsewhere has since been modified under continental pressures
of competition and predation.

In order to throw light on the question, it would be interesting to
know exactly what animals the prey eggs on Mauritius belong to, and
where and how they are taken by the ants. It may be that we shall
never find out, for the mountain forests of Mauritius appear to be
teetering on the brink of extinction.

Notes on other species of Proceratium

Since my summary of Proceratium (Brown, 1958: 241-248), the
genus has been found in additional parts of the world. Leston

82

Psyche

[March

(1971, J. Entomol. (B) 40: 118-119, figs. 1, 2, worker, queen)
described P. boltoni from Ghana; this species appears to be a fourth
member of the stictum group, perhaps annectant to the pergandei
group.

Borgmeier (1959, An. Acad. Brasil. Ci., 31: 309, worker) de-
scribed P. brasiliense , a close relative of the Central American-
Mexican P. micrommatum , from southeastern Brasil ; it is the first
representative of the genus from so far south in the New World.

I have found a lone Proceratium queen in rotten wood in wet
forest on the lower slopes of Mt. Klabat in northeastern Celebes.
The specimen is in the silciceum group and is fairly near P. carini-
frons and P . lombokense.

Roy Snelling has published a paper (1967, Contrib. Sci. Los An-
geles County Mus., 124: 1-10.) giving further data on P. calif orni-
cum of the West Coast of the United States, plus a key to the New
World Proceratimn species. The species of the micrommatum group
are, however, very close to one another, and I suspect that P . con-
vexiceps may eventually be shown to be synonymous with P. microm-
matum, because the chief character — presence or absence of a tri-
angular median lobe or tooth on the clypeus — involves this small
tooth that is very variable in size in the samples from Panama and
Mexico. P. micrommatum seems to be widespread in Mexico : Chi-
chicastle, Tabasco (F. Bonet) ; Pueblo Nuevo, Veracruz (E. O.
Wilson) ; Mesa de Chipinque, near Monterrey, Nuevo Leon (S. and
J. Peck) ; Antiguo Morelos, Tamaulipas, litter of palm- thorn forest
(S. and J. Peck) .

P. normandi Santschi (1929, Bull. Soc. Ent. Belg., 69: 138,
worker, queen; type locality La Caille, Algeria and Ain Draham,
Tunisia) is a new synonym of P. numidicum Santschi (1912, Bull.
Soc. Hist. Nat. Afr. Nord, 4: 1 72-1 73, figs. 1, 2, worker, queen;
type locality Ain Draham, Tunisia). In the Santschi Collection,
Naturhistorisches Museum, Basel, the types of P. numidicum are
callow in part, but I could find no other differences between them
and the types of P. normandi in the same collection.

Proceratium mancum Mann (1922, Proc. U. S. Nat. Mus., 61
(13): 6-7, worker, queen; type locality: Cecilia, Honduras) is a
new synonym of P. silaceum Roger (1863, Berlin. Entomol.
Zeitschr., 7: 172, worker; type locality “Nord-America”) . The
differences cited by Mann and listed in Snelling’s key are all very
variable in P. silaceum , as is body size. Specimens from the tropical
part of the range usually have the petiolar node thinner from front

1974]

Brown — Ant Genus Proceratium

83

to rear, and the posterior face is more or less concave and overhung
by the slightly produced posterodorsal nodal border, but a tendency
in this direction is seen in some northern specimens, for instance, in
a worker from the Ouachita Mts. of southwestern Arkansas, and
one from Schooler Lake, Choctaw Co., Oklahoma (W. L. Brown).
The geographical gap between Honduras and eastern U. S. is now
being filled in to some extent. Already in 1911, W. M. Wheeler had
collected specimens in Guatemala (Quirigua and without locality
except for the country), and now S. and J. Peck have sent samples
from litter berlesates in Veracruz, Mexico: near Cordoba, and in
the canyon of the Rio Metlac, near Fortin de las Flores. It should
only be a matter of time until we have samples from east Texas and
Tamaulipas. This case is paralleled by that of Cryptopone gilva,
another ponerine that ranges from the U. S. down through the up-
lands of Mexico and Central America as far as Panama.

P. goliath was taken at Zent, Costa Rica by N. A. Weber, the
collection being the first male known for the species, date 17 July
1956. A worker apparently belonging to this species, but a little
smaller than the types from Costa Rica, and with a very deep con-
striction between postpetiole and succeeding segment, comes from
Caves Branch, British Honduras (berlesate, S. and J. Peck).

References Cited

Brown, W. L., Jr.

1958. Contributions toward a reclassification of the Formicidae. II.
Tribe Ectatommini (Hyraenoptera) . Bull. Mus. Comp. Zool. Harv.,
118: 173-362.

1971. Characters and synonymies among the genera of ants. Part IV.
Some genera of subfamily Myrmicinae (Hymenoptera : Formici-
dae). Brev. Mus. Comp. Zool., 365 : 1-5.

Donisthorpe, H. St. J. K.

1946. The ants of Mauritius. Ann. Mag. Natur. Hist. (11) 13 : 25-35.

1949. A new Camponotus from Madagascar and a small collection of
ants from Mauritius. Ann. Mag. Natur. Hist. (12) 2: 271-275.
Kempf, W. W. and Brown, W. L. Jr.

1968. Report on some neotropical ant studies. Pap. Avuls. Zool. S.
Paulo, 22: 89-102.

SEXUAL BIOLOGY, CHROMOSOMES, DEVELOPMENT,
LIFE HISTORIES AND PARASITES OF BOREUS,
ESPECIALLY OF B. NOTOPERATES.

A SOUTHERN CALIFORNIA BOREUS.

II. (MECOPTERA: BOREIDAE)*

By Kenneth W. Cooper
University of California, Riverside
California 92502

Introduction

This, the second of three accounts devoted primarily to Boreus
notoperates Cooper, deals with aspects of its life history. The sub-
jects include, among others, sex ratio, mating, developmental stages,
a brief comparative account of its cytology, its host mosses and pecul-
iar adaptation to a life in their thin sods on diorite boulders in a
region annually subject to long periods of drought and considerable
heat. All of this is placed in a framework of what is now known
for the other species of Boreus on these topics.

Though some fifty or so articles are referred to or discussed that
are devoted to, or comment upon, Boreus , there are an additional
sixty to seventy more papers and accounts which, though all have
been consulted, either do not bear directly on the topics treated, or
seem wholly derivative. My choice of references is based upon their
substantial treatment of a subject, their content of new observations
or ideas, or the need to correct a standing error. To these sources,
I have added my own unpublished observations on B. brumalis Fitch
and B. nivoriundus Fitch where relevant. The overall aim, then, is
to draw together, to compare, and, where possible, to generalize
what is known of the species of Boreus on the topics presented and
on which I have information derived from B. notoperates.

Among the matters given emphasis are: the likelihood that species
of Boreus have a sex ratio approaching equality (a debated topic) ;
the fact that B. notoperates is the only species (of eight for which
information is known) that deviates in its mating; the curious re-
ciprocal intromission at mating by male and female alike, and the
anatomical relations of their parts; the conservative nature of the
chromosomal cytology of Boreus and other scorpion flies as compared
with Panorpa; the prevalence of a larval epistomal suture, which

* Manuscript received by the editor February 10, 1974.

84

1974]

Cooper — Borens

85

bears upon the tenability of the order Neomecoptera that specially
sets Boreus apart from other scorpion flies; the likelihood that Boreus,
like other Mecoptera, has four larval instars; the occurrence of both
pupal and adult pharate stages in development; that whatever limits
the distribution of a species of Boreus , it is not its host mosses; and
finally, that Boreus species, despite their peculiar life cycle, may have
an obligate hymenopterous parasite, a matter which puzzled Withy-
combe. The third and final article will deal with the range of habi-
tats species of Boreus have entered, the mosses with which Boreus
species are associated, interpretation of Boreus as a winter insect,
and a reconsideration of its distribution in relation to drift and glaci-
ation.

Occurrence and Span of Adult Activity

Boreus notoper ates has so far been collected at two localities only,
both being NW faces of steep canyons, at 1219m altitude (Cold-
water Canyon) and at 1645m (Black Canyon), on Mt. San Jacinto,
Riverside Co., California. The first is a yellow pine — chaparral
ecotone ( Pinus ponderosa Dougl. — A denostoma fasciculatum Hook,
and Arn.) that is Upper Sonoran, or Upper Austral in Merriam’s
( 1898) original sense, the mean temperature for the six hottest weeks
(July through mid-August) being — 24°C. The second is a mixed
woods of yellow pine, incense cedar, white fir and canyon oak ( Pinus
ponderosa , Libocedrus decurrens Torr., Abies concolor Gord. &
Glend., and Quercus chrysolepis Liebm.), fringed and penetrated by
chaparral elements; it is Transitional, the corresponding high mean
temperature being about 2i°C. Over the period 1955 — 71 the mean
annual precipitation (close by at the Idyllwild Fire Station, alt.
1645m) was 59.6 cm (23 in.), with 74% falling in the months of
November through March during which adult B. notoperates have
been collected. The adults have so far been found only on mosses
growing on boulders and large jointed blocks of diorite, or on snow
about them, in both fairly open (chiefly Coldwater Canyon) and
shaded (chiefly Black Canyon) situations. Even during the period
of winter precipitation the mosses are only periodically dampened
and luxuriant, for precipitation is widely scattered, and melts rapidly
when snow. The activity of Boreus at all its stages coincides with
those periods, frequently brief and generally spaced out, when the
mosses and their roots are damp. At other times the relative humidity
is low, the mosses are dry to the touch, even friable, and their sods

86

Psyche

[March

crumbly or powdery. In periods of desiccation Boreus are found, if
at all, with difficulty, unless a special method is used.

The earliest that adult Boreus notoperates have been taken was
mid-November, several days following the second snowfall of an inch
or more, although it is likely that a snow or a soaking rain in Octo-
ber would bring them forth to judge from laboratory emergences of
pupae collected in mid-August. Thereafter adults have been found
during the damp periods in all months until mid-March, when they
become rare. Air and surface temperatures have been mild when
adults have been collected, ranging from a low of 6.i°C to i8.3°C,
though in the November through March interval below freezing
temperatures are not uncommon, mean low temperatures for these
months ranging from — 4.2°C to 3.2°C, with absolute lows to
— I2.2°C.

Sex Ratio and Spanandry

The sex ratio among adult B. notoperates from field collections,
though slightly skewed toward an excess of males early in the season,
of females late in the season, does not depart from equality; thus:
I3icf d\ 150$$ (of which 32 of each sex were collected as mated
pairs) ; chi-squared gives 0.3 > P > 0.2. Nor is there a departure
from equality among immature stages collected in mid-August and
early September. For 47 such larvae and pupae we have:

29 late instar larvae and pharate pupae of which :

19 sexed by dissection of gonads, giving: I2cfcf, 7?? J
10 pupated, giving: 6 cf cf > 4$$ J
18 pupae when found, of which: 7 cT cT , n?$ ;
which gives a sex ratio of 25 cf cf : 22 $ $ , for which 0.7 > P > 0.5.

Now Striibing (1950) has shown from extensive field and lab-
oratory observations that males of Boreus hyemalis (L.) do not tend
to emerge earlier than females in the autumn. They do, however,
tend to die off somewhat earlier than females, and this has been
claimed or inferred by others both from field and laboratory observa-
tions ( e.g Withycombe 1922, Syms 1934, Cotton 1971; and for
B. westw'oodi Hagen, Brauer 1855; for an undescribed species?,
Kolenati 1847)1. My own observations on B. notoperates are in

1Striibing (1950) gives strong reason to believe that Brauer’s species is
almost certainly B. westwoodi, and not B. hyemalis as he and others have
thought. Furthermore, Pliginsky (1930) states that Kolenati’s specimens,
from the glacier of Aar (Kazbek, Kaukas), no longer can be found,
and are very likely an undescribed species. From Kolenati’s comments,
they fall among the species with a reduced antennal joint number (~20).

1974]

Cooper — Boreus

87

agreement. But it has also been claimed, as early as Hardy’s (1848)
and MacLachlan’s (1868) notes, that the sex ratio of Boreus is
spanandrous, but only loosely in Marchal’s (1911) sense, namely
that males regularly make up a minority of the population. Not
surprisingly, small collections of Boreus may show a predominance
of one sex. In general, however, collections of two dozen or more
specimens at a locality, but not made at the season’s close when
females do tend to predominate, give sex ratios which approximate
equality, as do those for B. notoperates. There are, however, excep-
tions. But for none of the following records is there a significant
departure (namely, P < 0.05) from equality of the sexes:

1. B. brumalis Fitch

27cf c?

26$$

(Hanover, N. H.,
'Cooper, unpubl.)

2. B. coloradensis Byers

21

20

(Byers 1955)

3. B. unicolor Hine

64

57

(Chapman 1954)

4. B. hy emails ( L. )

39

4i

(sifting, Druet & Le-
gros, ex Lestage 1941)

33

45

(table 1, to Dec. 20,
Strubing 1950)

103

no

(table 4, not text p. 84,
Strubing 1950)

19

1 1

(Schiirmann, ex Strub-
ing 1950)

38

45

(pitfall traps, Cotton
1971)

5. B. westwoodi Hagen

60

64

(Martynova 1954)

24

25

(Martynova 1954)

12

14

(Martynova 1954)

6. B. bey-bienkoi Tarb.

56

49

(Tarbinsky i960)

As exceptions, with probabilities

< 0.02

to < 0.00 1 as random

departures from equality of the sexes, we have:

7. B. nivoriundus Fitch

42 cf cf

23??

(Hanover, N. H.
Cooper, unpubl.)

8. B. hy emails (L.)

398

123

(Steiner 1937)

9. B. westwoodi Hagen

97

67

(3 collections, data

homogeneous, Fjellberg
& Greve 1968)

Judging by records 1-6, and from the proportions among im-
mature stages which have been sexed, the records 7-9, all of which
show a significant preponderance of males, may reflect a tendency for

88

Psyche

[March

males to wander widely at times in search of females, or true pecu-
liarities of the particular populations, or perhaps of the mode or
circumstances of collecting. Strubing’s (1950) laboratory rearings
of B. hy emails gave 40c? cf, 36?$, and Fraser’s (1943) account
implies that his collection (in September) of pupae of B. hy emails
consisted of 296" cf , 21 (for which 0.3 > P > 0.2). My own
collection of larvae, pharate pupae, and pupae of B. nlvoriundus in
August, at the same site in Hanover, N. H., at which the adults
scored in 7 above were later collected (December to mid-April),
gave 30 cf cf , 33 ?$ . Pupae of B. brumalis collected (from October
to mid-November) at Princeton, N. J., likewise do not depart from
an equality of sexes: 40 cf cf , 46 $ ? . As Striibing (1950) concluded,
the sex ratio in Boreus appears to be close to unity for both immature
stages and adults, as is the case for B. notoperates. The answer to
Lestage’s (1941) question “is there spanandry in Boreus?” must be:
“not so far as known, and perhaps not at all,” as Lestage suspected.
Indeed, the only significant departures so far recorded are in fact
spangynous , not spanandrous.

Mating

Mated pairs of B. notoperates are found chiefly on patches of damp
moss, free of snow, from early in November to near the middle of
March. Although Fraser (1943) claimed B. hyemalis to be cre-
puscular, there seems to be no special time of day that is favored for
mating by B. notoperates if the temperature is mild ; nor is light a
requirement, for B. notoperates mates readily in the dark (in an
incubator, at 9°C). In but one case (of 33) has a mated pair been
found on the snow, and that mated pair had most likely fallen a foot
or so out onto the snow from a steep, moss-covered rock-face. Cer-
tainly the suggestion that Boreus occurs on snow because it is easier
to find mates there is implausible; the rule seems to be that mating
generally occurs on or in moss, where they congregate when it is
available to the insects.

Nine complete matings of B. notoperates have been observed,
namely from the first attempts of the male to gain a partner to the
completion of intromission, as well as a good many partial sequences
from all stages in the routine. B. notoperates is without a courtship,
just as in the three species for which the course of events of mating
have been described, namely B. westwoodl (Brauer 1855, 1863,

1974]

Cooper — Borens

89

Sauer 1966, perhaps Svensson 19662), B. hyemalis (Withycombe
1926, Syms 1934, Steiner 1937), and B. brumalis (Cooper 1940,
Crampton 1940). From its very onset the affair is between a “coy”
female acting as though bent on escape, and an aggressive, though
not necessarily persistent, male. An ardent male, when within some
millimeters range, springs at the female, ensnaring her with his tong-
like wings while seizing whatever he can of her extremities with one
or both of his genital claspers (or gonostyles). If he fails to gain a
hold, as he occasionally does, the female leaps away and is not directly
pursued. Thereafter the male either takes a waiting stance on a
sprig of moss, or courses about the moss, in both cases twitching his
wings and opening and flexing his gonostyles from time to time.3
When chance again presents another or the same female, the male
attempts once again to gain a firm hold of the female.

When a mating spring has been successful, the male may have seized
a female by a posterior femur (5 cases), a mid-tibia (2 cases), a pro-
tarsus (1 case), or the antennae (1 case). In some other instances,
in which only a part of the mating routine was followed, males had
gained holds simultaneously of both a mid- and fore-tarsus, or a fore-
tarsus and antenna, or a mid- and hind-tibia, and so on. The initial
hold thus seems fortuitous and not limited to a particular appendage
or to but one appendage at a time. Depending upon the particular
grip of the gonostyles, and the appendage (s) seized, a male may
either face opposite to the captured female (7 cases), and may even
be chiefly behind her, or face in the same direction as his partner.
When a female’s femur or tibia has been grasped, a male, without
loosening or losing his hold, can generally draw his own body about
to a position at right angles to that of the female by forcibly rotating
her appendage, but he cannot wholly reverse the direction in which
he faced without obtaining an entirely new hold, as he must when
initially facing 180° away from his partner.

Once seized, the female’s response is immediate and energetic, as
though designed to free her from the clinging, intermittently passive,
male. She drags the male on his back, his side, or even vertically on
his hypopygium, over and through obstacles presented by the moss
and debris. Occasionally the male acts to resist, splaying his legs
outward as though a drag-anchor, or clutching at the moss, offering

2? Also Svensson (1966), whose account must refer to B. westwoodi, to
B. hyemalis, or to both.

Terminology referring to the external male “genitalia” follows Michener
(1944).

90

Psyche

[March

such resistance to the female as he can. From time to time the female
rests, and may even pause to feed at rosettes of young moss leaves.
At such times the male becomes enlivened, rights himself, turns the
length of his body at right angles to the female’s while still holding
his original grip with his gonostyles, and swiftly (and repeatedly)
arches backward over the female in an endeavor to gain firm grasp
of her body with his wings. If he fails and rests, the rebellion of the
female returns unabated and continues, as just described, until the
male succeeds in subduing her, or she finally rids herself of him.4

Should the male succeed in gaining a firm hold with his stiff,
spined and hooked wings, he deftly changes grip with his gonostyles
so that he now faces in the same direction as the female. Once a
position has been attained from which he can rear backward and to
the side of the female, and grasp her between her head and mesono-
tum with his wings, holding her body parallel to his own, he again
quickly moves his gonostylar grip forward. Once a sufficiently for-
ward grip with the gonostyles has been gained, the head of the female
now being behind the male’s forebody, he rears strongly backward a
number of times and rakes and manipulates the female’s rostrum
and antennae with his spined wings as he falls forward again. Should
he gain hold of the female’s pronotum with his wings, he may move
his gonostylar grip from the female’s legs to clutch one or both
antennae, and then briefly but smartly, drag her about by the anten-
nae. Mauling of the antennae is quickly followed by a wholly passive
state on the part of the female who thereafter stands as though mes-
merized. Surprisingly, on the initial assault in one case, the male
seized the female’s antennae with his gonostyles ; that female there-
upon became submissive without a struggle.

When the standing female has become passive, the male (still
anterior to the female and gripping an appendage with his gonostyles)
again bows backward repeatedly until his groping wings gain a secure
hold behind her head, to each side of the pro- or mesonotum, with
the female’s head pressed sharply and to the side by his flank. The
gonostylar grip is then moved as far posteriorly as the male can
manage, and the wing grip released. Once the male has the female
again firmly gripped with his wings, and the gonostyles reach suffi-
ciently far backwards, he tries to seize the ovipositor with his gono-
styles; ordinarily several attempts are required, and after each failure

4Marechal (1939) misremembers Lestage’s (1920) account when he states
“c’est la 9 qui saute sur le £, celui-ci cherchant a fuir et a s’en debaras-
ser!”, and goes on to tell still more of female sexual aggressiveness.

1974]

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91

Fig. 1. Mating of Boreus notoperates. A, mated pair preserved in female
perpendicular position; the medial tooth of the gonostylus has disengaged
(at death) from the lateral basal notch of the gonapophysis. B, full union
of male and female; medial tooth of gonostylus seated into the lateral basal
notch of the gonapophysis ; C, semi diagrammatic representation of mated
specimens cleared in clove oil ; as in A, the gonapophysis is partially with-
drawn from the endandrium, and the lateral basal notch of the gonapophysis
(at arrow) is clearly visible; dorsal proctigeral plate (closely, transversely
lined) lies within epandrial notch; gonapophyses stippled; aedeagus (widely,
transversely lined) partially inserted in common oviduct — note flap at
anterior margin of female gonopore and paired ejaculatory ducts entering
base of aedeagus; the sclerotized lamina that forms floor of endandrium is
cross-hatched (see text). D, male terminalia with everted aedeagus, apical
nipple of which bears the gonopore; upper left — inner face of right gono-
style to show median tooth and stylocavernula. All from camera lucida
sketches; scale: for A and B equals 1 mm, for C and D equals 0.5 mm.

92

Psyche

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the gonostyles are returned to a grip on the posterior legs, near the
coxae, and the male rests. When at last the ovipositor has been
grasped, the gonapophyses are pried down and slightly apart, the
female holding them, as though plastic, in whatever position they
were released by the gonostyles. The gonapophysis of each side is
then engaged between the inner tooth and the tip of the corresponding
gonostyle which, on adduction, directs and thrusts the tip of the gona-
pophysis into a pocket ventral to the epandrium (see below). Ratchet-
like, the two gonapophyses together are worked down and in by the
gonostyles. Finally each gonostyle becomes securely seated on the cor-
responding gonapophysis, for its inner tooth (figs. iD; 2A, B)5
engages the lateral notch of the gonapophysis near its base (fig. iB,
compare with iA and C, arrow).

The female is then released from the clasp of the male’s wings.
Thereupon she rocks backward 90° or more, with rostrum and an-
tennae folded between the forelegs, femora drawn up to the sides,
tibiae adducted, tarsi drooping — the “death-feigning” posture that
concludes the leap of a startled Boreus. Surprisingly, the female
thereafter remains vertically in that posture, though her legs relax,
and the male folds his wings to their usual rest position over the back,
fig. iA). In side view it can be seen that the tip of the aedeagus has
been fully inserted into the common oviduct (fig. iC). Despite the
unbalanced appearance of a male and female united thus at a right
angle, without any noticeable difficulty the male may run, climb,
feed, and hop, landing without loss of balance. If startled, the male
may leap several centimeters or more, landing in (or assuming) the
death-feigning posture, motionless and resting on his flank; the female
too death-feigns as before; after a few moments, the male returns
to his feet.

In all, ten cases have been timed from the first assault of the male
to intromission; these took from 6 minutes to 18, with a mean of
13 minutes (all at i7°-20°C). The total duration of intromission
has not been timed, but I have observed cases in which it was less
than an hour, others of more than several hours. Both sexes mate
repeatedly, with the same or different partners.

°Esben-Petersen (1921) was unable to see this median tooth in specimens
before him of B. brumalis and B. nivoriundus ; Lestage (1940) took the
supposed absence of a median gonostylar tooth as presenting a cardinal
character for Euboreus, a genus he erected for all American species. For
discussion of Euboreus, see Cooper (1972).

1974]

Cooper — Boreus

93

Discussion of Mating

The events just described are different in a number of significant
features from those in the matings of B. hyemalis , B. westwoodi and
B. brumalis, which are alike. Once a female of those species has
been seized by a male, it ordinarily becomes passive (but wot always,
see Syms 1934, Aubrook 1939, Sauer 1966, Crampton 1940). The
rested male then grips the female across the midbody with his wings
(as does B. notoperates) , and with them and his gonostyles works
her body over his dorsum so that it is axially symmetrical and paral-
lel to his own. The gonapophyses are then pried down and inserted
into the subepandrial pocket, the wing-hold is released and the female
rocks back until perpendicular to the male. Intromission very likely
occurs at this point, as it does in B. notoperates. The relative station
of the two sexes just prior to intromission, or the “pose” (Lamb
1922), in all four species is truly a “female vertical pose” with (pre-
sumably) an inverse correlation of the genital conduits of the two
sexes.6

Now Lamb (1922) used “vertical” as a contrasting term to
“linear” (or tail to tail) to denote arrangements of partners in which
one partner is above the other. In the overwhelming majority of such
cases, the body of the upper insect lies more or less parallel to that
of its partner. Consequently I shall call the pose common to the
four Boreus a “female perpendicular pose” to distinguish it. The
final copulatory attitude of B. notoperates does not depart from the
pose; accordingly it is a “female perpendicular position” (fig. iA)..
In contrast, B. hyemalis, B. westwoodi , B. brumalis, B. calif ornicus
(=var. fuscus Carpenter?) (Cockle 1908, 1914), B. nivoriundus
Fitch (Carpenter 1936; Cooper, unpubl.), B. unicolor Hine (Byers
1954) and B. vlasovi Martynova (Vlasov 1950), namely in all other
species for which the copulatory position has been recorded, the final
attitude differs from the pose, being a female vertical position.7

6On intromission, the apparent dorsal wall of the aedeagus of Boreus lies
in contact with the ventral wall of the common oviduct, hence in “inverse
correlation”, which is unusual (fig. 1C). In most insects correlation is
“direct”, or symmetrical, for contact is dorsal-dorsal and ventral-ventral ;
in many insects having a vertical position, direct correlation is brought
about by rotation of the male genital tract (see Lamb 1922).

Tor original drawings, or photographs (reference in boldface), of copu-
lating pairs in their final female vertical position, see: for B. hyemalis
fig. 3, pi. 8, Withycombe (1922), fig. 1, Steiner (1937), fig. 7, Striibing
(1958) ; for B. westwoodi, fig. 5, pi. 3, Brauer (1855), fig. 2, Sauer (1966) ;
for B. brumalis, fig. 9, Cooper (1940), fig. 1, Crampton (1940).

94

Psyche

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On the basis of its morphology, B. notoperates (along with the
similar B. brevicaudus Byers) has been adjudged the least primitive
of all known species of Bore, us (Cooper 1972). Interestingly, its
mating stands apart from that described for the other three studied
species: at its onset (the enduring resistance by the female), at its
midpoint (submission by the female only after antennal “abuse”),
and at its very end (a female perpendicular position derived without
change from a similar pose). In that last attribute, it differs from all
seven other species of Boreus for which the position has been re-
corded and, notably, that position is alike in all seven. The long
period of female coyness and the need for manipulation of the fe-
male’s antennae go hand in hand, but I cannot decide whether these
are primitive aspects or not. But the female perpendicular position is
almost certainly a specialization, the marked change from pose to
position that occurs in other species of Boreus most likely being primi-
tive. I suspect that the presumed loss of change is related to the
unusually shortened ovipositor of B. notoperates and the relative
depth to which the gonapophyses are inserted into the male; if that
is so, B. brevicaudus may be expected to have a female perpendicular
position, and possibly also the remote B. chadzhi-gireji Plikinsky
as well.

Because most Boreus follow closely similar mating patterns, as do
B. brumalis and B. nivoriundus , with poses and final positions that
are alike, the question quite naturally arises as to whether cross-
matings occur or are even attempted. My own trials on this score
were with smallish B. nivoriundus males and B. brmnalis females,
two species quite frequently found within common areas in New
Hampshire. They showed no mutual interest whatsoever. This was
not to be attributed to lack of potential competence or sexual aggres-
siveness on the part of the males for, when female B. nivoriundus
were added, the males soon paired, or attempted to do so, with the
females of their own species. Perhaps the scent or secretion claimed
for female Boreus (Hardy 1848, Withycomb 1922, Marechal 1939)
has a role as a species specific mating pheromone.

The Anatomical Correlations in Mating

The female vertical position is widespread among insects, yet not
common. It has been recorded for Orthoptera (where it, or its
equivalent, the “false male vertical position” of Richards, is the usual
mode), Plecoptera, Mallophaga, Anoplura, Mecoptera, Siphonaptera,

1974]

Cooper — Boreus

95

Diptera (as Empis) , and Coleoptera ( Lomechusa ) (Meisenheimer
1921, Richards 1927), as well as for aradids (Heteroptera) (Usinger
and Matsuda 1959). For the majority there is little information on
the correlation involved, but it is probably generally inverse, with
the anus of the female unobstructed, as in Boreus . The similarities
cease here, however, for so far as I am aware the species of Boreus
are unique in their adaptation to, and practice of, a reciprocal intro-
mission — namely a concurrent passage of the female’s gonapophyses
into a specialized genital pocket within the male, and of the aedeagus
into the common oviduct (fig. iC). This curious arrangement was
first commented upon by Cockle (1908, 1914), and later by Steiner
(I937) and Cooper (1940), but it has not properly been described.

Projecting cephalad from the medio-dorsal aspect of the gono-
coxites of the male, there are two sclerotized laterally flattened straps,
the dorsal apophyses of the gonocoxite, that expand medially and
unite to forjn a thin, broad, pigmented roofing plate that lies below
the anus (figs. 2A, B ; fig. 4, Cooper 1972). The more dorsal anus
is itself enclosed by sclerotized dorsal and ventral proctigeral plates
(segments X + XI?) (fig. 2A; fig. 3, Cooper 1972). 8 Ventral to
the gonocoxital roofing plate, or dorsal portion of the gonobase, there
is a broad, sclerotized band overlying the dorsum of the aedeagus
(figs. iC, 2A, B; also fig. 4 Cooper 1972). A wide, enclosed, shal-
low but elongated pocket is thus formed between the dorsal gonobase,
the muscles of the gonocoxites, and the sclerotized band over the
aedeagus. In the unmated male at rest, this pocket (which I call the
endandrium ) curves downward under the epandrium (fig. 2B).

Just prior to insertion of the aedeagus, the gonapophyses (sternites-
VIII, or “ventral valves of the ovipositor”) are thrust by the male
into the endandrium. The main fields of caudally-directed spines on
the bosses to each side of the epandrial notch (fig. 2A) catch on the
outwardly directed denticles of the blades of the gonapophyses, pre-
venting slippage outwards as the gonapophyses are initially forced,
ratchet-like, more and more deeply into the endandrium. Once fully
inserted within the endandrium, the gonapophyses are locked in place
there by the gonostyles of the male. Not only is the blade of each
gonapophysis grasped tightly between the apex and inner tooth of a
gonostyle, but that inner tooth of each gonostyle is in turn engaged
by being seated into the lateral basal notch of its corresponding gona-
pophysis (fig. iB cf. A, C, arrow). That union so forcibly locks the

8The proctigeral plates (= segment-XI) in the female of Boreus notoper-
ates are similar to those of B. brevicaudus, described by Byers (1961).

Fig. 2. Genital anatomy of male Boreus notoperates. A, terminalia
viewed obliquely from the right side. The epandrium (ep), or tergite-IX,
has been separated from its articulations on the right, and reflected about
60° to the left. The anus, enclosed by proctigeral plates (cross-lined) lies
immediately below the epandrial notch, the spined epandrial bosses to each
side. Gonostyles abducted on right, displaying in profile the short, sharp
medial tooth which engages the lateral basal notch of the gonapophysis.
The dorsal gonobase (dg) extends anteriorly from the gonocoxites, flaring
out as the roof of the anterior end of the endandrium. Median plaque (stip-
pled) immediately over the aedeagus forms the floor of the endandrium;
aed — partially everted aedeagus, ejr — right ejaculatory duct, hyp — hy-
pandrium, r — rectum.

B, partially exserted aedeagus (aed), right lateral view. Epandrium, proc-
tiger, right gonocoxite and hypandrium have been removed. Above, dg,
the left dorsal half of the gonobase; below, vg, the left ventral half of the
gonobase. Sclerotized plaque (stippled) overlying the aedeagus joined by
a fan of muscles to U-shaped sclerite (stippled) partially encircling the
aedeagus ventrally. The genital chamber is the space enclosed between
the dorsal and ventral gonobases, the endandrium lies between the dorsal
gonobase and the sclerotized plaque, and the aedeagal chamber lies between
the sclerotized plaque and the ventral retractor muscle attached to the ventral
gonobase; ejr — right ejaculatory duct. Both figures from camera lucida
drawings; scale equals 0.5 mm.

1974]

Cooper — Boreus

97

mated pair together that, unless the male’s grip is loosened, the female
cannot free herself. In all but B. notoperates , the female is then
rocked forward into the final female vertical position. Although the
spines of the epandrial bosses may once again act against denticles of
the gonapophyses’ blades, preventing slippage, and the male’s wings
give an added grip, the change in position seems a supererogation so
far as security of the male’s mating grip is concerned.

The aedeagus, at rest, lies within a chamber that may be distin-
guished from the endandrium. The roof of the aedeagal chamber is
the band-like sclerotization over the aedeagus which forms the floor
of the endandrium; the side walls are in part defined by the internal
faces and musculature of the gonocoxites, and the floor of the chamber
by the cephalad-directed apodemal band from the antero-medial,
ventral angle of each gonocoxite that joins its homolog as a broaden-
ing, very thin strap, the ventral gonobase (fig. 2B) which, in some
specimens, encloses a minute medial sclerotized plaque. The space
enclosed between the dorsal and ventral gonobases, then, forms the
genital chamber proper, and the endandrium and aedeagal chambers
are thus dorsal and ventral subdivisions of it.

The aedeagus itself is a heavily muscled tube containing the un-
paired ejaculatory duct. At rest it is hook-shaped, with its hooked
tip wholly withdrawn into the aedeagal chamber, resting upon the
ventral adductor muscle (fig. 2B). The paired ejaculatory ducts
separately enter the caudal end of the aedeagus (figs. iC, 2A, B),
uniting within it. Above the aedeagus is the sclerotized band that
serves as floor to the endandrium. Below, near the point at which
the aedeagus bends back on itself in its distal third, there is a slender,
sclerotized hoop which partially encircles the aedeagus on the ventral
side, from which a fan of muscles extends dorsally and caudally to
insert on the dorsal sclerotized band. In B. borealis Banks,9 and
B. coloradensis Byers, there is both a dorsal and ventral sclerotized
longitudinal band between which the aedeagus lies.

The contraction of the muscles running from the ventral hoop to
the floor of the endandrium both enables the aedeagus to be held and
packed in retraction within the genital chamber and, when the
aedeagus is everted, to reduce the space of the aedeagal chamber and
thus force the aedeagus caudally outward. No doubt, to judge from
the anatomical relations of the endandrium, the initial downward

The Whitney’s field-notes suggest that Banks, when describing B, borealis,
had but two of a total of four specimens that were captured; see records
129 and 157, pp. 136-137, in McAtee (1923).

98

Psyche

[March

thrust and pressure of the gonapophyses in the endandrium are addi-
tional (perhaps necessary) factors in the eversion and erection of the
aedeagus. Contraction of the annular musculature of the aedeagus
itself additionally extends the organ and causes the tip to inflate
into a bulb, bilobed dorsally, which abruptly terminates in a conical,
or nipple-shaped tip upon which the ejaculatory duct opens (fig. iD).
The fully extended aedeagus runs between, and is guided by, the
smooth inner faces of the gonapophyses and the ventral surface of
the ovipositor to enter the common oviduct, from the anterior ventral
margin of the orifice of which extends a chitinous guiding flap that,
at other times, closes the aperture (fig. i'C). Once entered into the
oviduct, the apical bulb of the aedeagus is inflated within the vesti-
bule of the common oviduct which opens at the junction of segments
IX and X. In that position the ventral bulge of the bulb evidently
blocks off the common duct of the tubular accessory glands of the
oviduct, and the conical tip of the aedeagus enters the short sper-
mathecal duct, within which there is a minute sclerotization (fig. 9,
Cooper 1972). Insemination therefore takes place directly into the
duct of the spermatheca.

The account just given will very likely be found to agree in its
main features with the complex “reciprocal intromission” of other
B'oreus, even though Martynova’s (1954) figures 1-6 hint that the
endandrium may differ in depth and conformation among different
species. However it is at variance with Steiner’s (1937) description
(or inference?) for B. hy emails in which the tips of the gonapophy-
ses are said by him to be received directly and separately into the
pockets of the epandrium. That is not the case in still other mated
Boreus (B. hrumalis, B. nivoriundus) that I have studied, and in
which endandrial insertion occurs. Nor is it likely that the condition
observed by Byers (1954) in a preserved, mated pair of Boreus uni-
color Hine is a normal state of affairs; namely that the tips of the
gonapophyses are bent across one another, and are held thus by the
gonostyles of the male at right angles across his epandrium. Nor does
B. notoperates conform to Potter’s (1938b) description for B. hy-
ejnalis of a coiled or folded ductus ejaculatorius (see her fig. 30)
which, at rest, is thrown into three flexures within the genital cham-
ber, and which is evaginated during coition. As Steiner said, the
intromittant organ is strongly muscular, and as I have shown it is in
effect erected without evagination comparable, say, to that of a true
internal sac. The element which Potter figures in front of the first
distal flexure, appearing continuous in her figure with the “ejacula-

1974]

Cooper — Boreus

99

tory duct,” is very likely the large retractor muscle in my figure 2B
which occupies that position in B. notoperates. Brief comment on
the aedeagus of Boreus will be found in Fitch (1847), Kolenati
(1847), Cockle (1908, 1914), and Byers (1954), as well as in
Withycombe (1926) and Crampton (1918, 1920, 1923, 1931) who
have figured it in an everted state. To judge from these figures and
my own, species of Boreus may differ in the apical morphology of
the aedeagus.

Cytology

Boreus notoperates has a diploid number of 20 in the female and
19 in the XO male. At spermatogenesis, the autosomes are conjoined
by chiasmata (nearly always but one per bivalent) that are interstitial
or proterminal at diakinesis, but for the most part terminalized by
metaphase-I. There is only a moderate premetaphase stretch.

The acrokinetic X is the largest chromosome of the set (from
7-9 jx long), being respectively two to five times the lengths of the
largest and smallest autosomes. The X possesses a nucleolus organ-
izer proximally in the long arm. Spermatogenesis is orthodox, the X
segregating precociously and reductionally at anaphase-I. At ana-
phase-II, as at anaphase in both spermato-and oogonial mitoses, the
separating chromatids of X stretch and span the entire length of the
elongating spindle before their distal tips disjoin.

The chromosome number of B. notoperates is the lowest so far
known for Boreus (B. bru?nalisJ, ncf — 11+ XiX2,Y; B. hy emails,
ncf ' — ' 14 + X,0; B. nivoriundus ncf — 15 + X,0; Cooper 1951
and unpubl.) ; except possibly for a cytological form of Nannochorista
dipteroides Till, (n = 9 ?, Bush 1967), it is indeed the lowest num-
ber so far claimed for Mecoptera. A second strain of N. dipteroides
(n — ca 14), Chorista australis Klug. (ncf = 14 + X,0) (both
in Bush 1967), and three species of Bittacus (ncf — 13 + X,0;
ncf — 14 + X,0; ncf = 15 + X,0) (Matthey 1950; Atchley
and Jackson 1970), all lie within the range of Boreus. My prepara-
rations of a single male of Merope tuber Newm. display chiefly very
drawn-out early diakinetic chromosomes, and are sufficient only to
establish that it is XO, chiasmate, and has a haploid number that is
less than ncf = 12 + X,0 — possibly ncf — 10 + X,0. Though,
at this time no more definite statement can be made for our most
primitive mecopteran, it is nevertheless clear evidence that the chro-
mosome number of Merope is not a relatively high one. On the other

IOO

Psyche

[March

hand, the seven species of Panorpa so far reported upon by Naville
and de Beaumont (1934), Kichijo (1943), Ullerich (1961), and
Atchley and Jackson (1970) stand apart, having the highest chro-
mosome numbers known for Mecoptera; they are: ncT = 20 + X,0,
ncf =21 + X,0, and ncf — 22 + X,0 (4 species, plus one with
n? = 23).

The cytology of Panorpa is thus characterized by two evolution-
ary differences from most other Mecoptera now known: absence
of chiasmata at meiosis in the male, as Ullerich (1961) first demon-
strated, and unusually high diploid numbers. Whether or not male
Nannochoristids and Choristids have chiasmata is not known, al-
though Bush’s (1967) account indicates that male Nannochorista
dipteroides , like Panorpids, may have a meiosis without chiasmata.

Despite the strongly derived nature of the external morphology of
Boreids, and especially of Boreus notoperates, therefore, their chro-
mosome numbers and possession of chiasmata at meiosis in the male
appear conservative. Congruent also with a primitive state is their
possession of panoistic ovarioles, as is known also for B. hrumalis
(Cooper 1940) and B. nivoriundus (unpubl.). It is likely that this
is the case also for B. hy emails; Steiner (1937) could find no nurse
cells in its ovaries, but was undecided as to whether the ovarioles
are panoistic or meroistic.

OviPOSITION

As commented above, both male and female Boreus mate repeatedly
and with varied partners. In B. notoperates the spermatheca of the
female has some 24 sperm receptacles (fig. 9, Cooper 1972), all of
which come to be filled with spermatozoa. The natural promiscuity
may be seen as tending to reduce close inbreeding within the small,
scattered populations. In the laboratory, females lay within two
weeks following their first mating, and perhaps within a few days as
is the case for B. hy emails (Striibing 1950, 1958).

The large majority of the eggs at laying (which measure from
0.5 to 0.6+ X 0.3 + mm, length by width)10 by B. notoperates are
placed singly, vertically, and ordinarily at such a depth that the apex
of the egg lies from 0.1 to 0.3 mm below the surface of the sod of
the moss, amid its rhizoids. Sixteen females, over a period of ten
days at 2i°C, laid 186 eggs that I was able to locate — assuredly a

10The egg widths in Cooper (1972), p. 275, should of course be 0.31 mm
and 0.30 mm respectively.

1974]

Cooper — Boreus

IOI

minimal number — giving an average of 1.16 eggs per day per fe-
male. The span of life of a female in such cultures was no more
than four weeks. Were such a rate of laying maintained, a labora-
tory-held female would be expected to lay about 32 eggs. At death,
these females possess mature eggs in their ovaries, and so their poten-
tial fecundity is higher than just estimated. These experiences are in
line with those of Withycombe (1922) and Striibing (1950, 1958)
for B. hyemalis , although their estimates that the average female lays
a maximum of ten or so eggs in a lifetime I suspect to be much too
low, considering the hazards of egg development, larval life, and the
full two-year life cycle that must be met. Females of B. brumalis
and B. nivoriundus which I have dissected at the close of their sea-
sons in late February and March, still have a few mature eggs in
their oviducts, as Striibing (1950) found for B. hyemalis but in
some there is widespread involution of follicles within the ovarioles,
the epithelium of the egg chambers incomplete, with many nuclei
lying within large protoplasmic masses that simulate nurse cells.

I have not witnessed oviposition, but Brauer (1855, see fig. 5,
plate III), Aubrook (1939) and Marechal (1939) have described
females, nearly perpendicular, with ovipositors buried in the soil.
Only Svensson (1966), however, has witnessed actual oviposition
(perhaps in B. westwoodi, as in Brauer’s case), and described the
passage of the egg along the incomplete tube formed by the gona-
pophyses and tergite-X. When the egg reaches the tip, the cerci are
flexed ventrally, forcing the egg out onto the ventral surface of the
gonapophyses. The egg is thereupon set down upon a moss stem, the
whole process taking 3 to 4 minutes. It is to be noted that the female
he observed laid 4 eggs within a period of two hours, lending some
credence to the belief that females of Boreus may actually lay at least
again as many eggs as Striibing estimates for the total production of
a female B. hyemalis. Indeed Striibing (1950) records one clump
of seven eggs, presumably laid by a single female. Certainly it is
reasonable to assume that laboratory conditions are not ideal for ovi-
position, and that such estimates do represent minima.

Hatching

Hatching occurred in laboratory samples of eggs of B. hyemalis
within 8 to 10 days, at 8.8 °C, with about 50% mortality in Withy-
combe’s (1922) series; in contrast, Striibing’s (1950) samples took
from 3 weeks at approximately 2Q°C to as long as one and a half

102

Psyche

[March

Fig. 3. Immature stages of Boreus notoperates. A, freshly laid egg.
B, egg just before hatching — note larval head, transverse band, margins
of reflexed end of abdomen. C, rent chorion following hatching. D, first
instar larva. E, first instar larva dissected from egg at stage comparable
to B. F, last instar larva — especially note conformation of pigment under-
lying stemmata (KOH preparation). G to I, pharate pupal heads — note
change in both position and conformation of eye pigment, and its relation
to the development of the imaginal eye. J, pharate adult female — note
connections between imaginal fixed setae of gonapophyses and corresponding
setae of pupal exuvium. Scale = 0.5 mm, applies to figs. F through I;
scale = 1 mm for J ; for dimensions of eggs and first instar larvae, see text.

months at 7°C, and 44 of 46 eggs of her series hatched. In the case
of B. notoperates, two of four eggs kept at 20°C hatched in a period
of 24 days; the other two were at the stage of hatching at that time,
but the larvae died without perforating the chorions.

The four freshly laid eggs of B. notoperates which I followed
swelled, in the first 15 days, from 0.5 X 0.3 mm to 0.66 X 0.46 mm
— 0.68 X 0.50 mm, at which time the larval head, pigmented eyes,
jaws (mostly held open, but moving from time to time), and reflexed
abdomen could be seen, as well as a grayish, transverse band that had
appeared on or near the chorion below the larval head (figs. 3B, E).

1974]

Cooper — Boreus

I 03

By the next day (day 16) the larvae moved occasionally within the
chorions. By day 18 the larval gut appeared swollen and gray. Over
the next five days the larvae became very active, even reversing their
positions within the eggs; at times the chorion of an egg would col-
lapse inwards, only to reinflate once again. On the 24th day two of
the larvae hatched by means of great rents torn through the chorion
(fig. 3C). I did not observe the emergence, or how the chorion was
torn open. As there is no egg burster such as Gassner (1963) has
described for the first instar larva of Panorpa nuptialis Gerst., it is
possible that infolding of the chorion, as earlier observed, enables the
larva to seize and rip the chorion with its jaws, or that the gray band
on the chorion is associated with a local weakness. Striibing (1950)
has described the swelling of the developing egg (which also occurs
in other Mecoptera, see Currie 1932, Setty 1940, Byers 1963), and
the hatching of B. hyemalis which takes but 10-12 minutes; the
events prior to hatching are similar to those of B. notoperates}1

The newly hatched larva of B. notoperates (fig. 3D) is dead
white, except for the pale amber head capsule, darker jaws, and pig-

"Some 127 developing eggs of B. notoperates were at hand when this
passage was written (Feb. 1974). Subsequently they were divided into two
lots of which one (of 88 eggs) was followed daily to hatching. All had
been kept at 9°C, and were at a mean age of approximately 30 days when
removed to 20°C for observation. They had all undergone swelling, but no
other sign of development was discernible. The smallest egg was 0.58 X
0.44 mm, the largest 0.78 X 0.52 mm, and the most distended in girth was
0.64 X 0.56 mm. In what follows, day 0 is the day of removal (Jan. 14,
1974) to 20°C.

At day +15 eyes were visible in nearly all eggs; at +21 heads pale
testaceous, eyes black, mandibles very dark and active ; most eggs alike in
stage — despite this, hatching spread out over a period of 36 days.

Hatch was 93% (82 of 88 eggs; 2 did not mature, 3 failed to hatch but
had had active larvae, 1 killed by mold). First egg to hatch was at +25
days, half of the eggs had hatched at +31, three-quarters at +33, 90% by
+ 38, and the last on +61.

Unlike the mandible of the last larval instar, that of the unhatched larva
has a sharp, falcate, apical tooth, a moderately long, sharp, subapical tooth,
and two successively smaller, sharp denticles. When hatching commences,
the turgid chorion is abraded, then penetrated, by the mandibles, whereupon
it collapses somewhat and folds inward to a varying degree, owing to loss
of fluid from the egg. Within an hour, or as long thereafter as several
days, the larva emerges. It does so by cutting through the fold, and else-
where, to produce a broad apical flap, or even a free cap to the egg. By
peristaltic movements, especially of the thorax, the larva forces the flap
and emerges through the gaping hole in the chorion in a matter of ten to
twenty minutes.

104

Psyche

[March

merited eyes. It has a head width of 0.28 mm, widest girth across
the thorax of 0.3 mm, and a body length of 0.9 mm. It is not pro-
vided with abdominal prolegs, as described by Brauer (1855, fig. 7,
pi. Ill; 1863, fig. 2, pi. XIV) for B. westwoodi, but the terminal
abdominal segment does serve, and very effectively so, as a holdfast
as he believed.

The two hatched larvae were placed on a small mat of damp moss
into the sod of which they speedily disappeared. Seventeen days later
the larvae were torpid, stretched out and turgid, and remained so for
the next four days. At that point the head capsule was 0.3 mm wide,
the metanotal girth 0.38 mm (the broadest portion of the thorax),
and the total body length 1.04 mm in one, 1.42 mm in the other. If,
at this point, they were ready for the first larval moult, then the first
instar is completed within 16 to 17 days at 20°C. Unfortunately
observations had to be discontinued, and the larvae were preserved.

Striibing (1950, 1958) has shown that the rate of development in
B. hyemalis is strongly affected by temperature and, unlike the adult,
eggs and larvae develop only very slowly, if at all, at the low tem-
peratures of winter. Indeed the egg remains dormant through the
winter months, normally hatching in the period from the end of
March to mid-April. Hatching is almost certainly earlier in the case
of B. notoperates for, by the close of January and February, air
temperatures above 20°C and warm sunshine are not uncommon in
its habitat on Mt. San Jacinto.

Larva

During the last two weeks of August and in September, larvae,
pharate pupae (Hinton 1971, 1973), and pupae of B. notoperates
are found in the sod of host mosses, amid the rhizoids. The apparent
larvae are of two size classes : ( 1 ) with a mean head length from the
tip of the labrum to the occiput of 0.49 mm, range 0.47-0.52 mm,
and a coefficient of variation (V) of 5.7 (6 specimens), and (2)
with a mean head length of 0.68 mm, range 0.59-0.87 mm, and a
surprisingly large V of 9.7 (23 specimens). Though smaller larvae
of the second class proved to be males, and the larger females, the
size distribution is not bimodal. No doubt their high coefficient of
variation is in part a consequence of two factors: a sexual difference
in size at the last instar, and the consequences of the larval-pupal
apolysis that closes the larval stage and which I did not detect until
later. They proved a heterogeneous lot of last instar larvae and
pharate pupae.

1974]

Cooper — Boreus

105

Fig. 4. Head of last instar Boreus notoperates. A, frontal view. B,
mouthparts in ventral aspect. For details see text. Full scale = 0.4 mm
for A, = 0.2 mm for B.

The number of larval ecdyses is not known for any species of
Boreus , but it is very likely four as in other species of Mecoptera
(Currie 1932, Setty 1940, Byers 1963). The ratio of the mean head
length of the larger larvae and pharate pupae to that of the smaller
larva is 1.38. This gives estimated head lengths for the two im-
mediately preceding larval stages as 0.36 mm and 0.26 mm respec-
tively. As the head length of the first instar larva is known to be
O.32 mm, the series ends there and the number of instars is thus
very likely four. The probable Dyar constant of 1.29 (which is low
compared with Byer’s figure of 1.46 for Panorpa nuptialis) gives a
calculated head length of 0.41 mm for instar-2, of 0.49 mm for
instar-3, and of 0.68 mm for instar-4. Striibing (1950) left the
matter of the number of larval instars of B. hyemalis open. Withy-
combe (1922), also unable to find larval exuvia, judged from head
capsule growth that there are at least four instars. Though Striibing’s
measurements of larval head capsules are in fact heterogeneous, they
are reasonably well satisfied by a Dyar constant of 1.34 and four
instars.

The full grown larva of B. notoperates is remarkably similar to
the first instar larva (compare figs. 3D and F). The head is pale
amber in color, the body white to pale yellow, and in life it ranges
from 2. 6-3. 7 mm in length. In shape it is “scarabaeiform,” to use
Peterson’s (1951) characterization of the larva of B. brumalis. As

io6

Psyche

[March

illustrated (fig. 3F), the larva of B. notoperates has localized patches
of denticles on the dorsum of the pronotum, a transverse row of
setae on the mesonotum, two rows of setulae enclosing a row of setae
on the metanotum, an oblique scattering of denticles on the flanks of
the metanotum, and patches of denticles on each of the indistinctly
3-segmented legs. Abdominal tergite-i has a dorsal transverse patch
of denticles behind which stands a row of setae. Abdominal tergites-2
through 5 have dorsal, transverse patches of denticles, and each seg-
ment thereafter possesses only a transverse row of setae, except for
segment- 10 which is encircled by a row of setae. Though not ap-
pearing unusually modified, the apical abdominal segment serves, as
in the first instar larva, as an adhesive holdfast. Denticles and setae
alike are pale testaceous. As usual for Mecoptera, there are 9 spira-
cles: one on the flank of the pronotum, and one on each of abdominal
segments- 1 to 8.

The larval head (fig. 4A) is convex and more or less ovoidal in
face view. At the bases of the mandibles, lower halves of the antennal
sockets, on the transverse sclerotized plaque on the proximal portion
of the clypeus, the head is bright castaneous, as are the mandibles
(except for their piceous tips and the cervical sclerites), all contrast-
ing markedly from the pale ground color. The eyes possess three
stemmata, the lower two of which have very convex lenses while the
uppermost is smaller, flat and somewhat atrophied. Underlying and
surrounding the stemmata is a distally attentuated patch of black
pigment. The antennae have two obvious segments, and terminate
in a bristle-like prolongation that may or may not be a true segment.
Coronal, frontal and epistomal sutures are well marked and complete.
The clypeus is roughly trapezoidal, the labrum apically emarginate;
together they conceal the biting edges and tips of the closed mandibles.

Almost all of the setae described and designated by Boese (1973)
for the larval head of Panorpa species are to be found, similarly situ-
ated, in B. notoperates. The differences between B. notoperates and
Panorpa species lie in the absence in Boreas of a pair of setae (SO)
immediately dorsal to the eye, the presence of but a single pair (in-
stead of 2 pair) of setae (SI) on the labrum which, in addition, are
not marginal as in Panorpa , the presence of but a single pair of setae
on the mandibles (instead of 3), and the possession of an extra pair
of setae, lacking in Panorpa , on the proximal edge of the medial
third of the sclerotized clypeal band, just below the epistomal suture.
In view of the notable differences between Boreas and Panorpa in

1974]

Cooper — Boreus

107

other aspects of larval and adult anatomy, the complex setation of
the larval head appears to have been remarkably stable.

The mandibles of B. notoperates (fig. 4B) are massive, triangular,
strongly sclerotized, and thrown apically into three, poorly demar-
cated, blunt teeth. Maxillae and labium are membranous. The max-
illary cardo is triangular, the stipes large and trapezoidal, the palpi-
fer (which seems a basal palpal joint) cylindrical and surmounted by
the subequally 2-jointed maxillary palps, the terminal joint of which
is cylindrical and densely papillate apically. A brush of closely set
long hairs is directed medially from each maxillary lobe. The 2-
jointed labial palps are slender and very elongate. Like the maxillary
palps, they are testaceous, and bear a cluster of small pegs at their
tips. The prementum, which is strap-like and sclerotized, bears two
stout, convergent setae medially at its anterior margin. The sub-
mentum is large, trapezoidal, and broader posteriorly.

The numbers and arrangements of setae on the submentum, pre-
mentum, basal joint of the maxillary palpi, and possibly of the cardo,
are similar to those figured by Potter (1938a) for B. hyemalis. B.
notoperates , however, appears to have one less seta on both the palpi-
fer and stipes, and the conformation of palps, maxillary brushes,
prementum, and mandibular teeth differ strikingly from that species.
It also differs from B. hyemalis in shape of head, the presence of a
complete frontal suture, a distinct epistomal suture, a sclerotized
clypeal band, emarginate labrum, and details of setal pattern. Brauer’s
(1855, 1863) descriptions and figures of B. westwoodi do not permit
a comparison with B. notoperates, but I agree with Potter (1938a)
that there is no spinneret on the labium as Brauer considered possible
and as Withycombe (1922) claimed. If Brauer is correct that the
labial palps of B. westwoodi are 3 -jointed, that is a very striking dif-
ference from B. hyemalis and B. notoperates. Peterson’s (1951)
brief description and figures are sufficient to show that the larva of
B. brumalis has notable differences from both B. hyemalis and B.
notoperates in head shape, labrum, sutures (only an epistomal suture
is figured), and setal distribution and lengths on both head and body.

It is important to emphasize that presence of a distinct epistomal
suture in the larvae of B. notoperates, B. brumalis, and B. westwoodi,
because its supposed absence in Boreus is one of the characteristics
emphasized by Hinton (1958) as a feature of Crampton’s (1930)
suborder Neomecoptera, only diffidently suggested by Crampton,
which Hinton elevated to ordinal status. Nor is it the case, as Hin-
ton supposes, that the eleventh abdominal segment of adult female

io8

Psyche

[March

Bore us differs from that of other Mecoptera by lacking cerci; Byers
(1961) showed their presence in B. brevicaudus , and I confirm their
presence in B. notoperates as well. As Hepburn (1970) has already
said, attempts to remove Boreidae to an order separate from the
Mecoptera seems unjustified at this time.

Pupa

The metamorphosis to the pupa occurs in mid-August and in Sep-
tember in B. notoperates as in the other Boreus for which pupation
has been recorded. Its onset is first recognizable by a separation,
elongation, hypertrophy and migration of the pigment of the larval
eye. The pigment becomes aligned and compacted as three nodes
which, at 20°C, over a period of a week or more, move as a unit
posteriorly and obliquely to the orbital region of the head (figs.
3F-I). For most of this period the larval jaws are functional, al-
though feeding does not occur, and the larva moves about readily if
disturbed.

During migration of the pigment, the purplish compound eye de-
velops as though its posterior edge, defined by the three nodes of
black pigment, were its developmental origin of growth. By the time
that the imaginal eye is nearly fully outlined (fig. 3I), the larval-
pupal apolysis has been completed, and the pharate pupa no longer
can move the larval mandibles. Evidently this remarkable sequence
occurs in at least some beetles as well. The migration of the pigment
of the larval eye to the rear of the head in Duvalius mallaszi subsp.
chappuisi Jean., described and figured from preserved specimens by
the puzzled Jeannel (1926), is certainly an example of the same
phenomenon ; his specimens in which the pigment had separated from
the stemmata are clearly pharate pupae.

It is during the migration of the components of the developing
pupal eye that spermatogenesis occurs in B. brumalis (Cooper 1940)
and B. nmoriundus, and not in the last instar larva as I stated earlier.
In B. notoperates , however, only spermatogonial divisions and early
prophase stages of the first meiotic division occur in the pharate pupa,,
the meiotic divisions and spermiogenesis taking place in the late pupa
and pharate adult.

Within one to several days following apolysis, the larval-pupal
ecdysis ensues. Initially the pupa, in contrast to the greasy-yellow,
late pharate pupa, is a translucent white with the tips of the mandi-
bles brown, and the eyes a purplish brown. Within a week the body-

1974]

Cooper — Boreus

109

yellows, the moveable mandibles become testaceous to their bases, the
eyes brown, yet with the three contrasting black nodes at their caudal
margins distinct, and ocelli purplish, the malpighian tubules brown,
and the bristles amber-brown. Within two weeks the eyes have be-
come black, the three pigment nodes are still discernible, the ocelli
are dark sepia, and the malpighian tubules are a dark purplish-brown.
At five weeks the jaws and body are still moveable, but the pupal-
adult apolysis has occurred. It is interesting that in the pharate adult
female, the heavy black fixed setae of the gonapophyses attenuate
near their midpoints; their threadlike prolongations (which are lost
at ecdysis) connect with the corresponding setae of the pupal cuticle
(fig- 3 J)- The pharate adult stage, during which the entire body,
wings and legs of the imago darken, may last from ten days to three
weeks. The pupal to adult ecdysis thus takes place (in the labora-
tory) from a month and a half to nearly two months after the onset
of pupation. Unlike B. hyemalis (Withycomb 1922, Fraser 1943,
Striibing 1950) and B. westwoodi (Brauer 1857, 1863), in which
the newly eelosed adult requires from a number of days to as much
as a week to reach full color, but like B. brumalis (Williams 1916),
B. notoperates is fully colored within a half day following eclosion.
Just how long the true pupal and adult stages last in other Boreus
is not known. Until Hinton (1971, 1973) straightened out the
matter, and called attention to it, the importance of dating the stages
from their apolyses, rather than from their ecdyses, had not been
appreciated. Striibing (1950, 1958) gives a duration for the “pupa”
of B. hy emails (as from) 40 to 59 days.

Habitat

The larvae and pupae of B. notoperates are found in the sods of
mosses growing on diorite boulders from moderate (2 ft. dia.) to
gigantic size. The immature stages occur in a very scattered distribu-
tion, and I have not been able to forecast, when the moss is of an
appropriate species, whether or not a given moss-covered boulder
will be found to be inhabited by Boreus. Ordinarily specimens are
few amid the moss of a given boulder but, when found, tend to be
clustered.

Initially I was unable to find immatures in the dry summer
months. Because the mosses and their sods are very compact, desic-
cated and crumbly at that time, and frequently at a. temperature
(when sunned) of 30°!C or more, it seemed likely that, following

no

Psyche

[March

melting of the snow, the young stages were washed to the ground,
thereafter burrowing into the soil. Following that, they would forage
as saprophiles, since Withycombe (1922) has found that larvae can
feed on leaf mold. Promising moss-covered boulders, standing on a
loose, fine and sandy soil, were therefore trenched on the run-off side
to a depth off a foot or more, and the soil carefully sifted through
Tyler screens down to 16 meshes to the inch (0.99 mm openings).
This seemed a hopeful attack at the time because Strubing (1950)
had found that B. hyemalis, during periods of drought, descend to a
depth of as much as 20 cm below their mosses, which grow on soil,
to reach a suitably humid surrounding. No Boreus were found.

During one collecting period in mid-August there was a brief but
heavy shower which, within 20 minutes, brought the mosses on the
boulders to a fresh, expanded green state, and soaked their sods
through. When the damp sods were broken or sliced open, there
amid the rhizoids were the larvae and pupae within small ellipsoidal
and cylindrical spaces that had readily cleaved open ! During dry
periods these cells, which are only slightly larger than the stages
enclosed, harden and act thereafter as a coherent whole, appearing
as no more than larger, compact particles of the sandy soil and
organic debris of the desiccated sod. When such sods are broken and
crumbled, these cells remain intact, even under the stresses of sifting.
Their inner walls are very smooth, and are perhaps cemented by
salivary secretions, but not by silken threads as has been suspected
or claimed for the pupal chambers of B. westwoodi (Brauer 1863)
and B. hyemalis (Withycombe 1922, Syms 1934). It is possible that
salivary secretions also make their walls impervious to water loss.
Thereafter search became routine during the predominantly dry
periods for the desiccated mosses on their boulders need only to be
soaked with water.

It is clear that B. notoperates is well adapted by its earthen cham-
ber for survival during periods of drought. T he larva must do most
of its foraging during the spring before the moss turfs become dry,
and thereafter as infrequent opportunity permits following scattered
storms on the mountain.

Associated Mosses

Records of the mosses with which Boreus is associated in one way
or another have been made 'for only seven or eight of the twenty-
seven or so species of Boreus known (species list in Svensson I972)>

1974]

Cooper — Boreus

1 1 1

and for but four species are the mosses known upon which the larvae
feed. I shall bring these records together and discuss them at another
time. Suffice it to say that more than forty species of moss are in-
volved, and that these are distributed among 9 orders, 19 families,
and 28 genera. The families of mosses for which a. member (or mem-
bers) is known to support the life history of a species of Boreus are
Dicranaceae, Bryaceae, Mniaceae, Thuidaceae, and Polytrichaceae.
Records which follow for B. notoperates now add two additional
families: Grimmiaceae and Orthotrichaceae. The generic and specific
names are those recognized by Crum, Steere and Anderson (1973).

B. notoperates uses as food and habitat for its larval stages: Grim-
mia apocarpa Hedw., G. montana B.S.G., Rkacomitrium sudeticum
(Funck.) B.S.G., and Orthotrichum rupestre Schleich. ex Schwaeger.
Each is widely distributed, being found in Europe, Asia, and North
America. Adults have been collected on all of the above, as well as
on the following: Grimmia laevigata (Brid.) Brid.; Tortula prin-
ceps De Not. and T. ruralis (Hedw.) Gaertner (Pottiaceae) ;
Homalothecium aeneum (Mitt.) Lawt., IT. nevadense (Lesq.) Ren.
& Card., II. pinnatifidum (Sull. et Lesq.) Lawt., and Camptothe-
cium amesiae Ren. & Card, (all Thuidaceae). The identifications of
all of these mosses have been made by Professor Lewis E. Anderson
of the Department of Botany, Duke University, Durham, N.C., and
voucher specimens have been retained in the herbarium at Duke
University.

Adult B. notoperates will browse in the laboratory on all of the
mosses listed above. But of the mosses upon which only adults have
been collected, only Grimmia laevigata and the two species of Tortula
are widely distributed (Eur., Asia, N. A.), and by the nature of
their rhizoids and sods are likely candidates as larval habitats and
food. The sods of the mosses in which larval and pupal B. notoper-
ates have been found are compact, but surprisingly thin, falling
within the range from 5— 10 mm when dry, and from nearly 6-12
mm when thoroughly damp. The Thuidaceae, on the other hand, are
all endemic species. Because their clumps are coarse, open, and gen-
erally without a suitably fine rhizoid mat and sod, they are probably
not used as larval habitat and food mosses.

It is of particular interest that the mosses which serve as larval
food for B. notoperates are very widespread; also that B. notoper-
ates is not limited to a single species or genus of moss but in fact
utilizes members of two families representing different orders (Grim-
miales and Isobryales). Whatever limits the geographic distribution

1 12

Psyche

[March

of B. notoperates , it is certainly not the availability of a local or
special moss. That generalization holds as well for those other species
of Boreus for which associations with mosses have been specifically
recorded; nearly all of the mosses concerned are widespread in Eu-
rope, Asia and North America, and indeed quite a number have a still
more extensive range than that.

Variation Among Adults

When B. notoperates was described (Cooper 1972), my series of
29 specimens was nearly uniform with respect to antennal joint
number, namely 19 joints — a feature used as a specific character.
The only apparent exceptions that I noted I did not take to be such;
they were a male and female, each of which had an incomplete sep-
aration of joints-3 from 4, but these joints were otherwise demar-
cated by their apical swellings. Since then I have found that indi-
viduals do occur with antennal joint numbers other than 19—19.

In a series of 86 males and 83 females held to determine their
relative lengths of life in the laboratory (me? = 16 days, m$ = 22
days; 0.05 > P > 0.02), the distributions of antennal joint numbers
were :

18-18 18-19 19-19 19-20 20-20 Total

cT c? 12 4 68 1 1 86

9 9 9 3 70 o 1 83

Thus about 12% of B. notoperates have a different antennal joint
number than 19-19. When numbers of 19-19 individuals are com-
pared with the lumped totals of the others, there is no evident dif-
ference between male and female (0.8 >P >0.7). Males with
antennal joints fewer than 19-19 do tend to die somewhat earlier
than the other males (m = 11 days), the Wilcoxon rank sum test
giving 0.02 > P > 0.01. Such females also die somewhat earlier
(m = 19 days), but not significantly so, for 0.5 > P > 0.4.

Attention is also called to new information on patterns of ab-
dominal fusions in males (Cooper 1 973 ) •

Predators, Parasites and Life Cycle

Very little has been recorded regarding the use of Boreus as prey.
Withycombe (1922) was of the opinion that the adult would be
expected “to be speedily devoured” by birds, unless distasteful. Greve

1974]

Cooper — Bore us

113

(196) holds Boreus to be acceptable to small birds as welcome sup-
plement to their winter fare, and that Boreus ’ escape leap and thano-
taxis perhaps offer it some protection against predation. No one seems
to have observed birds feeding on Boreus, however, and McAtee’s
extensive analysis of the stomach contents of birds provides no records.
Lestage (1940), on the other hand, cites a unique record of the
remains of Boreus from the stomachs of trout, even adding “[les]
Truites ont montre celles-ci friandes des Boreus”! Other possible
predators that have been mentioned are spiders, often abundant on
snow during warmer days, and rapacious insects (Vlasov 1950, Greve
1966) — but none have been shown to be such. Definite information,
however, can be given on actual and probable parasites of species of
Boreus.

In my own experience, I have but once found mites on Boreus.
Four adult individuals of a species of Pediculaster (Pyemotidae;
determined by Prof. Earle A. Cross) were in neat parallel array,
lengthwise to abdominal sternites-i and 2, nearly concealed between
the abdomen and the dorsal faces of the metacoxae, on which they
were seated, of an adult male of B. brumalis (Hanover, N. H.,
Jan. 8, 1961, air temp. i.7°C). The record is of special interest for
it is a likely, occasional parasite of Boreus. Insofar as the habits of
species of Pediculaster are known, they are ordinarily phoretic on a
number of diverse flies, and Vitzthum found that P. mesembrinae
(Canestrini) drop off their hosts at oviposition sites, attacking there
the developing fly larvae (see Cross 1965).

A not surprising second parasite of Boreus is a Cor dy ceps (Sphae-
riales, Clavicipitaceae) that in one case of 5 such emerged from the
intersection of the frontal and coronal sutures, as well as from be-
tween the 9th and 10th abdominal tergites of a mummified 4th instar
larva of B. brumalis found erect in its vertical burrow just beneath
the basal stems of the host moss, Dicranella heteromalla (Hedw.)
Schimp. (Princeton, N.J., Nov. 19, 1939). The portion of the
fungus from the head bifurcated into two fruiting bodies (each about
3-4 mm long), extending vertically upwards and nearly parallel.
The limb from the abdomen bent sharply and vertically upwards,
and was nearly 5 mm long. In all, the 5 Cordyceps-iniected larvae
were among 25 + larvae collected that day from one large turf of
the moss growing at the base of a tulip tree. Of 35 pupae collected
at the same site and time, none were afflicted by the fungus.

The only parasites that have been recorded from B. hyemalis are
hymenopterous. The braconid Dyscoletes lancifer Haliday was shown

Psyche

[March

114

by Aubrook (1939) and Fraser (1943) to be a parasite of the larva.
Withycombe (1922) had earlier reported hymenopterous parasites of
the larva, but it is by no means certain that they were also Dyscoletes.
Indeed, of the single larval parasite that Withycombe endeavored to
rear, but which was destroyed by mold as a pupa, he says that it
appeared to be that of an apterous form. At Princeton, N. J., I too
obtained hymenopterous larvae from larval B. brumalis in the au-
tumn, but was unable to rear them.

From 13 B. notoperates larvae collected at Coldwater Canyon on
Mt. San Jacinto (Aug. 22, 1973), 17 hymenopterous larvae emerged
(3 from 1 larva, 2 from each of 3 larvae, and 1 from each of 8
larvae) ; one additional larva was obtained from a larval cell of
Boreus containing the remains of the mecopterous larva. By Febru-
ary 1, 1974, only 5 specimens (4?, 1 cf ) had transformed to adults.
All are a megaspilid (Ceraphronoidea) , and are tentatively identi-
fied as an undescribed species of Conostigmus.12 It is an interesting
association, for so far as I am aware no ceraphronoid is known to
have a mecopterous host, although there are records of neuropteroid
hosts ( e.g Muesebeck 1959, Dessart 1967).

Withycombe (1922), who thought — like many others — that
Boreus has an annual cycle, was puzzled as to what alternative host
a hymenopterous parasite of Boreus might attack from August to
December. This, so that it could have a brood appearing early in the
new year that could once again parasitize a new generation of Boreus.
Syms (1934) and Aubrook (1939), however, conclude from their
finding in the autumn, of two stages of the larva of B. hyemalis,
that Boreus probably has a 2-year life cycle, and Syms actually showed
this to be so for at least some larvae. That it does in fact have a
2-year cycle has been proven conclusively by Strubing (1950), and it
is very likely that all Boreus have a cycle that normally takes 2 years
— * assuredly that is so for B. notoperates , B. brumalis, and B. nivo-
riundus, which I have studied. The problem that Withycombe posed
thus vanishes, for the larvae of a second generation of Boreus are
available at any time that the hymenopterous parasite emerges and
is ready to oviposit. Such parasites of Boreus , therefore, could be
specific parasites, having no other host.

In the case of the larva of B. notoperates , confined as it is to hard
earthen cells for perhaps half of its life, it is likely that the parasite

12Dr. Paul Dessart, Chef de Travaux, Entomologie, Institut Royal des
Sciences Naturelles de Belgique, Bruxelles, has very kindly confirmed the
generic placement of Conostigmus (March 11, 1974).

1974]

Cooper — Boreus

1 15

can attack its host only during periods of dampness when the larva
of Boreus is free to move about; since the parasite pupates, or may
enter diapause, enclosed within the cell of its host larva, parasite and
host are thereby synchronized to the same periods of activity. The
very means by which the larvae of B. notoperates withstands drought
thus entrains the parasite to the host’s timetable.

Acknowledgment

I wish to thank those who have helped me in various ways during
the course of this work. Especial thanks are extended to Prof. Lewis
E. Anderson, of Duke University, for his cheerful acceptance of the
numerous samples of mosses sent him over the years for identification.
Other plants, still to be mentioned, were collected and identified by
Mr. J. W. Stubblefield to whom I owe special thanks. Both he, and
my wife, Dr. Ruth S. Cooper, gave me much help with field work
that, at times, I could not otherwise have carried out. Prof. Earle
A. Cross, of Northwestern State College, La., made the identification
df the phoretic mite. Prof. G. W. Byers, of the University of Kan-
sas, obtained for me Tarbinsky’s and Vlasov’s articles, not available
in this country, from a Russian colleague. Prof. Peter Jorgensen, of
the University of California at Riverside, helped me with the Scandi-
navian articles, and as before I have been assisted with Russian works
by both Prof. Anatole Forostenko, also of U. C. R., and my son
Geoffrey K. Cooper. Errors of interpretation, if any, are however my
sole responsibility. Finally, it is a pleasure to acknowledge my con-
siderable indebtedness to Prof. Frank M. Carpenter, of Harvard
University, whose unsolicited kindnesses in sending me specimens and
literature otherwise unavailable to me, and in other ways, have been
of such help in the work on Boreus and other studies.

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119

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1927. Sexual selection and allied problems in the insects. Biol. Rev. 2:
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81-83.

EMERGENCE PATTERNS OF THE SUBALPINE
DRAGONFLY SOMATOCHLORA SEMICIRCULARIS
(ODONATA: CORDULIIDAE)

By Ruth L. Willey*

University of Illinois at Chicago Circle

Introduction

The genus Somatochlora is circumpolar and many of its species
reach high into the subarctic and subalpine regions. S. semicircularis
occurs in the Canadian and Hudsonian zones of the North American
Western Cordillera from Alaska to Colorado. Towards the southern
part of its range, this species is found at high altitudes in Colorado
where adults have been recorded at 3000 to 3700 m (Gloyd, 1939;
Walker, 1925; Carpenter, 1873). Larval habitats range from 2700
to 3600 m in shallow ponds and water meadows. These may well be
the highest breeding localities for any odonate known.

The alpine and subalpine habitats offer extremes in temperature
and insolation during the short growing season which follows the late
snow melt (Mani, 1968). The subalpine regions, particularly, pre-
sent relatively the highest day and lowest night temperatures (Bill-
ings and Mooney, 1968). Coupled with these factors, the frequent
periods of drought and general unpredictability of climate contribute
to the rigor of the high altitude habitat. Studies of life history strat-
egies of insects which have adapted to high altitudes have revealed
cold resistance and lowered temperature thresholds, a predominance
of diurnal activity, a seasonal delay and shortening of the flight
period, and a general increase in the length of the life cycle coupled
with overwinter hibernation or diapause (Mani, 1968). In the
dragonflies, the increased life cycle is accomplished by elongating the
larval period to as much as three or four years (Paulson and Jenner,
1971; Robert, 1958). Somatochlora semicircularis larvae are also
resistant to dessication when their ponds and water meadows dry up
annually (Willey and Eiler, 1972). An ecologically sensitive part
of the life history for a dragonfly is the emergence period during
which the “aquatic” larva leaves the water, transforms into the

^Department of Biological Sciences, University of Illinois at Chicago
Circle, Chicago, Illinois 60680; and The Rocky Mountain Biological Labora-
tory, Crested Butte, Colorado 81224.

Manuscript received by the editor February 7 , 1974.

2

122

Psyche

[March

Figure 1. Pond Nine (3416 m) on Galena Mtn. in the Mexican Cut
Research Area owned by The Nature Conservancy. 7 September 1970.

winged adult, and flies away as a reproductively immature (teneral)
adult. The observations reported here are intended to correlate a late
emergence season, a possible lowered temperature threshold, a com-
plete shift to diurnal behavior, and an abbreviated, synchronized
emergence period with the rigors of the high altitude habitat in
Colorado.

Habitat

Two subalpine ponds were selected for study areas. They repre-
sent the typical habitat where larvae of Somatochlora semicircularis
are found. The emergence areas of both ponds are restricted to the
shallow, sedge-filled margins.

Pond Nine (3416 m) is one of a group of small ponds in the
Mexican Cut Research Area on Galena Mountain (Gunnison Co.,
Colorado) owned by The Nature Conservancy and administered by
the Rocky Mountain Biological Laboratory. It is 10 m in diameter
and 1 m deep in the center. About 50% of its shoreline is lined with
Carex rostrata and C. aquatilis in the shallow bays which make up
the emergence areas for the dragonflies. The center of the pond is

1974]

Willey — Subalpine Dragonfly

123

lined with Isoetes bolanderi on the bottom. The pond is surrounded
by a belt of forb-sedge meadow which is surrounded by an open
association of spruce-fir ( Picea engelmanni and Abies lasiocarpa )
(Fig. 1). In the past eleven years of observation, Pond Nine has
never totally dried up.

Irwin Pond South (3141 m) lies near the edge of Lake Irwin
(Lake Brennan) near Kebler Pass in Gunnison Co., Colorado. Its
basin is 45 m in diameter, 1.2 m deep in June and dries up in late
August and early September (observed 1968-1973). The shoreline
and shallow areas where emergence takes place are lined with C.
rostrata and C. aquatilis. The deeper areas support Potamogehon
foliosus and P. illinoisensis and thin mats of Drepanocladus uncinatus.
This pond has been described further by Willey and Eiler (1972).

Materials and Methods

The emergence of Somatochlora semicircularis was studied from
1968 to 1972. At first, the transforming larvae were marked by
surveyor’s flags so that the transformation of each individual could
be accurately timed. Considerable trampling of the habitat was un-
avoidable so that the studies were repeated in 1971 by making daily
collections of the cast skins left behind by the teneral adults. Fewer
larvae were damaged during their transformation by this method.
However, more larvae were lost through bird predation so that their
cast skins could not be recorded. In order to verify the diurnal
pattern of the teneral flight in an undisturbed habitat, continual
observations by binoculars of the ponds were conducted from 100
feet away from the shoreline in 1972 during the peak of the emer-
gence period.

Water temperatures were measured 3 cm below the water surface
in areas of highest emergence activity. Solar radiation was measured
by a recording Belfort pyrheliograph with a 12-hour gear. Environ-
mental air temperatures were measured with a YSI telethermometer
shielded from radiation by thin, flat, double-layered shields, 46 cm
square, which were set horizontally 8 cm above and below the tem-
perature probe. The inner surface of each was painted flat black and
the outer surface was glossy white (Platt and Griffiths, 1964).

Observations

About one month after the ice leaves the ponds, the first larvae of
S. semicircularis leave the water, undergo transformation, and fly

124

Psyche

[March

Table 1. The annual emergence periods for S. scmicircularis

1st day of Total Emergence Days until

Location

Year

emergence

adults

Total days

50% emergence

Pond Nine

1968

15 July

543

14

3

1969

7 July

166

15

3

Irwin Pond S.

1971

3 July

1528

21

6

away from the ponds as teneral adults (maiden flight). At Pond
Nine, emergence began on 15 July 1968, 7 July 1969, and 6 July
1970. At Irwin Pond South, slightly lower in altitude and more
sheltered by spruce-fir, emergence began slightly earlier — 30 June
1970, 3 July 1971, and 27 June 1972.

The total period during which emergence occurs each year is short
(Table 1). The total annual production of adults occurs over an
average of 16 days during which 50% of the adults emerge within
the first three to six days. The 1969 emergence curve (Fig. 2)
demonstrates a typical, highly synchronized pattern. The 1968 curve
(Fig. 3) shows the effect of stormy weather and low temperatures
on 17 and 18 July which delayed the emergence of some of the larvae
and extended the emergence period. No secondary peak of emer-
gence was ever observed later in either season.

The distance flown during the initial flight of the teneral adults is
unknown. However, they remain away from the ponds, presumably
in the forest near the ponds or at lower altitudes, for one to two
weeks. In 1968, the first tenerals left Pond Nine on 15 July and
the first mature males were observed patrolling the edges of the pond
on 22 July — 7 days later. By this time, 95% of the adults had
emerged and flown away (Fig. 3). In 1969, the first tenerals left on
7 July and the first patrolling male was sighted on 22 July- — 2
days after the last teneral adult had flown (Fig. 2). These mature
adults are individuals which emerged at the beginning of the emer-
gence period, have matured, and returned to the pond for mating.
Teneral adults and reproductively mature adults are considered here
to be all part of the same emerging population. They are just tem-
porally separated. On two occasions mature males were observed
patrolling a pond when a newly emerged teneral flew up from the
sedges on its maiden flight. On both occasions, the mature male
harassed and tried to couple with the soft teneral before the latter
could leave the pond. One of the tenerals was probably injured by

1974]

Willey — Subalpine Dragonfly

125

10 15 20
July, 1969

Figure 2. Seasonal emergence pattern of S. semicircularis at Pond
Nine, 1969. Vertical arrow marks day on which first reproductive male was
observed patrolling the pond.

Percent Total

126

Psyche

[March

the encounter because its subsequent flight was very erratic. The
Somatochlora males are very aggressive and will try to grasp any
other dragonfly — its own species, Cordulia shurtleffi / or even the
much larger A eschna interrupta.

Water and air temperatures are relatively low in ponds at high
altitudes although the diurnal water temperatures are not as low as
might be expected due to the heat absorbing qualities of the pond
bottoms and the shallowness of the water. The water temperature
averages I2°C. (varies from 90 to I4°C.) when the larvae first
emerge in the morning, whereas the air temperatures, which rise very
rapidly once the sun is up, average 9°C. (vary from 40 to I2°C.).
The night air temperatures drop below freezing and, on many morn-
ings, frost still lay on the ground when the first larvae appeared.
The larvae climb up the sedges in areas where the water is seldom
more than 30 cm deep and usually 3 to 8 cm deep. The larvae start
to leave the water after sunrise (0700 to 0800 MDT).

The maximum number of larvae leave the water between 0800
and 1100 hours (Fig. 4). By 1000 to 1700, they are ready to fly.
In 1972, observations of undisturbed ponds indicate that 75% or
more of the tenerals actually leave the ponds between 1100 tnd 1230
if the weather is fair and the air temperature varies between 160
and i8°C. The initiation of the flights appears to be temperature-
dependent. On 1 July 1972, flights began at 1100 with an air tem-
perature of i8°C. The flights stopped abruptly at 1230 when clouds
covered the sun and the air temperature dropped to I4°C. Flights
did not resume until 1430 when the sunshine returned and the air
temperature rose to i6°C. All flights ended for the day at 1500
When a thunderstorm started and the air temperature dropped to
I4°C. and below. Any adults which had not flown by this time had
to remain on the sedges until the following morning.

Birds are the main predators of Somatochlora during the actual
transformation process. Spiders and ants will attack larvae near the
shore but generally account for a very small proportion of the loss.
Specific data are being assembled for a separate report on bird preda-
tion and will not be presented here. However, a general summary is
essential for the present argument. In general, bird predation does
not begin until the second or third day of emergence. Under normal
weather conditions, daily loss does not exceed 30 to 50% of the

*In the studies on drought resistance in Somatochlora larvae, Willey and
Eiler (1972) misidentified Cordulia shurtleffi larvae for those of Libellula
quadrimaculata.

1974]

fVilley — Subalpine Dragonfly

127

Figure 3. Seasonal emergence pattern of S. semicircularis at Pond Nine,
1968. Vertical arrow marks day on which first reproductive male was
observed patrolling the pond.

Percent Total Emergence

28

Psyche

[March

dragonflies until the second week of emergence. On cold and stormy
days, loss may approach 100% at any time during emergence.
Adults which have transformed but have been prevented from flying
by low temperatures or rainy weather may remain clinging to the
sedges for a day or more. Most of these adults fall prey to birds.
Once the tenerals fly, they are reasonably free from bird predation.
Only one teneral has been observed to be caught by a bird (Western
flycatcher) during its maiden flight.

Discussion

Somatochlora semicircular is is a western cordilleran dragonfly
whose adults normally appear between the middle of June and the
middle of July (Walker, 1925). The Colorado ponds under in-
vestigation occur at the southern limit of the species and yet the
adults emerge in early July and appear as reproductives in mid-July
near the end of the general emergence season. Such a relatively late
season is probably correlated with the late snow melt and low tem-
peratures of its high altitude habitat. Schiemenz (1952) has reported
a similar delay of season correlated with altitude for Libellula quad-
rimaculata.

Dragonfly emergence has been shown to be temperature-dependent
with the initiation of emergence primarily dependent on water
temperature (Trottier, 1973; Corbet, 1963). The larvae of S.
semicircularis in Colorado first leave the water when the water
temperatures average I2°C. and the air temperatures average 9°C.
Comparative data for other corduliid dragonflies are not available.
The only data available come from the studies of the aeschnid dragon-
fly Anax. Corbet (1957) found that A. imperator in England nor-
mally left the water when the temperature reached 190 to 22°C.
(air temperature, 12° to I5°C.). Trottier (1973) found that A.
junius in Ontario, Canada, normally initiated emergence when the
water temperature reached 24°C. (air temperature, 150 to 20°C.).
At water temperatures below I7°C., the Anax larvae which had left
the water would return to the water and delay emergence until the
following day. The Somatochlora larvae were able to leave the water
and undergo transformation at average temperatures 5°C. below the
minimum for A nax larvae. Whether this difference is a specialization
to high altitudes or is only a general family difference must be further
investigated.

In general, teneral corduliid dragonflies at lower altitudes and/or
latitudes are ready to fly early in the morning. However, the emer-

1974]

Willey — Subalpine Dragonfly

129

c

Figure 4. Daily emergence pattern of S. semicircularis on 2 July 1970
at Irwin Pond South. Lower curve shows the number of nymphs leaving
the water (solid line) and transformed teneral adults ready to fly (dashed
line) at successive hours during the day. Middle curve shows water tem-
perature 3 cm below the surface at two separate stations, 1 m apart, and
air temperature 45 cm above the surface of the water in the emergence
area. Top curve shows solar radiation. Nymphs recorded here completed
the entire transformation process successfully.

130

Psyche

[March

gence pattern is variable and freshly emerged adults have been col-
lected throughout the day and even in early evening on cold, cloudy,
or rainy days (Lutz and McMahan, 1973; Corbet, 1963; Lutz,
1961; Taketo, i960; Kormondy, 1959; Robert, 1958; Lieftinck,
1932). The larvae of S. semicircular is, under normal conditions, do
not leave the water until after sunrise and the tenerals generally fly
at noon. A similar shift of the emergence pattern to the middle of
the day has been described for Aeschna interrupta at high altitudes
in California (Kennedy, 1925) and for Aeschna subarctica on the
moors near Kiel, Germany (Schmidt, 1968).

The emergence curve of Somatochlora in Colorado is characterized
by brevity (16 days) and a high degree of synchrony (Fig. 2,
Table 1). Similar synchrony has been described for other dragonfly
species which have been categorized by Corbet (1963) as “spring”
species. Only one of these species, Oplonaeschna armata, exhibits a
shorter time for the total period ( 10 days) and for the 50% emer-
gence time (2 days). Oplonaeschna inhabits the water-pocket streams
of New Mexico and Arizona and is subject to flash floods and severe
dessication (Johnson, 1968). The lower altitude corduliid, Tetra-
gone.uria cynosura, averages a 23-25 day emergence period with 50%
of the adults emerging by the 6th or 7th day in Michigan (Kor-
mondy, 1959) and North Carolina (Lutz and McMahan, 1973;
Lutz, 1961 and 1962). Tetragoneuria in Michigan also exhibits a
bimodal curve in which a second peak of emergence occurs 10 days
after the end of the first peak. This bimodality has been described
for other dragonfly species (see Corbet, 1963) but is missing in the
Somatochlora curve (Figs. 2 and 3).

Selective pressures contributing to the synchrony of the single peak
of the Somatochlora emergence curve may well involve the brevity
of the season. Both in 1968 and 1969, 10 days after the end of the
initial emergence peak at a time when T. cynosura (Kormondy,
1:959) and Anax imperator (Corbet, 1957) normally exhibit a small
second peak of emergence, the shallow areas at Irwin Pond South
and Pond Nine had largely dried up. Somatochlora larvae leave the
water by climbing up the emergent sedges. They do not crawl out
on the land. Lack of emergence sites or competition for the few
remaining sites can be expected to be a factor in selecting for a tem-
poral shift toward early emergence.

Another contributing factor may involve intraspecific interference.
The diurnal emergence pattern which is necessitated by the climatic
character of the habitat creates problems which normally do not affect

1974]

Willey — Subalpine Dragonfly

13

nocturnal or early morning emerging populations. Teneral adults
which fly up during the middle of the day are subject to harassment
and possible damage from the reproductive males. However, by the
time the first Somatochlora reproductives return to the Colorado
ponds, 95 to 100% of the annual production of tenerals have already
left the pond (Figs. 2 and 3). The diurnally temporal separation
of the reproductives and the nocturnal or early morning tenerals of
the lower altitudes has been replaced by the seasonal separation of a
1 to 2 week teneral maturation period during which time the dragon-
flies fly in the forest away from the ponds while the remainder of the
population is emerging. The high altitude ponds of Colorado seem
to be sufficiently isolated from each other to keep recruitment of
reproductives from lower, and therefore earlier, ponds to a minimum.

The diurnally transforming dragonflies are also exposed to preda-
tion by birds. The highly synchronized emergence of Somatochlora,
however, provides that the relatively large number of larvae which
emerge during the first few days ensure that many tenerals get away
from the pond. To what extent a synchrony of emergence provides
selective advantage to the dragonfly population by ( 1 ) partially fore-
stalling the results of a revised predator searching image and/or
shift in prey preference or (2) saturating the environment with prey
with a resulting early satiation of the predators is as yet undeter-
mined. It is likely that both predation tactics are involved and
thwarted by this mechanism.

Summary

Somatochlora semicircular is has been able to adapt to the rigors of
the subalpine and lower alpine habitats of Colorado. The corduliid
tendency toward a flexible, early morning emergence pattern has con-
tributed to its shift to a full diurnal emergence pattern. In addition,
it may be able to initiate emergence at relatively low water and air
temperatures. The emergence pattern has been strongly synchronized,
shortened and restricted to a single seasonal peak, possibly as a result
of ponds drying up early in the season at high altitudes, predation
pressure, which may be the same or worse than that at lower alti-
tudes, and the effects of potential interference between the aggressive
reproductive males and the soft-cuticled teneral adults.

Acknowledgements

I wish to thank Dr. Robert B. Willey for constant advice and
encouragement. I appreciate the valuable critical reviews by Drs.

132

Psyche

[March

P. S. Corbet and T. Poulson. Dr. H. A. Miller kindly identified
the moss for me. The National Science Foundation provided the
following excellent students under the N.S.F. — U.R.P. Program :
Philip Myers, Daniel Ewert, David DeHufif, Paul Deaton, and
William Davis. The University of Illinois Graduate Research Board
aided with two research grants, one of which funded the assistance of
David Mucha.

Literature Cited

Billings, W. D. and Mooney, H. A.

1968. The ecology of arctic and alpine plants. Biol. Rev., 43 : 390-401.
Carpenter, W. L.

1873. Report on the alpine insect-fauna of Colorado. Ann. Rep. U. S.
geol. geogr. Survey of the Territories embracing Colorado, pp.
539-542.

Corbet, P. S.

1957. The life history of the emperor dragonfly, Anax imperator
Leach (Odonata: Aeschnidae). J. Animal Ecol., 26: 1-69.

1963. A Biology of Dragonflies. Quadrangle Books, Chicago. 247 pp.
Gloyd, L. K.

1939. A synopsis of the Odonata of Alaska. Entom. News., 50: 11-16.
Johnson, C.

1968. Seasonal ecology of the dragonfly Oplonaeschna armata Hagen
(Odonata: Aeschnidae). Amer. Midi. Nat., 80: 449-457.
Kennedy, C. H.

1915. Notes on the life history and ecology of the dragonflies (Odonata)
of Washington and Oregon. Proc. U. S. Nat. Mus., 49: 1-345.
1925. The distribution of certain insects of reversed behavior. Biol.
Bull., 48: 390-401.

Kormondy, E. J.

1959. The systematics of T etragoneuria, based on ecological, life his-
tory, and morphological evidence (Odonata: Corduliidae) . Misc.
Publ., Mus. Zool., Univ. Mich., 107: 1-79.

Lieftinck, M. A.

1931. The life-history of Procordulia artemis Lieft. (Odon., Cordul.),
with comparative notes on the biology of P. sumbawana (Forster).
Int. Revue ges. Hydrob. Hydrogr., 28 : 399-435.

Lutz, P. E.

1961. Pattern of emergence in the dragonfly T etragoneuria cynosura.
J. Elisha Mitchell Sci. Soc., 77: 114-115.

1962. Studies on aspects of the ecology and physiology of T etragoneuria
cynosura (Say) as related to seasonal regulation (Odonata:
Cordulinae). Ph.D. Thesis, Univ. North Carolina, Chapel Hill
(University Microfilms, 63-3502). 64 pp.

Lutz, P. E. and McMahan, E. A.

1973. Five-year patterns of emergence in T etragoneuria cynosura and
Gomphus exilis (Odonata). Ann. Entom. Soc. Amer., 66: 1343-
1348.

1974]

Willey — Subalpine Dragonfly

133

Mani, M. S.

1968. Ecology and Biogeography of High Altitude Insects. Dr. W. Junk
N. V. Publ., Hague. 527 pp.

Paulson, D. R. and Jenner, C. E.

1971. Population structure in overwintering larval Odonata in North
Carolina in relation to adult flight season. Ecology, 52: 96-107.

Platt, R. B. and Griffiths. J. G.

1964. Environmental Measurement and Interpretation. Reinhold, New
York. 235 pp.

Robert, P.A.

1958. Les Libellules (Odonates) . Delachaux et Niestle S. A., Paris.
359 pp.

SCHIEMENZ, H.

1952. Die Libellenfauna von Sachsen in zoogeographischer Betrachtung.
Wiss. Zeit. Techn. Hochsch. (Univ.) Dresden, 1: 313-320.
Schmidt, E.

1968. Das Schliipfen von Aeschna subarctica Walker, ein Bildbeitrag.
Tombo, 1 1 : 7-11.

Taketo, A.

1960. An ecological study of Somatochlora clavata Oguma (Cordu-
liidae). Tombo, 3: 8-15.

Trottier, R.

1973. Influence of temperature and humidity on the emergence be-
haviour of Anax junius (Odonata: Aeschnidae). Canaid. Entom.,
105: 975-983.

Walker, E. M.

1925. The North American dragonflies of the genus Somatochlora.
Univ. Toronto Studies, Biol. Ser., 26: 1-202.

Willey, R. L. and Eiler, H. O.

1972. Drought resistance in subalpine nymphs of Somatochlora semi-
circularis Selys (Odonata: Corduliidae) . Amer. Midi. Nat., 87:
215-221.

THE MILLIPED GENUS BOLLMANELLA
(DIPLOPODA, CHORDEUMIDA, CONOTYLIDAE )

By William A. Shear1
Department of Zoology, University of Florida,

Gainesville, Fla. 32611

Even while my recent (1970) revision of the milliped Family
Conotylidae was in press, new data on the family had begun to
accumulate. This paper is the first of a series of supplemental reports
designed to update our knowledge of the conotylids, a group of milli-
peds of considerable importance in biogeography.

Like many other milliped genera and species, the genus Bvllman-
ella , and its single species B. oregona, have remained enigmatic since
they were described from a single male specimen by R. V. Chamber-
lin in 1941. The description of the genus alluded mostly to body
form and color, and even contradicted the specific diagnosis of the
only included species. The description of B. oregona also contained
errors, and no illustrations were provided. As if this were not
enough, the type locality suggested an error of several hundred miles.

The type of B. oregona was in the Chamberlin collection in Salt
Lake City. The diplopod portion of the collection has remained
uncurated since Chamberlin’s death and is in a confused state, but
while my conotylid revision was in press, Mr. Thomas Lorenz, then
in charge of the collection, found the holotype of B. oregona and
loaned it to me.

In 1973, I was fortunate in receiving a large number of Berlese
extraction samples from Mrs. Ellen Benedict, Portland State Uni-
versity, Portland, Oregon, and from Dr. David Malcolm, Pacific
University, Forest Grove, Oregon. These samples were rich in
millipeds of many groups previously known only from a very few
specimens, and included literally hundreds of individuals of the
related genera Taiyutyla and Bollmanella. Members of these genera
must be among the more common humus animals in the area of
coastal Oregon. As I mentioned earlier in reporting on part of these
collections (Shear, 1973), northern California and the state of Wash-
ington are much in need of the kind of thorough exploration by

^art of the work for this paper was done while a Richmond Fellow at
Harvard University.

Manuscript received by the Editor February 6, 1974.

134

1974d Shear — Milliped Genus Bolhnanella 135

Berlese extraction which Benedict and Malcolm have carried out in
Oregon.

It is now possible to do a number of things about Bolhnanella ,
which in 1970 I had to treat as a nomen dubium. 1) The genus can
be characterized as fully distinct and valid, separate from, but related
to, Taiyutyla and Conotyla, and more distant from Austrotyla and
Achemenides , 2) some clear ideas of the distribution of the genus
can be gained, and 3) seven new species can be described.

The distribution of the eight known species of Bollmanella ranges
from southern coastal Oregon to just above the Columbia River in
Mason Co., Washington, with one species reported here from the
Wallowa Mountains in extreme northeastern Oregon. The scattered
nature of the localities suggests that species of Bollmanella , except
for B. oregona, have small ranges, and we might expect many more
to be discovered. In particular, the isolated mountain ranges of
eastern Oregon need exploration.

Ecologically, Bollmanella species have been collected exclusively
from litter and duff derived from deciduous trees. Conifer duff and
litter support species of Taiyutyla , though members of this latter
genus are also found in deciduous duff, often syntopically with Boll-
manella species. Although B. oregona ranges from sea level to around
2000' elevation, most of the other species seem to have been collected
from forests or parkland between 900' and 1500' elevation. This
contrasts again with some species of Taiyutyla , which (unpublished
data) are known from quite high elevations. Species of an unrelated
chordeumid genus, Rhiscosomides (Shear, 1973), are found at low
elevations and seem to favor coniferous forests. This ecological data
also represents information from many collections which did not
contain specimens of Bollmanella , thus clearly suggesting which habi-
tats are most favorable.

All type specimens of new species have been deposited in the
Museum of Comparative Zoology, Cambridge, Mass.

Superfamily Heterochordeumatoidea Pocock
Family Conotylidae Cook
Genus Bollmanella Chamberlin

Bollmanella Chamberlin, 1941, Bull. Univ. Utah Biol. Ser. 6(5): 12. Type
species, B. oregona Chamberlin, by monotypy and original designation.

Diagnosis: With the characters of the family. Small (ca. 6-7 mm
long) conotylid millipeds with typical form, distinct in gonopod
morphology. Anterior gonopods simple, usually acuminate, but may

136

Psyche

[March

be apically laminate, or may bear a basal mesal branch. Anterior
gonopod sternum bandlike and well-sclerotized. Posterior gonopods
three-segmented, coxal segment with large, prominent colpocoxites
characterized by a basal flagellar branch usually more or less sheathed
by a development from the main part of the colpocoxite. Apical two
segments typical for family. Posterior gonopod sternum with a T-
shaped process between the coxae in most species. Third and fourth
legpairs of males enlarged, strongly bowed, but with, at most, small
mesal femoral knobs. Legpairs 5-7 either normal in size and form,
or decreasing from slightly enlarged to normal in size. Prefemora of
male legpair 1 1 with mesobasal processes. Female genitalia without
any useful taxonomic characters.

Remarks: Bollmanella is clearly related to Taiyutyla, with which
it is entirely sympatric, and often syntopic. However, the uniformly
smaller size, the flagellar branch of the posterior gonopods, and the
T-shaped process of the posterior gonopod sternum clearly set Boll-
manella species apart as a distinct phyletic line.

I would like to comment further on the Family Conotylidae as
a whole, but several factors preclude it at this time. The genus
Taiyutyla remains to be revised, and is affected by, as are any con-
clusions about Conotylidae at this time, the description by Loomis
and Schmitt (1971) of several extremely interesting new higher taxa
related to conotylids, including new subfamilies and a new family.
The Taiyutyla material I have examined so far will also enable me,
in this projected future paper, to comment in detail on the functional
aspects of the gonopod complex.

I am still unable to find any taxonomically useful characters that
will allow reliable separation of females of the various species of
conotylids, and members of Bollmanella are no exception. However,
in the hope that some usable characters might turn up in the future,
I have designated bona fide female paratypes where possible. I do not
present a key to the eight known species, as they can readily be sep-
arated by reference to the illustrations of the gonopods. The gono-
pods of these species are so small they must be mounted on slides
temporarily (in glycerine) to see detail.

Bollmanella oregona Chamberlin
Figs. 1, 2

Bollmanella oregona Chamberlin, 1941, Bull. Univ. Utah Biol. Ser. 6(5) : 12,
no illustrations.

Type: Male holotype from “John Day Creek,” Douglas Co.,
Oregon, collected by J. C. Chamberlin, 19 November 1939. In

1974] Shear — Milliped Genus Bolbnanella 1 37

Figs. 1, 2. Bollmanella oregona. Fig. 1. Right anterior gonopod, posterior
view. Fig. 2. Posterior gonopods, anterior view. Figs. 3, 4. B. reducta.
Fig. 3. Left anterior gonopod, posterior view. Fig. 4. Right posterior
gonopod, posterior view. Fig. 5. Sternal process between posterior gono-
pods of B. laminata.

138

Psyche

[March

Chamberlin collection (now at U.S. National Museum?), examined.
There is no John Day Creek in Douglas Co., but there is a town
named Days Creek, and a stream of that name, a tributary of the
South Umpqua River. Probably this is the type locality, and not the
John Day River of Oregon’s semiarid northeast.

Description: Male from io mi east, 6 mi north of Gold Hill, Jack-
son Co., Oregon: Length, 7.0 mm. Ocelli 14 on each side (a series of
males from several localities had ocelli numbers from 14-17). Legpair
3 much enlarged, femora swollen and bowed mesad, without obvious
processes. Legpair 4 enlarged, but smaller than legpair 3, femora not
strongly curved, bearing small fungiform basal knobs mesally. Leg-
pairs 5-7 only slightly larger than postgonopodal legs, without pro-
cesses. Anterior gonopods simple, subtriangular, acuminate, curved
posteriolaterad (Fig. 1). Posterior gonopod colpocoxites appearing
3-branched (Fig. 2), flagellar branch (/) long, prominent; sheathing
structure (s) reduced, completely separated from anterior process
( c ), which is simple and without branches. Posterior gonopod ster-
num with a large T-shaped process (T). Pigmentation typical.

Females similar to males, slightly larger and more robust.

Distribution: (all collections by E. M. Benedict) OREGON:

Josephine Co., 0.3 mi S, 2.5 mi E O’Brien, T4oS/R8W/Sec 28,
elev. 1400', 18 December 1971, d cf ??• Coos Co., 3 mi N, 2 mi
W North Bend, T24S/Ri3W/Sec 27, sea level, 15 January 1972,
cf. Douglas Co., Island Campground, 0.5 mi S, 1.0 mi E Steamboat
on Ore. Rt. 138, T26S/RiE/Sec 5, elev. 1200', Berlese of maple,
dogwood dubb, 20 October 1971, d d ; Boulder Flat Campground,
3 mi E, 10 mi S Steamboat, T26S/R2E/Sec 13, elev. 1700', Berlese
red alder, vine maple duff, 30 October 1971, d ; 4 mi S, 9 mi E
Steamboat on Ore. Rt. 138, T26S/R2E/Sec 23, elev. 1600', Berlese
conifer and oak duff, 30 October 1970, d d $?. Jackson Co., 10 mi
NW Central Point on Ore. Rt. 234, T35S/R2W, elev. 1200', 22
January 1973, d d ?. Lewis Buckley Farm, 1 mi S Ruch on Ore.
Rt. 238, T38S/R3W, elev. 1700', Berlese litter, duff, soil, 13 No-
vember 1971, d ; Upper Applegate Grange, 6 mi S Ruch, T39S/
R3W/Sec 15, elev. 1600', Berlese mixed litter, 13 November 1971,
d d $$; Buckley County Park Rest Area, 3.5 mi S Ruch on Upper
Applegate Rd., 2 mi off Ore. Rt. 238, T39S/R3W/Sec 15, elev.
1600', Berlese white alder, willow duff, 13 November 1971, d d ;
French Gulch, For. Ser. Rd. 420, 3 mi N Copper, T40S/R4W/
Sec 36, elev. 1900', Berlese Oregon oak duff, 13 November 1971,

1974]

Shear — Milliped Genus Bollmanella

139

c? $; 15 mi SW Ruch on Upper Applegate Rd., T4oS/R3W/Sec.
8, elev. 1800', Berlese duff, debris, moss, bark, 13 November 1971,
cf d ; 10 mi E, 6 mi N Gold Hill on Ore. Rt. 234, T35S/R1W/
Sec 17, elev. 1300', 22 January 1972, <$ c? 6 mi E, 3 mi N
Gold Hill, T35S/R2W/Sec 27, elev. 1200', 22 January 1972, <J* cf.

Notes: Chamberlin’s brief discussion of the leg modifications is
confusing. In the generic diagnosis leg 5 is described as being “nor-
mal” while in the specific diagnosis, leg 5 is described as “less thick-
ened, (the) fourth joint with a . . . subconical process on mesal side
toward proximal end . . .” Thus B. oregona is immediately excluded
from the very genus it is being described under! Chamberlin saw
14 ocelli on each side, and gave the size as 7-7.5 mm. The male
holotype is in poor condition.

Where ecological data is available, the species appears to occur in
duff from deciduous trees. Many samples of coniferous duff did not
yield Bollmanella oregona. The species has been collected at eleva-
tions from sea level to 1900', and thus appears to be primarily a
coastal or river valley species, since many samples from higher eleva-
tions did not contain B. oregona. This is the most widespread and
common species of Bollmanella.

Bollmanella reducta n. sp.

Figs. 3, 4

Type: Male holotype from 2 mi north, 6 mi east of Ashland,
Jackson Co., Oregon (T38S/R3E/Sec 27), collected at an eleva-
tion of 1200' by E. M. Benedict, 27 December 1971.

Description: Male holotype. Length, 5.0 mm. Ocelli 8 on each
side of head. Legpair 3 much as described for B. oregona. Legpair 4
less enlarged, lacking basal femoral knobs present in B. oregona.
Legpairs 5-7 of normal size. Anterior gonopods (Fig. 3) much as
in oregona, but gonopod tip blunter. Posterior gonopod colpocoxites
(Fig. 4) of reduced complexity When compared to following species;
flagellum (/) short; sheathing structure ( s ) fused anteriorly to body
of colopocoxite, which is broad and laminate, with minute teeth
distally. T-shaped process of sternum present, but broken in making
temporary slide of gonopods. Pigmentation light.

Distribution: Known only from the type locality. The holotype
came from a Berlese sample of Oregon oak and buckbrush litter on
a steep slope.

Psyche

[March

140

Figs. 6, 7. Bollmanella laminata. Fig. 6. Right anterior gonopod, pos-
terior view. Fig. 7. Right posterior gonopod, posterior view. Figs. 8-10.
B. unca. Fig. 8. Left anterior gonopod, posterior view. Fig. 9. Left pos-
terior gonopod, posterior view. Fig. 10. Sternal process between posterior-
gonopods. Fig. 11. Anterior gonopods of B. bifurcata, posterior view.

1974]

Shear — Milliped Genus Bollmanella

141

Bollmanella laminata n. sp.

Figs. 5-7

Types: Male holotype, female paratype and other specimens from
0.3 mi west of Southern Pacific overpass on Rt. 26, 3 mi east of
Timber, Washington Co., Oregon (T3N/R5W/Sec ?) collected at
an elevation of 900' by E. M. Benedict, 27 November 1971.

Description: Male holotype. Length, 6.5 mm. 17 ocelli on each
side (a second male had 20 ocelli on each side). Legpair 3 much
swollen, femora enlarged and arched mesad, with sharp mesoposterior
tooth near each apical end. Legpair 4 normal in size, without pro-
cesses. Legpairs 5-7 of normal size and form. Anterior gonopods
(Fig. 6) erect, bearing lateral tooth near base; apex expanded into
thin lamina recurved and folded on lateral edge. Posterior gonopod
colpocoxites (Fig. 7) showing relationshhip to the previous two
species, flagellum (/) short, sheath (.r) completely separate from
anterior body of colpocoxite, as in B. oregona. T-shaped process of
sternum with arms sharply curved ventrad (Fig. 5). Pigmentation
light, much lighter than B. oregona but darker than in B. reducta.

Female similar to male in nonsexual characters, somewhat more
robust and larger.

Distribution: Known only from the type locality. The types came
from a sample of vine maple and Douglas fir duff.

Bollmanella unca n. sp.

Figs. 8-10

Types: Male holotype, juvenile female, and a second male from
4.5 mi east of Wells Creek Ranger Station, Douglas Co., Oregon
(T22S/R9W/Sec 15), elev. 300' collected 11 December 1971 by
E. M. Benedict.

Description: Male holotype. Length, 7.0 mm. 14 ocelli on each
side of head (second male from type locality had 12 ocelli on each
side). Legpair three greatly enlarged, femora strongly bowed mesad,
more so than in oregona , base of each femur so swollen and curvature
so extreme that appearance of large basal knob is presented. Legpair
4 much as in oregona , but femoral knob larger, more flattened. Leg-
pairs 5-7 of nearly normal size, lacking knobs. Anterior gonopods
(Fig. 8) broadly triangular at base, tapering to curved, sharply
aciculate apex, deeply excavated mesoposteriorly at base, with mesai
angle drawn ventrad into short, blunt process. Posterior gonopod
colpocoxites (Fig. 9) complex; flagellum (/) large, long, with mem-

142

Psyche

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Fig. 12. Right posterior gonopoid of Bollmanella bifurcata, posterior
view. Figs. 13, 14. B. bella. Fig. 13. Left anterior gonopod, posterior
view. Fig. 14. Right posterior gonopod, posterior view. Figs. 15, 16.
B . camassia. Fig. 15. Right anterior gonopod, posterior view. Fig. 16.
The same, lateral view.

1974] Shear — Milliped Genus Bollmanella 143

branous serrations on mesal edge; irregular chitinized membrane at
base ( fp ) ; sheath (s) large, enfolding flagellum, basally fused to
anterior part of colpocoxite. Near point of fusion is large basal pro-
cess ( b ) which is sharply elbowed and distally trifucate; colpocoxite
with fimbriate ridge mesally and irregular, apical, laminate teeth.
T-shaped process from posterior gonopod sternum much the largest
in genus, arms extending laterad (Fig. 10). Pigmentation typical,
but somewhat darker than in B. oregona .

Mature females not collected.

Distribution: In addition to the type locality: OREGON: Doug-
las Co., Mack Brown County Park on Umpqua R., T25S/R7W,
elev. 400', 7 February 1972, E. M. Benedict, cf .

Notes: Collections at both sites were from Oregon oak duff.

Bollmanella bifurcata n. sp.

Figs. 11, 12

Types: Male holotype, female paratype and other specimens from
2 mi west of Joseph along Hurricane Creek, Wallowa Co., Oregon,
collected 23 November 1968 by D. R. Malcolm.

Description: Holotype male. Length, 9.2 mm. 20 ocelli on left
side of head, 21 on right side. Legpairs 3 and 4 enlarged, approx-
imately same size, both with small mesal femoral knobs. Legpairs
5-7 decreasing in size, legpair 7 normal in size and form. Anterior
gonopods (Fig. 11) upright, curved, with small lateral spine and
large blunt mesal branch, presenting bifurcate appearance with telo-
podite. Posterior gonopod colpocoxites (Fig. 12) rather smaller than
usual, flagellum not observed, probably concealed by large sheath (s)
which is free from coxite for most of its length. Body of coxite with
large, spirally curved basal branch ( b ) bearing near its base a curved
lamina. Sternal process (T) much less prominent than in other
species. Pigmentation very light.

Females as usual, similar in general appearance to males, but
slightly larger and more robust.

Distribution: Known only from the type locality.

Note: This species is far removed from the other members of the
genus, being found in the foothills of the Wallowa Mountains in
extreme northeastern Oregon. This part of the state has not been
much explored by biologists, and other species of Bollmanella may
well ocur in other isolated mountain ranges. The region is also close
to that studied by Loomis and Schmitt (1971), and in which they

144

Psyche

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Fig. 17. Right posterior gonopod of Bollmanella comassia, posterior
view. Figs. 18-20. B. complicata. Fig. 18. Right posterior gonopod, pos-
terior view. Fig. 19. Anterior gonopods, posterior view. Fig. 20. Sternal
process between posterior gonopods.

found several highly unusual chordeumid species. Bollmanella bi-
furcata shows some intermediacy in size, pregonopodal leg modifica-
tions, and gonopod form between species of Bollmanella and Taiyu-
tyla, and this may suggest that the two genera could be difficult to
keep separate in eastern Oregon. Loomis and Schmitt (1971) de-
scribed Taiyutyla curvata, from Montana, but it does not appear
(from their illustrations) to be very much like B. bifur cat a.

The types and several other specimens of both sexes were taken
from cottonwood litter, in a riparian situation.

Bollmanella bella n. sp.

Figs. 13, 14

Types: Male holotype and a juvenile female (not designated as a

1974]

Shear — Milliped Genus Bolbnanella

145

paratype) from 11 mi east and 4 mi west of Allegany, Company
Road 5000 on the Weyerhauser Millicoma Tree Farm (T24S/
R9W/Sec 18), at the Douglas and Coos Cos. boundary, Oregon,
20 November 1971, by E. M. Benedict.

Description: Male holotype. Length, 6.0 mm. Ocelli 17 on both
sides of head. Femora of legpair three very much enlarged and bear-
ing small knobs mesally just above inflection point of curve. Legpair
4 somewhat smaller, femora with large capitate knobs just distad of
midpoint on mesal side. Legpairs 5-7 only slightly larger than nor-
mal. Anterior gonopods (Fig. 13) much as usual, curved laterad,
acuminate. Posterior gonopod colpocoxites (Fig. 14) somewhat
larger than in other species, flagellum (/), long, curved into sigmoid
sheath (f) ; body of coxite with mesal fimbriate ridge, apically lacini-
ate. Sternal process (T) small. Pigmentation dark.

Mature females definitely belonging to this species were not col-
lected.

Distribution: In addition to the type locality, a single female from
6 mi east of Allegany on the Millicoma Tree Farm may also be this
species.

Notes: The type was taken in a Berlese sample of myrtle and
Rhododendron litter and duff; the second female referred to above
also came from myrtle litter.

Bollmanella camassia n. sp.

Figs. 15-17

Types: Male holotype and female paratype from 2 mi northeast
of Camas Valley on Oregon Rt. 42 (T29S/R8W), collected at an
elevation of 1400' by E. M. Benedict, 19 February 1972.

Description: Male holotype. Length, 7.0 mm. Ocelli 12 on both
sides of head. Legpair 3 much less enlarged than in other species,
with a small knob near distal ends of femora, which are evenly
curved and swollen. Legpair 4 only slightly larger than normal,
femoral knobs basal in position. Legpairs 5-7 normal in size and
form. Anterior gonopods (Figs. 15, 16) simple, acuminate, curved;
in lateral view, gland channel ( ?) originates in deep anteriolateral
depression. Posterior gonopod colpocoxites (Fig. 17) with long
flagellum (/) bearing irregular membranous process near base;
sheath (s) arising from midpoint of coxite; coxite bears a strong
decurved process apically and a small basal branch (b). Sternal
process not observed, perhaps absent. Pigmentation as in B. oregona.

146

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Female as usual, somewhat larger and more robust than male.
Distribution: Known only from the type locality.

Notes: The holotype came from a Berlese sample of vine maple,
willow, and Madrone duff near a grove of Douglas firs.

Bollmanella complicata n. sp.

Figs. 18-20

Types: Male holotype and paratype from 1 mi west of Bayshore,
4 mi north of Shelton, Mason Co., Washington, collected 25 Novem-
ber 1967, by E. M. Benedict.

Description: Male holotype. Length, 6.0 mm. 13 ocelli on each
side of head. Legpair 3 enlarged as usual, but less curved mesad.
Legpair 4 nearly normal in size and form, without femoral knobs.
Legpairs 5 and 7 of normal size and form. Anterior gonopods
(Fig. 19) sharply elbowed mesad after crossing {in situ ) laterad of
posterior gonopod colpocoxites; apically laminate. Posterior gonopod
colpocoxites (Fig. 18) most complex in genus; flagellum (/) with
two branches about equal in size, one of these may be the homolog
of the basal lamina seen in other species; sheath {s) free, apically
with complex series of hooked, bulbous and aciculate processes; body
of coxite (c) very broad basally, flared, tapering to hooked apex.
Sternal process (Fig. 20) large, arms searly horizontal. Pigmenta-
tion light.

Females not collected.

Distribution : Known only from the type locality.

Notes: The holotype and paratype were collected from a Berlese
sample of oak and pine duff.

Literature Cited

Chamberlin, R. V.

1941. New western millipeds. Bull. Univ. Utah, Biol. Ser. 6(5): 1-20.
Loomis, H. F., and R. Schmitt

1971. The ecology, distribution and taxonomy of the millipeds of
Montana west of the Continental Divide. Northwest Sci. 45(2):
107-131.

Shear, W. A.

1970. The milliped family Conotylidae in North America, with a de-
scription of the new family Adritylidae. Bull. Mus. Comp. Zool.
141 (2) : 55-97.

1973. The milliped family Rhiscosomididae. Psyche 80 (3): 189-203.

A NEW GENUS OF PRIMITIVE MELOIDAE
FROM WEST TEXAS (COLEOPTERA)1

By Floyd G. Werner

Department of Entomology, University of Arizona, Tucson 85721

The beetle described here is the most unusual meloid I have ever
encountered. When Henry Howden sent me a male, several years
ago, I examined it and placed it among the unidentified pedilids.
But re-examination in relation to the pedilids convinces me that it
is indeed a meloid, and probably one of considerable evolutionary
interest. Since I first saw a specimen, Richard Selander (1966) has
published a comprehensive study of the meloid subfamily Eleticinae,
which is generally unfamiliar to students of North American beetles.
The Eleticinae have some characteristics that are different from those
of conventional meloids; part of these are shared by the new genus
described here. The Eleticinae are so different from other Meloidae
that Selander suggests that relationship would best be indicated if
they were set up as a separate family, or as a subfamily of weight
equal to one combining all other meloids. But, in the interest of
avoiding major changes before relationships are certain, he has (1964,
1966) followed a course of recognizing three subfamilies of Meloi-
dae: Eleticinae, Meloinae, and Nemognathinae, with the Eleticinae
considered the most primitive. The group has a disjunct distribution
in the Old World (mainly Africa but including a few species in
Southeast Asia) and South America, both of the tribes he recognizes
in his 1966 paper having members on both sides of the Atlantic. This
relict type of distribution certainly supports his opinion that this is an
ancient group. Unfortunately, there are no observations on the be-
havior of the subfamily, and the larval associations that have been
made are based on supposition rather than observation. So only adult
structures can be used in classification.

Perhaps the single thing that is most meloid-like about the insect
described here is the shape and conformation of the head (Fig. 2, 3).
The only thing that is unusual at all is that the eyes are more coarsely
faceted than is usual for the family. The general body shape ( Fig. 1 )
is also not greatly different from that of some other meloids ; the main
points of difference are those of the Eleticinae: cuticle much firmer

journal Paper No. 2266 of the Arizona Agricultural Experiment Station.
Manuscript received, by the editor February 20 , 1974.

147

148

Psyche

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than usual and elytra conjointly rather than separately rounded. To
these features must be added a uniform brown coloration, unusually
small size, less than 5 mm, quite dense and deep punctures over the
whole dorsal surface, and moderately dense, fine pubescence, all un-
usual for the family but found in some Eleticinae.

The combination of these features and lack of a separate blade
on the tarsal claws would make identification of the genus as a meloid
almost impossible with existing keys to the family, even if one had a
good representation of other genera for comparison. In fact, I have
tried repeatedly to find reasons for excluding it from the family, but
have then been faced with the decision that it is more like meloids
than it is like any other family known to me. So I am describing it
as a meloid, possibly assignable to the subfamily Eleticinae, but not
definitely assignable to this subfamily nor to either of the included
tribes recognized by Selander (1966). It is my hope that the exis-
tence of a population in a relatively accessible location will ensure
that both bionomics and behavior will come under study and that
these will permit more exact assignment.

Since it is almost inevitable that comparison will be made with the
Eleticinae and that Selander’s 1966 paper will provide the basis for
comparison, I have attempted to follow Selander’s terminology and
his pattern of description. However, I have not attempted to review
the whole subfamily, and therefore may have strained too far to see
structures that are obvious in the subfamily and not necessarily in
this genus. The bases for measurement also follow Selander’s paper.

Genus Thambospasta, new genus

Characteristics. Small, less than 5 mm, slender beetles (Fig. 1)
of the family Meloidae and doubtfully of the subfamily Eleticinae
as defined by Selander (1966); monochromatic brown; head, pro-
notum, and elytra densely, moderately deeply and moderately finely
punctate; with simple slightly curved decumbent setation, also mono-
chromatic. Mentum, palpi, coxae, trochanters, and abdominal sterna
(Fig. 5) with single to double, long erect tactile setae, those on men-
tum and abdomen sublateral. Head just perceptibly longer than
broad (Fig. 2) ; vertex not inflated, its base subtruncate. Eyes mod-
erately large, length 1 1/3 frontal ocular distance in male, 1 1/10 in
female; anterior margin slightly excavated (Fig. 3); facets coarse
and separate, ca. 8 across narrowest portion. Antennae slender, fili-
form; segment X 1/5 as wide as long (Fig. 4) in male; less slender

1974]

W erner — Primitive Meloidae

149

in female, segment X 1/2 as wide as long. Clypeus separated from
frons by a depression only, with a narrow, thin, shiny shelf over base
of labrum; labrum flat, not emarginate at apex. Mandibles half as
long as head from top of vertex to anterior mandibular articulation;
shallowly indented on mesal margin just basad of middle; prostheca
moderately long, reaching beyond middle of mesal margin ; molar lobe
small but distinct; mesal margin without teeth but apex bluntly
double-pointed, chisel-like, the mesal face flat for a small zone before
apex; lateral margin almost evenly curved from base, with moderately
dense setae. Maxillary palpi (Fig. 6) four-segmented, basal segment
small, second and third subequal, last long, narrowly securiform.
Galea simple, of but one segment. Labium (Fig. 7) with ligula
apparently entire, blunt; mentum not strongly produced laterad.
Pronotum 1/5 longer than wide, ca. 9/10 as wide as vertex, feebly
subhexagonal in outline; disk feebly convex except for impressed
midline. Front coxal cavities open externally, closed internally.
Mesosternum obtusely angulate at anterior apex. Mesepisterna in
contact medianly; anterior marginal area with a complete curved
ridge from dorsal edge to near midline, subparallel to main ridge but
converging toward it near midline. Apex of elytra simple in both
sexes. Hind wings (Fig. 13) fully developed in both sexes; apical
region 1/3 wing length; stem of media (M) short but distinct;
crossvein r present and radial cell closed; base of 2A2 strongly de-
veloped, forming a closed cell (iA3) with a crossvein from iA; 3A1
not combined with 2A3, apparently present as a short branch which
with a crossvein from 2A3 forms a closed cell . Male fore tibiae with
two unmodified spurs (Fig. 10). Hind tibial spurs slender, essen-
tially equal. All spurs with fine microspines (Fig. 10). Tarsal
formula 5-5-4. Penultimate tarsal segment (Fig. 11) forming a flat,
truncate lobe, the last segment arising from near middle of its dorsal
surface. All tarsal claws with a very blunt basal tooth ; those of fore
and middle tarsi with a longer pointed tooth in addition (Figs. 8, 9,
12). There is no vestige of a ventrolateral lobe or blade. Abdominal
sternum III excavated for hind coxae, the excavation bordered by a
fine ridge. Male abdominal tergum VII weakly sclerotized, its apex
slightly curved, at least this tergum with a micropubsecent area that
is not divided on midline. All terga before VII weakly sclerotized;
at least part seem to have the same kind of micropubescence. Male
sternum VII truncate across apex, simple on disk; tergum VIII
evenly sclerotized from very near base, bluntly ogival; sternum VIII
sharply angular posterolaterally, its disk simple and its apex just

150

Psyche

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7

9

1974] Werner — Primitive Meloidae 15 1

perceptibly emarginate between the angulations; tergum IX small,
undivided and weakly sclerotized; sternum IX Y-shaped with broad
lateral arms. Male genitalia (Fig. 14) with gonostyli fused dorsally
from base to beyond middle; base of fused gonostyli produced pos-
tered on dorsal side in a process that surpasses the gonostyli and bears
three dorsal and ca. 9 lateral short, stout setae on each side; gono-
coxal plate cupped, ca. 1/2 as long as gonostyli. Aedeagus (Fig. 15)
long, symmetrical, downcurved toward base, movable in relation to
gonostyli; with two oblique lateral ridges on each side toward apex;
secondary gonopore apparently apicodorsal ; there is no indication of a
sclerotized rod or projection from the internal sac in this region;
dorsal surface of aedeagus apparently open for most of its length.
Female abdominal segment VIII retracted into apex of abdomen,
weakly sclerotized, its sternum with a long, slender apodeme (Fig.
17); ovipositor (Fig. 16) well-developed, long, with one-segmented
gonostyli.

Type species. Thambospasta howdeni new species; fixed by present
designation.

Remarks. This genus contains only the type species, described
below. It is known from both sexes.

Thambospasta howdeni, new species

Male. Entirely rufescent brown except for black eyes, only tips
of mandibles appreciably darker; pubescence concolorous, fine, and
therefore inconspicuous. Length 3. 6-4.4 mm> mean 3.88, SD 0.29,
holotype 4.4. Surface generally shiny, but deep punctures dull it
somewhat. Punctures on head uniform, flat-bottomed, ca. 0.04 mm
across, 0.05 center to center, even across midline; midline indicated
only as a notch on vertex. Antennal calluses distinct, grading into a
small elevation over base of antenna, with punctures like rest of head.
Head depressed across front between antennae, in region of fronto-
clypeal suture, which is not visible, punctures extending evenly across
whole region; clypeus ending in a smooth shelf ca. 0.04 mm wide
and flat, over base of labrum. Eye facets almost exactly 0.03 mm
in diameter, very regularly arranged. Punctures of pronotum just

Figs. 1-12. Thambospasta howdeni. 1, dorsal view of holotype male;
2, head of holotype in front view ; 3, same in oblique lateral view ; 4, an-
tenna of paratype male; 5, profile of abdominal sterna of holotype; 6, max-
illa of paratype male; 7, labium of same; 8-9, pieces of tarsal claws of
same; 10, tip of front tibia of same; 11, front tarsus of same; 12, detail of
front tarsal claws, ventral view.

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Psyche

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Figs. 13-17. Thambospasta howdeni. 13, hind wings of paratype male;
14, genitalia of paratype male in dorsal, lateral, and ventral views, with
inset of details of median process; 15, aedeagus of same in dorsal and left
lateral views; 16, ovipositor of allotype female; 17, anterior apodeme of
abdominal sternum VIII of same; Figs. 14-17 at 60X magnification.

1974]

W erner — Primitive Meloidae

153

perceptibly smaller, 0.03-0.04 mm across and not as distinctly flat-
bottomed. Punctures of elytra deep across base, shallower behind,
ca. 0.03 mm across and 0.05 center to center, round-bottomed, closer
to 0.04 mm across near base, evenly distributed. Pubescence of upper
surface slightly curved, decumbent, ca. 0.07 mm long, fairly incon-
spicuous because it is not at all flattened and is concolorous with
cuticle. Antennae with moderately dense suberect setae ca. 0.05 mm
long from segment 2 to apex, a few such setae on first. Punctures of
underside of thorax more rugulose, fairly shallow, ca. 0.02-0.03 mm
across; those on abdomen smaller and finer, mose distinct from each
other, ca. 0.02 mm across and 0.03 center to center.

Female. Known only from allotype, 4.6 mm long. Antennae gen-
erally shorter and less slender, without the suberect setae of the male.
Unfortunately, most of antennae lost after description. Measure-
ments in 0.01 mm, base to apex: 55/20, 34/16, 44/15, 50/15, 49/1 5>
46/15, 47/15, 42/15, 39/15, 32/15, 54/I5- Last segment with little
indication of the change of thickness seen in the male. Eyes dis-
tinctly smaller and noticeably narrower than in male. Maxillary
palpi smaller, last segment 0.24 mm long, 0.29 in male holotype,
which is slightly smaller.

Holotype male, allotype female and 1 1 male paratypes, Big Bend
Nfational] P[ark], TEX [AS], 1850', Boquillas, May 23, 1959,
light; Howden & Becker; 3 male paratypes same but May 13. Holo-
type, allotype, and 11 paratypes in Canadian National Collection, 3
paratypes in collection of author.

Discussion. If this beetle is really a member of the family Meloi-
dae, it is probably one of the most primitive yet discovered. Of the
features that Selander (1966) lists as common to all meloids, in-
cluding Eleticinae, Thambospasta conforms in only three: head with
a well-developed vertex and narrow neck; pronotum not bordered
laterally [with exceptions in Nemognathinae] ; and female gonostyli
one-segmented. It resembles Eleticinae but not the other subfamilies
in that the cuticle is more heavily sclerotized; abdominal sternum III
is excavated and margined to accommodate the hind coxae; the female
has a distinct ovipositor; and in the following characteristics of the
venation of the hind wings: vein 2A2 present and joined to iA by a
crossvein (also in Meloinae: Lyttini: Prolyttina) ; and 3AX con-
nected to base of 3A (in Eleticinae except Spasticini: Protomeloina
and in some non-eleticines, apud Selander). It is very similar to
Eleticinae : Spasticini : Anthicoxenina in that the male gonostyli are
fused dorsally at the base and have a median dorsal projection; this

154

Psyche

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feature is unknown elsewhere in the family. It is more like Meloinae
and Nemognathinae than Eleticinae in that the apices of the elytra
are not swollen and provided with a pit in the male (as they are in
all Eleticinae but Spasticini: Anthicoxenina : Iselma).

It would be difficult to associate Tha?nbospasta with Iselma in the
Eleticinae on the basis of the similarity of the male genitalia and
lack of elytral pits, because Thambospasta males have abdominal ter-
gum VIII heavily sclerotized and would therefore become associated
with Eleticini, the other tribe in the subfamily in Selander’s classi-
fication. It differs from all Eleticinae in that vein 2A2 of the hind
wings is joined to the base of 2 A, a condition approached in some
Eleticinae, however, and in being nocturnal rather than diurnal, as
the few Eleticinae for which records are available seem to be. If
Selander’s 2A3 of the Nemognathinae: Nemognathini : Zonitina:
Pseudozonitis were counted as a primitive vein, rather than a spe-
cialization as he argues (1966), that genus would also have 2A2
joined to the base of 2A.

Returning to the features that Selander lists as being shared by
all subfamilies of Meloidae, Thambospasta differs in several probably
important ways: fore coxal cavities open externally but closed in-
ternally (open in all others) ; female abdominal sternum VIII with
a long anterior apodeme (none at all in others) ; last segment of
maxilliary palpi narrowly securiform (never securiform in others) ;
and cross-vein r apparently present (never in others). It also is un-
like most Meloidae in that the male genitalia lack a sclerotized ejacu-
latory rod ( lacking also in Eleticinae : Eleticini : Eospastina : Eospasta
and in many Nemognathinae) ; and in that the tarsal claws lack a
process arising from the lateral side of the base (lacking also in
some specialized Meloinae and Nemognathinae). A possible third
feature is that the aedeagus lacks a hook on its ventral side at or near
the apex, a condition that is universal in Meloinae and present also
in part of the Eleticinae. A feature not mentioned by Selander that
is probably universal in Meloidae is the lack of microspines on the
tibial spurs such as are seen in Thambospasta.

References

Selander, Richard B.

1964. Sexual behavior in blister beetles (Coleoptera: Meloidae). I.
The genus Pyrota. Canadian Entomologist 96(8): 1037-82.

1966. A classification of the genera and higher taxa of the meloid
subfamily Eleticinae (Coleoptera). Op. cit. 98(5): 449-81.

POLYMORPHISM IN STELOPOLYBIA AREATA
(HYMENOPTERA, VESPIDAE)

By Robert L. Jeanne11 and Robert Fagen2

Polymorphism in the social Hymenoptera has been defined as the
occurrence within a single colony of two or more distinct morpho-
logical forms, or castes, belonging to the same sex (Wilson 1953).
Wilson (1953) has pointed out that polymorphism arises out of the
occurrence of allometry (differential rates of growth of two parts of
the body) over a sufficient range of intranidal size variation to pro-
duce morphologically distinct forms at the extremes of this size range.
The underlying size variation is due in most cases to differences in
larval nutrition (Michener 1961; Wilson 1971). Polymorphism is
the morphological adaptation to the functional division of tasks among
the members of a colony (Wilson 1953). The most fundamental
polymorphism is the separation of the reproductive caste (queens)
from the non-reproductive caste (workers). This is followed, most
notably in the ants, by the evolution of more or less distinct castes
among the workers.

Among the wasps polymorphism appears to be limited to the queen-
worker dimorphism, where it is most pronounced in the Vespinae
(Wilson 1971). Among many of the Polistinae there is no detectable
difference, either in size or in morphology, between functional queens
and workers (Richards and Richards 1951). The present paper
reports the occurrence of complete queen-worker dimorphism in
colonies of the polistine species Stelopolybia areata (Say) from
Mexico.

Materials and Methods

Stelopolybia areata ranges from Mexico south to northern South
America (Ducke 1910). The four colonies available for the present
study were collected near San Andres Tuxtla in southern Veracruz,
Mexico, in January and February 1973. During these months the
nests are occupied by adult females only. No brood or males are
present. Population size and composition for each colony are given

department of Biology, Boston University, Boston, Massachusetts 02215.

Aiken Computation Laboratory, Harvard University, Cambridge, Massa-
chusetts 02138.

Manuscript received by the editor February 20, 1974.

155

Psyche

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156

Table I. Size and Composition of Adult Populations of Stelopolybia
areata colonies.

Estimated

Colony

No.

Date

Collected

No. Adults
Collected

No. Queens
Collected

Percent

Queens

Total Adult
Population

707

2 Feb. 1973

5884

716

12.2

7000

710

15 Jan. 1973

7950

515

6.1

8500

725

23 Feb. 1973

4711

257

5.5

6000

731

21 Feb. 1973

4227

503

11.8

6000

in Table I. A description of the nest of this species and notes on
colony cycle are published elsewhere (Jeanne 1973).

In a mass of anaesthetized adults, queens can readily be picked
out by virtue of their larger size and distinctive morphology and
coloration. Random dissections of queens and workers always showed
that the former had well-developed ovaries and full spermathecae,
while the latter had undeveloped ovaries and empty spermathecae.

For the present study random samples of queens and workers from
each colony were measured and compared. Length of alitrunk was
chosen as a measure of body size. This was measured from the front
of the humeral collar to the furthest posterior extension of the pro-
podeal valves, as seen from the side. This dimension was preferred
as a measure of body size for several reasons. First, the alitrunk is
the longest rigid structure of the body, hence is not subject to error
due to varying degrees of distension of the gut, or to differences in
body position. Second, it is not likely to be an allometric growth
center, unlike in the ants, since both queens and workers are winged.
Any functional difference in use of wings by workers and queens is
less likely to be reflected by allometric differences in size of the thorax
than in differences in the wings themselves. Finally, it is preferred
to wing length, used in other studies, because wings are subject to
fraying at the tips in older workers, making them impossible to mea-
sure, and thus biasing the sample against older workers.

The most conspicuous morphologic difference between the two
castes is the more bulbous first abdominal tergite (petiole) in the
queens (Fig. 1). This is best reflected in its width at the widest
point, as seen from above; hence this dimension was used as a measure
of allometric growth.

All measurements were made on specimens pinned while fresh
using an ocular micrometer at a magnification of 25 X .

1974]

Jeanne & Fdgen — Stelopolybia

157

Fig. 1. Strlopolybia areata queen (left) and worker (right).

158

Psyche

[March

Standard methods and tests (Hays 1963) were used for computa-
tion and assessment of significance of regression slopes. Lewis (i960)
presents techniques for the analysis of intersample differences between
regression slopes; we used the following statistic to test the signifi-
cance of intercaste regression slope differences.

Let k(p = estimated regression slope for i*,h caste.

<7(i> = sample variance in ith caste of In (tergite width)
given In (alitrunk length).

o-i(i) = sample standard deviation of In (alitrunk length)
in ith caste.

ni = number of individuals of ith caste sampled.

i = 1 for workers and 2 for queens.

Under standard assumptions of regression analysis (normality,
homeoscedasticity, interindividual independence of measurements)
the statistic

k(i)q,Ki)\Avi ~ k(2)<ri (2) V ft 2

V2

/n <7(i) + n2o- (2)
\ ru -f n2 - 4

has a t-distribution with nt + n2 — 4 degrees of freedom.

Results

Coloration. Both castes are yellow with brown or black dark
markings ( Fig. 1 ) . The yellow of workers, however, is a bright
lemon yellow, while that of queens contains much more brown. The
dark markings of workers are mostly black, while in queens they are
dark brown, especially on the gaster. Thus, the dark and light colora-
tion of workers is much more contrasting than that of queens.

On average the dark markings are slightly more extensive on the
head and thorax of workers than of queens. On the other hand, the
dark bands on the gaster are proportionately wider in queens than
in workers. The dark band of the second abdominal tergite in
workers has a pointed median posterior extension that is lacking in
queens.

Body size. Results of the two measurements made are summarized
in Table II. There is no overlap in size (length of alitrunk) of
queens and workers in any of the colonies; thus the two castes are

1974]

Jeanne F age n — Stelopolybia

159

Table II. Queen/Worker Dimorphism in Stelopolybia areata .
A. Length of Alitrunk

Colony Queens Workers Queens/

No.

N

Average

Range

N

Average

Range

Workers

707

45

4.22

4.02-4.37

45

3.63

3.29-3.75

1.16

710

49

4.10

3.83-4.29

60

3.57

3.29-3.75

1.15

725

23

4.13

4.02-4.29

50

3.68

3.41-3.83

1.12

731

50

4.19

3.98-4.33

50

3.68

3.33-3.83

1.14

B. Width of First Abdominal Tergite

Colony

No.

N

Queens

Average Range

N

Workers

Average Range

Queens/

Workers

707

45

1.60

1.34-1.72

45

0.94

0.88-1.00

1.70

710

49

1.52

’hfO-1.68

60

0.95

0.84-1.00

1.60

725

23

1.49

1.42-1.57

50

0.95

0.88-1.00

1.57

731

50

1.58

1.42-1.76

50

0.93

0.88-1.00

1.69

completely dimorphic with respect to this character. This is shown
graphically in Fig. 2, where size-frequency distribution data for
three of the colonies are grouped. (Colony 710, whose queens and
workers are smaller than average, is omitted from Fig. 2 because its
smallest queens overlap in size with the largest workers of two of the
other colonies, obscuring the bimodality that exists within colonies.)
The ratio of average length of alitrunk in queens to that in workers
ranges from 1.12 to 1.16 for each of the four colonies sampled, with
an overall average of 1.15.

Allometry. The width of the first abdominal tergite is proportion-
ately much greater in queens than in workers. This is evident in the
large queen/worker ratios with respect to this dimension (Table II).
The regression of the width of the first abdominal tergite on alitrunk
length has been computed for queens and workers of each of the
four colonies, and the equilibrium constants (k, or slope of the re-
gression line) are presented in Table III.

In all colonies the allometry of the worker caste is negative. For
queens this shifts to strongly positive in all colonies except 725 (where
small sample size may have biased the result). The intranidal dif-
ferences in k for queens and workers are significant at the 1% level

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[March

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1974]

Jeanne & Fagen — • Stelopolybia

1.80

1.60

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£ 1.40

1.20

0.80

Queens

Workers

3.20 3.40 3.60 3.80 4.00 420 4.40

Alitrunk length (mm)

Fig. 2. Intranidal allometry and complete dimorphism in Stelopolybia
areata. Log-log plot of first abdominal tergite width against alitrunk length
for members of colony 731. Computed regression lines are shown. Bar
graph gives frequency distribution of alitrunk length among queens and
workers of three colonies (707, 725, 731). Numbers at the top of each bar
give number of individuals.

Psyche

[March

162

for two colonies, and not quite significant at the 5% level for the
other two (Table III).

A log-log plot of allometry in colony 731 is shown in Fig. 2. The
lack of alignment of the computed linear regression lines suggests
that the dimorphism in this species may have arisen out of triphasic
allometry with a dropping out of intermediates (Wilson 1953).

Discussion

The positive allometric growth of the first abdominal tergite in
queens probably has an adaptive significance for the reproductive role
of this caste. The expanded width of this segment reflects a general
increase in size of the gaster (Fig. 1), which is most easily explained
as an adaptation to accommodate the well-developed ovaries. Blackith
(1958) found a similar increase in body proportions toward the pos-
terior of queens compared to workers for two species of V espula.

The negative allometry of the first abdominal tergite in workers
must have other causes, however. It is possible that the narrower,
more petiole-like first segment in workers may permit a greater
maneuverability of the gaster and hence the sting, a feature that
could in turn be interpreted as an adaptation to the role of workers
in defense. Presumably the evolutionary divergence of the two castes
with respect to this character reflects an almost complete functional
sparation of the reproductive and defensive roles.

The variance of the width of the first abdominal tergite given the
length of the alitrunk is higher than that for ants (Wilson 1953).
It resembles more closely the variance for certain halictine bees
(Knerer & Atwood 1966). These differences may reflect a genetic
component to the variability observed. In mononogynous ants all the
workers of a colony are offspring of a single queen, and quite likely
of a single male. In S. areata , however, since the colonies are
polygynous, the individuals sampled do not all have the same parents,
and thus represent a larger pool of genetic variability.

A comparison of the degree of polymorphism shown by S. areata
with that of other vespids is of interest, though difficult because of
the lack of comparability of measurements used from one study to the
next. The Vespinae are biometrically the best known social wasps
(e.g. Wright, Lee, and Pearson 1907; Thompson, Bell, and Pearson
1910, 1911; Blackith 1958). They also show the greatest degree of
polymorphism. Blackith’s results for V espula germanica and V. rufa
are probably typical (Blackith 1958). For the sake of comparison let

1974]

Jeanne & Fagen — Stelopolybia

163

us choose his measurement of thorax width as a measure of body size.
Confidence that this indeed is a good indication of body size is gained
from the fact that the cube of the queen/worker ratio for this dimen-
sion closely approximates the ratio of the reduced body weights of the
two castes. The ratio of queens to workers for this character is 1.37
and 1.34 for V. germanica and V. rufa , respectively, and compares
with 1. 1 5 queen/ worker ratio for alitrunk length for S. areata
(Table IV). Thus there is considerably greater separation of queens
and workers with respect to average body size in Vespula rufa and
V. germanica than in S. areata.

In the subfamily Polistinae the degree of polymorphism appears to
range from virtually none to the degree of dimorphism shown by
S. areata. Almost nothing has been done biometrically with the sub-
family, with the exception of Richards and Richards’ analysis of
certain characters in some of the South American species (Richards
and Richards 1951). The primary aim of Richards and Richards’
analysis was to find reliable morphological characters that could be
used to distinguish queens from workers. Queens were defined as
having sperm in the spermatheca. In some species the two castes
differed most significantly in the number of hamuli, while in others
wing length, vertex width, mesonotum length, or shape of the first
abdominal tergite were more reliable. Of the characters measured
by Richards and Richards probably mesonotum length comes closest
to being an indicator of body size. At any rate, it is most comparable
to alitrunk length, used in the present study. If we use as an index
of queen/worker dimorphism the ratio of average mesonotum length
for queens to that for workers, the two species analyzed by Richards
and Richards have values considerably below the ratio of queen to
worker alitrunk length for S. areata (Table IV). Small but sig-
nificant differences with regard to at least one character were found
by Richards and Richards to exist between queens and workers in
Polybia bistriata, P. occidentals occidentals , Protopolybia pumila ,
P. minutissima, Brachygastra scutellaris, and Polistes canadensis
(Richards and Richards 1951).

Conclusion

Stelopolybia areata thus appears to have achieved a high degree of
queen/worker dimorphism relative to other species of Polistinae that
have been studied. Other species in the genus have been mentioned
in the literature in this connection. Queens of S. flavipennis , for

164

Psyche [March

^ vr>

ON

1974]

Jeanne & Fagen — Stelopolybia

165

example, figured by Evans and Eberhard (1970), have an enlarged
petiole. Both of these species form large colonies. This is consistent
with the conclusion of Richards and Richards (1951) that species
which form larger colonies tend to have more distinctive queens.

Acknowledgments

We are grateful to Dr. Antonio Lot Helgueras of the Department
of Botany, Universidad Nacional Autonoma de Mexico, for making
available the facilities of the Estacion de Biologia Tropical ‘Los
Tuxtlas’ and to members of the staff of the Estacion for their help.
Dr. Jose Sarukhan, Dr. Raul MacGregor, Dr. Arturo Gomez-
Pompa, and Dr. Carlos Marquez Mayaudon, all of UNAM, also
deserve thanks for their help in making arrangements for the field
work. Mr. Mark Winston helped in the field and with the measure-
ment of the wasps. Dr. O. W. Richards kindly confirmed the iden-
tification of the wasp. Dr. E. O. Wilson provided helpful com-
ments and advice in the preparation of the manuscript. The research
was supported by the National Science Foundation (GB-33619) and
by the Boston University Graduate School (GRS-303-B1 ) .

References

Blackith, R. E.

1958. An analysis of polymorphism in social wasps. Insectes Sociaux,
5 (2) : 263-272.

Ducke, A.

1910. Revision des guepes sociales polygames d’Amerique. Ann. Mus.
nat. Hungarici, 8: 449-544.

Evans, H. E. and M. J. W. Eberhard

1970. The wasps. University of Michigan Press, Ann Arbor, vi -f-
265 pp.

Hays, W. L.

1963. Statistics. Holt, Rinehart, and Winston, New York, xvi -f- 719 pp.
Jeanne, R. L.

1973. Aspects of the biology of Stelopolybia areata (Hymenoptera :
Vespidae). Biotropica, 5 (3): 183-198.

Knerer, G. and C. E. Atwood

1966. Polymorphism in some nearctic halictine bees. Science, 152(3726) :
1262-1263.

Lewis, D.

1960. Quantitative methods in psychology. McGraw-Hill, New York,
xii + 558 pp.

Michener, C. D.

1961. Social polymorphism in Hymenoptera. Symposium of the Royal
Entomological Society of London, 1: 43-56.

Psyche

[March

1 66

Richards, O. W. and M. J. Richards

1951. Observations on the social wasps of South America (Hymenop-
tera; Vespidae). Trans. Roy. Entomol. Soc. Lond., 102: 1-170.
Thomson, E. Y., J. Bell, and K. Pearson

1910. A second cooperative study of Vespa vulgaris . A comparison of
the queens of a single nest with queens of a general population.
Biometrika, 7: 48-63.

1911. A third cooperative study of Vespa vulgaris. Comparison of
queens of a single nest with queens of the general population.
Biometrika, 8: 1-12.

Wilson, E. O.

1953. The origin and evolution of polymorphism in ants. Quart. Rev.
Biol., 28(2) : 136-156.

Wilson, E. O.

1971. The insect societies. Belknap Press, Harvard University Press,
Cambridge, Mass., x + 548 pp.

Wright, A., A. Lee, and K. Pearson

1907. A cooperative study of queens, drones and workers in Vespa
vulgaris. Biometrika, 5 : 407-422.

BEHAVIOR OF THE NORTH AMERICAN TERMITE
TEN UIR OS TRI TERMES TEN UIROS TRIS,1
WITH SPECIAL REFERENCE TO THE
SOLDIER FRONTAL GLAND SECRETION,

ITS CHEMICAL COMPOSITION,

AND USE IN DEFENSE2

By W. L. Nutting,3 M. S. Blum4 and H. M. Fales5
Introduction

T enuirostritermes is a small genus of Neotropical termites with
two species ranging northward into warm temperate, semi-arid areas
of southwestern Texas \cinereus (Buckley)] or beyond into south-
eastern Arizona [ tenuirostris (Desneux)]. As a member of the
Nasutitermitinae, it is somewhat more primitive than the very large
and well-known tropicopolitan genus, N asutitermes. The subfamily,
containing the most highly specialized members of the Termitidae,
is the largest in the order Isoptera. The major specialization within
the subfamily has involved gradual reduction of the soldier mandibles
to non-functional stubs with concomitant development of a small
projection on the front of the head into a long, attenuated snout or
nasus. The frontal gland, occupying most of the bulbous posterior
of the head capsule, elaborates a defensive secretion which can be
forcibly ejected through the frontal pore at the tip of the nasus by
contraction of powerful mandibular muscles. This fontanellar gun
represents the apex of sophistication among the varied chemical de-
fense mechanisms of the termites. The zoogeography and affinities of
the nasutitermitine genera have been discussed by Emerson (1955)
and Krishna (1970) and their defensive behavior by Ernst (1959),
Moore (1969) and Wilson (1971). Light and Weesner (1955)

Termitidae, Nasutitermitinae.

journal Paper No. 2269 of the Arizona Agricultural Experiment Station.
The work on which this report is based was carried out as a part of the
U.S./I. B.P. Desert Biome, and was supported (in part) by National Science
Foundation Grant No. GB-15886.

department of Entomology, University of Arizona, Tucson 85721.

department of Entomology, University of Georgia, Athens 30601.

5Laboratory of Chemistry, National Heart and Lung Institute, Bethesda,
Md. 20014.

Manuscript received by the editor February 22, 1974.

167

Psyche

[March

1 68

and Weesner (1953, 1970) have made the only major contributions
on the general biology of T. tenuirostris , and Nutting (1970) has
summarized the information on its foraging behavior.

Materials and Methods

Seven collections of the termites were made in early September,
1972, on the Santa Rita Experimental Range, elevation between 945
and 1220 m, ca. 45 km south of Tucson, Arizona. The area is a
shrub-invaded desert grassland ecotone, characterized by scattered
trees, shrubs and cacti. Four groups were taken from under stones
during one afternoon and three nearly complete foraging groups were
taken on the soil surface near midnight. Termites and a minimum of
soil were quickly scooped up with a trowel and placed in a large
enameled tray. A few hours later the termites were separated from
soil and debris by sifting and hand-picking. To determine soldier/
worker ratios, these castes were counted in each of the two combined
collections, those from under stones, and those taken on the surface.
Several hundred of these soldiers and workers were reserved for
chemical analyses.

A small foraging group was collected later for behavioral studies.
They were supplied with fine, dry native vegetation and survived
reasonably well in the laboratory for about three weeks in a plastic
petri dish of moist soil. A few minor workers of the myrmicine ant,
P heul ole desertorum Wheeler, were also collected in the same area
and maintained for the observation of termite-ant encounters. Al-
though most species of Pheidole are reportedly seed gatherers rather
than habitual predators, this was apparently of no consequence in the
artificial situations described below.

No encounters between termite foraging groups and small arthro-
pod predators were seen in the field. A few random encounters were
staged in the laboratory by introducing individual ants into the dish
containing the small group of termites. The defensive behavior of
the soldiers was observed in detail under low magnifications of a
stereomicroscope, during individual soldier-ant encounters contrived
as follows: Single living ants were mounted in a natural position on
glass slides with a small drop of rubber cement. Slide and ant were
then placed on the soil in the observation chamber with the termites.
In this distinctly one-sided situation, one or more soldiers invariably
located and attacked the ant within a few minutes.

Attacks by five different soldiers were quite literally recorded on

1974]

Nutting, Blum , £sf Bales — Tenuirostritermes

169

separate slides in the form of discrete threads of frontal gland secre-
tion which they had fired in the general direction of the ants. The
volume of most of these shots was calculated from measurements
made of the threads with an ocular micrometer under a compound
microscope. Drawings to illustrate the variation of individual shot
patterns were made with the aid of a camera lucida on the same
microscope. The fate of a few ant victims was followed until their
death and compared with that of normal ants which were isolated
and allowed to die of starvation and desiccation.

For the chemical analyses 600 living soldiers were placed in re-
agent rc-pentane. Although it had been determined that the heads of
the soldiers were the primary source of the odorous defensive com-
pounds, 400 living workers were also extracted for comparative
purposes. The extracts were concentrated under vacuum and ana-
lyzed on a gas chromatograph interfaced to an LKB-9000 mass spec-
trometer. A 4-meter column of 10% SP-1000 on 80/100 mesh
Supelcoport operated isothermally at 65 °C was employed for all
analyses.

Results

Foraging behavior. — T. tenuirostris forages in the open on the
soil surface, mainly at night in southern Arizona as we have recently
established, but it is also active on cloudy days, at least in western
Mexico (Nutting, 1970). The following account of its general
foraging behavior is a synthesis of observations made on about 12
different groups at the Santa Rita Experimental Range during Sep-
tember-October, 1972.

Workers presumably open one or more access holes to the surface,
but the soldiers are generally the first out and the last to return
underground during bouts of surface activity. As many as 100 or
more soldiers and very few workers may congregate within a 5- to
10-cm radius of the hole prior to foraging. Little organization is at
first apparent except that many of the soldiers may be standing on
the alert with heads pointed peripherally. One or more discrete for-
aging columns eventually emerge from such unorganized, milling
masses, perhaps after the soldiers are joined by a critical number of
workers. Once in motion, columns up to six workers wide move
quickly in an ant-like manner, and show very tight trail-following
behavior. While soldiers move out with the workers, they eventually
take up stations at 1- to 2-cm intervals on either side of, and ca. 1 cm

170

Psyche

[March

from, the worker traffic. Columns occasionally branch and have been
observed as long as ca. 2 m.

Irregular craters of soil, up to 8 cm in diameter and i cm high,
are often thrown up around the holes during periods of surface
activity. Ground-level runways radiating from the hole may be
bridged over to transform a crater into a rough pile of soil 2 to 3 cm
high. Since individuals of this species are relatively small (length
of soldier, 3.0 mm; of worker, 4.5 mm), access holes as large as
5 mm in diameter probably provide for efficient, emergency re-entry
of large numbers of foragers in a very short time. Workers have
been observed to close the exposed holes shortly after the conclusion
of surface activity and, although a few soldiers remain in and around
the hole during closure, none was ever left outside. The same holes
are apparently used repeatedly, at least for periods of several weeks.

The high proportion of soldiers in the foraging groups indicates
the relative importance of this caste in those species of termites which
routinely forage on exposed trails. The four combined groups col-
lected from colonies beneath stones contained 545 individuals with a
soldier rworker ratio of 1:3.82 (20.7% soldiers). Three combined
foraging groups contained 1,549 individuals with a soldier :worker
ratio of 1:1.23 (44-9% soldiers). A single foraging group of 230,
perhaps recently organized, had a ratio of 1 soldier to 0.40 worker
(71-3% soldiers).

Soldier behavior. — Whenever the laboratory group of soldiers and
workers was disturbed by jarring or removing the cover of their
container, several soldiers, with antennae raised and waving, congre-
gated on the edges of objects and on elevated vantage points. In-
dividual workers of the ant, Pheidole desertorum (ca. 3.5 mm long),
introduced into this situation were quickly fired on at very close
range by several soldier termites in rapid succession. The appendages
of such victims were almost completely immobilized, within 10 sec or
less, by the viscous, glue-like threads thrown over them.

Further details of soldier defensive behavior are based on observa-
tions made during encounters with living ant workers fixed to glass
slides. Although blind, the soldiers presumably orient toward such
threats by olfactory or auditory cues. An alerted soldier closes rapidly
on any potential target and may actually touch it for an instant with
its antennae before firing — from a distance of ca. 0.5-3.0 mm). As
it fires, it may jerk forward or backward, in position, and may or
may not then oscillate to-and-fro one or more times. This apparently
insignificant bit of behavior greatly enhances the effectiveness of the

1974]

Nutting , Blum , & Fales — Tenuirostritermes

171

Fig. 1. Patterns of the threads of frontal gland secretion fired by five
different soldiers of Tenuirostritermes tenuirostris at living ants fixed to
glass slides. Arrows show direction of target and scale of 1 mm.

172

Psyche

[March

Table 1. Measurements of shots or threads fired by four soldiers of
T enuirostritermes tenuirostris at fixed, living ants in the laboratory.

Soldier

Number

Shot

Number

Av. Diameter
(mm)

Length

(mm)

Volume

(mm3)

1

1

0.020

19.650

0.0062

2

0.015

11.425

0.0020

3

0.015

10.475

0.0019

4

0.015

7.550

0.0013

5

0.015

5.100

0.0009

6

0.015

2.500

0.0004

2

1

0.020

9.025

0.0028

2

0.020

7.300

0.0023

3

1

0.020

7.200*

0.0023

2

0.020

4.625

0.0015

3

0.020

1.725

0.0005

4

1

*

2

0.020

0.900

0.0003

* Shot entirely on ant and

not traceable, or

partly on ant and

measurement

only approximate.

soldier’s shot by throwing a bight, or even one or more loops, in the
thread (Fig. i). Attacks by as many as six soldiers from different
directions were invariably fatal to an ant. Soldiers often examined
victims with their antennae for up to a minute, occasionally becoming
temporarily entangled themselves. They were never seen to wipe the
nasus on the substrate after any of this activity.

In unstaged encounters, soldiers usually fired but once at an ant;
however, by carefully prodding and exciting the fixed ants, individual
soldiers were induced to fire from one to six additional shots at them.
Data on the shots recorded on and near four such ants are summar-
ized in Table i. It was impossible to determine the order of shots
fired by individual soldiers, but those of greater diameter and length
are probably the ones fired at or near the beginning of an encounter.
Longer shots often began with a series of droplets, and occasionally
contained short breaks in the thread. It would have been interesting
to determine what percent of their glandular holding capacity had
been spent in these individuals, but drought conditions during seasons
of activity have since prevented further collections.

Although the ant victims were physically immobilized within a
few seconds, they also appeared to be adversely affected by some of the
chemical components of the soldier secretion, probably the terpenes.

1974]

Nutting , Blum, & Fales — T enuirostritermes

173

They lay very quiet, moving their appendages but little even when
prodded. Their struggles were very feeble after 1.5 hr, while only
occasional, slow contractions could be elicited after 4 hr. They ap-
peared essentially dead after 5-6 hr. By comparison, normal ants,
similarly confined without food or water, remained so for 5-10 hr,
showed postural difficulties after 10-12 hr, responded vigorously to
prodding for as long as 24 hr, and did not appear to be dead until
24-33 hr after confinement.

Chemical A nalyses. — Three compounds accounted for over 90%
of the low boiling volatile compounds which were detected in the
extracts of the soldiers. Based on the congruencies of their gas-
chromatographic retention times and mass spectra, the volatiles were
identified as a-pinene, myrcene, and limonene. The mole percentages
of the compounds were: a-pinene 62%, myrcene 27%, and limonene
11%. None of these monoterpene hydrocarbons was detected in the
extracts of the workers.

Discussion

Ernst (1959) was inclined to believe that the frontal gland secre-
tion of soldiers of a N asutitermes sp. from the Ivory Coast was
essentially non-toxic and that it functioned primarily to entangle its
enemies. He did not rule out the possibility of an insecticidal effect.
Considering that normal worker ants survived without food or water
4 to 6 times as long as those immobilized by the T. tenuirostris sol-
dier secretion, we feel that the terpenoid components of its secretion
are definitely toxic.

Ernst also observed that the secretion fired by one of his Nasuti-
termes soldiers stimulated nearby soldiers to fire, thus suggesting
that it contains volatiles which act as an alarm pheromone. On the
basis of his own work with several Australian species of N asutitermes,
Moore (1969) has suggested that a-pinene might serve as such a
pheromone. This compound probably serves a similar function in
T rinervitermes geminatus Wasmann (Nasutitermitinae) according
to Quennedey (1973). Although we have not investigated this point,
our observations suggest that any such alarm signal in the secretion
of T. tenuirostris must act very briefly indeed. In a number of
laboratory encounters between soldiers and free ants, several soldiers
might fire once each on a single ant within a few seconds; however,
late arrivals usually examined the resulting immobilized ants, but
never fired on them.

174

Psyche

[March

Moore (1968) determined that both limonene and a-pinene were
rather typical components of termitid soldier secretions whereas myr-
cene was detected only in one species of Amitermes. Vrkoc et al.
have recently reported the major volatile components in the soldier
frontal gland secretion of Nasutitermes rippertii to be a- and /3-
pinene and limonene, with myrcene among six minor constituents.
Similarly, with N. costcilis they found a- and /?-pinene and limonene
among six major constituents of the secretion, and myrcene again
among three minor components. However, although it is evident that
monoterpene hydrocarbons are rather characteristic defensive products
of nasute soldiers, these compounds do not appear to have a wide-
spread distribution in the Insecta. Indeed, with the exception of the
myrmicine ant Myrmicaria natalensis F. Smith, which produces
limonene in its poison gland (Griinanger et al., i960), monoterpene
hydrocarbons have not been identified in any other insect taxa.

By taking high speed motion pictures of Nasutitermes soldiers
attacking vestigial- winged Drosophila , Ernst (1959) determined that
they move rapidly forward and backward (only once?) in delivering
the jet of frontal gland secretion. He also observed one soldier to
move its head from side to side during an attack. Our tracings of
the shot patterns (Fig. 1) show that either or both of these move-
ments are also used by soldiers of T. tenuirostris to increase the
effectiveness of their defense. The basic oscillatory behavior probably
represents the jerking or rocking movements which termites com-
monly exhibit in mild alarm situations. We have also shown that
soldiers are capable of firing more than once, if sufficiently alarmed,
and certainly do not deliver the entire contents of the frontal gland
in a single discharge as Ernst suggested.

Many authors have stated that ants are the chief predators of social
insects, with the termites not the least among their prey (Wilson,
1971). There can be little doubt that the nasute soldier is a highly
effective defender of the colony against arthropod predators close to
the size range of the termites themselves. A large force of nasutes
would seem to be of critical importance in protecting the workers on
their food-gathering expeditions, particularly those that forage in the
open as does T. tenuirostris. However, this common assumption
apparently rests largely on casual, subjective observations and cir-
cumstantial evidence of the type presented here. Quantitative data
required to prove this assumption are not available and would be
extremely difficult to obtain.

1974]

Nutting , Blum, & Fales — Tenuirostritermes

175

The high proportion of soldiers maintained by T. tenuirostris, and
presumably many other free-foraging termites, certainly attests to
their great importance in the colony. Our figure of 20% soldiers in
combined collections from four different colonies is in fair agreement
with Light and Weesner (1955) who determined that incipient
colonies of this species usually contained between 25 and 33%
soldiers. Three of our foraging groups combined contained 44.9%
soldiers, while a single group contained 71.3%. Ernst (1959) re-
ported 70 to 90% soldiers in outer portions of his laboratory colony
of N asutitermes sp. from the Ivory Coast. Although demands on
the defensive capabilities of the soldiers underground must be trivial
or rare, indeed, these figures show that a very high proportion of
soldiers is mobilized for foraging parties.

This situation is in striking contrast to that in another subterranean
termitid, Gnathamitermes perplexus (Banks) ( Amitermitinae) ,

which is very common over the range of T. tenuirostris in southern
Arizona. This species forages widely on the surface under cover of
thin, soil sheeting which it builds over all types of dead plant ma-
terial (Collins et al., 1973). Based on 74 foraging groups, the
average soldier rworker ratio is 1:89 (1.11% soldiers). Individual
groups ranged from 1 to 1031 termites and contained from 0.0 to
9.10% soldiers (Nutting and Haverty, unpublished).

During the few hours of field observations on the above-ground
activities of T. tenuirostris , no encounters with ants or other small
predators were seen. However, the presence of an impressive ant
fauna in our study area lends further credibility to the importance
of the soldier caste in the economy of this termite. In a preliminary
study of the ants on the Santa Rita Experimental Range, Gaspar and
Werner (unpublished) list 33 species, with high populations of
Crematogaster coarctata vermiculata Emery and Forelius foetidus
(Buckley), and estimate 1753 nests/ha for all species. About one
fourth of the species are regular predators, while at least an addi-
tional third are occasional predators or at least feed on animal matter.
Many of them forage at night during the summer rainy season when
T enuirostritermes is also most active on the surface. Although we
have no estimates of colony size or density for the termite, its colonies
are fairly common in the area and must contain several thousand
individuals.

176

Psyche

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Acknowledgments

We wish to thank Dr. Ch. Gaspar, Laboratoire de Zoologie
generale et de Faunistique, Faculte des Sciences Agronomiques de
l’Etat, Gembloux, Belgium, for identifying the ant used for our
laboratory observations, and Mr. Michael I. Haverty, Research
Associate, Department of Entomology, University of Arizona, for
his assistance in collecting the termites.

Literature Cited

Collins, M. S., M. I. Haverty, J. P. LaFage and W. L. Nutting.

1973. High temperature tolerance in two species of subterranean ter-
mites from the Sonoran Desert of Arizona. Environ. Entomol.
2: 1122-1123.

Emerson, A. E.

1955. Geographical origins and dispersions of termite genera. Field-
iana: Zool. 37: 465-521.

Ernst, E.

1959. Beobachtungen beim Spritzakt der Nasutitermes- Soldaten. Rev.
Suisse Zool. 66: 289-295.

Grunanger, P., A. Quilico and M. Pavan.

1960. Sul secreto odoroso del formicide Myrmicaria natalensis Fred.
Accad. Nazion. Lincei 28: 293-300.

Krishna, K.

1970. Taxonomy, phylogeny, and distribution of termites, pp. 127-152.
In “Biology of termites,” vol. 2, (K. Krishna and F. M. Weesner,
eds.). Academic Press, New York.

Light, S. F., and F. M. Weesner.

1955. The incipient colony of T enuirostritermes tenuirostris (Desneux).
Insectes Sociaux 2: 135-146.

Moore, B. P.

1968. Studies on the chemical composition and function of the cephalic
gland secretion in Australian termites. J. Insect Physiol. 14:
33-39.

1969. Biochemical studies in termites, pp. 407-432. In “Biology of
termites,” vol. 1, (K. Krishna and F. M. Weesner, eds.). Aca-
demic Press, New York.

Nutting, W. L.

1970. Free diurnal foraging by the North American nasutiform ter-
mite, T enuirostritermes tenuirostris (Isoptera: Termitidae).
Pan-Pacific Entomol. 46: 39-42.

Quennedey, A.

1973. Observations cytologiques et chimiques sur la glande frontale des
termites. Internat. Union for the Study of Social Insects, Proc.
Vllth Internat. Congress, London, 10-15 Sept., 1973. pp. 324-326.

1974]

Nutting , Blum , & Bales — T enuirostritermes

177

Vrkoc, J., K. Ubik, L. Dolejs and I.Hrdy.

1973. On the chemical composition of frontal gland secretion in ter-
mites of the genus N asutitermes ( N . costalis and N. rippertii;
Isoptera). Acta Entomol. Bohemoslov. 70: 74-80.

Weesner, F. M.

1953. The biology of T enuirostritermes tenuirostris (Desneux) with
emphasis on caste development. Univ. Calif. Publ. Zool. 57:
251-302.

1970. Termites of the Nearctic Region, pp. 477-525. In “Biology of
termites,” vol. 2, (K. Krishna and F. M. Weesner, eds.). Aca-
demic Press, New York.

Wilson, E. O.

1971. “The insect societies.” The Belknap Press of Harvard Univ.
Press, Cambridge, Mass. 548 pp.

THE CASTIANEIRINAE OF MEXICO. I.
CASTIANEIRA DUG ESI (BECKER)
(ARANEAE: CLUBIONIDAE)*

By Jonathan Reiskind
Department of Zoology, University of Florida,

Gainesville, Florida 3261 1

Since the revision of the Castianeirinae of North and Central
America (Reiskind, 1969) new collections have been made and
studied. This is the first of a series of papers that will update the
Mexican component of this subfamily.

Castianeira dugesi (Becker) has been known from a single male
described almost a century ago. Recent collections from locations
within 50 miles of the type locality have produced additional speci-
mens including the first females. C. dugesi is the nominal species in
the dugesi species-group (which also includes C. nanella Gertsch and
C. alfa Reiskind), a group restricted to the dry regions in south-
western U. S. and northwestern Mexico and characterized by its
moderately small size, lack of plumose setae, and distinctive genitalia
(Reiskind, 1969). As is characteristic of many ant mimicking spe-
cies of Castianeira with no distinct morphological separation of
cephalic from thoracic region the cephalic region in C. dugesi is
marked off by color pattern which we can assume reflects the color
pattern of the model ant.

Castianeira dugesi (Becker)

Figures 1-3

Micariaulax dugesii Becker, 1879, Ann. Soc. Entomol. Belgium, 22: 83, pi. 2,
fig. 9, 10, $, from Guanajuato, Mexico; in the Brussels Museum.
Castianeira dugesii: Simon, 1897, Histoire Naturelle des Araignees 2(1) : 167.
Castianeira dugesii: F. P.-Cambridge, 1899, Biol. Centrali-Americana.

Arachnida 2 : 81.

Castianeira dugesii: Reiskind, 1969, Bull. Mus. Comp. Zool. 138(5): 223-
225, pi. 10, fig. 127, 128.

FEMALE

Measurements. Based on two females: carapace length 2.65-
2.70 mm; carapace width 1.70- 1.75 mm; carapace index (carapace
width carapace length X 100) 64; sternum length 1.15 mm;

* Manuscript received by the editor February 28, 1974

178

1974]

Reis kind — Castianeirinae

179

Figures 1-3. Castianeira dugesi (Becker). Fig. 1. Dorsal aspect of fe-
male. Pattern division of carapace shown by shading. (Vertical scale
line = 1.0 mm) Fig. 2. External epigynum. Fig. 3. Internal epigynum,
dorsal view. (Horizontal scale line for epigyma = 0.5 mm)

i8o

Psyche

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sternum width 0.95 mm; sternum index (width -i- length X 100)
80.

Femur IV length 1.75-1.85 mm; femur IV width 0.45-0.55 mm;
leg thickness index (femur IV width femur IV length X 100)
26-29; leg length index (femur IV length carapace length X
100) 67.

Abdomen length 4.00 mm; abdomen width 2.30-2.35 mm; abdo-
men index (width length X 100) 57-58.

Description. Carapace orange-brown and hairless, with cephalic
region dark brown (Fig. 1). Carapace moderately narrow in head
region (width at level of second eye row 0.59-0.61 times the maxi-
mum width of the carapace) and smoothly truncated anteriorly.
Eyes moderately small, equal with posterior row somewhat wider
than anterior. Distinct thoracic groove.

Abdomen oval, grey-brown, with a large anterior dorsal dark
red-brown scutum (about one-third the length of the abdomen)
(Fig. 1); covered with sparce, fine setae. Epigastric and small in-
framammiliary scuta red-brown. Both pairs of anterior abdominal
setae hairlike.

Sternum orange-brown with light, thin setae.

Chelicerae dark-brown (like cephalic region of carapace) with
two moderate retromargin teeth and two slightly larger promargin
teeth. Pedipalps with red-brown femur and rest yellow.

Coxa I yellow-brown, rest of coxae yellow. A small notch on
trochanter IV.

Femur I dark brown with yellow distal end, rest of leg I and
leg II and III yellow; leg IV dark yellow with orange metatarsus.
Tibia I ventral spination: 2 (prolateral) -2 (retrolateral) and small.

External genitalia with a pair of wide, flared openings (Fig. 2).
Internal structure with globose spermathecae drawn out into narrow
straight posterior necks ( Fig. 3 ) .

MALE

Measurements. Based on two males: carapace length 2.05 mm;
carapace width 1.25- 1.30 mm; carapace index 61-63; sternum length
0.95-1.00 mm; sternum width 0.75 mm; sternum index 73-77.

Femur IV length 1.40- 1.45 mm; femur IV width 0.35 mm; leg
thickness index 24-25 ; leg length index 69-71.

Abdomen length 2.45-2.60 mm; abdomen width 1.15 mm; abdo-
men index 45-47.

Embolus length 0.06-0.07 mm; genital bulb length 0.63-0.65 mm;
male genital index (embolus length bulb length X 100) 9.5-10.

1974]

Reiskind — Castianeirinae

1 8 1

Description. Similar to female with the following differences:
Only the anterior portion of carapace cephalic region dark-brown.
Abdomen with a full scutum, granulose at anterior end and gradu-
ally becoming smooth and shiny posteriorly. Tibia I ventral spina-
tion: i-O and small. Genital bulb drawn out into a long neck ending
with an embolus having a right angle near tip (illustrated in Reis-
kind, 1969).

Diagnosis. C. dugesi differs from other dugesi group members in
its genitalia (right angle at end of male embolus and straight sper-
mathecal necks) , distinct patterns of carapace and legs.

Distribution. Apparently this species is restricted to a small region
of central Mexico in northeastern Jalisco and Guanajuarto states.

Records. Mexico. Guanajuarto. Jallisco: 10.8 mi. S. Talpa de
Allende, 4900', 8-VIII-67 (R. E. Leech) ; 13 mi. SE. Lagos de
Moreno, 6450', 7-IX-67 (R. E. Leech, G. E. Ball).

The female of Castianeira dugesi would key out to couplet #36
in the Key to the Females of Castianeira in Reiskind ( 1969). It can
then be separated from the other species included in that couplet by
its dark abdomen and bicolored carapace.

I wish to thank Dr. Robin E. Leech for the generous loan of the
specimens studied in this paper.

Literature Cited

Reiskind, J.

1969. The Spider Subfamily Castianeirinae of North and Central
America (Araneae, Clubionidae) . Bull. Mus. Comp. Zool. 138(5):
163-325.

THE SOLDIER OF THE ANT
CAMPONOTUS ( COLOBOPSIS ) FRAXINICOLA
AS A TROPHIC CASTE*

By Edward O. Wilson
Museum o-f Comparative Zoology Laboratories,

Harvard University, Cambridge, Massachusetts 02138 U.S.A.

Complete dimorphism, defined as the coexistence of a minor worker
of ordinary proportions with a larger major worker or “soldier,”
represents the pinnacle of the subcaste system within the ants. The
soldier not only weighs more but also possesses a disproportionately
larger head. By definition, intermediate forms are lacking. Complete
dimorphism has originated independently at least seven times, within
the following genera: the myrmicines Pheidole, Oligomyrmex ,

Acanthomyrmex, Paracryptocerus; the dolichoderine Zatapinoma;
and the formicines Camponotus and Pseudolasius. The head shapes
of the major workers of these groups are clearly modified either for
fighting or for defense of the colony by blocking the nest entrances.
These functions have been confirmed by direct observations of col-
onies of Pheidole , Paracryptocerus , and Camponotus (Wilson, 1971).
The behavioral repertory of the major workers is otherwise very
limited in comparison with that of the minor workers, giving logic
to their alternate designation in the literature as soldiers.

The principal purpose of this article is to demonstrate that in at
least one species, Camponotus ( Colobopsis ) fraxinicola M. R. Smith,
the soldier caste also plays a key role in liquid food storage.

Materials and Methods

Colonies were collected at St. Mark’s Lighthouse and Tall Tim-
bers, near Tallahassee, Florida. The taxonomic identification re-
quires a brief note. Three forms of the southeastern United States,
fraxinicola , impressus , and pylartes , are very similar to each other
and may prove synonymous. The Tallahassee series fall closest to
fraxinicola in the seemingly best character states of the major worker:
a slightly more flattened mesonotum, more rounded posterior rim of
the truncated portion of the head, and in other, more subtle details
of head shape. But all of these characters vary widely within and
among series 'from widespread localities, so that eventually only one

*Manuscript received by the editor February 28, 1974.

182

1974]

Wilson — Camponotus fraxinioola

1 8 3

species might be recognizable, which would then take the name
impressus.

The colonies were transferred to glass tubes 15 cm in length and
3-6 mm in inner diameter. The tubes were each plugged at one end
with dry wads of cotton wool and placed in open plastic containers,
the inner sides of which were lined with fluon to discourage climbing
by the ants. Since the latter method does not always work with these
highly arboreal ants, the containers were also supported by glass
bottles set in petri dishes filled with heavy mineral oil. Water was
made available in moistened cotton plugs at the bottom of test tubes
placed next to the tubes housing the colonies. The ants were fed
daily with honey and freshly killed insects. Entire colonies quickly
habituated to strong light. They could be observed in toto with a
swing-arm dissecting microscope without being disturbed in their
new, highly simplified ( and fluon-lined ) universe.

The Soldier as a Defensive Caste

Since the time of Forel (1874) it has been known that soldiers of
the subgenus Colobopsis of Camponotus use their oddly cylindrical
heads to block the nest entrances. Minor workers returning from
foraging trips identify themselves to the soldiers, presumably by
colony odor. The soldiers then pull back to let them enter. C. frax-
inicola, like most other Colobopsis , nest in the cavities of dead twigs.
The nest entrances are neat, circular holes into which the heads of
the soldiers fit snugly. By plugging glass tubes containing colonies
with disks of cork, I was able to observe the excavation of the nest
entrances on repeated occasions. The task was performed exclusively
by the minor workers. This is somewhat surprising in view of the
fact that the holes are cut to fit the specifications of the soldiers and
not those of the minor workers. The soldiers also rested more con-
sistently near the entrances of the glass tubes, with their heads point-
ing outward in a high proportion of cases, even when they were not
actively engaged in blocking the entrances.

Undecane, stored in Dufour’s gland of the abdomen, is a general
formicine alarm pheromone (Wilson and Regnier, 1971). When
small quantities of this substance were allowed to evaporate near the
nest entrance, all members of the fraxinicola colonies were thrown
into the typical excited running movements of the fraxinicola alarm
response. But some of the soldiers moved to the nest entrances, fill-
ing those holes still unplugged at the start of the alarm reaction.

184

Psyche

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Figure 1. Portion of a laboratory colony of Camponotus ( Colobopsis )
fraxinicola housed in a glass tube. The large soldiers use their cylindrical
heads to block the circular nest entrances, which are cut by the minor
workers. This colony has been fed ad libitum, and consequently both soldiers
and minor workers are in replete condition. In the center, a soldier and
minor worker exchange food by regurgitation.

When twigs containing fraxinicola colonies are first broken open,
both minor workers and soldiers rush out. Many attack any accessible
alien object, such as the observer’s hand or a bit of cloth offered to
them, biting it and spraying it with formic acid. The same response
was obtained in the laboratory by permitting fire ant workers ( Sol -
enopsis invicta) to invade the nests. Individuals of both castes were
about equally aggressive and effective in repelling these invaders. On
the other hand, the total population of minor workers, by virtue of
its greater size, was more effective than that of the soldiers.

To sum up the results, the fraxinicola soldiers are indeed a defen-
sive caste, but their specialization makes them superior in only one
aspect of this role.

The Soldier as a Trophic Caste

The fraxinicola major worker is also anatomically distinguished
by its proportionately larger abdomen. All individuals dissected from
two laboratory colonies had large fat bodies and well developed
ovaries containing one to seven eggs, some of which were of very

1974]

Wilson — Camponotus fraxinicola

185

large size. A few minor workers with distended abdomens taken
from within the nest had similar abdominal contents, but the ma-
jority from both within and outside the nest had proportionately
much smaller fat bodies and reduced ovaries. Thus the soldiers store
substantially more food in the form of fat than do the minors. The
nature of their eggs has not been ascertained. Should they prove to
be trophic eggs, this form of storage would be primarily the province
of the soldier caste.

The following data reveal that the soldiers also store dispropor-
tionately large quantities otf liquid food in their crops. When colonies
are fed ad libitum with saturated sucrose solution or honey, most of
the workers in both subcastes become repletes. Repletism has been
defined for purposes of this analysis as distention of the abdomen to
the extent that the intersegmented membranes are exposed, permitting
the interior of the crop to be seen when light is transmitted from
below. It has been repeatedly observed in our laboratory colonies
that a higher proportion of the major workers — sometimes all of
them — become repletes when the colony is offered a superabundance
of liquid food. But when the colony is starved, the proportion of
repletes among the majors drops below that off the minors. Two
examples are presented in Figure 2.

The amount of liquid stored by each major is substantially greater
than that stored by each minor, on both an absolute and per-unit-
weight basis. This difference, which was first guessed by simple in-
spection, was proved by the following series of measurements. Twenty
minors and 11 majors were selected at random and weighed from a
colony anesthetized after two weeks of starvation. Then the colony
was fed to satiety over a 24-hour period and anesthetized again; 20
minors and 16 majors were next selected at random from among the
replete individuals and weighed. The mean weight of the starved
minors was 1.62 mg (range 1.29-2.42 mg) ; after feeding, their mean
weight was 2.44 mg (range 1.75-3.33 mg), a gain of 0.82 mg or
50.6 percent. The mean weight of the starved majors was 3.44 mg
(range 2.38-4.65 mg); after feeding, their mean weight was 5.62
mg (3.14-7.34 mg), a gain of 2.18 mg or 63.4 percent.

The disparity in storage capacity can be seen even more clearly by
examining the colony as a whole. The colony labelled No. 1 in
Figure 2 can be taken as typical; it contained 139 minors, 26 majors,
and a single queen. Using the data on weight gain and percentage of
repletism during a single experimental run, the following storage
capacities were estimated: the entire minor population stored 88.52

FREQUENCY OF REPLETES FREQUENCY OF REPLETES

1 86

Psyche

[March

Figure 2. The frequency of repletes among the soldiers and the minor
workers respectively in two colonies of Camponotus ( Colobopsis ) fraxini-
cola. The ants were fed to satiation with sucrose solution on three occa-
sions ten days apart and starved in the intervening periods. The soldier
caste consistently achieved a higher level of repletion when the colony was
fed but surrendered it to a greater extent when the colony was starved.

1974]

Wilson — Camponotus fraxinicola

187

mg, and the entire major population stored 54.59 mg. Thus although
the majors made up a little less than 16 percent of the population and
contained 28.43 percent of the wet weight in the non-replete condi-
tion, they stored 38.15 percent of the liquid at repletion.

A distinctive history of liquid flow unfolds when a colony dis-
covers a single rich source of food and then endures a period of
starvation — the circumstance simulated by the laboratory experi-
ments. When the food is first discovered, the flow during the first
one or two hours is from the foraging minor workers to other minors
and majors encountered back at the nest. After saturation is attained,
and 90 percent or more of the adults are replete, regurgitation con-
tinues at a high rate. The two castes participate at about the same
per- worker rate, with no apparent difference between the majors and
minors in the frequency of donation as opposed to that of reception.
Within 2-3 days after the food is cut off, majors have begun to feed
minors larger quantities than they receive. Data on exchanges show
that the major-to-major donations are fewer than would be expected
by chance alone. This is due at least in part to the fact that the
majors are relatively sluggish in their movements. The minors are
much the more active caste, passing from one nestmate to another to
collect and pass along the dwindling supply of liquid food.

Acknowledgement

This article is part of a continuing study on caste systems sup-
ported by Grant Number GB-40247 from the National Science
Foundation.

Summary

(1) The major worker, or “soldier,” of Camponotus ( Colobopsis )
fraxinicola , helps to defend the nest by blocking the entrance holes
with its head, a behavior pattern reported by previous authors in
other members of the genus. However, when the nest is breached and
combat ensues, the soldier is no more aggressive or effective than the
minor worker.

(2) The nest entrances are constructed entirely by the minor
workers, which cut them to fit the cylindrical heads of the majors.

(3) Major workers respond to undecane, a general formicine
alarm substance, by moving to block those nest entrances still open
at the beginning of the episode.

1 88

Psyche

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(4) Major workers also serve as a storage caste. Their abdomens
are partially filled with exceptionally large fat bodies. They also
store a disproportionate share of sugary liquids collected by the col-
ony. When this food is superabundant, the majors reach a higher
level of repletion. When the colony is subsequently starved, the
majors regurgitate the liquid back to the rest of the colony faster
than they receive it.

Literature Cited

Forel, A.

1874. Les fourmis de la Suisse. Societe Helvetique des Sciences Natu-
relles, Zurich, iv + 452 pp.

Wilson, E. O.

1971. The insect societies. Belknap Press of Harvard University Press,
x + 548 pp.

Wilson, E. O. and F. E. Regnier.

1971. The evolution of the alarm-defense system in the formicine ants.
American Naturalist, 105: 279-289.

CHEMICAL DEFENSE AND SOUND PRODUCTION
IN AUSTRALIAN TENEBRIONID BEETLES
( ADELIUM SPP.)*

By Thomas Eisner, Daniel Aneshansley, Maria Eisner,
Ronald Rutowski, Berni Chong, and Jerrold Meinwald

Section of Neurobiology and Behavior, and Department of Chemistry,
Cornell University, Ithaca,, N. Y. 14850

Introduction

Girdled eucalyptus trees, felled to make room for grazing pasture,
are a common sight in rural Australia. In the dead and decaying
stumps that are strewn through the countryside, many insects flourish.
Two of these, the congeneric tenebrionid beetles Adelium percatum
and A. pustulosum (subfamily Adeliinae) possess interesting defense
mechanisms that we here describe. Both have eversible abdominal
glands such as are commonly found in Tenebrionidae (Roth, 1945),
but some features of the chemistry and biology of the glands are
anomalous. Moreover, A. pustulosum has a stridulatory apparatus
that may function for acoustical reinforcement of the chemical de-
fense.

Materials and Methods

Both species of Adelium are black and, lacking hindwings, are
flightless. Several dozen specimens were taken singly and in small
groups in and under pieces of decaying eucalyptus wood in sheep
pasture near Gudgenby, Australian Capital Territory, in mid-April,
1973 (Fig. 2). They were maintained in plastic containers, and given
water, sucrose, and a variety of cereal-based foods. Initial behavioral
and electronmicroscopic studies were done at the laboratories of the
Division of Entomology, C. S. I. R. O., Canberra, where T. E. and
M. E. were guests. Secretion was shipped to B. C. and J. M. for
chemical analysis at the University of California, San Diego. Some
beetles survived travel to Cornell, where acoustical studies were made
by D. A., R. R. and T. E.

Scanning electronmicrographs were made with a JEOL (JSMU-
UE) instrument. Preservation of beetles with everted glands was

*Paper no. XXXIX in the series Defense Mechanisms of Arthropods.

Manuscript received by editor March 5 , 1974.

89

190

Psyche

[March

effected by first immersing them in cold ( — I95°C) liquid Freon
(while held in forceps to cause eversion) and then transferring them
to a tissue freeze-dryer for desiccation.

Instruments for chemical analyses included conventional gas chro-
matographs and an LKB-9000 gas chromatograph-mass spectrometer.

Sound recordings were made at 23-25°C with a condenser micro-
phone from a Holgate Ultrasonic Receiver MK.V [frequency re-
sponse ± 12 db from 1 to 100 kHz; sensitivity at preamplifier out-
put 268 mv (rms)//ibar at 60 kHz (determined by substitution)],
an Ithaco Amplifier Model 225 [variable amplification range from
O to 80 db; flat frequency response (within 3 db) in range of 0.5
Hz to 100 kHz] and a Lockheed Magnetic Tape Recorder/Repro-
ducer Model 419 [recording speed set at 30 inches/sec; flat fre-
quency response (within 3 db) in range from 0.2 to 100 kHz; input
sensitivity of 1.4 volt (1.0 volt rms) for ± 40% deviation]. Sound
intensity was measured with a Briiell and Kjaer Precision Sound
Level Meter Type 2203 and a Briiell and Kjar Condenser Micro-
phone Cartridge Type 4145. The intensity measurements were
limited to audio frequencies due to limitations in the frequency
response of the Briiell and Kjaer Microphone [flat frequency re-
sponse (within 2 db) in range from 2.6 Hz to 18.5 kHz].

Instruments used for displaying and recording temporal patterns
of the sound included a Tektronix storage oscilloscope (Model
4103N) with differential amplifier (Model 5A22N) and Polaroid
camera, and a Tektronix dual beam oscilloscope (Model 502 A) with
a Grass Kymograph camera (Model C4N). Timing marks were
generated on the oscilloscope with a Hewlett-Packard function gen-
erator (Model 3310A).

Sound spectograms were made with a Kay Elemetrics Sonograph
(Model 7029A). Low frequency background noise was filtered as
needed during playback of the recordings with a Krohn-Hite filter
(Model 3550).

Chemistry of the Secretion

It was clear from the outset that the secretion of both species was
quinonoid in nature. The beetles responded typically to rough hand-
ling by protruding their glands, and the oily golden-brown fluid that
visibly coated the everted structures had the unmistakable pene-
trating odor of benzoquinones, and the characteristic quinonoid

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0

0

0

0

a

b

c

Fig. 1. Compounds in defensive secretion of Adelium percatum and A.
pustolosum. a; 2-methyl-l,4-benzoquinone ; b; 2,3-dimethyl-l,4-benzoquinone ;
c; 1-pentadecene.

property of tanning human skin. Our fingertips were always darkly
mottled after handling a quantity of the beetles.

Secretion for analysis was obtained by seizing and squeezing in-
dividual beetles in forceps and wiping the fluid from the everted
glands with small pieces of filter paper. The chemical procedure and
the findings were the same for both species. Extraction of the papers
with methylene chloride, followed by gas chromatography of the
extract (1% OV-i on Gas Chrom. Q, 4' X 50° to 240°C),
revealed the presence of three major components. Analysis by gas
chromatography-mass spectrometry showed two of the components to
have mass spectra identical to the published spectra ( Budzikiewicz
et al., 1967) of 2-methyl- 1,4-benzoquinone and 2, 3-dimethyl- 1,4-
benzoquinone (Fig. 1 a, b). The third component had a mass spec-
trum matching that of i-pentadecene (Stenhagen et al., 1969)
(Fig. 1 c). Definitive confirmation of the position of the double
bond in this compound was obtained by ozonolysis. A small sample
of the third component was collected from the gas chromatograph
and dissolved in methylene chloride, and the cooled solution ( — 70°C)
was subjected to a slow bubbled stream of ozone in oxygen until the
blue color persisted. Addition of a small crystal of triphenylphos-
phine, followed by gas chromatography-mass spectrometry, showed
the higher molecular weight product to have a mass spectrum identi-
cal to that of tetradecanal (Stenhagen et al., 1969).

One interesting point in connection with these analyses was the
observation of a fourth component in the initial sample of secretion
obtained from both species. This component was guessed to be 1,2-
dichloropentadecane on the basis of its mass spectrum, and the iden-
tification was confirmed by direct comparison of the isolated material
with an authentic sample prepared by addition of chlorine to i-penta-
decene. However, since additional milkings taken from the same

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beetles four months later into carbon disulfide rather than methylene
chloride failed to show this component, it is apparent that the diha-
lide is an artifact derived from the olefin.

Structure and Defensive Operation of the Glands

Although the two beetles differ slightly in size and appearance —
A. percatum is less rotund and longer (i. 5-1.9 cm) than A. pustu-
losum ( 1 .3-1.5 cm) — their glandular apparatuses are essentially
identical. No dissections of fresh beetles were made, but judging
from what could be discerned from specimens somewhat inadequately
preserved in ethanol, the glands are anatomically similar in both
species and probably homologous to eversible glands such as have been
described for other tenebrionids (Lengerken, 1925; Roth, 1945;
Tschinkel, 1972). Such glands are basically cuticular sacs, ordinarily
withheld in inverted condition within the body cavity, and presumably
everted by blood pressure. What is unusual about the Adelium glands
is that they extrude to such great lengths. For example, gland length
in a specimen of A. percatum 1.8 cm long that was freeze-dried with
its glands everted, was recorded at 0.8 cm. Relative to body size, the
glands of these beetles must rate among the largest eversible glands
known for insects.

Examination of a glandular sac of A. percatum , treated with warm
aqueous potassium hydroxide to remove all soft cellular parts, re-
vealed a conventional membranous cuticular lining, with typical
cuticular organelles attached (Fig. 8). Such organelles are a diag-
nostic feature of many insect gland cells (references in Eisner, 1970),
and the ones of Adelium resemble closely those described from cell
type 1 in the defensive gland of the tenebrionid Eleodes longicollis
(Eisner et al.> 1964). The organelles were distributed along the full
length of the glandular sac, indicating that the secretory tissue is not
restricted to a limited portion of the gland.

Gentle handling of Adelium usually induced no gland eversion at
all. Only when the animals were more forcibly stimulated, as when
they were grasped by the body or an appendage and squeezed, did

Figure 2. Pasture land near Gudgenby, A. C. T., Australia, where both
species of Adelium were taken. Figs. 3-4. Adelium percatum responding
to pinching of individual hindlegs by everting the gland of the side stimu-
lated. Figs. 5-6. A. percatum responding to pinching of left midlegs. In
Fig. 5 the gland has been maximally everted and has come into contact
with the leg stimulated. In Fig. 6 the gland, on eversion, discharged part
of its content as an anteriorly-directed spray (reference bar = 5mm).

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eversion occur with some consistency. Reluctance to employ the
glands in these animals is undoubtedly related to their possession of
a particularly tough integument, which may in itself provide an ef-
fective first line of defense.

In order to observe more precisely the method of employ of the
glands, individual A . percatum were affixed to wire tethers with wax,
and stimulated by pinching their legs with forceps. The beetles were
Sometimes placed over filter paper impregnated with an acidified
aqueous solution of potassium iodide and starch, a mixture that dis-
colors darkly on exposure to benzoquinones (Roth, 1943) and pro-
vides a good visual means for detecting secretory emissions. The
results were identical with all nine specimens tested. Stimulation of
a single leg generally caused extrusion of only the gland of the corre-
sponding side (Figs. 3-6). Simultaneous stimulation of two legs
caused bilateral eversion. The extent and rapidity of extrusion varied,
and seemed to depend on the intensity of stimulation. A brief squeeze
of a hindleg or midleg, for example, usually induced only a partial
eversion of the ipsilateral gland. Protracted or more vigorous squeez-
ing elicited more complete and usually sustained eversion, and the
extruded gland commonly extended outward far enough to contact
the seized leg or forceps (Figs. 3-5). The target was usually visibly
wetted by the gland. The beetles can apparently exercise some con-
trol over the direction of extrusion. Thus, when distal parts of a
hindleg were stimulated, the gland usually extended outward at a
greater angle relative to body axis than when more proximal parts of
the leg were grasped. The gland is apparently aimed by flexion at its
base rather than along its length, as evidenced by the fact that its
shape is relatively fixed irrespective of its angle of protrusion.

When gland eversion occurred abruptly rather than gradually, as
was generally the case in response to vigorous pinching, the sudden
extension of the gland sometimes caused secretion to be ejected
forcibly as a spray. (Figs. 6, 7). Ejections ranged from 1 to 5 cm,
but in one instance droplets were shot to as far as 25 cm from the
beetle. Forelegs and such other parts on the front of a beetle as are
ordinarily inaccessible to the glands were inevitably doused by such
ejections. Only beetles with replete or nearly replete glands tended
to spray. However, animals with partly depleted glands could still
effectively administer secretion to their front end. They did this by
relaying secretion from one pair of legs to another, following initial
wetting of the hindlegs by the glands. Even the antennae and the
head itself were eventually wetted with secretion if they had been

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195

Fig. 7. Response of Adelium percatum to pinching of its right midleg.
The animal everted its right gland abruptly, causing secretion to be sprayed
forward. The droplets of secretion are shown as black spots on the indi-
cator paper that served as substrate (filter paper impregnated with acidified
aqueous solution of potassium iodide and starch). Far-ranging discharges
as shown here occurred only when beetles had fully replete glands. Fig. 8.
Cuticular organelles from secretory cells of gland of A. percatum , isolated
by treatment of gland with warm 10% aqueous potassium hydroxide
(Nomarski interference contrast; reference bar = 30/*). Fig. 9. Scanning
electronmicrograph of hairs on exposed surface of everted gland of A.
percatum (reference bar = 5/i) .

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seized in forceps. The initial contact of hindlegs and glands appeared
at times to be deliberate rather than fortuitous. Thus, when glands
were insufficiently everted to effect automatic contact with the hind-
legs, the legs sometimes reached back toward the glands and brushed
against them. Wiping of midlegs against hindlegs, and of forelegs
against midlegs, quickly followed. In cases where the beetles had
been placed on indicator paper, the legs left dark markings on the
substrate as evidence of their contamination. Some experiments
comparable to the preceding were also done with A. pustulosum, with
essentially similar results. However, forcible gland eversion with
spray ejection was never seen in this species.

Scanning electronmicroscopic examination of the surface of everted
glands of A. percatum revealed a covering of slender curved hairs
(Fig. 9). Whether these are spaced closely enough to act as a mat-
ting remains uncertain. But it is conceivable that they function in
this capacity, providing perhaps for the maintenance of uniform
secretory wetness over the surface of the glands when they are everted
and for spontaneous re-spreading of secretion over areas wiped clean
by defensive action of the glands. Dispensation of secretion onto
target surfaces might also be facilitated by the hairs.

Stridulatory Mechanism

When seized or pinched, A. pustulosum produced a distinctly
audible sound. The response took precedence over gland eversion,
and could persist for seconds if the disturbance was sustained. Two
serrate ridges on the last (visible) abdominal tergite (Figs. 10-13)
are responsible for engendering the sound. By rhythmic downward
deflection of the abdominal tip — a motion depicted in Figs. 10 and
11 — the beetles scrape the ridges back and forth across the sharp
overhanging elytral margins, thereby inducing a repetitive bi-syllabic
chirp. Intensified stimulation eventually caused gland extrusion,
which silenced the sound, since during gland eversion abdominal
deflection is apparently precluded. A. percatum lacks the ridges and
is soundless.

Stridulatory mechanisms are common in insects, and have been
repeatedly noted. In the leaf-cutting ant, Atta cehpalotes , which pro-
duces sound by scraping a file in much the same manner as does
Adelium, acoustical output and sound generation have been master-
fully analyzed (Markl, 1968). Since this analysis is essentially
applicable to Adelium, we will here describe only the properties of

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197

Figs. 10-11. Dorsolateral view of rear of body of Adelium pustulosum,
showing the abdominal tip in the undeflected (Fig. 10) and deflected
(Fig. 11) positions that mark the limits of the stridulatory motion. The
arrow points toward the left serrated ridge (reference bar = 1 mm).
Figs. 12-13. Scanning electronmicrographs of the left stridulatory ridge.
The elytral margin against which the ridge is scraped extends across the
top of Fig. 12 (reference bar Fig. 13 = .05 mm).

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198

the beetle’s sound, without detailed consideration of the physical basis
of sound production.

Only two live females and one male of A. pustulosum were avail-
able for sound studies. The animals were affixed to wire tethers
glued to their pronotum with wax, and induced to stridulate by
pinching their legs individually with forceps. Sound recordings were
made with the beetles held at 1 cm from the microphone. An oscillo-
gram of a typical chirp sequence is shown in Fig. 14A. The bi-
syllabic composition of each chirp, indicative of the bi-directional
movement of the ridges across the elytral margins, is clearly evident.
Measurements made from oscillograms, of chirp repetition rates, chirp
and syllable durations, and duration of inter-chirp and inter-syllable
intervals, are summarized in Table 1, together with measurements
of tooth counts and tooth spacings made from the serrate ridges.
These parameters are evidently remarkably constant for any one
beetle, although they differ somewhat between beetles. The Table
does not give an indication of the gradual decline in the chirp repe-
tition rate that characteristically took place with time following onset
of pinching of a given leg (e.g. in a female, the rate declined from
2.25 to 1.63 chirps/sec over a period of 4.5 seconds). The decline
occurred by a lengthening of the inter-chirp interval, rather than by
prolongation of the syllables or inter-syllabic interval of the chirp
itself. Accelerated chirping (or gland eversion) always occurred
when pinching was shifted to a new leg or increased in intensity.

As was to be expected by analogy with the stridulatory chirps of
Atta, and shown clearly by the oscillograms (Fig. 14B, C), the
syllables are essentially trains of sound pulses, generated by impact
of the elytral margins with the sequence of teeth on the ridges.
Figs. 14D and 14E show oscillograms of individual pulses from a
first and second syllable respectively.

Spectrograph analysis (Fig. 15) of the chirps revealed a frequency
composition ranging broadly from below 1 kHz to 60 kHz. The
frenquency distribution pattern was essentially similar in the first and
second syllables, and differed little in spectrographs made with broad-
band (Fig. 15 top) and narrow-band (Fig. 15 bottom) filters. Since
broad-band filtration essentially analyzed the pulses one at a time,
rather than in groups of three to five as in narrow-band filtration, it
follows that the pulses themselves contain essentially the full range
of the beetle’s sound frequencies.

Apparatus for measuring sound intensities became available only
after the beetles had died. Intensities therefore had to be measured

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199

Fig. 14. Sound oscillograms of Adelium pustulosum. A, chirp sequence
(2.8 sec. of $ no. 1) ; note bisyllabic composition of chirps (time interval
= 200 msec). B-C, Expansion of the first ( B ) and second ( C ) syllables of
chirp shown lettered in A ; the pulsed nature of the sound is clearly resolved
(time interval = 20 msec). D-E, Individual pulses of first (D) and second
( E ) syllables. (0.0625 msec/division).

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A B C

50 ~ ~ 100 i&0 200 250

TIME (msec)

Fig. 15. Sound spectrograms of one chirp of Adclium pustulosum (the
chirp is that lettered BC in Fig. 14 A, and shown expanded in Fig. 142?
and C) . In each spectrogram the first syllable is on left and second syllable
on right. Spectrograms were made from recorded sounds replayed at x/%
speed. Top spectrogram: wide band analyzing filter (effective bandwidth
2400 Hz, with effective time response of 0.375 msec); bottom spectrogram:
narrow band analyzing filter (effective bandwidth 360 Hz, with effective
time response of 2.75 msec). Beside the top spectrogram are shown energy
distribution profiles, corresponding to time transects lettered on the spec-
trogram itself.

indirectly, from the tape-recorded sounds of the beetles. This was
done by monitoring the playback of the recordings with a microphone
pickup and oscilloscope (the same microphone as used previously for
recording of the chirps), and adusting the intensity of the playback
until the induced oscilloscope signal matched that initially elicited by
live beetles held i cm from the microphone. The intensity of the
adjusted playback was then measured with the Sound Level Meter
positioned at the site of the microphone, and found to be 65 db
(relative to 0.0002 dynes/cm2).

Detailed analysis of the oscillograms showed that the beetles ap-
parently do not scrape both ridges with equal consistency during
stridulation. In fact, stridulation appears to involve predominant
scraping of only one ridge during downward deflection of the ab-

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201

dominal tip (first syllable), and of both ridges with somewhat
unequal force during the upward movement of the tip (second syl-
lable) . This conclusion rests on the observation that the syllables
contain, in addition to a consistent major pulse (which in total num-
ber per syllable corresponds to the number of teeth per length of file
estimated to be scraped during a syllable)1, a subsidiary pulse of
varying lesser amplitude, which appears sporadically in the first syl-
lable, and fairly consistently in the second syllable. These lesser
pulses, when they appear, result in a doubling of the pulse repetition
rate. Moreover they bear a shifting rather than fixed temporal rela-
tionship to the primary pulses, suggesting that they are not subsidiary
acoustical concomitants of single tooth scrapings, but a consequence
of scraping of a second set of teeth of somewhat unmatched spacing
relative to the first. The near absence of secondary pulses in the first
syllable suggests that the beetle might not press the abdominal tip as
vigorously against the elytral margins during downward abdominal
deflection as during the return motion.

Discussion

Defensive glands that produce quinones are widespread among
arthropods. They have been reported not only from other Tene-
brionidae (references in Jacobson, 1966; Roth and Eisner, 1962;
Weatherston, 1967; Weatherston and Percy, 1970), but also from
certain termites (Moore, 1968), earwigs (Eisner and Blumberg,
1959; Schildknecht and Weis, i960), cockroaches (Roth and Stay,
1958), grasshoppers (Eisner et al., 1971a), carabid beetles (Anes-
hansley et al., 1969; references in Moore and Wallbank, 1968;
Schildknecht et al., 1968), staphylinid beetles (Blum et al., 1971;
Brand et al., 1973), millipedes (references in Jacobson, 1966; Roth
and Eisner, 1962; Weatherston, 1967; Weatherston and Percy,
1970) and opilionids (Fieser and Ardao, 1956; Eisner et al., 1971b).
As is often the case with arthropod defenses, they exist as close
counterparts in plants: glandular hairs that secrete a benzoquinone
have been described for Primula obconica (Schildknecht et al., 1967).
Contrary to claim (Schildknecht et al., 1964), not all Tenebrionidae

JFrom photographs such as Figs. 10 and 11, which depict the motion of
the abdominal tip during chirping, it was estimated that this motion has a
maximum amplitude of ca. 2/3 the length of a serrated ridge. This cor-
responds to about 130 teeth, a figure in line with the counted maxima of
major pulses in the syllables.

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have defensive glands (e.g., Tentyriinae and Asidinae). But the
glands are frequently present, and when they are, they are either
exclusively abdominal as in Adelium , or supplemented by a second
pair of glands in the prothorax (Roth, 1945; Tschinkel, 1969).

Only relatively few tenebrionid secretions have been studied chem-
ically. Most produce benzoquinones, including />-benzoquinone, and
its alkylated derivatives 2 -methyl- 1,4-benzoquinone and 2-ethyl- 1,4-
benzoquinone (references in Jacobson, 1966; Roth and Eisner, 1962;
Weatherston, 1967; Weatherston and Percy, 1970). Naphthoqui-
nones have been reported in one species (Tschinkel, 1972) and in
another the prothoracic glands produce phenols instead of quinones
(Tschinkel, 1969). 2, 3-Dimethyl- 1,4-benzoquinone itself has not

been found in a tenebrionid secretion, but it is produced by the de-
fensive glands of certain opilionids (Fieser and Ardao, 1956; Eisner
et al.j 1971b). The presence of an alkene in Adelium is no novelty.
Hydrocarbons, both saturated and unsaturated, are known from
arthropod secretions, and although i-pentadecene as such has not been
reported, shorter chain homologs of the substance are present in
Tenebrionidae (Hurst et al., 1964).

We made no effort to evaluate the defensive effectiveness of the
secretion of Adelium. However, judging from the proven deterrency
of other quinonoid mixtures to predators (references in Eisner, 1970,
1972; Eisner and Meinwald 1966), there can be no doubt that the
glands of Adelium are protective in function. The presence of the
alkene may be of more than incidental significance. It has been
argued (Blum and Crain, 1961), and in one case proven (Eisner
et al 1961), that lipoidal additives promote spreading and penetra-
tion of defensive secretions on target.

Defensive glands of arthropods vary greatly in structure and oper-
ation, and several characteristic types have been recognized (Eisner,
1970). The glands of Adelium appear to combine features of two
of these types. Basically, by virtue of their extrusibility, they clasify
as conventional eversible glands, but since they occasionally eject their

Table 1. Adelium pustulosum: Temporal characteristics of the chirps

and morphological features of the stridulatory ridges. Temporal measure-
ments were made from oscillograms and are given as mean ± standard
error; sample size (N) is given in parenthesis. Morphological measure-
ments were made from microscopic preparations of the last abdominal
tergite of the beetles ; tooth spacing was calculated by dividing ridge length
by tooth number. (The tergite of $ no. 1 was damaged in preparation,
hence the omission of measurements from one ridge.)

Table

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203

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contents, they also qualify as spraying glands. We know of no other
tenebrionid glands that operate in this way. Eversible glands are
common, but they are generally shorter than those of Adelium and
do not eject spray on extrusion. Spraying does occur, but only from
non-eversible glands, as in Eleodes (Eisner, 1966). The peculiar
habit of Adelium of relaying secretion by way of its appendages is
not unique. Other tenebrionids are also said to use their legs for
administration of secretion (Tschinkel, 1972), and comparable be-
havior has been reported for opilionids (Eisner et al., 1971b) and
Hemiptera (Remold, 1962).

Many insects produce chirps or other “disturbance” calls when
they are molested. Cerambycid beetles, for example, very generally
show this property, as do other beetles, cockroaches, Hemiptera,
adult and immature Lepidoptera, and members of most other groups,
including non-insectan arthropods (references in Alexander, 1967;
Haskell, 1961; Roth and Hartman, 1967; Tuxen, 1967). The
sounds are generally believed to be defensive, but it is by no means
clear how precisely they act in this capacity. Convincing evidence
has been advanced to account for the function of the calls in arctiid
moths. These insects are chemically protected (Bisset et al., i960)
and hence presumably distasteful, and they emit sounds, rich in ultra-
sound, in response to the echolocating chirps of bats (Blest et al.,
1963). Bats turn away from the calls, presumably because they have
learned on the basis of previous experience with the moths that a
prospective meal that “protests” audibly is distasteful (Dunning and
Roeder, 1965; Roeder, 1967). The sounds, then, act as acoustical
aposematic signals, operative in the dark just as visual aposematic
adornments supposedly operate in the light. Defensive calls generally,
to the extent that they are produced by chemically or otherwise pro-
tected insects and induced by encounters with predators, may function
in this way. They might certainly do so in Adelium pustulosum, a
furtive animal most likely to meet its enemies in darkness. But the
calls need not only serve for acoustical reinforcement of a concomi-
tant defense. They could also be intrinsically deterrent to predators,
perhaps only to some, or they might function socially — as do “alarm”
sounds in termites and ants (Howse, 1964; Markl, 1967, 1969;
Markl and Fuchs, 1972) — to alert conspecifics to states of emer-
gency. But these possibilities, like so many other functional sug-
gestions that have been advanced for disturbance calls, remain to be
proven. One wonders what predators might be affected by the chirps
of Adelium, although criteria for compiling a list of enemies are lack-

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205

ing. It seems clear, however, that the chirps are of sufficiently broad
frequency composition to be at least potentially audible to virtually
any acoustically sensitive predator.

Summary

The black flightless tenebrionid beetles Adelium percatum and A.
pustulosum from Australia possess a pair of eversible abdominal
glands that secrete a mixture of 2-methyl- 1,4-benzoquinone, 2,3-
dimethyl-i,4-benzoquinone, and i-pentadecene. The tubular glands
are unusually large, and when everted may be nearly half as long as
the beetles themselves. In A. percatum , the replete glands may, on
extrusion, spray their contents forcibly, to a recorded maximum dis-
tance of 25 cm. Defensive administration of secretion is also effected
by the beetles’ legs. A. pustulosum , unlike A. percatum , produces an
audible “disturbance sound” when molested. The sound is engendered
by the scraping of two serrated ridges on the last abdominal tergum
across the elytral margins. The morphology of the stridulatory ap-
paratus, and the acoustical properties of the sound, which contains
frequencies ranging from below 1 kHz to 60 kHz, are presented. It
is argued that disturbance sounds, when produced by chemically or
otherwise protected animals, act as acoustical aposematic signals,
operative primarily in darkness.

Acknowledgements

We thank the staff of the Division of Entomology, C.S.I.R.O., for
much help and hospitality during the Eisners’ stay in Canberra. The
beetles, which were first shown to us in the field by Dart Linsley,
were identified by E. B. Britton. Electronmicrographs were made at
the laboratories of Barry Filshie, Colin Beaton, and Margaret
Kovaks, who were most generous with time and facilities. Robert
Capranica provided acoustical equipment and technical advice. The
study was supported in part by Grant AI-02908 from the National
Institutes of Health, and was carried out while T. Eisner was a
John Simon Guggenheim Memorial Foundation Fellow.

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teus (Lucas) (Arachnida: Pedipalpida). J. Insect Physiol. 6:
272-298.

Eisner, T. and J. Meinwald

1966. Defensive secretions of arthropods. Science 153 : 1341-1350.

Fieser, L. F. and M. I. Ardao

1956. Investigation of the chemical nature of gonyleptidine. J. Am.
Chem. Soc. 78: 774-781.

Haskell, P. T.

1961. Insect Sounds. Quadrangle Books, Chicago.

Howse, P. E.

1964. The significance of the sound produced by the termite Zooter-
mopsis angusticollis (Hagen). Animal Behav. 12: 284-300.

Hurst, J. J., J. Meinwald and T. Eisner

1964. Defense mechanisms of arthropods — XII. Glucose and hydro-
carbons in the quinone-containing secretion of Eleodes longi-
collis. Ann. Ent. Soc. Amer. 57: 44-46.

Jacobson, M. ■

1966. Chemical insect attractants and repellents. Ann. Rev. Entomol.
11: 403-422.

Lengerken, H. V.

1925. Vorstiilpbare Stinkapparate der Imago von T enebrio molitor L.
Biol. Zbl. 45 : 365-369.

Markl, H.

1967. Die Verstandigung der Stridulations-signale bei Blattschneider-
ameisen. I. Die biologische Bedeutung der Stridulation. Zeit-
schr. Vergl. Physiol. 57: 299-330.

1968. Die Verstandigung durch Stridulations-signale bei Blattschneider-
ameisen. II. Erzeugung und Eigenschaften der Signale. Zeit-
schr. Vergl. Physiol. 60: 103-150.

1969. Verstandigung durch Vibrations-signale bei Arthropoden. Natur-
wissenschaften 56: 499-505.

Markl, H. and S. Fuchs

1972. Kopfsignale mit Alarmfunktion bei Rossameisen ( Camponotus ,
Formicidae, Hymenoptera) . Zeitschr. Vergl. Physiol. 76: 204-225.

Moore, B. P.

1968. Studies on the chemical composition and function of the cephalic
gland secretion in Australian termites. J. Insect Physiol. 14:
33-39.

Moore, B. P. and B. E. Wallbank

1968. Chemical composition of the defensive secretion in carabid beetles
and its importance as a taxonomic character. Proc. Roy. Entomol.
Soc. (London) B37: 62-72.

Remold, H.

1962. Uber die biologische Bedeutung der Duftdriisen bei den Land-
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Roeder, K. D.

1967. Nerve Cells and Insect Behavior. Harvard Univ. Press, Cam-
bridge, Massachusetts.

Roth, L. M.

1943. Studies on the gaseous secretion of Tribolium Confusum Duval.
II. The odoriferous glands of Tribolium confusum. Ann. En-
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1945. The odoriferous glands in the Tenebrionidae. Ann. Entomol.
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Roth, L. M. and T. Eisner

1962. Chemical defenses of arthropods. Ann. Rev. Entomol. 7: 107-136.

Roth, L. M. and H. B. Hartman

1967. Sound production and its evolutionary significance in the Blat-
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Roth, L. M. and B. Stay

1958. The occurrence of />ara-quinones in some arthropods, with em-
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SCHILDKNECHT, H., I. BAYER AND H. ScHMIDT

1967. Struktur des Primelgiftstoffes. Z. Naturforsch. 22b: 36-41.

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SCHILDKNECHT, H. AND K. H. WEIS

1960. Zur Kenntniss des Pygidialdrfisensekretes vom gemeinem Ohr-
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Stenhagen, E., S. Abrahamsson and F. W. McLafferty

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Tschinkel, W. R.

1969. Phenols and quinones from the defensive secretions of the tene-
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1967. The chemistry of arthropod defensive substances. Quart. Rev.
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SYNCHRONOUS VISUALIZATION OF VIDEO-TAPED
SOUNDS AND MOTIONS OF INSECTS

By Paul A. Wussow, Robert B. Willey,
and June B. Steinberg1- 2
The University of Illinois at Chicago Circle,

P.O. Box 4348, Chicago, IL 60680;
and the Rocky Mountain Biological Laboratory,

Crested Butte, Colorado 81224.

High-speed cinematography with synchronized oscillography has
been used to analyze accurately the sound production by a rapidly
moving structure during communicative behavior of insects (Walker
& Dew, 1972; Willey, 1973, 1974). However, this method involves
expensive equipment and perhaps kilometers of film containing only
a few hundred feet of analyzable sequences. Nevertheless, the ac-
curacy and resolution of audio-visual synchrony by this method has
been exceeded only by the use of minute magnetometers using the
Hall-effect (Eisner, 1970; Eisner & Huber, 1973) and by an in-
genious method of flash-photography (Morris & Pipher, 1972). On
the other hand, studies needing an average synchrony accuracy of
only ± 5 msec (essentially equivalent to motion picture framing rates
of 60 pictures per sec) can utilize inexpensive or already available
video-systems used commonly as educational tools.

Video has several advantages over cinematographic methods:
1) sound and picture are routinely recorded synchronously, 2) video-
tape can be erased and reused, 3) some video-cameras have great
sensitivity and can be used even for nocturnal insects at very low
light levels, and 4) the system can be used to monitor behavior for
hours and then sequences can be selected, duplicated on another tape
and then the original tape can be used again. There are disadvan-
tages of video for high resolution work and these will be discussed
in the Analysis (q.v.). J

JMailing address for reprints: R. B. Willey, Assoc. Prof., Biological
Sciences, University of Illinois at Chicago Circle, P.O. Box 4348, Chicago,
IL 60680.

2P. A. Wussow is Data-Systems Analyst with NSF Grant GB-35594, and
Computer-Aided Instruction Specialist, Office of Instructional Resources
Development, University of Illinois; Mrs. J. B. Steinberg is doctoral candi-
date and Teaching Assistant in Biological Sciences.

Manuscript received by the editor March 1, 1974.

209

210

Psyche

[March

MONI TORS

Fig. 1. Block schematic diagram of the equipment connected for line-
by-line synchronization of a video field with a simultaneous full sweep of
the cathode ray gun of the oscilloscope. The CRO is laid on its side so
that the horizontal sweep passes from top to bottom of the superimposed
field in VTR2 (see text, sect. 8, Analysis). For ordinary horizontal sweep,
the CRO is in normal position and the external trigger on the vertical sync
is disconnected.

It seems strange that, with the exception of publications from our
laboratory (Steinberg & Conant, 1974; Steinberg & Willey, 1974),
we have found only a few studies of insect behavior which have
involved the analysis of video-taped sounds and motion (Loher &
Chandrashekaran, 1970, 1972; Hoeg-Guldberg, 1972). Moreover,
even these studies have neglected to capitalize on the audio-recording
and electronic potential of the video-tape system. We have devised a
simple method to superimpose the oscillographic trace of the sound
already recorded on any pre-existing video-tape on a duplicate record-
ing. Extensions of this method to synchronous oscillography trans-
duced from any mode will make video monitoring of insect behavior
amenable to more versatile and precise analysis.

Method

A sequence was video-recorded with the following sound-related

1974]

W ussow et al. — Sounds and Motions of Insects

211

parameters noted : i ) distance from the microphone to the subect,
and 2) the average temperature of air between subject and micro-
phone. Then we played the recorded sequence on a compatible
video-tape reproducer (VTR.!) which was linked by a multichannel
synchronizing circuit (SWITCHER) to another video-recorder
(VTR2) which preferably used at least one-inch tape. Figure l
diagrams the set-up schematically. The audio circuit of the VTRa
was linked also to the input of an oscilloscope (CRO), the screen
(CRT) of which has an image persistence on the order of micro-
seconds (Pi i phosphor coating). A video-camera (CAM) focused
on the CRT image which was positioned so that it would not mask
the image of the subject in the superposition. If the trace was to be
positioned along the bottom of the frame, the horizontal sweep of
the CRO was set at io msec/cm so that a single sweep of 12.5 cm
will encompass at least eight video fields (q. v., sect. 7 & 8, Analysis
and Fig. 1 for further explanation and an alternative method). We
also masked the negative potentials with opaque tape. The video-
camera is connected with the VTR2 by the vertical interval switcher
and the CRT screen is shielded by an improvised hood from the
room lights. The results of superposition are simultaneously video-
monitored.3

The resulting video-tape, minus all the inactive time, can be played
on a VTR which has stop-action and single-field forwarding capabil-
ity (or the tape can be moved forward by hand). The audio signal
can still be heard as well as seen as an oscillo-trace when the VTR
is running at normal speed. At low speed the audio circuit auto-
matically turns off and, although an adustment can be made in the
machine’s circuitry to enable the sound to be heard, with the oscillo-
graphic trace this is neither necessary nor desirable. With the VTR
stopped on a single field, we marked with a wax crayon the positions
of the moving parts on the projection screen of the monitor and made
measurements of the angle changes with a protractor ( a better method
will be outlined later in section 9, Analysis).

Equipment used: “Quiet Room” (Suttle, Inc., Chicago), Audiometric
Room, double-walled (Industrial Acoustics, New York), Sony EV-310 25 mm
videocorder, Scotch Brand MT 20568 video-tape, General Electric model
4TE33D1 TV camera with a Soligor 10 cm, lens, Sony AVC 3200 TV
camera with Angenieux 7.5 cm lens with a +1 diopter auxiliary lens,
AKG shotgun microphone model D900, vertical interval switcher-sync
generator with sync lock capability (Shintron 360), Tektronix 502A os-
cilloscope with Pll phosphor, Panasonic video-monitors (3-inch), one 50W
electric spot bulb.

212

Psyche

[March

Analysis

Analysis of audio-visual synchrony must consider the following
factors :

1 ) A video field is scanned from top of the screen to the bottom
once in 16.67 msec at a maximum resolution of 262.5 lines per field;
the horizontal sweep time for a line is 0.0635 msec. The vertical
blanking interval (retrace time for the electron-gun to move from
bottom of the screen to the top) requires up to 21 lines, and sub-
tracted from the maximum resolution leaves a net resolution of 241.5
lines per field with a net duration of 15.5 msec, and a vertical blank-
ing interval of 1 to 1.2 msec.

2) A video frame consists of two interdigitating video fields with
a total net duration of 31 msec and a total net resolution of 483
lines.

3) Transduction of light and electronic impulses can be regarded
as instantaneous, however the speed of sound in air is a variable.
Near sea level, at 37 C and no greater than 10 cm from the micro-
phone, one can calculate that the maximum time interval between
sound production and electronic transduction is no greater than 0.4
msec and can be neglected as an important factor.

4) A crucial discrepancy in visual-acoustic synchrony and visual
analysis of movements is the time taken by the electron-gun of the
video camera to trace any action within a field. This factor causes
three time-related distortions in the image which must be considered
in the analysis, namely (a) vertical loss of resolution, (b) excessive
curvature of a rotating image, and (c) inability to record events
which occur after a scanning trace has passed until the next field scan.

5) Vertical loss of resolution: the image formed during a single
field has a maximum vertical resolution of only 241.5 lines per field
whereas a 16 mm optical-film frame has 50-60 lines of resolution per
mm (at the center) or about 700 to 800 lines of resolution overall
(allowing for loss of about 20 lines of resolution at the front and rear
edge of the frame). On the other hand, the horizontal resolution of
a good video system probably approaches that of a comparable optical-
film system. However evaluation of information content of a video
field and an optical film frame encompassing 15 msec is obscured by
the next considerations.

6) Excessive curvature of rotating image: the time required for
scanning a single field naturally produces a distorted image in any
movement having lateral components. Plate ia-e illustrates this

1974]

W ussow et al. — Sounds and Motions of Insects

213

PI. 1. A portion of a stridulation (chirp) by Chortophaga viridifasciata,
showing a pulse of sound in five photographs of consecutive fields (a-e).
The male which is chirping is outlined in (a), (b) and (c) show the right
femur blurred as it makes the downward stroke (left femur is missing).
The two vertical black lines drawn on the trace in (d) bracket the portion
of the oscillotrace which was scanned at the time the leg was moving in
this field. To the right of the lines is the base of a trace which really was
much higher, but its peaks will not be scanned until the next field (e).
The image persistence is due to retention by the video-camera pick-up tube
and is about equal to one field (= 16 msec). The image in (f) is the
result of triggering and synchronizing the CRT sweep with the “front
porch” of the vertical field (set up as in Fig. 1). The field is identical to
(e), but only the most intense peaks were registered. (See text, sect. 8)

214

Psyche

[March

problem with the oscillographic trace and a barely detectable clock-
wise curvature of the femur tip relative to the base in ib and ic.
This is caused by the fact that a video field is a metachronic series of
241.5 lines each representing a “shutter speed” of 1/15,750 sec. The
scanning time of the femur image in PI. ia is about 45 lines of 2.86
msec. In other words, the tip of the femur, which is moving in an
arc to the left, is imaged nearly 3 msec earlier than that of the femur
base. Further magnification on the screen would result in greater
temporal displacements and greater artifact in the shape of the organ.
Any attempt to measure angles of rotation or points of displacement
must take into account the video scanning time of the specific image
and its vector relative to the vertical trace.

7) Inability to record events after passage of the scanning trace:
it is usually desirable to superpose the oscillotrace along the bottom
of the video field (PI. ia-e) where the action is less likely to be
obscured. However, the lower portion of the field will be scanned
milliseconds after the action in the center. Nevertheless, visual-
acoustic synchrony can be computed as shown in PI. id. The selec-
tion of the horizontal sweep setting of the CRO is determined by the
duration of the longest acoustical signal and should be about 40 or
50 msec longer, so that the oscillotrace of the entire action can be
visualized in one sweep, and traces in successive frames can be cross-
referenced. The additional time allows some time before and after
the signal on the screen so that the beginning or the end of the signal
and its causative motion aren’t lost in the CRT retrace time, the
vertical blanking interval, nor at the sometimes distorted edges of the
monitor screen. Sometimes traces are badly reproduced or oriented,
but constant inspection of the VTR2 monitor will allow such mistakes
to be corrected immediately by duplicating the sequence again.

8) An alternative method to achieve visualization of motion-sound
synchrony is shown in Fig. 1 and PI. if. Here the CRO is laid on
its side so the oscillotrace will sweep from top to bottom of the
camera pickup tube. The external trigger of the CRT electron-gun
is triggered on the electronic transition from the vertical sync to the
start of the first horizontal scanning line of the video field. The hori-
zonal sweep of the CRT is set to equal the time constant of the video
field (15 msec per sweep or 1 cm/msec), thus only the sound re-
corded during the video scan will be oscillographed. The oscillotrace
peaks appearing at the same horizontal plane as the video-recorded
movement will be relatively synchronous with that movement, with

1974]

W ussow et at. — Sounds and Motions of Insects

215

allowance for the time lag induced by the speed of the original sound
in air (PL if, arrows).

9) Close examination of CRTs can be a health hazard and the
thickness of the protective cover can induce parallax error. There-
fore, if exact angle and displacement measurements are desired the
fields should be photographed with careful focussing on the video
lines. Such photographs will form a permanent working record of
key fields of the rather fragile video-tape which can be damaged or
destroyed by the constant friction of scanning in the stopped position.

Discussion and Conclusions

This method allows relatively synchronous visualization of motions
and the sounds produced by the movements if the actions are rela-
tively slow. For example, it is possible to determine whether the stri-
dulation by a single up-down motion of a grasshopper’s hind femur
rubbing against the fore-wing is produced during the up-stroke or
down-stroke, provided the entire motion has a duration of >60 msec.
In the case of oedipodine grasshoppers, the chirps produced by males
during courtship usually are produced by motions which last at least
60 msec and often 100 msec (Willey, 1974). Since many such sig-
nals are complicated by being segmented into pulses within a single
up-down motion, normal-speed cinematographic and audiospectro-
graphic investigations of Oedipodinae (Otte, 1970; Willey & Willey,

1969) have concluded that the first pulse must be produced on the
up-stroke. However, high-speed motion analysis (Willey, 1974)
shows that Arphia sulphur ea produces the definitive chirp on the
downstroke only. A very weak signal was produced on the up-stroke
but was of such low amplitude that it had never been audiospectro-
graphed. Video-tapes of Chimarocephala (Loher & Chandrashekaran,

1970) and Chortophaga (Steinberg & Willey, 1974) also show that
stridulations are produced only on the downstroke of the femur in
these species. However, the mechanism for production of two-pulsed
chirps must be analyzed by the higher speed methods (Willey, 1974).

Further refinements of this system can involve direct synchrony of
oscillotrace with the initial video-tape recording. Also the use of a
light-emitting diode (LED) and a small high speed digital clock in
the background, synchronized with each video field, would simplify
identification of particular fields. However, the method we have
reported can be used for improving the analysis of already existing
tapes.

2l6

Psyche

[March

Summary

Oscillographic traces of sounds recorded on videotapes can be
superimposed easily and synchronously on a duplicate of those tapes.
The audio recording remains intact in the duplicate and the oscillo-
trace allows rather precise analysis of slower movements and the
sounds they produce. This system was tested on pre-existing tapes
of courtship in the grasshopper Chortophaga viridifasciata (De
Geer). The method promises to be useful in analysis of tapes used
for monitoring behavior and also as a teaching device.

Acknowledgements

We are grateful to Prof. Gershon Berkson for sponsoring our use
of the television facilities of the Small Groups Laboratory of the
Department of Psychology at the University of Illinois at Chicago
Circle. David Mucha helped prepare the photographic figures. Dr.
Ruth Willey gave helpful advice during preparation of the manu-
script. Technique development and publication were aided by NSF
Grant GB-35594 to RBW through the Rocky Mountain Biological
Laboratory.

Literature Cited

Elsner, N.

1970. Die Registrierung der Stridulationsbewegungen bei der Feld-
heuschrecke Chorthippus mollis mit Hilfe von Hallgeneratoren.
Zeitschr. vergl. Physiol., 68: 417-428
Elsner, N. and F. Huber

1973. Neurale Gundlagen artspezifischer Kommunication bei Orthop-
teren. Fortschr. Zool., 22: 1-48.

Hoegh-Guldberg, O.

1972. Pupal sound production of some Lycaenidae. Jour. Res. Lepid.,
10: 127-147.

Loher, W. and M. K. Chandrashekaran

1970. Acoustical and sexual behaviour in the grasshopper Chimaro-
cephala pacifica pacifica (Oedipodinae) . Entom. Exp. & Appl.,
13: 71-84.

1972. Communicative behavior of the grasshopper Syrhula fuscovittata
(Thomas) (Gomphocerinae) with particular consideration of
the male courtship. Zeitschr. Tierpsychol., 31: 78-97.

Morris, G. K. and R. E. Pipher

1972. The relation of song structure to tegminal movement in Metriop-
tera sphagnorum (Orth., Tettigoniidae) . Can. Entomol., 104:
977-985.

1974]

W ussow et al. — Sounds and Motions of Insects

217

Otte, D.

1970. A comparative study of communication in grasshoppers. Misc.
Publ., Mus. Zool., Univ. Michigan, 149: 1-168.

Steinberg, J. B. and R. Conant

1974. An informational analysis of the intermale behaviour of the
grasshopper, Chortophaga viridifasciata. Anim. Behav., 22: in
Press.

Steinberg, J. B. and R. B. Willey

1974. Visual and acoustical social displays by the grasshopper Chorto-
phaga ‘viridifasciata (Acrididae: Oedipodinae). In preparation.

Walker, T. J. and D. Dew

1972. Wing movements of calling katydids: fiddling finesse. Science,
178: 174-178 and cover picture.

Willey, R. B.

1974. Slowed motion studies of sound production in the grasshopper
genus Arphia (Acrididae: Oedipodinae). In preparation.

Willey, R. B. and R. L. Willey

1969. Visual and acoustical social displays by the grasshopper Arphia
conspersa (Orthoptera: Acrididae). Psyche, Cambridge, 76:
280-305.

Willey, R. B. and P. A. Wussow

1973. The flight dances of grasshoppers (Acrididae: Oedipodinae)
(motion picture). Amer. Zool., 13: Abstr. no. 2.

CAMBRIDGE ENTOMOLOGICAL CLUB

A regular meeting of the Club is held on the second Tuesday of
each month October through May at 7:30 p.m. in Room B-455,
Biological Laboratories, Divinity Ave., Cambridge. Entomologists
visiting the vicinity are cordially invited to attend.

BACK VOLUMES OF PSYCHE

Requests for information about back volumes of Psyche should
be sent directly to the editor.

F. M. Carpenter

Editorial Office, Psyche
16 Divinity Avenue

Cambridge, Mass. 02138

FOR SALE

Classification of Insects, by C. T. Brues, A. L. Melander and
F. M. Carpenter, Published in March, 1954, as volume 108 of the
Bulletin of the Museum of Comparative Zoology, with 917 pages
and 1219 figures. It consists of keys to the living and extinct families
of insects, and to the living families of other terrestrial arthropods;
and includes 270 pages of bibliographic references and an index of
76 pages. Price §14.00 (cloth bound and postpaid). Send orders
to Museum of Comparative Zoology, Harvard College, Cambridge,
Mass. 02138.

No, 2

"psyche

A JOURNAL OF ENTOMOLOGY

Vol. 81 June, 1974

CONTENTS

Social Carrying Behavior and Division of Labor During Nest Moving
of Ants. M. Moglich and B, Holldobler 219

Temporal Activity Patterns in Two Competing Ant Species (Hymenop-
tera: Formicidae). J. H. Hunt 237

Relationship of Larval Food-plants and Voltinism Patterns in Temper-
ate Butterflies. F. Slansky 243

Variations in Cleaning Between the Sexes of Sinella coeca (Collembola:
Entomobryidae) . E. S. Waldorf 254

Centennial of Entomology at Cornell University 257

Notes on Necrophoric Behavior in the Archaic Ant Myrmecia vindex
(Formicidae: Myrmeciinae) . C. P. Haskins and E. F. Haskins 258

The First Recent Species of Protomutilla (Hymenoptera : Mutillidae:
Myrmosinae) D. J. Brothers 268

Life History of Abedus herberti in Central Arizona (Hemiptera:
Belostomatidae) . R. L. Smith 272

Prey Capture by Drymusa dinora (Araneae, Scytodidae). C. E. Valerio 284

A New Cockroach Genus {Gurney a) Previously Confused with Pina-
conata (Blaberidae: Epilamprinae) . L. M. Roth 288

Merope tuber (Mecoptera) : A Wing-body Interlocking Mechanism.

T. F. Iilavac 303

The Unusual Web of Spilasma tubulofaciens, with Taxonomic Notes
of the Species. D. Quintero, Jr. 307

Wing-folding in the Paleozoic Insect Order Diaphanopterodea (Paleop-
tera), with a Description of New Representatives of the Family
Elmoidae. J. Kukalovd-Peck 315

Observations on the Nesting Behavior of Rubrica surinamensis (De-
Geer) (Hymenoptera, Sphecidae).

H. E. Evans, R. W. Matthews, and E. McC. Callan 334

A New Species of Opithes from Mexico, with a Key to the Species.

I. Moore

353

CAMBRIDGE ENTOMOLOGICAL CLUB

Officers for 1973-1974

President ,
Vice-President .
Secretary .

Treasurer .

Executive Committee

B. K. Holldobler
W. D. Winter, Jr,
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F. M. Carpenter
R. E. SlLBERGLIED

L. P. Lounibos

EDITORIAL BOARD OF PSYCHE

F. M. Carpenter (Editor), Fisher Professor of Natural History,
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J. M. Burns, Associate Professor of Biology, Harvard University
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and Associate in Entomology, Museum of Comparative Zoology
P. J. Darlington, Jr., Professor of Zoology, Emeritus, Harvard
University

B. K. Holldobler, Professor of Biology, Harvard University
H. W. Levi, Alexander Agassiz Professor of Zoology, Harvard
University

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E. O. Wilson, Professor of Zoology, Harvard University

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The Lexington Press. Inc.. Lexington. Massachusetts

PSYCHE

Vol. 8 1

June, 1974

No. 2

SOCIAL CARRYING BEHAVIOR
AND DIVISION OF LABOR
DURING NEST MOVING IN ANTS

By Michael Moglich* and Bert Holldobler
Biological Department, MCZ-Laboratories,

Harvard University, Cambridge, Mass., 02138

Social carrying behavior is one of the most remarkable social ac-
tivities in ant societies. Not only eggs, larvae and pupae, but also
adult workers, queens and males are frequently carried by worker
ants to various target areas. Although carrying behavior has been
observed in many ant species (see review in E. O. Wilson 1971),
only a few analytical investigations have dealt with the biological
significance of social carrying behavior in ants. Kneitz (1964)
reports that in Formica polyctena special “storage workers” are
passively moved between the summer nest and winter nest. Arnoldi
(1932) observed that during the slave raids Rossomyrmex proformi-
carum uses the carrying technique to recruit sister workers to the
nest of the slave ants. In Camponotus herculeanus social carrying
behavior serves as a “social timer” during the nuptial flight activi-
ties: males that tend to take off too early or too late during the daily
flight periods are carried back into the nest by their worker nestmates
(Holldobler and Maschwitz 1964).

Most frequently, however, carrying behavior is employed during
emigration from one nest site to another. If a nest becomes too small
and cannot be extended, or if the microclimatic conditions change,
the colony searches for a better site. Although the communication
signals used by different ant species to organize nest movings vary
considerably, adult transport seems to be the basic recruitment tech-
nique of most.

^Present address: Fachbereich Biologie, Frankfurt, W. Germany.

Manuscript received by the editor June 28, 1974.

219

220

Psyche

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Fig. 1. Primitive adult transport in A) Odontomachus (worker carries
worker) B) Pogonomyrmex badius (worker carries “soldier” caste).

1974]

Moglich &f Holldobler — Nest Moving of Ants

221

The Specificity and Stereotypy of Carrying Behavior

Escherich (1917) and Arnoldi (1932) pointed out that the be-
havioral patterns of adult transport in ants are very stereotyped and
are often specific for certain taxonomical groups. These assumptions
are confirmed by the following brief survey of adult transport be-
havior in ants.

Myrmeciinae :

In one of the most primitive genera of the myrmecoid complex, the
Australian Myrmecia Haskins and Haskins 1950 observed adult
transport. However, the behavioral patterns are apparently not
stereotyped. One worker grasps another at the mandibles or any
other part of the body and drags it over the ground.

Dorylinae :

According to Rettenmeyer (1963) workers of the New World army
ants (tribe Ecitonini) carry other adults like larvae and pupae slung
beneath the body and between the legs of the transporting worker.

Ponerinae:

An unstereotyped carrying behavior, similar to that described for
Myrmecia , was observed by the present authors in Bothroponera
tesserinoda. It occurs only rarely; nestmates are primarily recruited
by the tandem running technique (see Maschwitz, Holldobler and
Moglich, in press). Somewhat more advanced is the adult transport
behavior of Odontomachus. The transporter ant simply grasps a
nestmate on a leg, the petiole or some other part of the body, lifts it
up and carries it away. Although there is no general, stereotyped
response, the transported individual usually folds its appendages
tightly to the body (Fig. ia). Much more elaborate and stereo-
typed behavior is displayed by Rhyditoponera metallica. When the
transporting worker approaches a nestmate head on, it repeatedly
grasps the ant at the head and jerks it slightly forward. The move-
ment lasts only 1-2 seconds. The nestmate responds by turning its
bodyaxis slightly sideways, whereupon the transporting ant seizes the
nestmate with a firm grip. After turning through an angle of ap-
proximately 1800, the transportee is lifted and curled over the head
of the carrying ant. During transportation the gaster is bent in-
wards and the appendages folded tightly to the body.

Myrmicinae :

Although Escherich reports that in myrmicine ants the transported
ant is grasped at the petiole and carried with the head and legs

222

Psyche

[June

Fig. 2. Adult transport technique in myrmicine ants: A) Leptothorax
nylanderi (worker carries worker). B) Pogonomyrmex maricopa (worker
carries worker). C) Novomessor cocker elli (worker carries worker).

1974] Moglich & Holldobler — Nest Moving of Ants

223

pointed forward, in fact this seems to be the exception and has been
specifically observed only in Crematogaster (see Wilson 1971).
Generally, however, the stereotyped adult transport behavior of
many myrmicine species, which has been repeatedly described since
Escherich’s time, resembles that of Rhyditoponera. The transportee
is either seized at the mandibles or at the “neck” or “cheeks” and
curled over the head of the transporting worker (Fig. 2). But there
are a few other exceptions: we found, for example, that the harvest-
ing ants Pogonomyrmex badius , P. rugosus and P. barbatus employ
a rather primitive carrying technique. As in Odontomachus the
transportee is grasped at any part of the body, lifted up and carried
away. During transportation the carried individual folds its ap-
pendages to the body (Fig. ib). This primitive transporting behavior
is especially remarkable, because in some other Pogonomyrmex spe-
cies, for example P. maricopa, we found the typical highly stereo-
typed adult transport behavior described above for other myrmicine
ants (Fig. 2) .

Formicinae:

The most uniform and stereotyped social carrying behavior can be
observed in the subfamily Formicinae. In several species we have
analyzed the behavioral patterns that initiate and guide carrying be-
havior and found them to be virtually invariant. For the following,
more detailed description of this behavior we have chosen Campono-
tus sericeus, a common species on Ceylon. The description is based
on single frame analyses of slow motion pictures (Fig. 3).

When a recruiting ant faces a nestmate head on, it conducts a
jerking behavior for 2-3 seconds, grasping the nestmate at the mandi-
bles and subsequently pulling it forward. Usually the recruiting ant
responds by turning around for 180°. The nestmate is thereby slightly
lifted, and this elevation evidently causes it to fold its legs tightly to
the body and to roll the gaster inward. In this “pupal” posture it is
then carried to the target area.

Identical stereotyped adult transport behavior has been observed
in many formicine ants and has been described in particular detail in
Formica rufa (Zahn 1957), F. polyctena (Kneitz 1964, Moglich
I97I)> Cataglyphis (Wehner and Lutz 1969), Camponotus socius
(Holldobler 1971) and several other species.

Sex specific carrying postures

Reproductive females are sometimes carried like workers (Fig. 4).
But in cases where the size difference is too large, workers merely

224

Psyche

[June

Figure 3

1974]

Moglich & Holldobler \ — Nest Moving of Ants

225

grasp the females at the mandibles and pull them to the target area
(Fig. 5). The first behavioral steps that lead to this pulling be-
havior are usually similar to those which initiate carrying behavior
in workers.

The methods by which males are transported are notably different,
however. These individuals are sometimes picked up at any part of
the body and dragged or carried to the target area. In a few species
workers apply specific stereotyped carrying methods for males. In
several Camponotus species we observed that most of the males are
grasped at the “neck” and lifted into an oblique position in which
they are carried away (Fig. 6a). During transportation the males
remain motionless with the antennae and legs folded to the body. As
Fig. 6 b and c shows, there are occasionally exceptions of this carry-
ing technique.

In some other species, as in N ovomessor cocker elli and Aphaeno-
gaster floridanus , workers grasp the males between the thorax and
gaster and carry them beneath their body between their legs (Fig. 7).

Division of Labor During Nest Movings

Social carrying behavior in ants can serve many purposes; it is,
however, most frequently employed during nest emigrations. To-
gether with the tandem running technique it can be considered to be
a rather primitive recruitment method. In both cases, each recruiting
ant can only recruit one nestmate at one time. This leads to the
question: Do all workers of a colony have the same carrier or tan-
dem-leader potential, or is there instead a group of specialists, who
organize nest movings. To investigate this matter we chose two
formicine species, one which moves almost exclusively by using the
adult transport method ( Formica sanguinea) and another which
primarily employs the tandem running technique ( Camponotus seri-
ceus) .

Nest emigrations can be induced in the laboratory by keeping the

Fig. 3. The behavioral sequences that initiate carrying behavior. 1. The
recruiter ant (black) approaches a nestmate (white) and conducts the
jerking response for 2-3 seconds. 2. The recruiter grasps the nestmate at
the mandibles and pulls it back for about 2-20 cm. 3. When the recruiter
turns, it holds the nestmate with a firm grip. The nestmate is thereby
slightly lifted. 4. The nestmate folds its legs and antennae tightly to the
body and rolls its gaster inward. 5. In this posture it is carried to the
target area. The arrows indicate the direction of the movements. These
sequences were based on a film analysis. (After Holldobler, Moglich,
Maschwitz 1974; illustrations by Turid Holldobler).

226

Psyche

[June

Fig. 4. Adult transport technique in formicine ants: A) Formica rufa
worker carries a sister worker. B) Formica rufa worker carries a queen.

1974]

M oglich <o Holldobler — Nest Moving of Ants

227

Fig. 5. If the difference of size between workers and queens becomes
too large, queens are either pulled or led along a short odor trail to the
target area. This is the case in young colonies of Camponotus pennsyl-
vanicus.

old nest under less than optimal conditions while at the same time
providing a new nest nearby with optimal conditions. Soon a scout
discovers the new nest, and after a short inspection returns to the
old nest. Usually it runs 2-3 times back and forth between the old
and new nest, before beginning to recruit nestmates to the new site.
Our intention was to identify and follow all individual workers of
an entire colony throughout the experimental procedures. In order
to do this we marked the ants individually by applying spots of one
or the other of four colors at one or more of four pre-selected posi-
tions on the body. In this way we were able to recognize up to 256
workers individually. With each of our test colonies we conducted
20 nest-moving experiments over a period of about 2 months, before
the entire colony was killed and each worker subsequently dissected.

During the nest emigrations we recorded the following parameters
in particular : 1 ) How often a worker was carried or led by tandem
running, respectively, during all nest movings; 2) How often a
worker functioned as a carrier or leader, respectively, during one
nest-moving episode and throughout the entire experimental period
of 20 movings. 3) We also distinguished between the recruitment
intensity and the continuity of the recruitment activity of workers.

228

Psyche

[June

Fig. 6. Male transport in Camponotus sericeus: A) Commonly used

technique. B) and C) Exceptionally used techniques.

1974]

Moglich & Holldobler — Nest Moving of Ants

229

Fig. 7. Male transport in A) A phaenogaster floridanus and B) Novo-
messor cocker clli.

230

Psyche

[June

The intensity is defined as the total number of recruitment perform-
ances during the entire experimental period, independent how these
performances were distributed over the nest movings. The continuity,
on the other hand, is defined as the percentage of nest movings during
which an individual worker acted at least once as a recruiting ant.
4) In order to rank degrees of specialization of different workers we
combined all these parameters in an index which gives us a measure
of the relative recruitment activity (RRA) of individual ants:

R

RRA = -=r2- X C. Ra is defined as the total number of active re-

cruitment acts performed by the individual ant; Rp is the total
number of recruitment episodes during which the ant acted as trans-
portee or tandem follower, respectively, and C is the continuity, as
defined above.

The quantitative records gathered for each individual worker dur-
ing 20 nest movings, are summarized in Fig. 8 and 9. From these
sociograms it can be seen that, although both species (F. sanguinea
and C. sericeus) use different recruitment techniques, the social or-
ganization of their nest movings is basically similar. The nest emi-
grations are organized by a pronounced system of division of labor.
There is a relatively small group of workers ( in our colonies 11%
in F. sanguinea and 6% in C. sericeus ) which were active in 80%
of all nest moving experiments, and some individuals of this group
recruited up to 31 nestmates during a single nest emigration.

This specialized group shows a significantly higher recruitment
activity in comparison with all the other workers (X2-test, p If 0.0 1).
The sociograms also reveal that even the mover specialists are
sometimes transported themselves. This is especially the case at the
beginning of a nest moving, when a single scout ant first recruits
workers of the specialists’ group. Those recruits subsequently become
recruiters themselves. For F. sanguinea we recorded at least 7 cases
where an ant has been carried to a target area and subsequently
returned to the old nest to become a recruiter itself. The same pro-
cedure was demonstrated 20 times in F. polyctena. From those data
we conclude that the carrying act can function as a complete recruit-
ment procedure, by which the recruited ant is not only brought to the
target area but also is stimulated to become an active recruiter itself.
The same, of course, is true for the tandem running technique as pre-
viously documented (Hblldobler, Maschwitz, Moglich 1974).

The degree of specialization of division of labor in different colo-
nies can be demonstrated by ranking the workers according to their

1974]

Moglich Holldobler — Nest Moving of Ants

23i

RRA-indices and by plotting the rank positions against the index
values (Fig. 10). In the test colonies of both species the curve
declines rather steeply indicating that the degree of specialization is
high. We were able to prove the crucial role of this small group of
specialists simply by removing them from the colony and observing
the performance of the workers left behind. Although some regula-
tive replacements by other workers appeared to be possible, the time
needed for moving was considerably increased. When we also re-
moved this second mover group, the colony was almost unable to
achieve an organized nest emigration, even when their nest conditions
became very unfavorable. Workers left the nest and dispersed, but
no organized nest emigration took place. From this we conclude that
a specialized group of nest movers exists and is important in leading a
colony in a relatively short period of time from unfavorable to favor-
able nest sites.

According to the investigations by Otto (1958) and Kneitz
(1964) we should expect that the moving specialists belong to the
so called outside workers (Aussendiensttiere) with reduced ovaries.
The dissection of all workers of the colonies showed that indeed all
movers had reduced ovaries, whereas the ovaries of many of the ants
being carried or led were relatively well developed. However, from
our invstigations it became also clear that only a small portion of the
“Aussendiensttiere” function as mover specialists. If they are re-
moved experimentally, they can be replaced by other workers with
reduced ovaries, whereas nestworkers (Innendiensttiere) with well
developed ovaries cannot substitute for lost mover specialists.

Summary

Adult transport behavior is a common social activity in many ant
species. It is employed most frequently as a recruitment technique
during emigration from one nest site to another. As verified in this
report, the behavior patterns vary among species and are correlated
with phylogenetic groupings.

The social organization of emigration, in particular the division
of labor among workers during the process, was analyzed in two
species, one of which recruits nestmates by carrying ( Formica san-
guinea) and the other by tandem running ( Camponotus sericeus) .
In both cases it was found that only a special group of workers
organizes nest emigrations. All these moving specialists have com-
pletely reduced ovaries whereas many of the carried or led ants have
well developed ovaries.

232

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total number of recruitment acts during which the ant led a nestmate by tandem running to the new nest. The
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dividual. In Rp the height of the columns expresses the total number of recruitment procedures during which the
ant was led, while the number of dashes indicates in how many nest emigrations this was the case.

234

Psyche

[June

Fig. 10. Individual ants were ranked according to their relative indices.
The rank positions (abscissa) are plotted against the index values (ordinate,
logarithmic). The steepness of the curve directly represents the degree of
specialization of the worker group.

Dotted line: Camponoius sericeus ; solid line: Formica sanguinea.

1974]

Moglich & Holldobler — Nest Moving of Ants

235

A cknowledgements

We would like to thank E. O. Wilson for critically reading the
manuscript. The research has been supported by a grant from the
National Science Foundation GB-38789X1.

Literature

Arnoldi, K.

1932. Biologische Beobachtungen an der neuen palaarktischen Sklaven-
halterameise Rossomyrmex proformicarum K.Arn., nebst einigen
Bemerkungen uber die Beforderungsweise der Ameisen. Z.
Morph. Okol. Tiere 24: 319-326.

Escherish, K.

1917. Die Ameise. Schilderung ihrer Lebenweise. Braunschweig, Friedr.
Vieweg & Son.

Haskins, C. P. and E. F. Haskins

1950. Notes on the biology and social behavior of the archaic ponerine
ants of the genera Myrmecia and Promyrmccia. Ann. Entom. Soc.
Amer. 43: 461-491.

Holldobler, B.

1971. Recruitment behavior in Camponotus socius. Z. vergl. Physiol.
50: 551-568.

Holldobler, B. and U. Maschwitz

1964. Die Hochzeitsschwarm der Rossameise Camponotus herculeanus.
Z. vergl. Physiol. 50 : 551-568.

Holldobler, B., U. Maschwitz and M. Moglich

1974. Communication by tandem running in the ant Camponotus seri-
ceus. J. Comp. Physiol. 90: 105-127.

Kneitz, G.

1964. Saisonales Trageverhalten bei Formica Polyctena Foerst (Formi-
cidae). Ins. Soc. 11 (2): 105-130.

Maschwitz, U., B. Holldobler, and M. Moglich

In press. Tandemlaufen als Rekrutierungsverhalten bei Bothroponera
tesserinoda Forel (Formicidae: Ponerinae). Z. Tierpsychol.

Moglich, M.

1971. Nestumzugs- und Trageverhalten bei Ameisen. Staatsexamens-
arbeit an der Universitat Frankfurt.

Otto, D.

1958. tiber die Arbeitsteilung im Staate von Formica rufa rufo -
pratensis minor Gossw. und ihre verhaltensphysiologischen
Grundlagen. Wiss. Abh. Deutsch. Akad. Landwirschaftswissen-
schaft. Berlin, Nr. 30.

Rettenmeyer, C.

1963. Behavioral Studies of Army Ants. Sci. Bull. Univ. Kansas.
44: 281-465.

236

Psyche

[June

Wehner, R. and P. Lutz

1969. Orientierungsmechanisraen beim Nestbauverhalten der Wiiste-
nameise Cataglyphis bicolor Fab. Natur und Museum 99: 177-
189.

Wilson, E. O.

1971. The Insect Societies. Belknap Press of Harvard University Press,
Cambridge, Mass.

Zahn, M.

1957. Temperatursinn, Warmehaushalt und Bauweise der Roten Wal-
dameise Formica rufa L. Zool Beitr. 3 : 127-194.

TEMPORAL ACTIVITY PATTERNS IN
TWO COMPETING ANT SPECIES
(HYMENOPTERA: FORMICIDAE)1

By James H. Hunt
Dept, of Zoology, Univ. of California
Berkeley, CA 947202 3

Most ant species are known to exhibit some degree of patterning
in foraging rate. Interspecific differences in foraging rate can be
noted both temporally and, in temperate latitudes, seasonally. Such
differences can contribute to an effective partitioning of resources
among coexisting species. The following set of observations documents
a difference in foraging pattern that may be the significant component
of coexistence of the two closely competing species observed. The
system will be described, and possible implications of the observations
will be discussed.

The species observed were Dorymyrmex antarcticus and Tapinoma
antarcticum* , both members of the subfamily Dolichoderinae. Obser-
vations were made at Fundo Santa Laura, near Til Til, Santiago
Province, Chile, during October and November of 1971 and 1972.
The site was at 1,000 m elevation on the east-facing slope of the low
coastal cordillera. Vegetation was mixed shrubs and annuals forming
the community known as matorral, which is characteristic of the
Mediterranean climate zone of central Chile.

On visits to the site in 1971 I noted that the two species- compete
strongly for baits. A bait of honey on a small wad of cotton would
attract Dorymyrmex antarcticus workers when it was placed on the
ground in the early morning. These workers recruited a. small
number of nestmates, and activity would continue until midmorning
when the first few workers of Tapinoma antarcticum appeared.
These workers, once they located the bait, would quickly recruit
many of their nestmates, and the many small T. antarcticum workers
aggressively repelled the fewer, larger, less aggressive workers of

1A Contribution of the Structure of Ecosystems Subprogram, International
Biological Program.

2Present address: Dept, of Biology, University of Missouri, St. Louis,
Missouri 63121.

3Generic placement of this species in Tapinoma is almost certainly in-
correct. Ecologically, the species is similar to Iridomyrmex pruinosum of
the southwestern U. S.

Manuscript received by the editor May 2, 1974.

237

23B

Psyche

[June

o

CO

c

c

o

o

sunrise

noon

sunset

Time of Day

Figure 1. Activity patterns of the two ant species as observed in 1971.
Solid line — solar intensity; dotted line — T apinoma antarcticum foraging
activity; dashed line — Dorymyrmex antarcticus foraging activity.

D. antarcticus. The Tapinoma workers continued on the bait
throughout the midday, whereas the Dorymyrmex workers retreated
to their nests, apparently to avoid the high midday temperatures.
Late in the afternoon, as temperatures dropped, the Tapinoma
workers would retreat to their nest leaving the bait to be reoccupied
by Dorymyrmex workers. Activity at the bait seemed in general to
parallel the apparent activity patterns of the species, which I illus-
trated as in Figure i. The apparent tolerance of cooler temperatures
granted an exclusive period of foraging activity to D. antarcticus ;
aggressive dominance yielded foraging success for T. antarcticum
during the period when both species were active simultaneously.

In 1972 I documented these patterns quantitatively. Closely adja-
cent nests of each species were chosen, and counts of workers passing
the nest entrance during a two-minute period each half-hour were
tallied with hand counters. Soil surface temperatures were monitored
with a bulb-type thermometer placed touching the soil near the nests.
Solar intensity values were taken from a pyroheliometer (Belfort
Inst. Co.) stationed 150 m from the study nests. The pattern docu-
mented for one nest of each species on 4 October is illustrated in
Figure 2a.

On the following day I returned to the same nests to document
the early morning activity missed the preceding day. The patterns
for 5 October are shown in Figure 2b. The day was cloudy and
cool, and differences in foraging activities of the ants are of interest.
The D. antarcticus continued active all day. The T. antarcticum
workers emerged later and were out in fewer numbers than on the

1974]

Hunt — Competing Ant Species

239

Figure 2. Activity patterns of the two ant species as documented in 1972;
A = 4 October, B ~ 5 October. Triangles = Dorymyrmex antarcticus ;
circles = Tapinoma antarcticum (both in ants per minute passing nest
opening); dotted line = soil surface temperature (°C) ; dashed line —
solar intensity (cal/cmVmin) .

Soil surface temperature (°C) ( *) Soil surface temperature (°C) (•

240 Psyche [June

Hour of day

Figure 3. Activity patterns of Dorymyrmex antarcticus on 19 October
with and without experimental shading of nests; A = an unshaded nest,
B — a shaded nest.

1974]

Hunt — Competing Ant Species

241

Hour of day

Figure 4. Activity patterns of T apinoma antarcticum with and without
experimental shading of the nest; A = 1 November (unshaded), B == the
same nest on 2 November (shaded).

Ants / minute

242

Psyche

[June

preceding day, with activity peaks corresponding to peaks of soil
warmth. It seemed that foraging rates of the species were mediated
by soil surface temperatures.

On 19 October I tested this hypothesis by using a Thermos Brand
Space Blanket (a highly reflective material) to shade a nest of
D. antarcticus on a hot, sunny day. Figures 3a and 3b show activity
patterns at two neighboring nests of similar size. Workers of the
unshaded nest illustrate the expected pattern, but workers from
the shaded nest continued foraging throughout the day. Figures 4a
and 4b show a similar test with T. antarcticum. Figure 4a illus-
trates activity of a single colony on 1 November; Figure 4b is the
same nest, shaded, on 2 November. The shading yielded a marked
reduction in foraging rate. The late afternoon peak is of workers
carrying larvae and pupae and emigrating to a new nest site under
unshaded stones about 1 m from the shaded site.

Discussion

The two species studied have similar food and foraging site prefer-
ences. The observed differences in temporal foraging pattern are
hypothesized to contribute significantly to the species’ coexistence.
Soil surface temperature seems to be the proximate factor by which
the foragers regulate their activity. Endogenous activity rhythms,
often keyed to environmental factors, are known for many species.
This study illustrates how variation in environmental factors can
yield alteration of the activity rhythms.

During the course of a year at this temperate latitude (33° S)
it seems probable that Dorymyrmex antarcticus would have more
foraging hours available to it, especially during spring, fall, and
winter days, than would Tapinoma antarcticum. Perhaps the aggres-
sive dominance of Tapinoma is a necessary requisite for survival
in competition with other species. The rapid recruitment of a large
number of rather small workers in Tapinoma may illustrate com-
ponents of a foraging strategy that must yield successful foraging
returns during activity periods that are more limited than those of
a coexisting competitor.

Acknowledgments

This research has been supported by NSF grant GB 31 195 to
R. K. Colwell. This paper is a contribution from the Structure
of Ecosystems Subprogram, International Biological Program. I
thank R. R. Snelling, who taxonomically determined the species
studied, and E. Reid, who prepared the illustrations. For review
of the manuscript I am grateful to R. K. Colwell, B. Holldobler
and E. O. Wilson.

RELATIONSHIP OF LARVAL FOOD-PLANTS
AND VOLTINISM PATTERNS IN
TEMPERATE BUTTERFLIES*

By Frank Slansky, Jr.**

Department of Entomology, Cornell University
Ithaca, New York 14850

An interesting aspect of phenology is the number of broods that a
species produces in the growing season. Even among a fairly uniform
group like the butterflies within a restricted geographical area, voltin-
ism patterns of the different species may vary considerably. Explana-
tions of possible causes of, and ecological implications of these pat-
terns have apparently seldom been attempted.

It might be expected that voltinism patterns are often genetically
determined and are regulated by some environmental clue (cf. Wig-
glesworth, 1967), but knowing the proximate reason does not reveal
the ultimate causes that brought the voltinism patterns under such
control. Obviously, the presence of a food source for the larvae (and,
although less well studied, for the adults as well) will have a major
influence on these patterns. The exclusive use of the vernal herb
Dentaria by larvae of the West Virginia white, Pieris virginiensk,
precludes any more than one brood per season. As an adaptation to
feeding on this ephemeral food-plant, the larvae upon entering the
pupal stage undergo obligate diapause, even though the genetic
mechanism for multivoltinism exists in this species (Shapiro, 1971).

A more subtle explanation of voltinism patterns may emerge from
examining larval growth rates on different plants (e.g. Dowdeswell
& Willcox, 1961 ; Hovanitz & Chang, 1962; Sharifi & Zarea, 1970).
Nutritional (including water) differences (Soo Hoo & Fraenkel,
1966; Waldbauer, 1968; Feeny, 1970; J. M. Scriber and P. P.
Feeny, ms. in prep.; Slansky, 1974) and differences in secondary
chemical content (Gupta & Thorsteinson, i960; Nayar & Thorstein-
son, 1963; Feeny, 1970) of different plant species and of the same
plant species at different stages of growth in large part determine the
growth rates of larvae feeding on the plants, and thus perhaps for

*This work was supported by Hatch Grant NYC-139413 and NSF Grant
GB-33398.

**Present address: Department of Zoology, University of Iowa, Iowa City,
Iowa 52242.

Manuscript received by the editor April 12, 1974.

243

244

Psyche

[June

some species, voltinism patterns as well. The purpose for this work
is to examine the hypothesis that, of the butterflies in particular
climatically similar areas, those which have larval food-plants that
are apparently nutritionally unsuitable for much of the growing sea-
son will exhibit in general fewer broods per season than those butter-
flies which have larval food-plants that are apparently nutritionally
adequate throughout the season.

Methods

In order to examine the hypothesis the number of broods produced
per year by, and the larval food-plants of, eighty-six species of but-
terflies were tabulated for three transition zone areas in eastern and
mid-western United States (Forbes, 1906; Saunders, 1932; Ebner,
1970). Additional food-plant records were obtained from Ehrlich
and Ehrlich (1961) and Shapiro (1966). The voltinism patterns of
selected species of these butterflies were then related to changes in
the nutritional suitability of their larval food-plants.

Results and Discussion

That differences in larval growth rates are reflected in the voltin-
ism patterns of butterflies is seen in the following example. Under
similar temperature and humidity conditions the duration of the
larval stage of the imported cabbageworm, Pieris rapae , is about 15
to 20 days on some of its food-plants (Slansky, 1974), while that of
the Eastern tiger swallowtail, Papilio glaucus, is about 35 to 45 days
on some of its food-plants (J. M. Scriber, unpublished data). As a
consequence, P . rapae generally exhibits one brood more than P.
glaucus in the areas where they occur together (Table 1).*

Further support for the hypothesis is found when the voltinism
patterns of selected butterfly species are examined in relation to sea-
sonal changes in the nutritional suitability of larval food-plants. For
example, species whose larvae feed on oaks (Quercus) , such as the
hairstreaks Satyrium edwardsii, S. liparops , and S. calanus , and the
skippers Erynnis brizo , E. horatius, and E. juvenalis, usually have

^Although P. glaucus is generally considered to have two broods in the
areas surveyed in this study, it may be that there is only one ‘true’ brood.
It appears that a high percentage of the pupae from the previous season
emerge in the early summer, constituting the ‘true’ first brood, and that
the remaining pupae emerge later in the summer, constituting an ‘appa-
rent' second brood (i.e., not the progeny of the first brood of that season)
(Scudder, 1889; R. Lederhouse and J. M. Scriber, unpublished data).

1974]

Slansky — Temperate Butterflies

245

TABLE 1. Number of broods, larval food-plants, and hibernating stages
for several butterflies in three transition zone areas. Data from
Forbes (1906), New England (A) ; Saunders (1932), Allegany
State Park, N.Y. (B) ; and Ebner (1970), Wisconsin (C).
E — egg; L = larva; P = pupa; A = adult.

Number Hiber-

of Broods nating

Species

A

B

C

Larval food-plants

Stage

HESPERIIDAE

Amblyscirtes samoset

l

_

1

Gramineae

?

A. vialis

2

-

1

Andropogon

P

Euphyes dion

-

-

1?

Scirpus, Carex?

?

E. conspicus

-

-

1

Car ex

?

Poanes massasoit

2

-

1

Carex

?

P. hobomok

1

1

1

Poa, Panicum

L,P

P. viator

-

-

1

Zizania, Phragmites?

?

tV allengrenia otho

-

1

1

Panicum, Digitaria

L

Polites coras

1

-

2

Gramineae

?

P. themistocles

-

-

1-2

Panicum

?

P. mystic

2

2?

1

Poa

L

Hesperia metea

1

-

1

Andropogon

?

H. sassacus

1

1

1

Panicum

?

H. leonardus

1

-

1

Panicum, Erogrotis

L

Ancyloxipa numitor

3

3

2-3

Poa

L,P

Pholisora catullus

2

-

1

Chenopodium, Amaranthus ,
Ambrosia, Marrubium

L

Pyrgus communis

-

3

2

Althaea, Malva

L,P

Erynnis icelus

1

1

1

Populus, Betula

L

E. brizo

1

-

1

Ouercus, Castanet

L

E. lucilius

3

-

2

Aquilejia

L

E. horatius

2

-

-

Quercus

?

E. juvenalis

2

-

1

Quercus

L

Thorybes bathyllus

1

-

1

Leguminosae

P

T. pylades

2

2?

1

Lespedeza, Trifolium,
Medic ago

P

Epargyreus clarus

1

2

1-2

Robinia, fVistaria

P

PAPILIONIDAE

Battus philenor

2

2

_

Aristolochia

P,A

Papilio polyxenes

2

2

2

Umbelliferae

P

P. glaucus

2

2

1-2

Fraxinus, Prunus, Betula,
Populus, others

P

P. troilus

2

2

-

Lindera, Sassafras, Persea

P

246

Psyche

[June

Number Hiber-

of Broods nating

Species

A

B

C

Larval food-plants Stage

PIERIDAE

Pieris protodicr

3

_

3

Cruciferae P

P. napi

3

-

3

Cruciferae P

P. virginiensis

-

-

1

Dentaria P

P. raPae

3

3

3

Cruciferae P

Colias eurytheme

3

3

3

Medicago, Trifolium P

C. philodice

4?

3

3

Trifolium L?

C. interior

1

-

1

V accinium ?

Anthocaris midea

1

Cruciferae, esp. buds, flowers, P
seeds

LYCAENIDAE

Harkenclenus titus

l

1

1

Prunus E

Satyrium liparops

1

1

1

Quercus, Salix, Rubus, Malus, E
Prunus, V accinium, others

S. calanus

1

1

1

Quercus, Carya, Castanea E,L

S. edwardsii

1

-

1

Quercus E,L

S. acadica

1

1

1

Salix E

Callophrys irus

1

—

—

Babtisia, Crotalaria, Lupinus, P
esp. flowers, fruit

C. henrici

1

—

1

V accinium, Prunus, Cercis, P

bores into flowers

C. augustinus

1

1

V accinium, Kalmia, Arbutus, P
Arctostaphylus, Ceanothus,
esp. flowers and berries

C. niphon

1

-

1

Pinus P

C. gryneus

1-2

-

2

Juniperus P

Strymon mclinus

2

2

Polygonum, Phaseolus, Malva, ?
Hypericum, Humulus, others,
bores into bud, fruit

Lycaena thoc

2

2

3

Rumex, Polygonum E

L. epixanthe

1

-

1

V accinium E

L. phlaeas

3

2

3

Rumex P,L?

Everes comyntas

3

3

3-4

Leguminosae, foliage, buds, L

flowers

Celastrina argiolus

2

2

2

Cornus, Rhus, V accinium, P

Cimicifuga, Ceanothus, others

NYMPH ALIDAE

Asterocampa celtis

2

-

2

Celtis L

A. clyton

2?

-

2

Celtis L

Limenitis arthemis

1-2

1-2

2

Salix, Betula, Populus, Tilia L

1974]

Slansky — Temperate Butterflies

247

Species

L. astyanax 1

L. archippus
Vanessa atalanta
V. virginiensis

V. cardui
Junonia coenia

Nymphalis vau-album i
N. milberti
N. antiopa
Polygonia

interrogationis
P. comma
P. faunas
P. progne
Chlosyne nycteis
C. harrisii
Phyciodes tharos
P. batesii :

Euphydryas phaeton
Botaria selene
B. toddi
Speyeria idalia
S. atl antis
S. cybele
S. aphrodite

DANAIDAE

Danaus plexippus

SATYRIDAE

Lethe portlandia
L. eurydice
Euptychia cymela
Cercyonis pegala
Oeneis jutta

Number Hiber-

of Broods nating

B C Larval food-plants Stage

-2 2? 2

2 2 2

2 2 2

2 2 2

2 2 2

2

2? 2? 2?

3 2 3

2 2 2

Prunus, Malus, Pyrus, L

Crataegus, Salix, Populus
Salix, Populus, Prunus, Malus L
Urticaceae P,A

Artemisia, Gnaphalium, P,A

Antennaria, others
Cirsium, Carduus A,P?

Plantago, Gerardia, Sedum, A?
Antirrhinum

Betula, Salix, Populus A

Urtica A

Ulmus, Celtis, Salix, Populus A

2

2

2

2

1

2

2?

1

3

3

1

1

2

2

1

2

1

1

2

1

1

3

3

1

1

1

1

2 Ulmaceae, Urticaceae A

2 Ulmaceae, Urticaceae A

1 Betula, Salix, Alnus, Ribes A

2 Ribes A

1 Helianthus, Actinomeris , Aster L

1 Aster L

2 Aster L

1 Aster L?

1 Chelone L

2-3 Viola L

2 Viola L

1 Viola L

1 Viola L

1 Viola L

1 Viola L

2 2 2 Asclepias

Migratory

1 1 1

1 1 1

1 1 1

1 1 1

1 - 1

Gramineae, Cyperaceae?
Gramineae, Cyperaceae?
Gramineae

Gramineae esp. Tridens
Gramineae, Cyperaceae

L

L

L

L

L

248

Psyche

[June

one, or at most two, broods, and most hibernate in the larval and/or
egg stage (Table 1). This may be due to the presence in the oak
leaves of tannins that restrict protein utilization and hinder larval
development, in many cases limiting consumption of the foliage to
spring and early summer when the protein content is highest and the
tannin content lowest (Feeny, 1970).

In addition, the leaves of many species of shrubs and trees contain
substantially lower percentages of water than the leaves of many
annual and perennial herbs (Way, 1853; Fagin & Watkins, 1932;
Soo Hoo & Fraenkel, 1966; Slansky, 1974; J. M. Scriber, unpub-
lished data). Since the percentage of water in most lepidopterous
larvae appears to be about 80 to 90% (Evans, 1939a; 1939b; Wig-
glesworth, 1967; Slansky, 1974; J. M. Scriber, unpublished data),
the growth of larvae feeding on plants with a percentage of water
lower than their own tissue may be limited by the availability of
water so that they exhibit slower growth rates in comparison to
larvae feeding on plants that have a percentage of water equal to or
greater than that of the larvae (Southwood, 1972; J. M. Scriber &
P. P. Feeny, ms. in prep.). The fact that a number of tree and
shrub feeders exhibit but one or two broods (Table 1) supports this
contention. Along these lines Feeny (1974) has suggested that plant
species that are abundant and/or persistent have apparently evolved
“quantitative” defenses (low nutrient contents, tough leaves, high
contents of unspecific chemicals like tannins) that act as significant
ecological barriers to phytophagous insects, in contrast to plant spe-
cies that are rare and/or ephemeral that have apparently evolved
“qualitative” defenses (specific secondary chemicals) that act as only
slight ecological barriers to adapted insects although having consid-
erable impact as evolutionary barriers to non-adapted insects.

The majority of grass- and sedge-feeders (all Satyridae and sev-
eral Hesperiidae) , with characteristically sluggish larvae (Scudder,
1889), exhibit but one brood (Table 1). In view of the low moist-
ure and high fiber content of many grasses (Way, 1853; Watson,
1951), it is not surprising that larval development might be slow on
such plants. The hibernation of many of these satyrids and hesperiids
as early instar larvae (Table 1) may be an adaptation for the avoid-
ance of nutritionally poor, mature grass plants and for the maximum
utilization of the spring flush of succulent growth when moisture
and nitrogen levels are high and fiber content low (Watson, 1951).

Thus, the voltinism patterns of several butterfly species are given
an ecological meaning, and the hibernation of many butterfly species
as eggs, larvae, and perhaps fertile adults (Scudder, 1889), when

1974]

S/ansky — Temperate Butterflies

249

viewed as a means of utilizing tender, nutritious plant tissue in the
spring, is seen as an adaptive strategy (c.f. Morse, 1971; Schoener,
1971). Of course, it is not suggested that the nutritional quality of
the larval food-plant is of sole importance in determining voltinism
patterns. For example, larvae of both the Tawny crescent, Phyciodes
batesii, and the Pearl crescent, P. tharos, feed on Aster; the former
exhibits one brood and the latter two or three (Table 1). Larvae
of two Bolaria species and of four Speyeria species all feed on
Viola; the former exhibit two to three broods, the latter one brood
(Table 1). There may be differences in the nutritional quality of
different species of plants within these genera, but other factors
appear to be causing these differences in voltinism patterns. The
Least skipper, Ancyloxipa numitor, larvae of which feed on what
might be characterized as ‘nutritionally poor’ grass plants, exhibits
two or three broods while the Eastern black swallowtail, Papilio
polyxenes, larvae of which feed on what might be characterized as
‘nutritionally adequate’ plants in the Umbelliferae, exhibits but two
broods (Table 1 ).

Obviously, the interaction of several other factors (e.g. larval size,
larval mortality rates, and the presence of adult nectar sources)
with the nutritive and secondary chemical contents of the larval
food-plants influences the voltinism patterns. For example, the slow
growth rates of larvae of P. glaucus may only be ‘permitted’ because
the combination of the patchy distribution and warning coloration
of the larvae may reduce mortality losses from predators and para-
sitoids (J. M. Scriber, personal comm.). On the other hand, the
rapid growth rates of larvae of P. rapae may be a ‘necessity’ because
of the nature of its food-plants (i.e., mostly early successional
species) and because of the high mortality losses of the larvae to
predators, parasitoids, and disease (Richards, 1940; Pimentel, 1961;
Dempster, 1969; Parker, 1970). The more rapid growth rates of
lepidopterous larvae from ‘geographical races’ occurring in regions
with short growing seasons in comparison to those of larvae from
regions with longer growing seasons (Goldschmidt, 1940) provides
another example of the ecological importance of differences in larval
growth rates.

Such considerations of the interrelationship of life history phe-
nomena and population dynamics (e.g. Cole, 1954; Murdock, 1966;
Istock, 1967; Gadgil & Bossert, 1970; Morse, 1971 ; Schoener, 1971 ;
Willson, 1971) raise a number of ecologically relevant and as yet
insufficiently answered questions, such as :

250

Psyche

[June

1 ) What are the advantages of monophagous and polyphagous
larval feeding habits (c.f. Brues, 1920; 1924; Buxton, 1923; Deth-
ier, 1954; Brower, 1958; Schoener & Janzen, 1968; Levins &
MacArthur, 1969) ?

2) Why do some butterflies exhibit a long-lived adult stage (e.g.
almost a full year in the Tortoise shell, N ymphalis vau-album ) (c.f.
Murdoch, 1966; Howe, 1967; Gadgil & Bossert, 1970)?

3) What selective forces cause the complex voltinism pattern ex-
hibited by a number of butterfly species in which part of a brood
becomes dormant while the remainder continues normal development
(Scudder, 1889; Oliver, 1972)? Perhaps this may allow these spe-
cies to exploit marginally favorable periods while maintaining a. re-
serve population for the usually favorable season and/or may aid in
reducing losses to temporally restricted predators and parasitoids
(c.f. Baltensweiler, 1968; Waloff, 1968).

4) Why do some butterflies exhibit multivoltinism ? Perhaps this
is a means of building up large populations to withstand high mor-
tality losses in the summer, especially because of biological causes,
and in the winter, especially because of physical causes, such as is
seen in several pierid butterflies that start out with low population
levels in the spring and become more abundant as the summer pro-
gresses (Scudder, 1889).

Detailed ecological studies of a taxonomically and geographically
well-known group like the butterflies will help to answer such ques-
tions and will aid in making meaningful predictions about such im-
portant phenomena as the population buildup of a potential or actual
pest species.

Acknowledgements

The author wishes to thank J. Mark Scriber and William Blau
for reading the manuscript and for their helpful suggestions.

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VARIATIONS IN CLEANING BETWEEN THE SEXES OF
SINELLA COECA (COLLEMBOLA: ENTOMOBRYIDAE)*

By Elizabeth S. Waldorf
Department of Zoology
Louisiana State University
Baton Rouge, Louisiana 70803

With the widespread and frequent occurrence of pheromones
among insects, it is easy to appreciate the significance of cleaning
and grooming behaviors. However these behaviors are as yet poorly
known. Among springtails two papers by Simon (1961, 1963) present
the most thorough reports. These describe the types of cleaning in
representatives of several families, noting the widespread, though
infrequent, use of cleaning droplets released from the mouth.

My work examines Sinella coeca (Schott) (Family Entomobryidae)
for behavioral differences between the sexes. In particular, I have
determined the frequency of cleaning and the duration of bouts of
cleaning behavior. The initial expectation is that the sexes will spend
about equal time cleaning. However, since males are more active,
moving about most of the time, their bouts of cleaning are probably
shorter. To compensate, I expected males to have more frequent
cleaning sessions. An experiment was carried out to test these
expectations.

Methods

Sinella coeca was reared in mass culture as described previously
for Sinella curviseta (Waldorf, 1971). Cultures were maintained
at room temperature (230 ± i°C) and supplied commercial yeast
as food. Males and females were isolated in individual glass vials
with plastic caps. The vials, measuring 18.1 mm in inner diameter,
contained a moist plaster of paris-charcoal substrate (to maintain
high humidity) and food. After 24-26 hours of isolation, each animal
was observed for five minutes, and the number of bouts of cleaning
and the total time (out of 5 minutes) engaged in cleaning recorded.
This procedure was followed for 50 males and 50 females. To
prevent bias the vials were mixed so that during the observation
periods the sex of test animals was usually unknown.

* Manuscript received by the editor May 20, 1974.

254

1974]

Waldorf — Sinella coeca

255

Table 1. Characteristics of male and female bouts of cleaning behavior.

Number of
bouts/animal

Length of
bouts (in sec)

X

n

SD

X

n

SD

Males

1.88

50

1.33

29.4

94

38.6

Females

1.38

50

1.23

23.8

69

29.5

Results

The data in Table i present the overall characteristics of males
and females. Comparison of the lengths of bouts using a t-test shows
that these are not significantly different (t = 1.049). However the
number of bouts is significantly larger in males than in females
(t = 1.949; .05>P>.0i). Rather than the expected differences,
it appears that male sessions of cleaning are of equal length and
more frequent than those of females.

At times of observations some individuals were pharate, some vials
contained exuviae and others eggs. These allowed the animals to be
subdivided into groups corresponding approximately to position in
instar. Among the males only two ecdysed within the 24 hour isola-
tion period, a sample too small to be meaningful. However among
the females, the categories were sufficiently large to permit evaluation.

Table 2 gives the data for females of different types. Of the
four pharate females, none exhibited any cleaning behavior. Com-
parison of recently ecdysed females and females with eggs to remain-
ing females revealed no significant differences in bout length. In
contrast, comparisons of cleaning frequency among these females
showed that females with eggs clean significantly more often
(t = 2.488; 25df; P .01) than other females.

Discussion

As Simon (1961) has noted, the antennae, the location of im-
portant sensory receptors, are cleaned most often. There is possibly
a positive correlation between frequency of cleaning and the required
sensitivity of receptors. In pharate females the old receptors have
presumably lost their nervous connections, so that there is no adaptive
advantage in cleaning them. In contrast, females that have recently
deposited eggs are sensitive to the required stalked spermatophores
on the substrate. If this species has alternate reproductive and non-

256

Psyche

[June

Table 2. Characteristics of cleaning bouts by females of various types.

X

Number of
bouts/animal

n

SD

X

Length of
bouts (in sec)

n

SD

Pharate females

0

4

0

—

—

—

Recently ecdysed
females

1.63

19

1.34

22.4

31

29.0

Females with eggs

2.1

9

1.05

20.8

19

28.9

Other females

1.05

18

1.00

29.1

19

31.7

reproductive instars with oviposition confined to early in the repro-
ductive instar as Sinella curviseta (Waldorf, 1971), newly ecdysed
females are possibly a mixture of two types: sensitive ones early in
the reproductive interval and indifferent, or less sensitive ones, early
in the nonreproductive instar. Although the distribution of numbers
of cleaning bouts per female is not bimodal, the mean of this category
is 1.63, approximately the average (1.57) of the sensitive group
with eggs and the remaining females. Note also that the variance
of the number of bouts of recently ecdysed females is 1.796 and
that of females with eggs and remaining females are 1.102 and .998
respectively.

Considering males, Mayer (1957) has reported that males of
Sminthurides aquaticus Bourlet clean more frequently when clasping
the antennae of females than when alone. This is consistent with
the hypothesis that these males are highly sensitive to olfactory
stimuli.

The data indicate that males engage in more total cleaning than
females. My recent studies of Sinella curviseta (Waldorf, in manu-
script) demonstrate that females of this species produce a sex
pheromone. This stimulates spermatophore deposition by males in
the reproductive instar. Of the males in the present study 48 of 50
were in the reproductive instar at observation time. By frequent
cleaning, these maintain their sensitivity to olfactory stimuli including
possible pheromones.

Literature Cited

Mayer, H.

1957. Zur Biologie und Ethologie einheimischer Collembolen. Zool.
Jahrb. abt. Syst. Okol. Geogr. Tiere 85: 501-570.

1974]

Waldorf — Sinella coeca

257

Simon, H. R.

1961. Beobachtungen zum Verhalten arthropleoner Collembolen (Aptery-
gota). Deutsch Entomolog. Z., N. F. 8: 216-221.

1963. Zum Putzverhalten arthropleoner Collembolen (Ins., Apterygota).
Entomolog. Z., 73 : 221-228.

Waldorf, E. S.

1971. The reproductive biology of Sinella curviseta (Collembola:
Entomobryidae) in laboratory culture. Rev. Ecol. Biol. Sol
8: 451-463.

Centennial of Entomology. Cornell University has announced
a Centennial of Entomology, celebrating John Henry Comstock’s
graduation and the founding of studies in entomology at the Uni-
versity. A symposium on Insects , Science and Society will be held
on October 14 and 15, 1974. The speakers at the symposium will
be as follows:

Howard E. Evans, Colorado State University: The Comstock
Heritage.

John J. McKelvey, Jr., The Rockefeller Foundation: Insects and
Human JV elf are.

Edward O. Wilson, Harvard University: Insect and Hu?nan
Societies.

John S. Kennedy, Imperial College Field Station, Ascot, England:
Insect Dispersal.

Richard D. Alexander, University of Michigan: Insect Communi-
cation — A coustical.

Wendell L. Roelofs, New York State Agricultural Experiment
Station, Geneva: Insect Communication — Chemical.

Mano D. Pathak, International Rice Research Institute, Philip-
pines: Patterns of Interaction between Plants and Insects.

T. R. E. Southwood, Imperial College Field Station^ Ascot, Eng-
land: The Dynamics • of Insect Populations.

Powers Messenger, University of California, Berkeley: Parasitoids ,
Predators and Population Dynamics.

Waldemar Klassen, U.S.D.A., ARS-Plant Industry Station,
Maryland: Pest Management — Organization and Resources.

L. Dale Newsom, Louisiana State University: Pest Management
— Concept to Practice.

Further information can be obtained from the Chairman, Depart-
ment of Entomology, Cornell University, Ithaca, New York 1485a.

— Editor

NOTES ON NECROPHORIC BEHAVIOR IN THE
ARCHAIC ANT MYRMECIA VINDEX
(FORMICIDAE: MYRMECIINAE)*

By Caryl P. Haskins and Edna F. Haskins
2100 M Street, N.W.

Washington, D.C.

Introduction

Ants of the Australian and New Caledonian genus Myrmecia
apparently include the most archaic living Formicidae. Brown
(1954) suggested that the genus may represent a relatively late
evolutionary offshoot, specialized but fundamentally conservative,
from a line of extremely generalized archaic forms represented as
fossils by the genus Prionomyrmex of the Baltic amber and by the
living Australian Nothomyr/necia macrops , of which only two workers
have ever been found. Species of Myrmecia , therefore, may well
illustrate the earliest patterns of Formicid social organization, and
embody the most archaic patterns of Formicid social behavior, that
we are likely to be able to study in detail in the laboratory or the
field.

The bodily habitus of Prionomyrmex and Nothomyrmecia — not
to mention of the far more archaic Mesozoic fossil genus Spheco-
myrma first described by Wilson, Carpenter, and Brown (1967)
which may well represent, as Wilson (1971) suggests, an antecedent
of the Myrmecioid complex of ants — all suggest active, epigeicallv
foraging insects: a characteristic virtually universal in contemporary
species of Myrmecia. This, combined with the fairly large size of
the communities of many species of Myrmecia — specific counts of
1586 workers and over 2000 in total colony personnel have been
made from larger colonies of M. gulosa (Haskins and Haskins,
1950) — raise the interesting question of how far these archaic forms
may have evolved any pheromone-mediated patterns of community-
integrating behavior so conspicuous in many higher ants. This is an
interesting and complex area of inquiry, the answers to which are
far from obvious, as the recent investigations of Robertson suggest.

A prior question may be significant in this context. Do ants of
the genus Myrmecia exhibit characteristic behavioral responses to
particular chemical substances normally encountered in the external

* Manuscript received by the editor March 28, 1974.

258

1974]

Haskins & Haskins — Myrmecia vindex

259

environment: substances which commonly evoke specific behavioral
responses in higher ants? And if so are the responses elicited in
Myrmecia essentially like those of higher ants?

We have recently described one such set of reactions: the stimulus
to attack behavior in Myrmecia gulosa elicited by formic acid, a
normal exocrine defense product of its common prey-genus Campo-
notus , but not, so far as can be determined, of M. gulosa itself
(Haskins, Hewitt, and Haskins, 1973). The present investigation
is concerned with the reactions of workers of a species of Myrmecia
to substances which commonly stimulate necrophoric behavior in
higher ants, such as oleic and related fatty acids.

Wilson (1958) and Wilson, Durlach, and Roth (1958) demon-
strated that when groups of workers of the Myrmicine ant Pogono-
myrmex badius come into contact with formic acid, ethylamine,
triethanolamine, phenol, n-butyric acid, n-valeric acid, n-caproic acid,
or n-caprylic acid, absorbed on centimeter-square patches of filter
paper, they exhibit weak to moderate alarm behavior, sometimes
passing into digging behavior, concentrated about the squares. In
the case of oleic acid — and of that substance alone — the ants
transported the paper squares away from the nest to the kitchen
middens. A fatty acid component — quite probably oleic acid —
obtained from the decaying bodies of P. badius workers also elicited
necrophoric behavior in Solenopsis saevissima.

Blum, Doolittle, and Beroza (cf. Blum, 1970) fractionated dead
workers of Solenopsis saevissima and determined that the releasers of
necrophoric behavior were restricted to the rich free fatty acid
fractions. Myristoleic, palmitoleic, oleic, and linoleic acids were
present in these fractions, and all these acids possessed necrophoric
activity in this species.

Such a reaction to chemicals like oleic acid has obvious adaptive
advantage for any ant, or indeed for any social insect inhabiting a
closed nesting situation. Such chemicals can serve as sensitive “indi-
cators” of objects in the nest — whether prey or remains of adult
or immature members of the community — which are dangerously
decomposed and should be removed. It seems plausible, therefore,
that this behavior pattern may have been established very early in
the social evolution of the Formicidae. Hence it seemed of particular
interest to investigate it within the Myrmeciinae.

Necrophoric Behavior in Myrmecia vindex

The species Myrmecia vindex was chosen for several reasons.
It is a common and wide-ranging form in western Australia, where

2bo Psyche [June

it prefers open, partially xerophytic woodland habitats, but ranges
much further, to South Australia and probably New South Wales
and Victoria as well. It is an unusually hardy species and its colonies
can be maintained in laboratory culture for many years. The pri-
mary reason for choosing it for this investigation, however, was that,
in the populations from which nests were selected for both field and
laboratory work, even mature colonies are typically quite small.1
This and other characteristics suggest that the species is one of the
socially less-evolved members of the genus and thus lend particular
interest both to the reactions of its workers to necrophoric substances
and to pheromones of higher ants.

Laboratory Tests

Reactions of workers of Myrmecia vindex to Oleic Acid

A portion of a colony of M. vindex consisting of approximately
30 workers, 20 cocoons, and 25 larvae, collected at Kings Park,
Perth, West Australia, on December 28, 1961, and long established
in the laboratory in an earth-containing Lubbock type glass nest of
dimensions 7" X 12" and Y\r in depth, was set up in a foraging
arena consisting of a rectangular plastic box, 48" X 24" X 6.5"
in inside dimensions, to which the ants had free access. The bottom
of the arena was covered with clean brown paper.

After the colony had become thoroughly conditioned to the en-
vironment, piles of brood which had been impregnated with 1 to 2
drops of various substances were placed on cardboard sheets, 6" X
8", at some distance from the nest entrance. There was early indi-

^ounts of four completely excavated apparently mature colonies from
this population were made in January, 1964, with the following results:

Colony 1

21

typical brood female

callow subalates (apparent female-worker intermediates)

211

workers

Colony 4

1

typical brood female

176

workers

Colony 6

1

typical brood female

187

workers

Colony 7

1

typical brood female

149

workers

This compares with a population count of a study nest of M. vindex reported

by Douglas

and

McKenna (1970) as follows:

1

typical brood female

38

alate virgin females

38

dealate virgin females

210

workers

224

males

1974]

Haskins Haskins — Myrmecia vindex

261

cation that larvae were only marginally suitable for this work.
Treated larvae were sometimes returned to the nest, as though larval
odor, even when overlaid by that of oleic acid, still dominated
behavior. With cocoons, however, worker reactions were more
consistent.

At 5:30 p.m. two piles of 10 larvae and 10 cocoons each were
placed on top of the Lubbock nest. The larvae were quickly restored
to the nest. By 6:15 p.m., 5 cocoons of the untreated group had been
transported to the nest, together with 3 from the fraction treated
with oleic acid. By 6:38 p.m. all cocoons from the untreated pile had
been transported to the nest, while no further oleic-acid-treated
cocoons had been removed. By 8:55 the following morning, how-
ever, 4 of the oleic-acid-treated cocoons had been deposited in refuse
middens in the arena, 2 being deposited 45" from the nest entrance
and 2, 10" away. Three were still in original position on the card-
board, but had been well covered with earth. Earth grains obtained
from inside the nest had also been deposited on spots of oleic acid
which had drained from the cocoons to the cardboard.

The experiment was then repeated more precisely in the pattern
of the work reported with P. badius. Ten squares of paper, y2n X
yy* , were soaked with ole’c acid and scattered at random over the
surface of the nest, together with 10 identical but untreated squares,
appropriately identified. Two larvae, likewise treated with oleic
acid, were included. The trial was begun at 9:10 a.m.

Workers coming into contact with the treated objects immediately
executed marked cleaning movements and a rubbing of the gular
surface of the head against the substrate. At 9 :20, 1 larva was picked
up by a worker after much hesitation and held in the mandibles
for 4 minutes, when it was grasped by a second, and then deposited
by both at the nest entrance, but not taken inside. The impression
of a conflict of drives was strong. One minute later this larva was
again picked up, carried about 3" from the nest entrance, and
dropped. By 9:53, it had been brought into the nest, while the
second larva was held within y^' of the nest entrance. Two minutes
later it also was brought into the nest.

At 10:00 a.m. a worker was beginning to deposit earth grains on
the oleic-acid-treated papers, and by 10:53, 4 papers carried earth
grains. By 1 :oo p.m., one of the treated larvae had again been
brought out of the nest, and a worker was preparing to drop it on
the midden located 45" from the entrance. By 5 :0O p.m. both
treated larvae were on this midden. Burying of the oleic acid squares
continued but they were not moved toward the middens.

262

Psyche

[June

At 5:10 p.ni., 2 Tenebrionid larvae, 1 treated with oleic acid
and the other untreated, were placed outside the nest entrance. At
5:13 p.m. the untreated larva was carried into the nest, and 1 minute
later the treated larva also was seized and carried in. At 5 130 p.m.,
however, the treated larva was brought out, carried to a distance of
34", and discarded. The untreated larva was retained within the
nest and shortly consumed.

These trials gave strong evidence that patterns both of burying and
of rejection of objects impregnated with oleic acid are exhibited by
M. vindex. It then seemed of interest to compare the reactions to
oleic acid with those of other substances tested by Wilson et al. in
their work with P. badius.

Reactions of workers of Myrniecia vindex to Methyl
benzylamine; Caproic acid; Oleic acid; Formic acid

Using the same colony and with the same experimental arrange-
ments as above, 14 cocoons, ranging from freshly spun to nearly
ready to eclose, were impregnated as follows:

Methylbenzylamine (3 drops)

Caproic acid (3 drops)

Oleic acid (3 drops)

Formic acid

2 cocoons were retained as controls.

All these cocoons were placed on cardboards, and mounted on
top of the Lubbock nest as before, at 2:04 p.m.

At 2:19 p.m. the 2 controls and the 3 cocoons treated with
formic acid had been permanently returned to the nest. At 5 140
p.m., 1 cocoon from the caproic acid group had been removed 40.5"
from the nest and deposited on a refuse midden. All other cocoons
remained untouched.

At 8:45 a.m. on March 31, no further cocoons had been removed
from the cardboards, but a few grains of earth had been deposited
on the cardboards of the caproic- and oleic-acid treated objects.
During the night the caproic-treated cocoon deposited on the refuse
midden had been retrieved into the nest, but at 8 155 a.m. it was
again brought out and carried to the midden, 44" from the nest
entrance. Almost immediaeely it was again retrieved and returned
to the nest. At 9:10 a.m., it was once more removed and dropped
about 9" from the nest entrance. By 2 103 p.m. this cocoon had
been carried to a midden 36" from the entrance, but 1 minute later

3 cocoons
3 cocoons
3 cocoons
3 cocoons

1974]

Haskins csf Haskins — Myrmecia vindex

263

it was returned to the nest once more, only to be finally removed
by 2:33 p.m. and dropped about 32" from the nest entrance. At
this time the 3 cocoons treated with oleic acid, the 3 treated with
methylbenzylamine, and the remaining 2 treated with caproic acid
were still in place on the cardboards, and a few grains of earth
had been deposited about each. They were permanently ignored.

Reactions of workers of Myrmecia vindex to
Triethanolamine: n- Valeric acid; Oleic acid

The same experimental arrangement was set up, using another
colony of M. vindex , taken from the same population in Kings Park,
Perth, West Australia, on January 10, 1965. Cocoons were treated
as before:

Triethanolamine (3 drops) 3 cocoons

n-Valeric acid (3 drops) 3 cocoons

Oleic acid (3 drops) 3 cocoons

3 cocoons were retained as controls.

The 12 cocoons were arranged radially about the nest entrance,
the control and the oleic acid groups furthest from it. Exposure
was begun at 2 :47 p.m.

Triethanolamine proved a powerful attractant, and the cocoons
treated with it aroused immediate and intense attention. Within
3 minutes, 1 was dragged into the nest. At 2:51 a second trietha-
nolamine-treated cocoon was dragged to the nest enerance and
dropped. At 2 :52 the third was carried to the oleic-acid-treated
group outside the nest and dropped there. Two control cocoons
were carried into the nest at the same time. A spot of triethanolamine
stain on the containing card was repeatedly examined by workers.

The second triethanolamine-treated cocoon, which was deposited
initially near the nest entrance, at 2 :56 p.m. was carried to a midden
45. 5" away; it was then picked up almost immediately and re-
deposited at a distance of 40". At 3 :02 the triethanolamine-treated
cocoon which had first been brought into the nest was taken out,
carried to a point about 9" away, and dropped. Earth grains had
been deposited about the n-valeric acid group. The 3 oleic-acid-
treated cocoons had not been moved nor buried. By 4:00 p.m. all
three triethanolamine-treated cocoons had been discarded in a group
39" from the nest entrance; 1 control cocoon was still in place;
and all 6 cocoons treated with n-valeric and oleic acid remained in
their initial position. At 4:50 p.m. there had been no further change
and the run was terminated.

264

Psyche

[June

On April 2, 1965, the same experiment was repeated, substituting
n-butyric for n-valeric acid. Results were essentially the same. Fifty-
five minutes after the start, 3 of the 4 triethanolamine-treated cocoons
had been transported for distances of 40", 21", and 25.5" from the
nest entrance and dropped ; although none of the n-butyric or oleic
acid cocoons had been moved, there was a considerable deposit of
earth particles about them. One control cocoon had been moved
about 27', but none had been carried into the nest. Twenty minutes
later, the triethanolamine group was still being moved at frequent
intervals, and the oleic acid group was being heavily banked with
earth grains. No attention was paid to the n-butyric acid groups.

The strong impression left by these trials, and others like them,
was that oleic acid, and to a lesser degree caproic acid stimulate in
Myrmecia vindex both a burying reaction and sometimes the trans-
port of objects so contaminated to refuse middens. No evidence of
overt digging was seen. Formic acid seemed without effect. Tri-
ethanolamine, however, functioned as an efficient excitant and at-
tractant in higher concentrations. In lower concentrations it stimu-
lated necrophoric behavior. The “conflict” behavior exhibited in
the handling of some larvae and cocoons treated with several of these
substances, when immatures were repeatedly taken into the nest,
brought out again and discarded on a midden, then returned to the
nest, sometimes through three or four cycles, was in several cases
striking.

Field Tests

Reactions of workers of Myrmecia vindex to Oleic acid ;

Caproic acid ; Formic tcid ; Methylbenzylamine
It was of interest to conduct essentially similar tests with wild
colonies in the field. A large and active colony of M. vindex in
Kings Park, Perth, West Australia (a part of the same population
from which the colonies had been taken for laboratory tests), was
selected. It included approximately 200 workers, had a single large
entrance-hole, and a well-defined crater. Since workers of this
population show a predominantly crepuscular and nocturnal foraging
pattern in warm weather, tests were begun in the evening.

At 7:25 p.m., as dusk was gathering, 5 white cards, of dimensions
3” X 5", were mounted on the nest crater, approximately equi-
distant from the entrance. Twenty-five cocoons, obtained from a
neighboring colony, were impregnated with 3 drops of test substance,
with 7 retained as untreated controls. The arrangement and treat-
ments were as shown below:

1974]

Haskins & Haskins — Myrmecia vindex

265

Card Marking

No. of Cocoons

Substances Impregnated

C

7

Caproic acid

F

6

Formic acid

M

7

Methyl benzylamine

O

5

Oleic acid

L

7

Control

Immediately on exposure a worker on the crater seized one control
and dragged it inside the nest. A second worker, encountering an
oleic-acid-treated cocoon, started away. At 7 132, 1 caproic-acid-
treated cocoon was dragged into the nest, followed at 7 :35 by a
second control cocoon. At 7 :37 a cocoon from the formic acid group
was taken to the nest. Between 7:37 and 7:40, 3 additional formic-
acid-treated cocoons were taken into the nest, and one-half minute
later the last control cocoon was taken in. One of the remaining
two formic acid cocoons was inadvertently toppled into the nest
by another worker, leaving 1 on the card. The other groups were
untouched. At 8:02 p.m., when darkness forced cessation of observa-
tion, all cocoons treated with oleic acid, methylbenzylamine, and five
of those with caproic acid remained in place on the cards. One
cocoon treated with caproic acid had been deposited with the oleic-
acid-treated group.

At 6:00 a.m. the following day, 3 of the caproic-acid-treated co-
coons had been removed, but it was not possible to determine whether
they had been taken into the nest or discarded. Some grains of
earth had been scattered on the caproic-acid-treated card.

Observations were continued throughout the day, but no further
attention was paid to the remaining cocoons. However, when the
cards were removed from the crater at 7 :oo a.m. on the third day,
it was found that a new entrance-hole had been excavated under
the card carrying the oleic-acid-treated cocoons.

It was clear that oleic acid, caproic acid, and methylbenzylamine
effectively inhibited return of the cocoons to the nest by the workers,
in marked contrast to those treated with formic acid and the controls,
which were brought in promptly. Digging behavior was observed in
the vicinity of oleic acid. In these experiments, however, the oleic
acid objects were not buried nor transported to kitchen middens — -
perhaps because such middens of M. vindex are often located near
the nest entrance. Indeed, it is an interesting behavioral characteristic
of M. vindex , at least in this population that, at the seasons of
maximum brood production, empty cocoons and pupal exuviae are
frequently piled near the nest entrance. That this disposition may
be more than accidental is suggested by the behavior of some colonies

266

Psyche

[June

in the laboratory (not those with which the above-mentioned tests
were conducted), where such exuviae, originally scattered about a
foraging arena, may be gathered into dense craters about the entrances
to Lubbock nests.

Summary and Conclusions

In a parallel series of experiments to those reported by Wilson
(1958) and Wilson, Durlach, and Roth (1958) with Pogonomyrmex
badiuSj the effects of oleic acid, caproic acid, methylbenzylamine,
n-butyric acid, n-valeric acid, formic acid, and triethanolamine as
behavioral releasers in the ant Myrmecia vindex were investigated
both in the laboratory and the field. The comparison was deemed
particularly interesting because of the archaic character of the
Myrmeciinae and their societies.

As with Pogowomyrmex and other higher Formicid genera, oleic
acid was found to act as a releaser of both necrophoric and digging
behavior, suggesting the early establishment of this set of reaction
patterns in Formicid social evolution — a not unexpected situation
in view of the highly adaptive character of this pattern in ridding
the nests of dangerous animal decomposition products, perhaps in
response to the bacterial production of oleic and related fatty acids.
In Pogonomyrmex , n-butyric acid, n-valeric acid, and n-caproic
acid stimulated weak to moderate alarm behavior, passing over into
digging behavior. In Myrmecia , caproic acid stimulated mildly
necrophoric behavior, associated with some digging and burying
behavior. Exposure to filter papers of cocoons impregnated with
11-butyric acid resulted in deposition of earth grains on the treated
object. No reaction was observed to n-valeric acid.

Thus responses to these substances were of the same general quality
as those reported for P. hadius. There were, however, very marked
differences in the responses of M. vindex to two presumed releasing
substances: formic acid and triethanolamine. Formic acid neither

stimulated necrophoric nor digging behavior nor prevented immediate
transport into the nest of formic-acid-treated cocoons. This result
was somewhat surprising to the authors in view of the highly positive
and specific attack reactions stimulated by formic acid in Myrmecia
gulosa mentioned earlier, which have been reported elsewhere
(Haskins, Hewitt, and Haskins, 1973). The most marked contrast
between behavior patterns of P. badius and M. vindex , however,
occurred with those released by triethanolamine. With M. vindex ,
vapors of this compound consistently stimuated the most conspicuous
necrophoric behavior of any substance tested, while also acting as a

1974]

Haskins £sf Haskins — Myrmecia vindex

267

decided attractant. In P. badius on the contrary, the reaction to
triethanolamine was reported to be virtually neutral.

In this connection it is interesting to note that, in the populations
studied, M. vindex commonly piles discarded cocoons and pupal
exuviae on the crater of the nest, near the entrance, where they
characteristically accumulate in some numbers, rather than trans-
porting them far afield. Similar behavior has been described for
P. badius (Wilson, Furlach, and Roth, 1958), though in the latter
case, removal of the discards is quickly accomplished by other foraging
ants. The situation with M. vindex, however, appears more specific.
Indeed established colonies in the artificial nest may collect discarded
cocoons and pupal exuviae scattered about the foraging area to form
a crater about the entrance. It may be that subtle behavioral balances
are involved, depending on the nature and “mix” of the included
decomposition products, as suggested by the reversal of behavior,
apparently in response to vapor concentration, released by trietha-
nolamine.

References

Blum, Murray S. The chemical basis of insect sociality. From Chemicals
Controlling Insect Behavior. Academic Press, New York, 1970, pp 61-94.
Brown, William L., Jr. Remarks on the internal phylogeny and subfamily
classification of the family Formicidae. Insectes Sociaux. 1, 1, 1954,
pp 21-31.

Douglas, Athol M. and L. M. McKenna. Observations on the bull-dog
ant, Myrmecia vindex. The IV estern Australian Naturalist, 11, no. 6,
pp 125-129, 1970.

Haskins, Caryl P., Richard E. Hewitt, and Edna F. Haskins. Release
of aggressive and capture behavior in the ant Myrmecia gulosa by
exocrine products of the ant Camponotus. Jour. Ent. (Royal Ent. Soc.,
Great Britain) (A), 47 (2), pp 125-139, 1973.

Haskins, Caryl P. and Edna F. Haskins. Notes on the biology and social
behavior of the archaic Ponerine ants of the genera Myrmecia and
Promyrmecia. Ann. Ent. Soc. of Am. 43, 4, December, 1950, pp 461-491.
Robertson, Phyllis L. Pheromones involved in aggressive behavior in the
ant Myrmecia gulosa. Jour. Insect Physiol. 17, 1971, pp 691-715.
Wilson, Edward O. A chemical releaser of alarm and digging behavior
in the ant Pogonomyrmex badius (Latreille). Psyche 65, 2-3, June-
September, 1959, pp 41-51.

Wilson, Edward O. The Insect Societies. Harvard University Press,
Cambridge, Mass. 1971, pp 31-32. (Origin of Ants, Sphecomyrma and
Myrmecioid complex).

Wilson, Edward O., Frank M. Carpenter, and William L. Brown, Jr.

The first Mesozoic ants. Science, 15 7, September, 1967, pp 1038-1039.
Wilson, Edward O., N. I. Durlach, and L. M. Roth. Chemical releasers
of necrophoric behavior in ants. Psyche, 65, 4, December, 1958 pp
108-114.

THE FIRST RECENT SPECIES OF PROTOMUTILLA
(HYMENOPTERA: MUTILLIDAE ; MYRMOSINAE)1

!: By Denis J. Brothers2

During a recently completed study of the higher classification of
the aculeate Hymenoptera (Brothers, 1974), my attention was drawn
to a female specimen from India representing a species of Myrmo-
sinae (considered a subfamily of Tiphiidae by many authors, e.g.,
Krombein, 1940) but showing marked similarities to Nanomutilla
(a member of the Mutillidae often included in the Myrmillinae) .
This situation is reminiscent of that found by Bischoff (1915) for
seven female specimens in Baltic amber. All of these have the well-
developed pro-mesonotal suture characteristic of the Myrmosinae,
but in addition “die Tiere weisen Beziehungen zu verschiedensten
Gattungen auf, so zu Myrmilla, Nanomutilla , Mutilla s. str. etc.”
Bischoff described the genus Protomutilla to include these seven
species, and considered it best placed in the “Mutillinae”, although
intermediate between this taxon and the “Myrmosinae” (which he
considered to be a member of the Mutillidae). He listed the most
characteristic features of the genus as: the distinctly developed pro-
mesonotal suture, the transverse first abdominal (metasomal) seg-
ment, the lack of a pygidial area, the usually longitudinally striate
sculpturing of the mesonotal region, and the absence of long pubes-
cence. In addition, all of his specimens have the posterolateral angles
of the mesosoma acute or dentate. The Indian specimen possesses
all these characters except for that of the mesonotal sculpturing
(absent in some of Bischoff’s species also). The form of the postero-
lateral angles of the propodeum differentiates this genus from the
others with a well-developed pro-mesonotal suture, so that the Indian
specimen seems best considered to represent a modern species of
Protomutilla. Placement of this genus in the Myrmosinae is sup-
ported by the almost straight and well-differentiated hind margin
of the pronotum and also by the presence of a well-developed carina
dorsally on the hind coxa (a character not mentioned by Bischoff),
although this is not as markedly lamellate as in most other myrmosines.

Contribution number 1538 from the Department of Entomology, University
of Kansas, Lawrence, Kansas 66045.

2Present address: Department of Entomology, University of Natal, Pie-
termaritzburg, South Africa.

Manuscript received by the editor January 21, 1974.

268

1974]

Brothers — Protomutilla

269

Figs. 1-2. Protomutilla microsoma, sp. nov., $. 1, dorsal view, legs

omitted; 2, head, lateral view, antenna and palpi omitted. Fig. 3. Nano-
mutilla microsoma Andre, 9 , dorsal view, legs omitted. Scale — 1.0 mm.

270

Psyche

[June

Protomutilla microsoma sp. nov.

(Figs. 1-2)

female : Length 2.6 mm. Body finely and evenly punctate with
sparse, short, pale pubescence ; black except clypeus, antennal tubercle,
antenna, mandible, ventral third of pronotum laterally, legs, fifth
and sixth metasomal segments and apical margins of other metasomal
segments testaceous, and palpi stramineous. Head almost prognathous,
1. 1 times as wide and 0.7 times as high as long, 1.1 times as wide
as mesomoma; sides weakly converging behind eye, about 0.3 times
length of eye ; ocelli absent ; eye broadly oval, with many short setae ;
frons evenly swollen, produced between barely differentiated antennal
tubercles; malar space 0.4 times as long as eye; weak genal carina
present; clypeus flattened, anteriorly produced with strongly convex
apical margin; antenna with scape 3.0 times as long as wide, pedicel
about as long as first flagellar segment which equals second in length,
flagellum widest at third segment; mandible approximately parallel-
sided, obliquely tridentate, the apical tooth strongest. Mesosoma
1.4 times as long (excluding anterior collar) as wide; width at
humeral angle 0.9 times that at propodeal angle. Pronotum (ex-
cluding collar) 0.3 times as long as wide; hind margin shallowly
concave; lateral margins slightly convergent posteriorly; humeral
angle pronounced. Mesonotum approximately parallel sided, 0.9 times
as wide as pronotum. Propodeum with lateral margins divergent
posteriorly, forming a strong dentiform angle on each side; disc and
declivity barely differentiated. Mesosomal pleura differentiated from
nota by a longitudinal carina; concave except convex posteriorly
forming propodeal angle. Legs unmodified ; foretarsus without spines
(i.e., pecten undeveloped) ; hind coxa with obtuse lamellate tooth
dorsally ; hind tibia without spines externally, only a very few minute
spines apically. Metasoma about 1.7 times as long as wide, 1.25 times
as wide as mesosoma; first tergum 0.4 times as long as wide, about
as wide as and 0.4 times as long as second ; sixth tergum sparsely
punctate, without differentiated pygidial area; first sternum with
single blunt anteriorly-directed midventral tooth.

male: Unknown.

holotype: ?, [India, Orissa Prov.], Nilgiri Hills, 3500 ft.
(H. L. Andrewes) ; in Hope Department, University Museum,
Oxford, England. (No date of collection given.) The specimen
is unfortunately damaged, and lacks the apical three segments of
the left foretarsus, the left mid- and hind and right mid-tibiae and
tarsi; the left antenna is glued to the card bearing the “minuten”

1974]

Brothers — Protomutilla

271

pin. Since the specimen has been remounted from a glued card
mount, some features of the venter and appendages remain somewhat
obscured by glue.

Although Bischoff’s (1915) descriptions are very short, P. micro-
soma seems clearly to differ from all the amber species. It may be
most similar to P. nana, but the head undoubtedly differs in form
from that species.

The resemblance of P. microsoma to Nanomutilla (Fig. 3) is
quite remarkable, the most basic difference being the form of the
pro-mesonotal suture. In addition to gross body form, these species
have highly similar coloration and puncturation, and all are rather
small in size. Nanomutilla even has a weakly developed carina on
the hind coxa as well as pubescent eyes, characters lacking in the
higher Mutillidae. These marked similarities tend to support my
conclusions (elaborated in Brothers., 1974) that the Myrmosinae
should be considered a subfamily of Mutillidae.

Acknowledgements

I should like to thank Mr. Christopher O’Toole of Oxford for
bringing this specimen to my attention, and also Mr. W. L. Overal
of the Univesitv of Kansas for his help with preparation of the
manuscript.

References

Bischoff, H.

1915. Bernsteinhymenopteren. Schrift. Phys.-Oekon. Ges., Konigsberg
56: 139-144.

Brothers, D. J.

1974. Phylogeny and classification of the aculeate Hymenoptera, with
special reference to Mutillidae. Univ. Kansas Sci. Bull, (sub-
mitted).

Krombein, K. V.

1940. Studies in the Tiphiidae (Hymenoptera Aculeata). IV. A re-
vision of the Myrmosinae of the New World with a discussion
of the Old World species. Trans. American Ent. Soc. 65: 415-
465, pi. 24.

LIFE HISTORY OF ABEDUS HERBERTI IN CENTRAL
ARIZONA (HEMIPTERA: BELOSTOMATIDAE)* *

By Robert L. Smith
Department of Zoology
Arizona State University
Tempe, Arizona 85281

Introduction

Torre Bueno in 1906 noted the meagerness of information on the
biology and immature stages of aquatic Hemiptera. Although this
condition has changed somewhat in the more than 60 subsequent
years, complete life histories for representatives of only two of the
five genera belonging to the subfamily Belostomatinae (Lauck and
Menke 1961) are available. Belostoma is represented by B. flu-
mineum Say (Torre Bueno 1906), B. oxyurum (Dufour), B. bi-
foveolatum Spinola (Schnack 1971), and B. malkini Lauck (Cullen
1969). The Old World genus Limnogeton is represented by L.
fieberi Mayr (Voelker 1968). Menke (i960) was successful in
bringing a single individual Abedus dilatatus (Say) to its fifth instar,
at which time it died. Life histories for representatives of the genus
Lethocerus include: L. americanus (Leidy) (Rankin 1935), L. cor-
dofanus Mayr (nec. L. niloticus Stal) (Tawfik 1969), L. mazzai
De Carlo (De Carlo 1962), and L. maximus De Carlo (Cullen
1969). Nothing is known of the immature stages of 1 Horvathinia
(subfamily Horvathiniinae, Lauck and Menke 1961). The tax-
onomy and distribution of Abedus herberti , as well as a behavioral
and ecological sketch of the genus, is provided by Menke (i960).

Methods and Materials

I collected adult bugs, including males encumbered with eggs, with
the aid of a gasoline lantern at night from the following central
Arizona localities: Sycamore Creek, Maricopa County, near Sun-
flower; Camp Creek, Maricopa County, near Carefree; and Tule
Creek, Yavapai County, in the Bradshaw Mountains.1 Plastic bags
containing stream water isolated individuals for transport to the

Henke’s 1960 revision recognizes polytypy in A. herberti. The central
Arizona specimens studied in this paper belong to “subspecies herberti ”
in contradistinction to “subspecies utahensis.,y

* Manuscript received by the editor July 1 , 1974.

272

1974]

Smith — Life History of Abedus herberti

273

laboratory. Plastic washbasins (15 by 26 by 30 cm) containing clean
gravel and deionized water housed bugs in the laboratory. Finger
bowls (10.5 by 4.5 cm), provided with aquarium gravel and deion-
ized water to a depth of about 4 cm, received hatchlings obtained
from eggs of field-captured encumbered males. Gravid females,
paired with unencumbered males, produced fresh eggs for incubation
studies. First and second instar nymphs were fed on cultured vesti-
gial winged Drosophila which were floated on the water’s surface.
Third and fourth instars received legless crickets ( Acheta domestica) ,
and fifth instars ate intact crickets. I renewed food and water every
other day, and recorded molts which had occurred each day. Cast
skins were preserved in seventy percent alcohol, one percent glycerin
for study and measurement. Open water temperature in the labor-
atory was a reasonably constant 18 C (± 0.5 C).

I observed hatching and used still and motion picture photography
to record this event for analysis. I measured eggs, first, second, and
third instar nymphs and the legs of fourth and fifth instars using a
stereoscope ocular micrometer, and the bodies of fourth and fifth
instars with a dial caliper.

Eggs, Incubation, and Hatching

eggs

The eggs of Abedus are always laid on the back of the male
(Fig. iA). Oviposition, which continues for 12-48 hours in the
laboratory, alternates with copulation. Encumbered males are found
throughout the year in Arizona. A detailed study of courtship and
male brooding behavior in A. herberti will be presented elsewhere
(Smith, in prep.) .

A mucinous glue is secreted by the female prior to the deposition
of each egg starting at the apex of the male’s hemelytra and pro-
ceeding forward, uniformly covering the male’s back up to and some-
times including the pronotum. It is usually the case that males carry
the eggs of only one female, but polygynous matings apparently do
occur (Menke i960). I inferred from data on hatching, that of
seven encumbered field captured males, only one individual had con-
sorted with two females to acquire his load of eggs. In this instance,
72 eggs hatched over a five day period, this followed by three days
during which no eggs hatched. The remaining 15 eggs hatched over
a 24 hour period. In all other cases, hatching occurred more or less
continuously over a three to five day period.

Freshly laid eggs begin swelling immediately. Swelling has the

274

[June

Psyche

Figure 1. Encumbered male Abcdus herberti and hatching of eggs.

A. Encumbered male Abedus herberti.

Inset. Enlargement of hatching nymphs.

B. Rupture of the chorion around cap.

C. Resting position.

D. Newly emerged nymph.

1974]

Smith — Life History of Abedus herberti

275

ultimate effect of compacting the mass, enhancing its structural in-
tegrity and conformity to the male’s back. The resulting gelatinous
plate with its embedded eggs is strong, flexible and adheres tena-
ciously to the male’s hemelytra so long as it is kept moist. If exposed
to air for 60 minutes or longer, the gelatinous pad becomes brittle,
loses its adhesive properties and on prolonged exposure will eventu-
ally fall off.

Eggs increase in size from mature oviducal (3.064 db .022 mm
by 1.732 ± .017 mm, n = 50) to those containing nymphs ready
to hatch (4.983 =±= .036 mm by 2.001 d= .017 mm, n = 50). This
represents a 59 percent mean increase in length and a 15.5 percent
mean increase in width. The initial swelling of the egg (during the
first 48 hours) may be due to imbibition of water, however later
increase in size seems to be due to the development of the embryo.
Both Hungerford (1925) and Tawfik (1969) noted dramatic in-
creases in egg size during the development of Lethocerus eggs which
are laid above the water on emergent vegetation. Cobben (1968)
provides a synthesis of what is known about belostomatid eggs and
embryology.

Mature oviducal and freshly laid eggs are variable in shape,
usually imperfect oval with one nearly straight side, yellow in color
and irregularly hexagonal microreticulate. Soon after occlusion, eggs
darken to tan. The cap (upper 3/3) differentially darkens and re-
mains several shades darker until just prior to eclosion of the nymph
at which time it becomes grayish white in color. As development
proceeds the shape of the egg comes to resemble an elongate “printed
comma” ; that is broad and rounded at the free end, narrow and
pointed at the attached end, with one convex and one concave side.
The concave side seems always to face the posterior of the mass.
Cobben (1968, Fig. 232) shows the head of the embryo (of Sphae-
rodema — Diplonychus ) facing the attached end, but embryonic
rotation must occur prior to hatching. I observed one instance of
egg failure due to nonrotation of the embryo.

INCUBATION

Male brooding of eggs, a complex of behaviors, is necessary for
achievement of the greater than 95 percent survival observed for
eggs of field-captured encumbered males. I never observed any type
of egg parasitism. Egg failure, in every case but the one, seemed due
to insemination failure; usually a few eggs on each plate did not de-
velop. Open spaces in the laying pattern in Figure iA are undevel-
oped eggs. Harvey (1906) reported that infertile eggs of Pedino-

2j6

Psyche

[June

1974] Smith — Life History of Abedus herberti 277

Figure 2. Individual molt records for 31 laboratory reared nymphs of
A. herberti. Open circles indicate molts. Closed circles indicate the death
of that nymph.

coris macronyx (= Abedus indentatus Haldeman) drop off and are
replaced (presumably by the original or another female) by others
as late as the sixth or eighth day of incubation, but in every instance
I observed, nonviable eggs were retained in the nidus through incu-
bation and hatching of the viable eggs. Incubation periods from lay-
ing of the first egg to hatching of the first ranged from 21 to 23
days in the laboratory. Voelker (1968) has shown developmental
time for eggs of Limnogeton fieberi to be directly correlated with
temperature, the optimal temperature being 32-33 C. He reports an
8 day development time at optimal temperatures, which seems to be
more in agreement with the results obtained by the majority of other
workers for belostomatids (Torre Bueno 1906, Harvey 1906, Hun-
gerford 1925, and Tawfik 1969). Menke (i960) found the incu-
bation time for A. dilatatus to exceed 30 days in the laboratory, but
we both (Menke, personal communication) suspect that development

278

Psyche

[June

of eggs for Abedus could be substantially accelerated by increasing
the temperature of the incubation water. Open water temperature
in my laboratory (18 C) fell within the range of spring, fall and
winter stream temperatures in Arizona.

HATCHING

Twenty-four hours prior to hatching, the free end of the egg
swells dorsally. Pressure from within eventually causes a rupture in
the chorion around the cephalic cap (Fig. iB). Invariably, nymphs
hatch facing the brooding male’s posterior. I can’t explain this con-
stant orientation of hatching nymphs, but its effect is that adjacent
eggs can hatch simultaneously with a minimum of interaction be-
tween eclosing nymphs. The cap is lifted by the head as the nymphal
thorax emerges, but remains hinged to the egg on the side of the
latter which faces the ventral surface of the emerging nymph. The
nymph’s head slips from under the operculum as peristaltic contrac-
tions free one half of the body. When the nymph is two thirds re-
moved, it rests in a position perpendicular to the plane of the egg
plate (Fig. iC). Moments later it arches backward and stretches its
legs which are employed in final extraction when the nymph again
leans forward. A newly emerged bug generally lingers on the back
of the male for several minutes to inflate its laterally constricted
abdomen (Fig. iD). This accomplished, it moves off in search of
cover among aquatic plants or debris where it assumes the predatory
stance characteristic of nymphs and adults alike. Freshly emerged
nymphs are light honey yellow in color and translucent, but darken
within one hour.

Nymphal Development and Nymphs

DEVELOPMENT

Of 31 nymphs reared in the laboratory, 26 survived to adulthood.
Individual developmental records are illustrated in Fig. 2 and mean
developmental periods are presented in Table 1. These results com-
pare favorably with those of Voelker (1968) for Limnogeton and
probably reflect critical minimum periods required for morphogenesis
between molts under optimal conditions. That availability of food
is an important factor related to intermolt time for these sedentary
hunters was demonstrated in a simple experiment. Ten of 20 sibling
first instars, fed Drosophila daily, all molted in under eight days
while the other ten, fed every fourth day, did not begin to molt until
the nineteenth day. Thirty additional sibling first instars were di-

1974]

Smith — Life History of Abedus herberti

279

30 -

25-

10 20 30 40 50 60

MEAN MOLT TIME (days)

Figure 3. First through fifth instar nymphs of A. herberti. Mean length
and mean duration of instars illustrated graphically.

vided into three groups of ten individuals. Each group was kept at a
different temperature (18 C, 22 C, and 31 C) and all nymphs re-
ceived Drosophila daily in numbers exceeding that which they could
consume. All molted at approximately the same time (on the sixth
or seventh day), suggesting that temperature is relatively unimpor-
tant. These experiments were not intended to be conclusive and it
is probable that a food-temperature interaction could be detected
with more sophisticated experimentation.

NYMPHS

There are five nymphal instars for A. herberti. Dorsal views of
the five are figured (Fig. 3) and mean morphometric data for fifteen
characters of each instar are presented in Table 2. The wide (.001)
confidence limits around each mean should encompass interpopulation
variation for central Arizona. Mean size of bugs seems to be stream
specific, perhaps due to low dispersal of these animals and reduced
gene flow between their populations. A. h. herberti intergrades with

Table 2. Confidence limits* on mean nymphal instar morphometry of Abedus herberti.

280

Psyche

NO o N no OO
OO ON NO

rH

C^ tO ^ On
i— 1 vo ON 1— 1
ON NO CM y-4

00 to 00 O
to O to
00 to o

+1 +1

o 00
CM
ON

On no
CM i-H

+1 +| +1 +1

ON ,4- 00 O

CM — ^ CM 1-H

cm 00 *0

+1 +1 +1 +1

t-H 00 NO

+1 +1 +1 +1

o CM NO CM
On On no vo
^ ON ON CM
o © to t-5

o o o mo
NO U-, O to
to ^ o

OO N CM +
vo 10 O0 NO
no CM O

NO no o OO
N CM ^ ON
NO ON tO O

+1 +1 +1 +1 +| +1 +1 +1 +1 +1 +1 +1 +1 +1 +1

NO tj- OO
no 00 CM O
to 10 O NO

to O NO

00 CM o to

00 ON ©

NO rt- CM

NO NO to 00

O O NO

to to s 00
t'' CM

©

VO O 00 NO
to CM H o

CM NO vo CM
to h 00 N
no tj- CM O

+1 +1 +1 +| +| +| +| +| +| +| +| +| +| +| +|

NO O
On On

i-H 00 CM NO

y-i ON CM 1-1

On NO OO rj-

to CNJ

OO OO NO tO
N OO CM 00
i— < VO tt" NO

-h to 1—1

On to -t NO
00 On

v-J <M O NO
vo vo CM

•H- to
00 CM

ON NO

t-- 1 — 1 CM — J-

H tO OO O
CM 00 O 1— 1

to CM O vo

to *-)- i— 1 UN

CM i-H h o

CM O to

1-1 CM r*- to
to CM 1-1 O

+1 +1 +1 +1 +1 +1 +| +1 +1 +1 +1 +1 +1 +1 +1

NO Ti- no

© <o o

On ON vo vo

00 00

NO 00
CO CM

t''. *t-

UN NO
00 !>.

H NO tJ" o
On ts -t un
O 00 O O

1— I H NO NO

i- 1 to CM CM

H i-l y* ©

CM VO NO ON
N N N if
1-1 CM O O

+1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1 +1

NO rj- CM OO

VO NO 1— I ON

It vo NO CM

•f NO NO *t-

O ^ 1-t o

-t- O On to

CNJ CM

N N t t

O N H rf
On ON CM to

b D -a

ji £

3

s 3

t* h H u

3 .2 M k.
S -Q J: a
aj y y —

fe H H u

I .2 « *

C _Q CJ ct
i> y — <

fe H E-1 u

>1

-o

o

CO

fa/D

[June

I

^Confidence level = 99.999 around means of measurements from 10 laboratory reared specimens.

1974]

Smith — Life History of Abedus herberti

281

a smaller subspecies A. h. utahensis in northern Arizona and south-
ern Utah.

The first through the fourth instars all have two segmented an-
tennae and one segmented pro-, and two segmented meso- and meta-
tarsi. The beak is four segmented from the first instar to imago.
One antennal segment is added in the fifth instar and usually another
in the adult. All tarsi add one segment in the adult molt.

The most noteworthy changes through instars are the allometric
development of the wing pads (Fig. 3) and of the protarsal claws.
Torre Bueno (1906) states that there is a progressive diminution in
size (through the instars) of the anterior (external) protarsal claw
of Belostoma flumineum and that the external claw finally disappears
in the adult. This statement is erroneous on at least one count, but
nevertheless focuses attention on an ontological aspect of phylogenetic
significance. It is now known (Lauck and Menke 1961) that both
Belostoma and Abedus have vestigial external protarsal claws. The
other three genera in the Belostomatinae have two well developed
protarsal claws as does the genus Lethocerus. In Abedus herberti and
probably Belostoma as well, only the magnitude of the size difference
between the external and internal protarsal claws increases with each
molt due to nondevelopment of the external claw and continued
growth of the internal. In the first instar relative length of the ex-
ternal claw is 50 percent of the internal. This is reduced to about
38 percent in the second, 30 percent in the third, 23 percent in the
fourth, and 19 percent in the fifth instar. Curiously, in the adult
the external is proportionally slightly longer than in the fifth instar
and the internal claw is shorter and broader in the adult. This re-
sults in an increase in the relative length of the external claw to
about 28 percent that of the internal in the adult bug.

Longevity

A. herberti is a relatively large animal and is a sedentary hunter,
both of which characteristics suggest that it might be long lived. This
seems to be the case; several specimens have lived in the laboratory
for up to one year and one in particular, a female collected on Octo-
ber 14, 1972, lived until November 30, 1973 during which time she
produced four sets of eggs totaling 344. Nymphs from the first
through the third instars are probably subject to predation by other
aquatic organisms. I have found first instar nymphs in the stomachs
of trout taken at three forks of the Black River, Apache County,
Arizona. Nymphal instars seem to play an important role in the

282

Psyche

[June

food economy of the species during periods of low stream produc-
tivity; I commonly find nymphs and adults feeding on smaller
nymphs of their own species. After the second instar, cannibalism
seems to be far the most common cause of fatality among nymphs.
Ecdysal failure and physical catastrophes (flood and drought) also
regulate populations. Lethocerus adults, which are seasonally common
but never abundant in Sonoran streams, may regularly take adult
Abedus but I have observed only one instance of this. In general,
A. herberti adults are at the pinnacle of the aquatic food chain, and,
having reached adulthood, stand an excellent chance of surviving to
produce or brood several clutches of eggs.

Discussion

The life history of Abedus herberti seems to follow closely the
pattern reported for other genera belonging to the subfamily Belo-
stomatinae. Allometric development of the protarsal claws closely
relates Abedus to Belostoma, and suggests their common ancestry
with the two clawed genera. Five instars is the number characteris-
tic of acquatic Hemiptera (Hungerford 1920).

Acknowledgements

I wish to thank Dr. M. Parsons, Dr. A. S. Menke, and Dr. J.
Alcock for review.

Literature Cited

Cobben, R. H.

1968. Evolutionary trends in Heteroptera. Part I Eggs, architecture
of the shell, gross embryology and eclosion. Center for Agricul-
tural Publishing and Documentation, Wageningen, Netherlands.
Pp. 206-209.

Cullen, M. J.

1969. The biology of giant water bugs (Hemiptera: Belostomatidae)
in Trinidad. Proc. R. Ent. Soc. Lond. (A) 44: 123-137.

De Carlo, J. M.

1962. Consideraciones sobre la biologia de Lethocerus mazzai De Carlo.
Physis, 23 : 143-151. •

Harvey, G. W.

1906. A ferocious water-bug. Proc. Ent. Soc. Wash., 8: 72-75.
Hungerford, H. B.

1920. The biology and ecology of aquatic and semiaquatic Hemiptera.

Kansas Univ. Sci. Bull., 11: 1-328.

1925. Notes on the giant water bugs. Psyche, 32: 88-91.

1974] Smith — Life History of Abedus herberti 283

Lauck, D. R. and A. S. Menke

1961. The higher classification of the Belostomatidae (Hemiptera).
Ann. Entomol. Soc. America, 54: 644-657.

Menke, A. S.

1960. A taxonomic study of the genus Abedus Stal (Hemiptera, Belo-
stomatidae). Univ. Calif. Pub. in Entomol., 16: 393-440.

Rankin, K. P.

1935. Life history of Lethocerus americanus (Leidy) (Hemiptera, Belo-
stomatidae). Kansas Univ. Sci. Bull., 15: 479-491.

Schnack, J. A.

1971. Las ninfas del genero Belostoma Latreille (Hemiptera, Belosto-
matidae) (I) Belostoma oxyurum (Dufour) y Belostoma bifoveo-
latum Spinola. Rev. Soc. Ent. Arg., 33: 77-85.

Tawfik, M. F. S.

1969. The life history of the giant water-bug Lethocerus niloticus Stael
(Hemiptera: Belostomatidae). Bull. Soc. Ent. Egypte, 53: 299-
310,

Torre Bueno, J. R.

1906. Life history of North American water bugs. I. Life history of
Belostoma flumineum Say. Can. Ent., 38: 189-197.

Voelker, J.

1968. Untersuchungen zu Ernahrung, Fortpflanzungsbiologie und Ent-
wicklung von Limnogeton fieberi Mayr (Belostomatidae, Hemip-
tera) als Betrag zur Kenntnis von natiirlichen Feinden tropishen
Siisswasserschnecken. Entomologishe Mitteilungen, 3 (60): 1-24.

PREY CAPTURE BY DKYMUSA DINORA
(ARANEAE, SCYTODIDAE)

By Carlos E. Valerio*

Departamento de Biologia.

Universidad de Costa Rica

Introduction

The family Scytodidae includes at present three genera, of which
Scytodes and Loxosceles are well known because of their specialized
prey-capturing strategies. Scytodes species eject a sticky substance
(perhaps similar to the silk from the spinnerets) from the chelicerae
at a considerable distance to trap the prey (Bristowe, 1931; Mc-
Alister, i960). The species of Loxosceles have developed a very
effective venom capable of subduing strong prey almost instantly
(Hite et al ., 1966). This venom affects even vertebrate tissues,
including those of man (Bucherl, 1961).

The genus Drymusci, a small and poorly studied group, is morpho-
logically more closely related to Loxosceles than to Scytodes. It
lacks the high carapace, and possesses a colulus ; also the male bulbus
is located at the tip of the tarsus (Valerio, 1971). The forest-dwell-
ing species of D. dinora Valerio, which lives exclusively under logs
utilizing crevices and horizontal tunnels in the decomposed wood
(Valerio, 1971), exhibits highly specialized behavioral patterns never
observed in other spiders. The permanent web, composed of a few
tangled threads, seems to alert the spider to the presence of prey and
to restrict the movement of prey. Clearly, this type of construction
represents a very primitive conditon in the phylogeny of the web
(Kaston, 1966).

Materials

Several mature and immature specimens of D. dinora , of both
sexes, were collected in a wet lowland forest in southwestern Costa
Rica and kept individually isolated in 12-dram vials ( 100 X 22 mm),
at IOO percent humidity and 24.5 zb 0.2°C.

Observations

This species is remarkable in two aspects of its attack behavior,
departing from all known patterns: for large prey the spider spins

*1 wish to thank Dr. Herbert W. Levi (Museum of Comparative Zoology,
Harvard University) for his critical review of this manuscript.

Manuscript received by the editor January 5, 1974.

284

1974]

Valerio — Drymusa dinora

285

a trap after the prey’s arrival in the web, and prey-wrapping is car-
ried out exclusively by movements of the abdomen and without using
the appendages. Prey is treated in a different manner according to
size and perhaps other qualities, as also occurs in other groups of
web-building spiders (Eberhard, 1967; Robinson, 1969; Shear,
1969).

Attack on small prey: The spider rests in the center of the web
( B in figure 1 ) near the superior edge of the crevice or tunnel.
When small prey (less than % the size of the spider) enters the
tangled threads, the spider moves and attacks directly with the cheli-
cerae and holds on firmly until the prey stops moving. Usually the
prey is then carried to the resting site and feeding starts immediately
without previous wrapping in silk. The arrival of a second item of
prey does not elicit a response from the spider.

Attack on large prey: When large prey penetrates at one side
(C in Figure 1), the spider runs to A and starts immediately spin-
ning a horizontal partition. The prey then moves through the
tangles of the center towards the trap web. If the prey cannot cross
this barrier of dry silk and starts heading back, the spider moves
ahead of it to C where it builds another vertical web, thus enclosing
the prey in a silken trap. Then, the direct attack begins. The spider
approaches its victim with certain caution and suddenly strikes five
or six times with the chelicerae at intervals of one second. Some-
times, some chasing is involved. After the envenomation the prey
may move around the web but the spider usually ignores it. Once
the prey slows down (apparently due to the effect of the venom), it
is wrapped in silk.

The silk is distributed by oscillatory movements of the whole body,
reinforced by more pronounced side movements of the abdomen (with
conspicuous flexions of the pedicel), changing position at intervals
to deliver silk to different parts around the prey. No appendages
(other than the spinnerets) are involved in the process. The prey-
is carried in the chelicerae to the upper portion of the web where
wrapping continues for a few seconds. This post-immobilization
wrapping seems to facilitate transportation of the prey to the resting
site and attachment to the web for later feeding (Robinson et ah,
1969). Once the prey is wrapped, the spinnerets are carefully
cleaned by back and forth movements of the distal third of the
fourth metatarsus. Later, the fourth metatarsi, are, in turn, cleaned
by the chelicerae.

Very large or very strong prey items entering the web do not
produce an aggressive response from the spider. It simply lies flat

286

Psyche

[June

Figure 1. Spider Drymusa dinora in the resting area (B) in the center
of the web. A trap is built at site A after prey enters through site C;
later a second trap web is built at C to corral prey.

against the substrate while the prey passes through the area and
moves on.

Females carry the egg-sac in their chelicerae much like the species
of the genus Scytodes (and the structure of the sac itself resembles
that of Scytodes also). The sac is temporarily abandoned when a
suitable prey enters the web.

Discussion

In the attack on small prey the species behaves like the very
primitive spiders, including their relatives of the genus Sicarius
(Sicariidae) (Levi, 1967), attacking and subduing the prey solely
by the use of the chelicerae (Eberhard, 1967).

Large prey is caught by trapping webs and is subdued by biting,
but neither holding nor wrapping is involved in the immobilization
process. The trapping is a remarkable adaptation to the species’
habits, since the web is frequently exposed to prey too large to be
captured (e.g., passalid beetles). An extensive capturing web, often
destroyed without reward for the spider, would represent a significant
loss of energy (through the production of silk).

During the post-immobilization wrapping the spinnerets are ap-
plied directly to the prey in a fashion similar to that observed in the
diguetids (Eberhard, 1967).

The capturing behavior of Drymusa dinora suggests the presence
of an effective venom mdicating a closer relationship with Loxosceles.

1974]

Valerio — Drymusa dinora

287

The three genera in the family Scytodidae share in common the
small size of the permanent web and the specialized technique for
subduing of prey.

Conclusions

The species should be considered very primitive since no wrapping
is involved in the immobilization of prey. There seems to be a
tendency for the economy of silk through the reduction of the perma-
nent web and the overcoming of small prey without the use of trap
webs.

These behavioral observations, along with the morphological evi-
dence, indicate that it might be best to keep the three genera
( Drymusa , Loxosceles and Scytodes) within one family, the Scyto-
didae.

References

Bristowe, W. S.

1931. Spitting as a means of capturing prey by spiders. Ann. Mag. Nat.
Hist. 10(8) : 469-471.

Bucherl, W.

1961. Aranhas do genero Loxosceles e loxoscelismo na America.
Ciencia y Cultura 13 (4): 213-224.

Eberhard, W.

1967. Attack behavior of diguetid spiders and the origin of prey
wrapping in spiders. Psyche 74(2): 173-181.

Hite, J. M., Gladner, W. J., Lancaster, J. L., and Whitcomb, W. H.

1966. Biology of the brown recluse spider. Bull. Agr. Exp. Sta. 711:
1-26.

K ASTON, B. J.

1966. Evolution of the web. Nat. Hist. 75 (4): 27-32.

Levi, H. W.

1967. Predatory and sexual behavior of the spider Sicarius (Araneae:
Sicariidae). Psyche 74(4): 320-330.

McAlister, W. H.

1960. The spitting habit in the spider Scytodes intricata Banks. Texas
J. Sci. 22(1-2) : 17-20.

Robinson, M. H.

1969. Predatory behavior of Argiope argentata (Fabricius). Am.
Zoologist 9: 161-173.

Robinson, M. H., Mirik, H., and Turner, O.

1969. The predatory behavior of some araneid spiders and the origin
of immobilization wrapping. Psyche 76(4): 487-501.

Shear, W. A.

1969. Observations of the predatory behavior of the spider HyPochilus
gertschi Hoffman. Psyche 76(4): 407-417.

Valerio, C. E.

1971. The spider genus Drymusa in the New World (Araneae,
Scytodiadae) . Florida Ent. 54(2): 193-200.

A NEW COCKROACH GENUS (GURNEY A)
PREVIOUSLY CONFUSED WITH PINACONOTA
(BLABERIDAE: EPILAMPRINAE ) .*

By Louis M. Roth
Pioneering Research Laboratory
U.S. Army Natick Laboratories
Natick, Massachusetts 01760

Princis (1967) listed 2 species of Pinaconota , viz. bifasciata (Saus-
sure) and obliqua (Walker). Ischnoptera (?) sicca Walker was
synonymized by Kirby (1904) with bifasciata, but I (1973a) showed
that Kirby was incorrect in his interpretation of the species and
placed sicca in the new genus Alphelixia. The male genitalia and
other morphological characters of obliqua indicate that this species
is not a Pinaconota and a new genus is erected for it (see below).

In describing Pinaconota, Saussure (1895) simply stated that the
hind metatarsi were very short, with a fringe of long hairs, and a
large apical pulvillus. Shelford’s (1910) characterization of the
genus is as follows: Form depressed. Pronotum trapezoidal, an-

teriorly and posteriorly sub-truncate, deeply punctate. Scutellum
exposed. Tegmina and wings fully developed, extending beyond
the apex of the abdomen. Femora moderately spined beneath. Tarsi
very short, fimbriate, and entirely unarmed beneath ; posterior meta-
tarsus equal in length to the two succeeding joints, its pulvillus large.
Arolia very large. In this generic description, Shelford was appar-
ently influenced by his belief that Ischnoptera ( ?) obliqua Walker
was a Pinaconota.

Pinaconota can be recharacterized as follows:

Pinaconota Saussure

Type-species: Blatta bifasciata Sauss., Monotypy (1895, p. 337)

Both sexes with tegmina and wings abbreviated, or extending
slightly beyond the end of the abdomen. Ventro-anterior margin of
front femur with 2 or more, small, heavy spines, followed by a row
of small uniformly spaced slender setae, terminating in a large distal
spine; ventro-anterior margins of mid and hind femora with 2 or 3
widely spaced spines, with or without distal spines; hind metatarsus
short with 2 rows of fine ventral setae; pulvilli and arolia very

* Manuscript received by the editor January 15, 1974.

288

1974]

Roth — Cockroach Genus Gurneya

289

Figs. 1-5. Pinaconota bijasciata. 1-4. Male, from Brazil. 2. Pronotum.
3. Head. 4. Terminal abdominal segments (dorsal). 5. Male from S. Cath.,
Brazil, (scale: figs. 1 and 5 = 5 mm; fig. 2 = 2 mm; figs. 3 and 4 =
1 mm).

290

Psyche

[June

large. Supra-anal plate rounded with a mesal indentation. Cerci
short, reaching to about the hind margin of supra-anal plate. Sub-
genital plate ( cf ) asymmetrical, broadly, but weakly concave on
right side; styli about equal. Male genitalia: L2d a small irregu-
larly shaped sclerotized plate separated from L2vm ; prepuce densely
spicular in part, with or without large spines on the free margin
(Figs. 15, 17, 20, 23, 29, 30; see Fig. 29 for naming of parts), a
light sclerotized plate present ventral to the L2vm. The membrane
lying above the L2vm may have a number of small spines (Fig. 15)
but in the flattened KOH preparations are not shown in their normal
position (Figs. 20, 23). R2 unique in having a finger-like basal
projection directed toward the inner curved margin of the hook
(Figs. 16, 18, 21, 31). Right paraproct considerably more developed
than the left (Figs. 13, 27-28).

The presence of large spines on the prepuce is, as far as I know,
unique for a member of the Epilamprinae, and has only been found
in the Blaberinae. However, in that subfamily the spines usually
closely surround the L2d (Roth, 1970a). Because of the unusual
genital structures, I place Pinaconota in the tribe Pinaconotini.

The following is a redescription of Pinaconota bifasciata supple-
menting that of Saussure:

Pinaconota bifasciata (Saussure)

(Figs. 1-23)

Blatta bifasciata Saussure (Rev. Mag. Zool., 1862, p. 165; Mem. Hist. nat.

Mexique, Geneve, 1864, 4, p. 98, $ 9).

Phyllodromia bifasciata (Saussure) (Brunner, Nouv. Syst. Blatt. Vienne,
1865, p. 94).

Epilampra bifasciata (Saussure) (Miss, scient. Mexique, Rech. zool. Paris,
1870, pt. 6, p. 84, pi. 2. Figs. 44, 44A).

Pinaconota bifasciata (Saussure) (Rev. suisse Zool. 1895, 3, p. 337).

cf. (Figs, i, 5, 7). Face smooth with scattered small setae,
vertex exposed. Pronotum semi-circular, somewhat flattened but with
sides slightly deflexed, hind margin subtruncate or weakly curved.
Tegmina abbreviated, reaching to about the middle of fourth ter-
gite (Fig. 7) or fully developed extending beyond the end of the
abdomen (Figs. 1, 5), rounded apically; plical furrow strongly
arched, the furrows from both tegmina meeting to form a semi-ovoid
anal field, anal veins of left tegmen shallowly punctate; veins of
right tegmen punctate only in that portion not covered by the over-
lapping left tegmen; many veins on other parts of the tegmina also
shallowly punctate. Abdomen broad. Supra-anal plate rounded,
mesally indented. Cerci not extending beyond hind margin of supra-

1974]

Roth — Cockroach Genus Gurneya

291

Figs. 6-12. Pinaconota bifasciatia. 6. Brachypterous female. 7. Bra-
chypterous male. 8. Pronotum of female shown in fig. 6. (both $ and $
from Ilha das Alcatrazes, S. Paulo, Brazil.) 9. Pronotum of macropterous
$ taken at quarantine on a plane in Miami, Florida (with bromeliads),
Jan. 20, 1972. 10-12. Three nymphs: 10 and 12. From Brazil, in plane
at San Francisco, Nov. 17, 1970 (with bromeliads). 11. From Brazil, at
Hoboken, Oct. 30, 1940 (with orchids), (scale: figs. 6-7 = 5 mm; figs,
8-12 = 2 mm).

292

Psyche

[June

anal plate (Figs. 4, 13), flattened dorsally with few scattered setae,
somewhat convex ventrally covered with many short and a few long
setae. Subgenital plate asymmetrical, apex of hind margin subacute,
the right side shallowly concave, styli about equal, not extending
beyond the apex of the plate (Fig. 14). Leg armament as described
for genus: Antero-ventral margin of front femur with 2-6 small
but stout spines followed by a row of very small setae. Pulvilli
very large, apical on the metatarsi, and covering practically the
entire ventral surfaces of the other tarsal segments; arolia very
large. Genital phallomeres shown in figures 15-23: L2d an irregu-
larly shaped sclerotized plate separated from L2vm (Figs. 15, 17,
20, 23); preputial membrane modified into a densely spicular area
with 3 or more large spines; a pale sclerotized plate continues beyond
the dense spicular area (Fig. 15) ; the membrane above L2vm with
many small spines; Li with a deep, heavily sclerotized, upcurved
cleft, the upper lobe smaller than the lower lobe which lacks setae
(Figs. 19, 22). R2 strongly curved, the apex partly membranous,
enlarged, a large finger-like projection extending from the base
toward the inner margin of the curved portion (Figs. 16, 18, 21).

Measurements (mm): Total length, 15-16.5; pronotum length
X width, 4.1-4.2 X 5. 9-6. 2 ; tegmen length, 12-12. 5 (macropterous) .
7.6 (brachypterous) .

Coloration: Face (Fig. 3) brownish to black, followed by a broad
testaceous band just above the antennal sockets, then a broad brownish
to black band connecting the tops of the eyes and followed again by
testaceous; basal half of clypeus testaceous. Antennae more or less
unicolorous throughout, or with basal segments yellowish brown, the
remainder darker. Pronotum and tegmina semi-hyaline to subopaque,
the less transparent specimens whitish-yellow, the others testaceous.
Pronotum with an irregularly shaped, arched, longitudinal, light to
dark brownish band on each side of the mid-line, not reaching the
borders, but converging posteriorly; surface sprinkled with a few
to many brownish dots of variable size (Figs. 1, 2, 5, 7). Humeral
vein with a brownish stripe of variable length, the stripe solid
(Fig. 5) or pale with dark lines on either side (Fig. 1); right
tegmen dark on the portion covered by the left tegmen (Fig. 1).
Tergites mostly dark, either solidly dark brown or interspersed with
some pale areas, laterally with a broad testaceous or yellowish-white
border, sprinkled with small brown dots; in the subopaque c? the
seventh segment has a round brownish spot on each side (Fig. 4).
Sternites dark brown with or without a lateral testaceous border

1974]

Roth — Cockroach Genus Gurneya

293

Figs. 13-23. Pinaconota bifasciata. Male structures. 13-14. Supra-anal
plate (dorsal) and subgenital plate (ventral; left style missing), re-
spectively. From male taken at Miami (with orchids from Brazil), 26 July,
1971. 15-23. Genital phallomeres: 15-16. L2vm, L2d, and prepuce (15) and
R2 (16), from $ shown in fig. 5. 17-19. L2d and prepuce (17), R2 (18),
and LI (19) from $ shown in fig. 1. 20-21. L2d, and prepuce (20), and
R2 (21), from $ shown in fig. 7. 22-23. Ll (22), and L2d and prepuce
(23), of male whose supra-anal and subgenital plates are shown in figs.
13 and 14. (scale: figs. 13-14 = 0.5 mm; all others = 0.3 mm; all figures
from KOH cleared, flattened preparations).

294

Psyche

[June

(if present, continuing around the margin of the subgenital plate).
Legs testaceous with some brownish areas, especially on front femora.

9- (Figs. 6, 8, 9). Similar to male except that the subgenital plate
is symmetrical and rounded.

Measurements (mm): Total length, 17-21; pronotum length
X width, 4. 2-5. 5 X 6. 5-8.2 ; tegmen length, 14-16 (macropterous) ,
8.9 ( brachypterous ) .

Nymphs: Nymphs are brownish, and yellowish brown, the mark-
ings differing with different instars (Figs. 10-12). Young and older
stages have the characteristic head markings of the adult. The pro-,
meso-, and metanotum are sprinkled with variable sized raised dots,
a row of several elongated ones along the mid-posterior border just
above the margin. A similar row of elongated granules along the
hind margins of the abdominal tergites, the granules decreasing in
size laterally, and smaller on the posterior segments.

Material examined (All specimens from the U.S. National Mu-
seum) : cf, 9 nymph, 2 cf nymphs, Brazil (at San Francisco, 17.
XI. 1970, with bromeliads) ; 9, cf nymph, Brazil (at Hoboken,
29, 30. X. 1940, with bromeliads); cf and 9. Corupa (Hansa
Humboldt), S. Cath., Brazil, XI. 1944 (9)> XI. 1945 ( cf ) (A.
Mailer leg) ; 9, Brazil; 9, Brazil 20. I. 1972 (on plane at Miami,
with bromeliads); cf, Brazil, 26. VII. 1971 (at Miami, with or-
chids) ; brachypterous cf and 9> Ilha das Alcatrazes, S. Paulo,
Brazil.

I was not able to obtain Saussure’s type 9 but the available
macropterous specimens in general fit his description and figure. The
pronotal markings of the brachypterous specimens were of a much
lighter brown than the long-winged forms and the male preputial
modification differed slightly (Fig. 20) from the others (Figs. 17,
23). It is possible that more than one species is represented in the
above material but additional specimens, particularly of brachypterous
forms are needed before this can be decided.

Walker (1868) described the male of Panchlora inaequalis which
Kirby (1904) listed under Tribonium, and which was placed in
Proscratea by Princis (1958). The male genitalia of Panchlora
(Roth, 1971a), Tribonium spp. (Roth, 1970) \ and Proscratea com-

The following changes should be made in the explanation of figures in
my Zetoborinae paper (Roth, 1970) : p. 223, Fig. 18, Tribonium conspersum
(Guerin) should be Tribonium spectrum (Eschtz.) ; Fig. 19 should be 20;
Fig. 20 should be 19 and Tribonium sp. should be Tribonium conspersum
(Guerin); page 234, shown in Fig. 19 should read 20; Tribonium
conspersum should read Tribonium spectrum; p. 235, Tribonium sp. should
read Tribonium conspersum and shown in Fig. 20 should read 19).

1974]

Roth — Cockroach Genus Gurneya

295

planata (Perty) (Roth, 197 3; type of genus, Princis, 1964), are so
different from inaequalis that it is unwarranted to place this species
in any of these genera. The most unusual features of the genitalia
of inaequalis are the R2 (Fig. 31) with a basal projection extend-
ing toward the inner margin of the hook, and the densely setose
preputial modification (Figs. 29, 30). The R2, L2d, and Li
(Figs. 29-32) of inaequalis are remarkably similar to these structures
of Pinaconota bifasciata (Figs. 15-23). The prepuce in both have
a large faintly sclerotized plate which lies under the L2vm; the pre-
puce of inaequalis differs in lacking some large spines on the border
and smaller ones on the membrane above L2vm. Although the color
markings and size of Panchlora inaequalis differs from Pinaconota
bifasciata (cf. Figs. 1 and 24), the marked similarity in their
genitalia and other characters indicates that the former species
should be placed in Pinaconota.

The following is a redescription of Panchlora inaequalis supple-
menting that of Walker.

Pinaconota inaequalis (Walker), n. comb.

(Figs. 24-32)

Panchlora inaequalis Walker (1868, 33, $ not 9 as indicated).

Tribonium inaequalis (Walker) (Kirby, 1904, 157).

Proscratea inaequalis (Walker) (Princis, 1958, 70, $).

cf holotype (Fig. 24). Interocular space about twice the width
of one eye. Prothorax widest below middle, arched anteriorly, disk
flat, slanting laterally; lateral angles rounded, posterior margin
slightly convex. Supra-anal plate broadly rounded with a distinct
mesal invagination. Cerci reaching to about the hind margin of supra-
anal plate (Fig. 25). Subgenital plate asymmetrical, roughly tri-
angular, rounded on the left side and broadly incised on the right;
styles absent, but these apparently had been broken off (Fig. 26).
Costa of tegmina slightly and regularly curved. Anterior margin
of front femur with 5 or 6 heavy spines, more or less uniform in
size, followed by about a dozen small uniformly spaced piliform
setae, terminating in a large distal spine (Type B) ; posterior
margin with 1 stout spine about distance from apex; distal spine
present; genicular spine absent; anterior margins of mid and hind
femora with 2 or 3, and posterior margins with 3 widely spaced,
small stout spines; genicular spines present as well as distal spines

296

Psyche

[June

Figs. 24-32. Adult $ of Pinaconota inaequalis (Walker) (holotype of
Panchlora inaequalis Walker). 24. Adult. All structures shown in the
following figs, are from this $. 25. Supra-anal plate and cerci (dorsal).

26. Subgenital plate (ventral). 27. Right paraproct. 28. Left paraproct. 29-
32. Parts of the genitalia ; fig. 30 shows the entire preputial modification,
which is only partly shown in fig. 29. (L2vm = ventromedial sclerite (onl)r
apex is shown) ; L2d = dorsal sclerite of second left phallomere; P =
preputial membrane). 31. R2 (second sclerite of right phallomere). 32.
LI (first sclerite of left phallomere). (scale: fig. 24 = 10 mm; figs. 25-26
= 1 mm; figs. 27, 28, and 32 = 0.5 mm; fig. 29 — 0.4 mm; figs. 25-32
from KOH cleared, flattened preparations on 2 slides labeled 66 BMNH).

1974]

Roth — Cockroach Genus Gurneya

297

on both margins of mid and hind femora.2 Tarsal claws equal,
arolia and pulvilli well developed. Right and left paraprocts shown
in figures 27 and 28. Genitalia as shown in figures 29-32; L2d is
a flat unevenly sclerotized plate separated from L2vm (Figs. 29-30) ;
prepuce highly modified with an area of densely packed small setae
leading into a large smooth, banana-shaped (when flattened) sclero-
tized plate (Fig. 30). R2 with an elongated process extending
upward toward the curved inner margin of the hook (Fig. 31);
Li with a curved, heavily sclerotized cleft, lower lobe lacking
setae, with an unevenly broad, sclerotized marginal area (Fig. 32).

Measurements (mm) : Overall length about 21 ; length and width
of pronotum, 5.6 X 8.6.

Coloration: Head black. Pronotum brownish yellow, semi-hya-
line; a broad black inverted Y-shaped band extends from the anterior
arched border to the disk; a broad black, undulating band extends
along the posterior border between the two lateral angles. Abdominal
tergites and sternites piceous. Tegmina reddish tawny, costal area
testaceous and corciaceous towards base, with a dark humeral line;
right tegmen with a large blackish region where it is covered by
the left tegmen. Wings brownish.

Material examined : Holotype cf of Panchlor a inaequalis Walker,
locality unknown. From Mr. Argent’s Collection. Type in British
Museum (Natural History).

Shelford (1907) stated that Ischnoptera (?) obliqua Walker
“appears to be undoubtedly referable to the genus Pinaconota, Sauss. ;
it can be distinguished from the only other species in the genus
P. bifasciata, Sauss., by its much larger size.” P. obliqua differs
markedly from bifasciata in femoral armature, relative length of
cerci, bulging eyes, and cf genital phallomeres. Because of the
morphological differences, especially in the 3 genital phallomeres, I
am placing obliqua in a new genus characterized as follows:

Gurneya n. gen.3

Type-species: Ischnoptera (?) obliqua Walker (1869, p. 148)
(present designation).

2Princis (1958) placed inaequalis in Proscratea. The femoral armament
of a female of Proscratea complanata was as follows: Anterior and posterior
margins of front femur unarmed except for 1 distal spine on each ; genicular
spine absent. Both margins of mid femur unarmed; distal spines lacking;
genicular spine present. Anterior margin of hind femur unarmed; hind
margin with 3 widely spaced heavy spines; distal spines absent, genicular
spine present.

The genus is named in honor of Dr. Ashley Gurney, orthopterist at the
United States National Museum, who has helped me considerably in my
taxonomic studies of the Blattaria.

298

Psyche

[June

cf. Vertex, face, and pronotum deeply punctate (Figs. 34-35).
Entire ventro-anterior margin of fore femur armed with a row of
small, bimarginally serrated, closely spaced spines, lacking a large
distal spine (Fig. 36) ; ventro-anterior margin of mid femur similarly
armed but the spines fewer and not uniformly spaced. Tarsi very
short, fimbriate, unarmed beneath. Supra-anal plate rounded, entire.
Cerci long, extending well beyond hind margin of supra-anal plate
(Fig. 38). Subgenital plate asymmetrical, with a broad shallow
incision on right side (Fig. 39). Styles equal, not extending beyond
hind margin of subgenital plate (Fig. 39). Genital phallomeres;
L2d absent, prepuce spiculate, otherwise unmodified (Fig. 40). R2
with a deep subapical incision which extends to about the middle of
the hook (Fig. 41); Li with the upper lobe (above the cleft)
elongate and more narrow than the lower (Fig. 42).

In obliqua, the absence of an L2d and shape of the spicular sur-
face of the prepuce (though lacking some long “silky” hairs on one
side) resembles the same structure in Alphelixia (see Figs. 21 and
23, in Roth, 1973a). The hooked phallomere (R2) resembles this
structure in certain genera of Epilamprinae (e.g. Litopeltis, see
Fig. 37, in Roth, 1971) and Blaberinae (e.g. Blaptica, see Fig. 57>
in Roth, 1970a). The shape and sclerotization of the Li differs
from that found in the other genera with similar appearing R2’s.

Gurneya obliqua (Walker), n. comb.

(Figs. 33-42)

Ischnoptera (?) obliqua Walker (1869, 148, $).

Pinaconota obliqua (Walker) (Shelford, 1907, p. 496, $ ).

cf . (Fig. 33). Vertex and face (Fig. 34) deeply punctate, clypeus
nearly smooth. Eyes large, bulging, their upper margins raised above
the margin of the vertex, wide apart, the distance between them only
slightly less than the distance between the antennal sockets (Fig. 34).
Pronotum with scattered deep punctures, transversely elliptical, an-
terior margin almost straight, not covering vertex of head, posterior
margin nearly straight, sides deflexed (Fig. 35). Tegmina with basal
fourth punctate, covering the abdomen. Scutellum exposed, punctate.
Supra-anal plate entire, rounded, its surface and hind margin covered
with small fine setae; cerci more than twice the length of supra-anal
plate (Fig. 38). Subgenital plate slightly asymmetrical, the right
side weakly indented; styli about equal, short, not extending beyond
hind margin of subgenital plate (Fig. 39). Legs short, front and
mid tibiae shorter than their corresponding femora. Front femur:
entire ventro-anterior margin armed with about 30 small, bimargin-

1974]

Roth — Cockroach Genus Gurneya

299

Figs. 33-42. Gurneya obliqua (Walker). Holotype $ of Ischnoptera (?)
obliqua Walker. All structures shown in figs. 34-42 are from the type.
33. Adult. 34. Head (frontal view). 35. Pronotum. 36. Front femur (an-
terior surface). 37. Tarsal claws and arolium of prothoracic leg (end
view). 38. Supra-anal plate and cerci ; the dark structures are paraprocts
seen through the cleared tergite (dorsal). 39. Subgenital plate (ventral).
40-42. Genital phallomeres: 40. Apex of L2vm, and preputial membrane.
41. R2. 42. LI. (Specimens shown in figs. 38-42 are KOH treated, cleared,
flattened preparations on a slide); (scale: fig. 33 — 5 mm; figs. 34-35
~ 1 mm; figs. 36, 38-39 = 0.5 mm; fig. 37 = 0.25 mm; fig. 40 — 0.1 mm;
figs. 41-42 = 0.2 mm).

300

Psyche

[June

ally serrated, close-set row of spines (Fig. 36) ; enlarged distal spine
absent; posterior margin with 5 larger bimarginally serrated spines,
2 of them relatively close together basally, the others much wider
apart; distal spine present; genicular spine absent. Mid femur:
entire ventro-anterior margin with about 13 bimarginally serrated
spines, those on the distal third somewhat closer together than the
others; distal spine obsent; posterior margin with 5 spines plus a
small distal spine; genicular spine present. Hind femur: anterior
ventral margin with 7 spines; enlarged distal spine absent; ventral
posterior margin with 3 or 4 larger spines; distal spine absent;
minute genicular spine present. Tarsi short, unarmed beneath, both
tibiae and tarsi fimbriated ; all metatarsi shorter than the remaining
segments combined, of their respective tarsi. Tarsal claws simple,
about equal or one slightly smaller (possibly broken or worn) than
the other; arolia large, roughly triangular in end view (Fig. 37) ;
pulvilli very large and, except for those on the metatarsi, cover the
entire ventral surface of the segments. Genitalia shown in figures
40-42. Li with a deep sclerotized cleft; the upper lobe (above cleft)
elongate, relatively narrow, roughly rectangular extending slightly
beyond margin of lower lobe; lower lobe much larger than upper,
and without setae (Fig. 42). L2d absent, preputial membrane spicu-
lar, unmodified (Fig. 40). R2 broadened at base, a deep subapical
incision extends to about the middle of the hook (Fig. 41).

Measurements (mm): Overall length, 21; body length, 18; teg-
men length, 17; pronotum length X width, 3.9 X 6.2.

Coloration: Pale testaceous. Head castaneous, fuscous band be-
tween eyes; clypeus, mouthparts and antennae testaceous. Two
angulate black stripes on the pronotum extend from anterior to
posterior margins; anteriorly the stripes are narrow and converge;
posteriorly the stripes are broader and diverge (Fig. 35). Tegmina
with a short humeral castaneous stripe. Scutellum marked with
castaneous. Abdomen and legs pale testaceous.

9. Unknown.

Material examined: Holotype d* of Ischnoptera (?) obliqua
Walker. Brazil. Type in Hope Department of Entomology, Oxford
Museum, England.

Summary

The genus Pinaconota is redescribed together with 2 species, P.
bifasciata (Sauss.) and P. inaequalis (Walker). A new genus
Gurneya is erected for Ischnoptera (?) obliqua Walker which
Shelford had incorrectly assigned to Pinaconota.

1974]

Roth — Cockroach Genus Gurneya

301

Acknowledgements

I thank Dr. David Ragge, British Museum (Natural History),
London for the loan of the type of Panchlora inaequalis Walker,
Dr. M. W. R. de V. Graham, Hope Department of Entomology,
Oxford, for the loan of the type of Ischnoptera (?) obliqua Walker,
Dr. Ashley Gurney, U.S. National Museum for specimens of Pina-
conota bifasciata (Saussure), and Mr. Samuel Cohen for taking the
photographs.

References

Kirby, W. F.

1904. A synonymic catalogue of Orthoptera. Vol. I. London, pp. 61-
205.

Princis, K.

1958. Revision der Walkerschen und Kirbyschen Blattarientypen im
British Museum of Natural History, London II. Opusc. Entomol.
23: 59-75.

1964. Orthopterorum Catalogus. Pars 6: 173-284. ’s-Gravenhage.

1967. Orthopterorum Catalogus. Pars 11: 616-710. ’s-Gravenhage.
Roth, L. M.

1970. The male genitalia of Blattaria. III. Blaberidae: Zetoborinae.
Psyche 77: 217-236.

1970a. The male genitalia of Blattaria. IV. Blaberidae: Blaberinae.
Psyche 77: 308-342.

1971. The male genitalia of Blattaria. VII. Galiblatta, Dryadoblatta,

Poroblatta, Colapteroblatta, Nauclidas, Notolampra, Litopeltis ,
and Cariacasia (Blaberidae: Epilamprinae) . Psyche 78: ISO-

192.

1971a. The male genitalia of Blattaria. VIII. Panchlora, Anchoblatta,
Biolleya, Pelloblatta, and Achroblatta. (Blaberidae: Panchlori-
nae). Psyche 78: 296-305.

1973. The male genitalia of Blattaria. X. Blaberidae. Pycnoscelus ,
Stilpnoblatta, Proscratea (Pycnoscelinae) , and Diploptera (Di-
plopterinae) . Psyche 80: 249-264.

1973a. Brazilian cockroaches found in birds’ nests, with descriptions
of new genera and species (Dictyoptera : Blattaria: Blaberidae
and Blattellidae) . Proc. Entomol. Soc. Washington 75: 1-27.
Saussure, H. de

1862. Orthoptera Nova Americana. Diagnoses preliminares, III series
Rev. Mag. Zool. 14: 163-171.

1864. Orthopteres de L’Amerique Moyenne. Mem. Soc. Phys. et d’Hist.
Nat. Geneve, 279 pp.

1895. Revision de la Tribu des Panesthiens et de celle des Epilampriens.
Insectes Orthopteres de la famille des Blattides. Rev. suisse Zool.
3: 299-364.

302

Psyche

[June

Shelford, R.

1907. Studies of the Blattidae (continued). Trans. Entomol. Soc.
London. Part IV.: 487-519.

1910. Genera Insectorum. Orthoptera. Fam. Blattidae. Subfam. Epi-
lamprinae. Fasc. 101: 1-21.

Walker, F.

1868. Catalogue of the specimens of Blattariae in the collection of the
British Museum. London. 239 pp.

1869. Supplement to the Blattariae of the British Museum. London,
pp. 119-156.

MEROPE TUBER (MECOPTERA) :

A WING-BODY INTERLOCKING MECHANISM

By T. F. Hlavac1
Museum of Comparative Zoology
Harvard University
Cambridge, Massachusetts 02138

As an insect pushes its dorsal surface against obstacles while
moving through a substrate, the wings will tend to be forced apart.
In many Coleoptera, such divergence is prevented by a complex
of devices interlocking the elytra with the thorax and abdomen. A
common interlocking mechanism involves intermeshing of parallel
arrays of setae angled towards the potentially disrupting force
(Fig. 2). A similar high friction binding system on the mesojugum
and metascutellum of Merope tuber provides additional evidence for
the ground dwelling habits of this rare mecopteran.

The fore-jugal lobe of Merope is highly modified relative to its
counterpart on the hind wing and as compared with the jugal regions
of other panorpoids (Fig. 1 ; JL2, JL3). It does not bear setae and
does not appear to function in wing coupling. More importantly, the
mesojugal lobe is much thicker, more heavily sclerotized than the wing
proper, is quite rigid, yet capable of slight movement about the third
anal vein, but does not fold over as the wing comes to rest. The
ventral surface is completely covered with uniformly small (.007
X .003 mm) flat topped carinae, angled posteriorly and organized
in rows perpendicular to the long axis of the wing (Figs. 1, JL2;
3, 4). The metascutellum bears two patches of anteriorly projecting
ridges about twice as long (.017 X .003 mm) but about as high as
those on the fore wing (Figs. 1, R, S3; 5).

In rest position, the jugal lobes lie close together and directly above
the striate metathoracic areas. When so placed, a small ventral
movement will cause the slanting nearly parallel ridges to intermesh.
Two functions, which may be combined, may be served by the jux-
taposition of parallel carinae: stridulation and interlocking. Stridu-
lation is an unlikely function for these structures. As the wing
moves medially to rest, the ridges move parallel to one another, not

Research supported by NSF grant GB 31173, F. M. Carpenter, Harvard
University, Principal Investigator. Manuscript completed at Cornell where
E. H. Smith kindly made necessary facilities available.

Manuscript received by the editor June 15, 1974.

303

304

Psyche

[June

Figs. 1-2. Wing-body interlocking mechanism of Merope. Fig. 1. Left
half of pterothorax, dorsal view. As the fore-wing moves medially (a) ;
parallel ridges, indicated by two heavy lines, on the jugal lobe (JL2), and
on the metascutellum (R, S3) are brought into contact. Fig. 2. Model of
interlocking system, posterior view. Small arrow (a) indicates direction of
emplacement as wing comes to rest, see Fig. 1 ; large arrow (b) the direc-
tion of intermeshing of the slanted ridges. 1A, 2A, 3A, anal veins; JL2,
JL3, meso- and metajugal lobes; R, ridge bearing patch of metascutellum,
S3; a, direction of wing movement; b, direction of interdigitation.

perpendicularly as in many sound producing structures (Fig. 2, a).
Stridulatory mechanisms are not known in Mecoptera (Riek 1967).
The geometry of ridge interdigitation, however, is precisely that
required to resist postero-lateral wing displacement during locomo-
tion within a substrate or interstitial spaces. The unique jugal lobe
and metascutellar structures of Merope may serve then as a wing-
body interlocking mechanism which increases structural integrity
during crawling within a matrix.

Other structural features and meagre biological data suggest that
Merope is a substrate inhabitant not a surface dweller as are other
winged Mecoptera. The body is dorso-ventrally flattened ; the pleural
axes of the undifferentiated thoracic segments are sharply inclined
from the vertical (Mickoleit 1967). The fore wing is somewhat
tegmenized, being thicker and more heavily sclerotized than the hind
wing. Antennae and legs are short; pterothoracic appendages may
operate almost entirely within the outlines of the flattened wings.
Little membrane is exposed between tagmata and between abdominal
segments. These characters result in greatly reduced cross section area

!g!&»

1974]

Hlavac — Merope tuber

305

<" bb

D

4h

o o

<Vj ~

^ 2

S ^
.2 bb

« fa
ja

zone of metascutellum, ca. .65K.

3°6

Psyche

[June

during locomotion and improved structural integrity relative to other
panorpoids. Some specimens have been taken under logs or stones, as
were the only known examples of Austromerope. The photographs
in the original description (Killington 1933) show that the jugal
lobe of this Australian form is similar to that of Merope in size and
shape. Several lines of evidence point towards a marked divergence
of meropeids from surface dwelling, loosely constructed configura-
tions of related taxa.

The transformations required to derive Merope from a generalized
mecopteran are parallel and analogous to those hypothesized for the
very early evolution of substrate dwelling Coleoptera (Hlavac 1972).
The ability to keep flexible wings in place under stress through inter-
locking devices is an obvious precursor for the development of elytra.
Meropeids may then represent an attempt, albeit a feeble one, by
another holometabolous lineage to occupy the substrate adaptive zone.

References Cited

Hlavac, T. F.

1972. The prothorax of Coleoptera: origin, major features of variation.
Psyche. 79: 123-149.

Killington, F. J.

1933. A new genus and species of Meropeidae (Mecoptera) from
Australia. Ent. Mon. Mag. 69: 1-4.

Mickoleit, G.

1967. Das thoraxskelet von Merope tuber Newman (Protomecoptera) .
Zool. Jb. Anat. Bd. 84: 313-342.

Riek, E. F.

1967. Structures of unknown, possibly stridulatory, function on the
wings and body of Neuroptera with an appendix on other En-
dopterygote orders. Aust. J. Zool. 15: 337-348.

THE UNUSUAL WEB OF SPILASMA TUBULOFA CIENS,
WITH TAXONOMIC NOTES ON THE SPECIES
(ARANEAE: ARANEIDAE)*

By Diomedes Quintero, Jr.

Museum of Comparative Zoology,

Harvard University, Cambridge MA 02138

During December 1972 I had the opportunity to visit French
Guiana to study some of its little known amblypygid fauna. While
searching for amblypygids I came across the unusual web of an
araneid, Spilasma tubulofaciens , and was able to make some ob-
servations in the field.

I would like to acknowledge the gracious help and encouragement
given by Dr. Herbert W. Levi, who also provided the laboratory
facilities to complete this work. I would like also to thank Dr. J. J.
de Granville for his hospitality and guidance in Cayenne, for the
use of orstom facilities in Saul, and for many other favors.

Web of Spilasma tubulofaciens
Figures 6-7

Spilasma tubulofaciens females were found in a forest patch clear-
ing occupied with mixed growth of shrubs (mainly Rubiaceae and
Leguminoseae), 2-3 m from the muddy banks of Crique Limonade,
French Guiana. The suspension lines of the web were attached
between leaves of a rubiaceous shrub, about 1 .2-1.4 m from the
ground. In the middle they inserted at the apex of the conical re-
treat. The snare region was below, hanging obliquely, and had 16
radii and more than 28 turns, with no distinct frame threads. It
showed numerous torn spaces and repairs, giving a general appear-
ance of long use and disorganization. No major changes were intro-
duced by the spiders during the night before they were collected next
morning. The suspension lines were quite taut, forming nearly a
straight horizontal line (Fig. 6). The outer surface of the conical
retreat was rough, made up of diverse materials, predominantly re-
mains of insect prey (small coleoptera, flies, etc.) and vegetable mat-
ter (protonema of mosses, small bark fragments, vegetable debris).
This debris was firmly fixed to the surface by underlying, fine threads
of silk. The inside of the retreat was smooth, lined with silk. The

* Manuscript received by the editor June 2, 1974,

307

Psyche

[June

308

Figs. 1-5. Spilasma tuhulofaciens (Hingston), female. 1. Dorsal view,
appendages not represented. 2. Lateral view of carapace. 3. Ventral view
of abdomen. 4. Head and part of chelicerae from in front. 5. Epigynum.
Size indicators: Figs. 1-4, 0.5 mm. Fig. 5, 0.2 mm.

1974]

Quintero — Web of Spilasma tubulofaciens

309

Fig. 6. S. tubulofaciens (Hingston), frontal view of web and retreat
(entrance of retreat closed), drawn from photographs. Suspension lines
shaded.

Size indicator: 10 mm.

broad, lower end of the retreat had a transverse, single opening with
two lips, the posterior lip being the longer. The lining of the anterior
lip appeared thinner and more flexible. The supporting lines were
strong, reinforced with silk, and contained numerous moss fragments,
particularly close to the insertion on the retreat. There were also
remains of insects and vegetable matter. The spider remains inside
the retreat, and waits head down for its prey, a characteristic position
of the orb-weaving spiders. When I offered living ants, which I
placed on the lower corner of the catching web, the female dashed
out of the retreat rapidly, wrapped its prey (Fig. 9), bit it, and
promptly returned to the retreat, without eating the prey.

Discussion. By changing the position of the hub (coincident in
this case with the retreat) to a peripheral, uppermost position, an
additional load of tension has been placed on the suspension lines.
Additional tension has also been brought to these lines by the taut,
almost straight line, to which they have been drawn. Consequently
the spider has carefully reinforced the lines with extra layers of silk,
enclosing different building materials, and they no longer function
in prey capture but to support the rest of the web and the retreat.
It differs from S. artifex Simon, which Simon, in 1896, described as
hanging the retreat and catching web from a single perpendicular

3io

Psyche

[June

Fig. 7. S. iubulofaciens (Hingston), closer frontal view of the retreat,
open. Posterior lip not shaded.

Fig. 8. S. artifex Simon, schematic representation of a cross section of
the retreat. Posterior lip of the retreat hangs loose, enclosing mass of eggs
(Modified from Simon, 1896, pi. 12, fig. 4).

Size indicators: Figs. 7-8, 5 mm.

1974]

Quintero — Web of Spilasma tubulofaciens

3ii

suspension line (Fig. 8) and having a catching web parallel to the
floor, with an empty segment.

The numerous repairs in the web of S. tubulofaciens, the thickness
of the suspension lines, and the elaborate construction of the retreat
all suggest that this web is long lasting (relative to the life span of
the spider), and with time is repaired and modified. The retreat of
the younger female had more space for occupancy than the retreat of
the older female (as determined by body length as a percent of the
length of the retreat), the space perhaps serves to accommodate ad-
ditional growth of the spider without having to enlarge the retreat.
Additional collections might verify this observation.

Nielsen (1928, pp. 534-535) has excellent photographs of Achaea-
ranea saxatile (Koch), a theridiid, building a similar type of conical
retreat on the ground and later scaffolding it to the desired height.
It is likely that a similar (energy saving?) strategy is used by S.
tubulofaciens, instead of making numerous trips to carry and assemble
the materials above the ground, although above the ground modifica-
tions and repairs might be introduced at a later date.

The adaptive convergence of webs has been discussed recently by
several authors (see Kullmann, 1972). Strangely, emphasis in their
discussions has been placed on considering orb webs as indivisible
entities which appear to have evolved as a unit. Although interac-
tions must likely occur among the major structural components of
the web (hub, retreat, and capture threads — snare), my impression
is that each has distinct adaptations to cope with in response to
environmental and physiological cues (e.g. the retreat to best hide
the spider). The selective forces which might bring about evolution-
ary changes in these components, by selecting particular motor pat-
terns of the spider, must differ and thus these components have
evolved as distinct characters, although not totally independent.

Taxonomic notes

Spilasma tubulofaciens (Hingston), comb. nov.

Figures 1-5

Epeira tubulofaciens Hingston, 1932, A naturalist in the Guiana Forest, E.
Arnold & Co., p. 366. From Guyana, Essequibo River. Web illustrations,
pp. 150-153; no type material left.

Araneus tubulifaciens, Bonnet (1955, Bibliographia Araneorum, 2: 620, 80),
makes emendation of specific name which is of Latin derivation.

Records . 2 $ $ (one immature) from French Guiana, Crique Limonade,
3-4 km S. of Saiil, 21-22.VIII.1972. Adult $ and retreats deposited in the
MCZ, Harvard collection.

312

Psyche

[June

In his 1932 work, Hingston described 27 new species from eight
different genera collected during an expedition to Guyana in 1929.
Hingston left no type material for his new species and his descrip-
tions were very general, lacking diagnostic characters (Levi, 1963,
p. 493). Nevertheless, Hingston included descriptions and detailed
sketches of the webs of his new species, which may help to validate
some of his names, in some instances. Lehtinen (1967, p. 201) con-
sidered that the only way to treat the species described by this author
would be to propose all of them as “nomina dubia.” I am convinced
that the specimens which I have collected belong to one species de-
scribed by Hingston as E. tubulofaciens. I base my assertion mainly
on the unique abdominal and leg coloration patterns, and the remark-
able architecture of the retreat and the catching web, which matches
Hingston’s descriptions. Although there is the possibility of sibling
species, I have decided for the sake of stability in nomenclature to
retain Hingston’s name.

Description. Carapace, chelicerae (except claws), pedipalps, and
dorsal surface of legs brownish yellow, with the two posterior legs
a darker brown. Inner margin of endites white. Sternum and labium
reddish brown. Carapace covered with fine-pointed bristles which
become longer toward the anterior margin. Carapace with a well-
defined cephalic suture. Ventral coloration of the two first legs is
reddish orange but this pigmentation washed away after preserving
the specimens in alcohol. Abdomen is longer than wide and slightly
wider in its anterior half. Dorsum of abdomen with a broad, white
H-shaped mark over black background, positioned transversely on
the anterior half. Near its middle, the posterior bar of the H mark
has an additional smaller triangular white mark pointing anteriorly
and two dorsal black sclerites. On its posterior third, latero-ventrally,
two elongated white marks occur, one on each side of the abdomen.
Venter with a light colored epigastrium and colulus. Anterior median
eyes are largest and closely positioned, 0.7 diameter apart. Posterior
medians one diameter apart. Laterals closely set on a common raised
black-pigmented carapace projection. Total length 4.9 mm. Cara-
pace 2.6 mm long, 2.3 mm wide. First femur, 2.1 mm; patella and
tibia, 2.0 mm; metatarsus, 1.1 mm; tarsus, 0.9 mm. Second patella
and tibia, 1.9 mm; third, 1.3 mm; fourth, 1.7 mm.

Diagnosis. Spilasma males are not known. Females of S. tubulo-
faciens have distinct body coloration and web construction, different
from the only other South American species: S. tridecim guttata
Simon 1896 from the Amazonas and S. artifex Simon 1896 from
San Esteban, Venezuela. A third species, S. africana Simon 1903,

1974]

Quintero — Web of Spilasma tubulofaciens

313

Fig. 9. Female of S. tubulofaciens wrapping prey on web.

314

Psyche

[June

is only known from Equatorial Guinea. Hingston (1932, p. 366)
described a species, E. sacculifaciens , with similar conical retreat and
web architecture to S. tuhulo facie ns, which, I suspect, will eventually
be ascribed to the genus Spilasma when new material is collected.
Its reported size, 1.5 mm, suggests it might be an immature specimen
as the other known species of Spilasma range from 3 to 6 mm in
length.

References Cited

Bonnet, P.

1955. Bibliographia Araneorum, 2: 620, 80. Toulouse.

Hingston, R. W. C.

1932. A naturalist in the Guiana forest. E. Arnold & Co., London.
Kullmann, E. J.

1972. The convergent development of orb-webs in cribellate and ecri-
bellate spiders. Am. Zoologist, 12: 395.

Levi, H. W.

1963. American spiders of the genus Theridion (Araneae: Theridiidae) .
Bull. Mus. Comp. Zool., 129(10): 493.

Lehtinen, P. T.

1967. Classification of the cribellate spiders and some allied families.
Ann. Zool. Fennici, 4: 201.

Nielsen, E.

1928. De Danske Edderkoppers Biologi. Denmark (pp. 521-535).
Simon, E.

1892-1895. Histoire Naturelle des Araignees. I. Paris.

1896. Etudes Arachnologiques. Ann. Soc. Entomol. France, 65: 465-510,
1 pi.

WING-FOLDING IN THE PALEOZOIC INSECT ORDER
DIAPHANOPTERODEA j( PALEOPTERA ) , WITH A
DESCRIPTION OF NEW REPRESENTATIVES
OF THE FAMILY ELMOIDAE

By Jarmila Kukalova-Peck* 1
Department of Geology, Carleton University,

Ottawa, Ontario, Canada

The order Diaphanopterodea is closely related to two other Paleo-
zoic paleopterous orders, the Palaeodictyoptera and the Megasecoptera,
all three having elongate, haustellate mouth-parts. In the history of
the study of paleopterous insects, the Diaphanopterodea represent
the most intriguing group. The order combines the basic character-
istics of the Paleoptera, fluted wings and very long, multisegmented
cerci, with the ability to fold the wings backwards over the abdomen,
which is the principal character of the Neoptera. Since that resting
position of the wings indicates a major development in the evolution
of insects, as well as major achievement in adaptation to more varied
environment, its occurrence within the Paleoptera is of great interest.

At present there is general agreement among students of insect
evolution that wing-folding ability arose at least twice in insects —
in the Neoptera and in the Diaphanopterodea among the Paleoptera
(Rohdendorf 1962, Carpenter 1963, Sharov 1971). It has been
assumed that the wing-folding mechanism in the Diaphanopterodea
was probably different from that in the Neoptera, but the structure
of the axillary region of the diaphanopteran wing has not actually
been known. While examining all known specimens of Diaphanop-
terodea from the Lower Permian deposits of Czechoslovakia, Kansas
and Oklahoma, I succeeded in finding a dozen wings, belonging to
the families Elmoidae, Martynoviidae and Asthenohymenidae, with
the bases of the wings at least partially preserved. The present paper
is concerned only with the family Elmoidae; the two remaining

This research has been supported in part by the President’s Research
Grant (Canadian National Research Council) from Carleton University,
and in part by NSF Grant G.B. 39720, F. M. Carpenter, Harvard Uni-
versity, Principal Investigator.

I am deeply indebted to Professor F. M. Carpenter for placing at my
disposal the Diaphanopterodea in the Museum of Comparative Zoology
which he had collected in Kansas and Oklahoma, and for his critical review
of this manuscript.

Manuscript received by the editor June 20, 1974.

315

3 1 6

Psyche

[June

families and the wing base morphology of the Palaeodictyoptera and
Megasecoptera will be discussed elsewhere.

The new material includes six new genera and ten new species
of Elmoidae from the Lower Permian beds near Obora in Moravia,
Czechoslovakia. The base of the wing is well preserved in one
specimen only, Permodiapha carpenteri n.sp. (No. 8/1974), and
incompletely preserved in seven other specimens.

Since no definitive studies of the wing bases of the extant Paleop-
tera [Odonata and Ephemeroptera] have been made, some difference
of opinion exists about the interpretation of the basal sclerites. The
interpretation of the diaphanopteran structures used in this paper
should be considered only as tentative. The terminology employed is
derived from the account of the wing base of Odonata by Tannert
(1958), since the fore wing base in the Ephemeroptera has never
been described in functional terms. For further comparison, the
wing base structures of the related family Martynoviidae (un-
described material) and the related order Palaeodictyoptera (Kuka-
lova i960, 1969, 1970) have been used.

Family Elmoidae

Elmoidae Tillyard 1937: 82; Carpenter 1943: 56; Carpenter 1947: 27;

Rohdendorf 1962: 69; Carpenter 1963: 249.

Parelmoidae Rohdendorf 1962: 71.

The family Elmoidae was originally assigned by Tillyard to the
paleopterous order Megasecoptera because of the similarity in the
venation pattern. This point of view was followed by Carpenter
in his emendation of Elmoidae in 1943 and 1947, although he sub-
sequently (1954) separated the Elmoidae, along with the Diapha-
nopteridae and related families, into a distinct suborder (Parame-
gasecoptera) , on the basis of the folded, resting position of the
wings. Later, Rohdendorf (1962) and Carpenter (1963) independ-
ently reached the conclusion that these families had to be referred
to the separate order Diaphanopterodea, established by Handlirsch
(1919). A detailed evaluation of the phylogenetic position of Elmoi-
dae and Diaphanopteridae within the order, and of Diaphanopterodea
within the Paleoptera has been given by Carpenter (1963).

Until now, the family has been known by only three genera, all
from the Lower Permian of Kansas and Oklahoma — Elmoa,
Parelmoa and Pseudelmoa, represented by five species. The new
fossils from Moravia contribute additional and varied morphological
features, which broaden the family characteristics formerly used. In
the Moravian material thickened cuticular spots are sometimes pres-

1974]

Kukalova-P eck — Diaphanopterodea

317

Fig. 1. Elmodiapha ovata n.sp., specimen 1/1974, fore wing. Holotype.
Fig. 2. Elmodiapha ovata n.sp., specimen 2/1974, fore wing. Fig. 3. Pro-
todiapha maculifera n.sp., specimen 3/1974, hind wing. Holotype. Fig. 4.
Protodiapha maculifera n.sp., specimen 4/1974, hind wing.

3i8

Psyche

[June

ent on the wings, between the veins, and the main veins are covered
with a series of setae. Costa is flattened and in larger wings some-
times serrated ; it often carries a series of long setae, which are more
pronounced basally. Subcosta consistently forms a conspicuous bend
at the very base, probably related to the wing-folding process. Radius
is always fused basally with the stem of the media. M, after diverging
from R, immediately coalesces or joins either the stem of Cu and
CuA, or directly CuA, for a short distance. MA and Rs are con-
nected in North American Elmoidae by a cross vein. In Moravian
Elmoidae the cross vein is present in only three species. In others,
MA almost touches Rs or MA joins Rs at a point, or for a short
distance, or both veins are anastomosed for a short distance. Cubitus
runs closely to, or touches, R. The stem of Cu is always ribbon-like,
deeply concave, broad and flattened. CuA and CuP originate near
the point at which M diverges from the common R + M stem. Soon
after diverging from M, CuA approaches CuP and is connected
with it by an oblique cross vein. This feature seems to be a stable
character of the family. Anal veins start from the posterior part of
the sclerotized region, which is the thickened part of the wing
membrane posteriorly from Cu and distally from the basal fold
(fig. io,S). It undoubtedly served as a reinforcement of the wing
base while folding the wings backwards. The basal plate is repre-
sented either by the enlarged cubitoanal plate, or by the fusion of
the median, cubital, and anal plates. It is partly homologous with
the fused subcostoanal plate of the Palaeodictyoptera, the fused basal
plate of the Ephemeroptera, and the radioanal plate of the Odonata.
The basal fold runs between Sc and R + M, crosses R + M where
it is flattened and weakly sclerotized, follows closely the distal
margin of the fused basal plate, and crosses the anal part of the
sclerotized region (fig. io,B). The axillary sclerite articulates with
the proximal margin of the basal plate and with R. Its size and
shape are not known.

The family Elmoidae, as long as it was known only by North
American species, was well separated from the related Upper Car-
boniferous family Diaphanopteridae. The family Diaphanopteridae,
represented by Diaphanoptera (France) and Philiasptilon (Soviet
Union, Kuznetsk Basin), was characterized by the presence of the
anastomosis between MA and Rs. In the Permian material from
Moravia, however, this feature is highly variable. In Elmodiapha
and Protodiapha (new genera), MA and Rs are connected only by
a cross vein, as in the Elmoidae from Kansas and Oklahoma. In the
other Moravian genera, MA and Rs are either touching each other

1974]

Kukalova-Peck — Diaphanopterodea

319

Fig. 5. Protodiapha lineata n.sp., specimen 5/1974, fore wing. Holo-
type. Fig. 6. Diapha Candida n.sp., specimen 6/1974, fore wing. Holotype.
Fig. 7. Diapha Candida n.sp., specimen 7/1974, hind wing. Fig. 8. Per-
modiapha carpenteri n.sp., specimen 11/1974, fore wing.

320

Psyche

[June

at a point, or anastomosed as in Diaphanopteridae. Since the pres-
ence or absence of the MA-Rs anastomosis is no longer useful as the
family character, the status of the Elmoidae as a separate family is
questionable. However, since Diaphanoptera, the type genus of the
family, is from Upper Carboniferous strata, and since its body
structure is entirely unknown, it Seems advisable to consider the
families distinct at least until more is known about Diaphanoptera.

The generic classification used here is based mainly upon the
presence or absence of MA-Rs anastomosis, the general shape of the
wings, and the character of the Rs area, since these features have
been found relatively stable in the related order Palaeodictyoptera
(Kukalova-Peck, 1971). All specimens available are figured. The
lettering of the veins is shown in figures 9 and 10.

Genus Elmodiapha, new genus2

Type species: Elmodiapha ovata n.sp., Lower Permian of Mora-
via.

Fore wing: MA and Rs not anastomosed, connected by a cross
vein. Wings elongated, oval in shape. Sc terminating on R shortly
beyond mid-wing. Rs area large, with about 6 branches. R sending
off a series of long branches beyond the end of Sc. Hind wing not
known.

Elmodiapha differs from Parelmoa in having more elongate wings
and richly branched Rs. From Pseudelmoa it differs in the shorter
Sc, and in MA approaching more closely to Rs.

Elmodiapha ovata n.sp.

Figure 1, 2

This species is represented by two detached fore wings with a
richly branched Rs.

Fore wing: length 1 7.5-21 mm, width 5. 4-6. 6 mm. Costa ser-
rated. Rs with 6 long, densely arranged branches. MA simple, MP
twice forked. CuA and CuP simple. iA usually with a series of
branches, anal veins sometimes branched.

Holotype: No. 1/1974 (fore wing fragment, length 18.5 mm,
width 6.6 mm, obverse) ; specimen No. 2/1974 (fore wing fragment,
length 16 mm, width 5.4 mm, obverse and reverse). Paleontological
Institute of Charles University, Prague, Czechoslovakia.

2The latter part of this generic name (and that of the other genera which
are proposed in this paper) is derived from the Greek adjective diaphanes
(distinct) ; it is treated as a noun and is considered feminine

1974]

Kukalova-Peck — Diaphanopterodea

32i

Genus Protodiapha, new genus

Type species: Protodiapha maculifera n.sp., Lower Permian of
Moravia.

Fore wing: MA and Rs connected with a very short cross vein.
Wings short and broad, broadest at about mid-wing. Posterior
margin convexly bent, forming a lobe. Sc terminating on R shortly
beyond mid-wing. Rs area small, with about 5 branches. Hind wing
similar, but shorter, with the lobe on the posterior margin probably
more pronounced.

Protodiapha differs from all the other genera which lack the
anastomosis between MA and Rs by the presence of the lobe on the
posterior margin.

Protodiapha maculifera n.sp.

Figure 3, 4

This species is established upon hind wings having a prominent
lobe on the posterior margin and circular, cuticular spots on the
wing membrane.

Hind wing: length 14 mm, width 4.5-5 mm. Wing broadest
before the level of the end of Sc. The lobe on the posterior margin
extending to the end of CuP. C with a series of long setae. Rs
almost straight, with 5 branches. MA simple, MP with a fork.
CuA simple, CuP sometimes with short branches. Anal veins with
few branches. Thickened cuticular spots present.

Holotype: No. 3/1974 (hind wing, length 13.5 mm, width 4.5
mm, obverse); specimen No. 4/1974 (hind wing fragment, length
8.3 mm, width 5 mm, obverse). Paleontological Institute of Charles
University, Prague, Czechoslovakia.

Protodiapha lineata n.sp.

Figure 5

This species is monotypic and based on the proximal half of a
fore wing. However, the unique S-shaped cross veins accompanied
by dark spots separate the species distinctly from all other Elmoidae.

Fore wing: length of the fragment 11.4 mm, width 5.8 mm.
Wing broadest beyond the level of the end of Sc. The small lobe
on the posterior margin extending to the end of CuP. Veins darkly
colored. CuA with a short branch, CuP simple. Anal veins with
few branches. Cross veins long, S-shaped, surrounded by dark
spots. Circular cuticular spots absent.

322

Psyche

[June

Fig. 9. Permodiapha carpenteri n.sp., specimen 8/1974, hind wing.
Holotype. Fig. 10. Permodiapha carpenteri n.sp., specimen 8/1974, hind
wing. Holotype. Ax = axillary sclerite ; B == basal fold; FP1 == fused
basal plate; S — sclerotized region.

1974]

Kukalova-Peck — Diaphanopterodea

32 3

Protodiapha lineata differs from maculifera in having the wings
widest beyond mid-wing, a smaller lobe on the posterior margin and
long, S-shaped cross veins.

Holotype: No. 5/1974 (fore wing fragment, length 11.4 mm,
width 5.8 mm, obverse). Paleontological Institute of Charles Uni-
versity, Prague, Czechoslovakia).

Genus Diapha, new genus

Type species: Diapha Candida n.sp., Lower Permian of Moravia.

Fore wing: MA anastomosing with Rs for a short distance.

Wings slightly broadened beyond mid-wing. Posterior margin con-
cave. Sc terminating on R shortly beyond mid-wing. Rs with 5-6
branches. Hind wing shorter and probably narrower, with anal
veins less branched. Diapha differs from the other genera described
herein by the shape of the wings.

Diapha Candida n.sp.

Figure 6, 7

A detached fore wing and a hind wing are referred to this species.
The hind wing differs slightly in the outline of the posterior margin
and in the thinner membrane. It may belong to a different species.

Fore wing: length 19 mm, width 5 mm. Wing broader in the
distal half. C basally with a series of more prominent setae. Stem
of R distinctly swollen next and distally from the basal fold. Rs
concavely bent in the apical part, with 6 branches. MA simple,
MP forked. CuA simple, CuP simple or with a branch. iA with
several branches, anal veins sometimes branched.

Hind wing: length 14 mm, with 4.2 mm. Wing membrane very
thin.

Holotype: No. 6/1974 (fore wing, length 18.7 mm, width 5 mm,
obverse); specimen No. 7/1974 (hind wing, length 14 mm, width
4.2 mm, obverse and reverse). Paleontological Institute of Charles
University, Prague, Czechoslovakia.

Genus Permodiapha, new genus

Type species: Per?nodiapha carpenteri n.sp., Lower Permian of
Moravia.

Fore wing: MA touching Rs or anastomosing with Rs for a
short distance. Wing very broad. Posterior margin evenly con-
cave. Sc terminating on R shortly beyond mid-wing. Rs with 3-4
branches. Hind wing similar to fore wing, but broader.

324

Psyche

[June

Fig. 11. Permodiapha lata n.sp., specimen 10/1974, fore wing. Fig. 12.
Permodiapha lata n.sp., specimen 9/1974. Hind wing, fore wing outlined.
Holotype.

1974]

Kukalova-Peck — Diaphanopterodea

325

Permodiapha differs from all other genera with MA-Rs anasto-
mosed by its very broad wings.

Permodiapha carpenteri n.sp.3

Figure 8, 9, 10

A detached fore wing and a hind wing are referred to this
species. The hind wing No. 8/1974 has been chosen as the holotype
because of its well preserved base. The structure of the wing base
are discussed above with the family characters.

Fore wing: length about 16 mm, width 5.3 mm. Wing large,
with relatively strong membrane, veins and cross veins. Apex prob-
ably rounded as in the hind wing. Rs sending off 4 branches,
slightly S-shaped distally. MA simple, MP with a long fork. 'CuA
simple, CuP simple or forked. Anal veins with several branches.
Fore wing about equally long but narrower than the hind wing.

Hind wing: length 16.3 mm, width 6.8 mm. Similar to the
fore wing but broader, and narrowing more towards the base.

Holotype: No. 8/1974 (hind wing, length 16.3 mm, width 6.8
mm, obverse and reverse) ; specimen No. 11/1974 (fore wing frag-
ment, length 11 mm, width 5.3 mm, reverse). Paleontological
Institute of Charles University, Prague, Czechoslovakia.

Permodiapha lata n.sp.

Figure 11, 12

The species is represented by the holotype, probably a male, con-
sisting of a vaguely preserved fore wing, well preserved hind wing
and part of the abdomen, and by a detached wing, probably a fore
wing.

Fore wing: length about 12.6-15.5 mm, width 4.9-5. 2 mm. Wing
membrane very delicate, cross veins almost invisible. Apical part
of the wing somewhat narrower than in carpenteri. Rs curved
posteriorly, sending off 4 branches. MA and CuA simple, MP and
CuP with a long fork. Anal veins with few branches. Fore wing
longer and narrower, with the apical part more pointed than the
hind wing. Hind wing: length about 14.5 mm, width 6 mm. An-
terior margin strongly curved towards the apex.

Permodiapha lata differs from carpenteri in having a very thin
membrane and narrower fore wings.

3This species is named for Dr. Frank M. Carpenter (Harvard Univer-
sity), who has published extensively on the Diaphanopterodea.

326

Psyche

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Fig. 13. Permodiapha specula n.sp., specimen 13/1974, fore wing. Holo-
type. Fig. 14. Permodiapha sp., specimen 14/1974, hind wing. Fig. 15.
Stenodiapha angusta n.sp., specimen 17/1974, fore wing. Holotype. Fig. 16.
Stenodiapha angusta n.sp., specimen 18/1974, fore wing.

1974]

Kukalova-Peck — Diaphanopterodea

327

Holotype: No. 9/1974 (fore wing, length about 15.5 mm, width
5.2 mm; hind wing fragment, length 13 mm, width 6 mm; abdomen,
1.-10. segment, length 13.5 mm; hind tibia, length 3 mm; obverse) ;
specimen No. 10/1974 (fore wing fragment, length 12.6 mm, width
4.9 mm, obverse). Paleontological Institute of Charles University,
Prague, Czechoslovakia.

Permodiapha specula n.sp.

Figure 13

This species is monotypic and based upon a fore wing.

Fore wing: length 13.3 mm, width 4.8 mm. Wing relatively
small, with very thin wing membrane. Cross veins almost invisible.
Proximal half of the wing much narrower than the distal half.
Venation very simple, seldomly branched. MA only touching Rs
at a point. Rs sending off 4 branches, Rs S-shaped distally. MA,
MP, CuA and CuP simple. Anal veins mostly simple. Circular
cuticular spots present.

Permodiapha specula differs from carpenteri and lata in having
the smallest wings with the most simple venation; in having the
wings narrowed in the proximal half; and in the presence of the
circular cuticular spots.

Holotype: No. 13/1974 (fore wing, length 13.3 mm, width 4.8
mm, obverse and reverse). Paleontological Institute of Charles Uni-
versity, Czechoslovakia.

Permodiapha sp. specimen No. 14/1974. (fig. 14) (hind wing
fragment, length 11 mm, width 4.5 mm, obverse and reverse) might
represent a separate species within the genus Permodiapha , but the
fragment does not provide enough features for specific diagnosis.

Genus Stenodiapha, new genus

Type species: Stenodiapha angusta n.sp., Lower Permian of

Moravia.

Fore wing: MA anastomosed with Rs for a short distance.

Wing narrow, elongate, with main veins obliquely oriented. Sc
terminating on R shortly beyond mid-wing. Rs with 3-4 branches,
rs area long and narrow. Hind wing similar to the fore wing, but
shorter and broader.

Stenodiapha differs from all other genera with MA-Rs anastomosed
by the longest and narrowest wings, with the veins most obliquely
oriented.

328

Psyche

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Fig. 17. Stenodiapha angusta n.sp., specimen 19/1974, fore wing. Fig. 18.
Stenodiapha angusta n.sp., specimen 20/1974, hind wing. Fig. 19. Steno-
diapha moravica n.sp., specimen 21/1974, fore wing. Holotype. Fig. 20.
Stenodiapha moravica n.sp., specimen 22/1974, fore wing.

1974]

Kukalova-Peck — Diaphanopterodea

329

Stenodiapha angusta n.sp.

Figures 15, 16, 17, 18

Three detached fore wings and one hind wing are referred to
this species.

Fore wing: length 17.2-20 mm, width 3. 8-4. 3 mm. Wing broadest
at mid-wing, narrowing proximally. Wing membrane very thin. Rs
with 4 long branches. MA simple, MP simple; CuA simple, CuP
simple or forked. Anal veins with few branches or simple. Hind
wing: length about 16.5 mm, width 4.4 mm. Similar to the fore
wing, but shorter. In hind wing No. 20/1974, the richly branched
MP is reminiscent of the Carboniferous genus Diaphanoptera.

Holotype: No. 17/1974 (fore wing fragment, length 18.2 mm,
width 4.3 mm, obverse); specimen No. 18/1974 (fore wing frag-
ment, length 15.7 mm, width 3.8 mm, obverse and reverse) ; specimen
No. 19/1974 (fore wing fragment, length 14.6 mm, width 3.8 mm,
obverse and reverse); specimen No. 20/1974 (hind wing fragment,
length 15 mm, width 4.4 mm, obverse and reverse). Paleontological
Institute of Charles University, Prague, Czechoslovakia.

Stenodiapha moravica n.sp.

Figure 19, 20

This species is based on two detached wings, probably fore wings.

Fore wing: length about 20.7 mm, width 5.3 mm. Wing broad-
ened slightly at mid-wing. C serrated, basally carrying a series of
setae. R sending off anteriorly several long twigs beyond the end
of Sc. Rs with 3 long branches. MA simple, MP forked. CuA
and CuP forked. Anal veins with few branches or simple.

Stenodiapha moravica differs from angusta in having the wings
broader in the basal third and in the presence of long twigs on
R beyond the end of Sc.

Holotype: No. 21/1974 (fore wing fragment, length 19.2 mm,
width 5.3 mm, obverse and reverse); specimen No. 22/1974 (fore
wing fragment, length 11.5 mm, width 5.3 mm, obverse). Paleonto-
logical Institute of Charles University, Prague, Czechoslovakia.

Paradiapha, new genus

Type species: Paradiapha delicatula n.sp., Lower Permian of
Moravia.

Fore wing: MA joining Rs for a short distance. Wing broadest
at about the end of the second third, narrowing abruptly at the
base. C serrated. Sc terminating on R shortly beyond mid-wing.

330

Psyche

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Fig. 21. Paradiapha delicatula n.sp., specimen 15/1974. Holotype. A.
right fore wing, B. left hind wing, C. right hind wing.

1974]

Kukalova-Peck — Diaphanopterodea

331

Rs area small, with about 3 Rs branches. Hind wing similar to the
fore wing, slightly shorter and broader proximally.

Paradiapha has a similar wing shape to ParelmoaJ from which
it differs in having a shorter Sc and by having MA almost anasto-
mosed with Rs. From all other genera it differs in the wings being
very broad in the distal third.

Paradiapha delicatula n.sp.

Figure 21, 22

The species is based upon two specimens. The holotype consists
of the fragment of the right fore wing, of an almost complete left
hind wing, and of the basal third of the right hind wing, all in
partly folded position. Specimen No. 16/1974 is represented by the
fragments of overlapping fore and hind wings, broad pterothorax
and a remnant of the hind leg, with prominent setae on the tibia.

Fore wing: length about 12.9 mm, width about 4.3 mm. Wing
membrane very thin. Anterior margin strongly bent apically, apex
shifted posteriorly. C serrated, with a series of long setae. Rs
strongly curved posteriorly, sending off 3 branches. MA, MP, CuA,
CuP and anal veins simple. Hind wing almost identical with the
fore wing, slightly shorter.

Holotype: No. 15/1974 (right fore wing fragment, length 12.6
mm; left hind wing fragment, length 12.8 mm, width 4.3 mm;
right hind wing fragment, length 7.4 mm, width 4 mm; reverse);
specimen No. 16/1974 (fore wing fragment, length 8.7 mm; hind
wing fragment, length 6.2 mm; hind tibia, length 1.8 mm; obverse).
Paleontological Institute of Charles University, Prague, Czecho-
slovakia.

Discussion

On superficial examination, the wing base in Permodiapha does
not differ from the generalized neopterous type. The axillary region
penetrates deeply into the wing, from which it is separated by the
basal fold. However, careful study reveals that instead of the
second and third axillary sclerites, which would be located proximally
from the basal fold in true Neoptera, there is a large basal plate
which is characteristic of the Paleoptera. The true axillary sclerite
(fig. 9, Ax), perhaps homologous with 2Ax in Ephemeroptera
( sensu Tsui and Peters, 1972), is located much more distally, in the
“paleopterous” position.

332

Psyche

[June

Fig. 22. Paradiapha delicatula n.sp., specimen 16/1974. Fore wing and
hind wing, hind leg.

With the subcostoanal basal plate of advanced Palaeodictyoptera
(Kukalova, 1970), the basal plate of Diaphanopterodea is only par-
tially homologous. It is apparently formed only by the fused cubi-
toanal plate or eventually by the mediocubitoanal plate in which
subcostal and radial basal plates do not participate. This condition
is reminiscent of the “prefusion” stage of basal plates, known in
primitive Palaeodictyoptera (Kukalova, i960), from which Dia-
phanopterodea are supposedly derived.

The diaphanopteran wing base developed, in all probability, from
the ancient paleopterous base, at the stage when the basal plates of
the main veins were still separated, as indicated in some ancient
Palaeodictyoptera. While the subcostal and radial plate became
completely reduced to give place to the basal fold, the cubital and
anal (or median, cubital, and anal) plates gradually fused and
became separated from the wing by the basal fold.

The ability to penetrate narrow spaces for feeding and protection
was undoubtedly a major factor in the success of Neoptera. The
Diaphanopterodea, although having a wing-folding mechanism of
their own, had highly specialized, haustellate mouthparts, as did
their relatives, the Palaeodictyoptera and Megasecoptera. None of
these paleopterous orders were apparently able to compete success-

1974]

Kukalova-Peck — Diaphanopterodea

333

fully with the more diversified Neoptera and none have been found
so far in strata formed after the close of the Permian Period.

References

Carpenter, F. M.

1943. The Lower Permian Insects of Kansas. Part 9. The Orders
Neuroptera, Raphidiodea, Caloneurodea and Protorthoptera
(Probnisidae), with additional Protodonata and Megasecoptera.
Proc. Amer. Acad. Arts Sci., 75 (2): 55-84.

1947. Lower Permian Insects from Oklahoma. Part 1. Introduction
and the Orders Megasecoptera, Protodonata, and Odonata. Proc.
Amer. Acad. Arts Sci., 76(2) : 25-54.

1954. Extinct Families of Insects. In Classification of Insects (C. T.
Brues, A. L. Melander and F. M. Carpenter). Part III. Bull.
Mus. Comp. Zool., 108: 777-824

1963. Studies on Carboniferous Insects from Commentry, France.
Part V. The Genus Diaphanoptera and the Order Diaphanop-
terodea. Psyche, 70(4): 240-256.

Handlirsch, A.

1906. Die fossilen Insekten und die Phylogenie der rezenten Formen.
Leipzig. 430 pp.

1919. Revision der palaeozoischen Insekten. Denkschr. Akad. Wiss.
Wien. Math. Naturw. Kl., 96: 82 pp.

KUK ALOVA, J.

1960. New Palaeodictyoptera of the Carboniferous and Permian of
Czechoslovakia. Sbornik UUG, 25: 236-251.

1969. Revisional Study of the Order Palaeodictyoptera in the Upper
Carboniferous Shales of Commentry, France. Part 1. Psyche,
76(2): 163-215; Part 2. Psyche, 76(4): 439-486; Part 3. Psyche,
77(1) : 1970: 1-44.

Kukalova-Peck, J.

1971. The Structure of Dunbaria (Palaeodictyoptera). Psyche, 78(4):
306-318.

Rohdendorf, B. B., et al.

1962. Principles of Paleontology. Akad. Nauk SSSR, Moscow. 535 pp.
Tannert, W.

1958. Die Fluegelgelenkung bei Odonaten. Deutsche Ent. Zeit., (N.F.),
5 (5) : 394-455.

Till yard, R. J.

1937. Kansas Permian Insects. Part 17. The Order Megasecoptera
and Additions to the Palaeodictyoptera, Odonata, Protoperlaria,
Copeognatha, and Neuroptera. Amer. J. Sci., (5) 33: 81-110.

OBSERVATIONS OF THE NESTING BEHAVIOR
OF RUBRIC A SURINAMENSIS (DEGEER)
(HYMENOPTERA, SPHECIDAE)

By H. E. Evans,1 R. W. Matthews,2 and E. McC. Callan3

Introduction

Rubric a surinamensis (DeGeer) is one of the most familiar and
well-studied digger wasps of South America. Yet the published
accounts of its nesting behavior present a number of puzzling fea-
tures. For example, Brethes (1902) and Llano (1959) reported
that females clean the cells of uneaten parts of flies and other debris,
but others have not confirmed this. There are many records of
females preying upon flies, but Callan (1950) has reported the
occasional use of Lepidoptera and Odonata, a striking exception to
the usual prey constancy of solitary wasps. Also Brethes (1902)
reported that the cocoons contain only two pores, although other
Bembicini in this size range make 5-10 pores per cocoon. A fuller
summary of published observations has been presented by Evans
(1966).

In the hope of clarifying these and other problems relating to this
widely distributed and apparently very successful species, the three
of us have pooled our notes on three widely separated populations.
One of us (EMC) resided in Trinidad for several years, and had
an opportunity to observe a nesting aggregation in the hard, sandy
driveway of his house near the Imperial College of Tropical Agri-
culture at St. Augustine, a residential area about 13 km east of Port
of Spain. Two of us (HEE and RWM) visited Colombia and
Argentina in January and February, 1972, and encountered the
species in some abundance in both countries. Studies in Colombia
were conducted on a large aggregation in a hard, gravel road on a
denuded and eroded hillside just west of the city of Cali. In Argen-
tina, we found three smaller aggregations in diverse situations within
one day’s drive of the city of Tucuman. Thus if any of the puzzling
features of the behavior of this species bear a relationship to geog-

department of Zoology and Entomology, Colorado State University, Fort
Collins, Colorado 80521.

department of Entomology, University of Georgia, Athens, Georgia
30602.

division of Entomology, C.S.I.R.O., Canberra, A.C.T., Australia.

Manuscript received by the editor August 13, 1974.

334

1974] Evans j Matthews t & Callan — Rubrica surinamensis 335

raphy, to substrate, or to population density, the following synthesis
of our notes from these various localities may serve to clarify this
relationship. We also include a description of the larva and of the
cocoon, in the hope of shedding light on the relationship of Rubrica
to other genera of Bembicini.

Distribution and Ecology

R. surinamensis ranges from Trinidad and southern Mexico to
central Argentina, being absent only from forested areas, higher
mountains, and the friable sand of dunes and beaches. Because of its
propensity for nesting in roads and paths, often in populated areas,
it is one of the most commonly observed of solitary wasps, and its
large size (17-24 mm) and pattern of yellow maculations against a
rufous and fuscous background render it especially likely to attract
attention. Since this species is subject to much variation in color,
it should be pointed out that the populations we studied were all
“typical of the species”, that is, without discal yellow markings on
the mesocutum or other unusual features. We found no consistent
differences in color or structure between Argentinian wasps and
those from northern South America, despite behavioral differences
to be pointed out below.

As noted above, the aggregations at Cali, Colombia, and at St.
Augustine, Trinidad, both occupied hard-packed roads. At Cali, an
estimated 150 females nested along the road for a distance of about
25 m, many nests being separated by 0.5- 1 m, but some clustered so
that entrances were only 3-15 cm apart. The soil here is best de-
scribed as clay-sand containing many small stones. At St. Augustine,
the nesting aggregation varied seasonally from about a dozen to an
estimated 50 individuals. These occupied an area of about 50 m2,
with nest entrances 25 cm to 1 m or more apart. The wasps occu-
pied the same site at least from 1941 to 1944. The aggregation at
Cali is also known to have persisted at the same site for several years.

On Trinidad, R. surinamensis has also been found nesting at the
Erin Savanna, an area of open, virtually uninhabited country sur-
rounded by evergreen seasonal forest, typical of the sandy soils of
the south of the island. Here it nested in company with Stictia
pantherina ( Handlirsch) . At other sites, R. surinamensis has been
found nesting essentially alone, as few other bembicine wasps are
adapted for digging in so hard a substrate.

The three small aggregations near Tucuman, Argentina, were in
diverse situations. At Yacochuya, 8 km northwest of Cafavate, Salta,

336

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some 20-30 females nested in a bare area measuring about 4x8m,
which had been cleared of rocks and vegetation, possibly for pro-
jected agriculture, in scrubby woodland in the foothills. The soil
was a moderately friable, sandy loam. One additional nest was
located in much harder and coarser soil in a schoolyard 0.5 km
away. At Santa Maria, Catamarca, several nests were discovered
in bare, firm, sandy loam in vacant lots in the city. At Las Termas
de Rio Hondo, Santiago del Estero, six nests were found in coarse,
hard-packed, sandy clay close to the banks of the Rio Dulce.

The three sites in Argentina were all in areas of low rainfall, and
were notably dry during the period of study. Collecting records
suggest that Rubrica is active here only during the warm summer
months (chiefly December-April) . In Trinidad, wasps were col-
lected at almost all times of the year. Nesting activities, however,
occurred mainly toward the end of the wet season and during the
dry season, that is, December through May.

Male Behavior

In Argentina, males were frequently seen visiting various flowers,
along with many females, but we observed no mating here. At
Yacochuya, two males were seen consistently in the nesting area, and
these males made several attempts to mate with females that were
digging or opening or closing their nests. Much of the time these
males flew about close above the ground and perched here and there
with their wings partially extended. We saw no males at all in the
large aggregation at Cali, Colombia.

At St. Augustine, Trinidad, males were occasionally seen flying
in the immediate vicinity of the nesting site. More usually they were
seen resting and apparently sunning themselves on nearby vegetation.
Similar observations were made by Vesey-Fitzgerald (1940), who
also reported that males and females rest on vegetation at night, the
two sexes separately, the males “in groups of many individuals to-
gether.” Bodkin (1917) also reported large sleeping clusters, of
unspecified sex, in Guyana. At Cali, Colombia, we searched sur-
rounding vegetation carefully in the evening, but were unable to find
such clusters. It was clear, however, that females do not spend the
night in the nests and males do not dig “sleeping burrows,” as is
the case in Bembix.

Nest Construction

Females dig effectively through the firm and often coarse substrate,

1974] Evans , Matthews £sf Callan — Rubrica surinamensis 337

using the mandibles to cut through the soil and to haul out small
stones, the front legs to scrape loosened soil beneath the body
(Fig. 1). Apparently the better part of one day is required for
completion of a nest. When the burrow and cell are complete, the
female partially disperses the mound of soil and constructs a rounded
pile of soil over the entrance. This pile serves as a perch to which
she returns intermittently as long as the nest is active (Fig. 3).
Since this “pile-building” behavior is, so far as we know, unique,
we present in some detail our notes on a typical individual at Cali
(RWM, note C6).

This female was observed digging steadily from 1030 to 1100
hours on 11 January 1972. She then came out, made a closure, and
flew off, leaving the accumulated soil intact. She returned more
than an hour later, opened the nest, and resumed digging. She spent
much time in the burrow, from time to time emerging and scraping
soil onto the mound, then re-entering to dig further. At 1358 she
came out, closed the entrance, and walked slowly to the end of the
mound, where she turned in a circle several times while scraping
sand in all directions. After spending 3 minutes leveling in this
manner she flew off, returning 5 minutes later with a small fly. In
25 seconds she emerged from the nest, turned around outside and
immediately re-entered, remained inside 35 seconds, then came out
again. This sequence of going in and coming out was repeated 1 1
times. (Several other females were seen to exhibit similar behavior,
which we assume to play a role in familiarizing the wasp with the
new nest.)

Following this the female made a strong outer closure, packing the
soil with vigorous blows of the tip of the abdomen, and then resumed
leveling. She did this by walking to the end of the mound while
scraping soil back toward the entrance, then turning around and
walking to the entrance without scraping; then turning around again
and walking to the end of the mound scraping soil. This was re-
peated many times over a 12-minute period, the result being removal
of most of the soil to a heap over the entrance. The wasp dragged
several pebbles onto this pile and periodically tamped it with the tip
of her abdomen. Eventually she made several short flights ; each time
she returned adding a little more soil to the pile, which now meas-
ured 35 mm X 25 mm X 7 mm deep in the center.

We observed similar behavior on many occasions at Cali. Often
the pile was surrounded by a ring of smooth soil, where the loose
surface soil had been scraped upon it. The original mound, resulting
from the digging of the nest, was sometimes partially in evidence,

338

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Figs. 1-3. Rubrica surinamensis females nesting in a hard-packed gravel
road near Cali, Colombia. (1) Digging the nest. (2) Entering the nest
through the entrance pile, carrying a paralyzed fly beneath the abdomen.
(3) Resting on the entrance pile during an inactive period.

1974] Evans, Matthews & Callan — Rubrica surinamensis 339

often forming a rather flat, fan-shaped area as much as 16 cm wide
and extending as much as 14 cm from the entrance. The pounding
movements, made with the tip of the abdomen when making the pile
and at each nest closure, were sometimes audible up to 3 m away.
The entrance piles were used as periodic perches to which the females
returned off and on throughout the day. We rarely observed aggres-
sion between females on adjacent piles, and we do not regard them
as territorial perches.

Many details of this behavior were confirmed in Argentina, but
some differences were noted. At Yacochuya, leveling of the mound
seemed not to occur, the wasps merely scraping soil from the im-
mediate vicinity of the hole when making the pile over the entrance.
Thus each nest came to be marked by a small pile surrounded by a
smooth area 2-4 cm across, and finally, on one side, the remains of
the mound, usually 7-8 cm wide X 10-13 cm long X 1-2 cm deep.
We noticed several nests here that had short grooves or “back bur-
rows” into the mound, only 1-2 cm deep. These were made at the
time of temporary or final closures, and appeared to represent quar-
ries for obtaining soil for fill. One nest at Las Termas, still active
and containing a half-grown larva, had a well-defined accessory bur-
row 4 cm long, about 1.5 cm from the true burrow and at a right
angle to it.

Nest Structure

The nest is a simple, oblique burrow with a terminal cell. All who
have worked on this species have found the burrow to be very short
(summary in Evans, 1966). Our studies confirmed this. One nest
at St. Augustine had a burrow about 15 cm long ending in a cell
about 3.5 cm long. Burrows at Cali averaged even shorter than this,
although burrows at all three localities in Argentina were somewhat
longer (Table 1). We attribute this to the fact that the soil at the
Cali site was unusually hard. We found that cells when first made
measured about 15 mm in diameter by 25-30 mm in length, but as
the larva grew tended to be lengthened to as much as 4 cm.

In contrast to previously published observations, we found that
nests were often multicellular. That is, after completing the pro-
visioning of the initial cell, the female often made another cell from
a short side-burrow of the same nest. Of the 22 nests excavated at
Cali, 8 had one cell, 8 had two, 5 had three, and 1 appeared to have
five (but we are not certain that all five belonged with this nest).
The cells of any one nest were at about the same depth and were

340

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Table 1. Nest structure of Rubrica surinamensis

Locality

No.

nests

% multi-
cellular

Burrow
length (cm) *

Cell

depth (cm)*

Cali, Colombia

22

64

11.8 (9-16)

7.3 (5-9)

Cafayate, Argentina

12

17

16.8 (15-18)

9.5 (8-11)

Santa Maria. Argentina

3

0

16.0 (15-18)

9.2 (8.5-10)

Las Termas, Argentina

6

0

16.3 (14-22)

10.5 (8-13)

* Mean and range of variation.

separated by only 1-4 cm of soil. It is, of course, possible that the
unicellular nests we dug out might have had more cells added (ex-
cept for one that had received the final closure) . Four of the 2-
celled nests and two 3-celled nests had received the final closure
when excavated. In multicellular nests, the earlier cells invariably
contained cocoons, but it was usually possible to trace the filled bur-
rows to these cells because the fill was not as compact as the sur-
rounding soil.

However, only 2 of 21 nests in Argentina were bicellular and none
had more than two cells ; one of the 2 bicellular nests was in the hard
soil of the schoolyard at Yacochuya. There is a general tendency for
wasps nesting in firm soil to make shallower nests and to prepare
more than one cell per nest. Thus it would be rash to assume that
these differences between the Colombian and Argentinian populations
had a genetic basis.

There is no evidence that an inner closure, separating cell from
burrow, is maintained at any time. However, an outer closure is
normally made whenever the female leaves the nest. We noted no
exceptions to this in Colombia or Argentina, but occasional individu-
als in Trinidad did omit the outer closure. This was also noted in
Trinidad by Vesey-Fitzgerald (1940), who believed that closure was
usually omitted when the larva was large.

Provisioning the Nest

There are many published records of Rubrica surinamensis preying
upon flies, and our own records confirm the fact many kinds of flies
of small to moderate size are accepted (Table 2). The flies are
evidently killed by the sting. The egg is laid erect on the side of
the first fly placed in the cell, and provisioning is progressive.

The flies taken are apparently those readily available near the
nesting site. Belt (1874) observed females pursuing horseflies around

1974] Evans, Matthews J & Callan — Rubrica surinamensis 341

livestock and humans, and Callan (1945) reported the capture of
engorged stable-flies, one nest having been provisioned entirely with
stable-flies. The frequent use of stable-flies and Tabanidae in Trini-
dad probably reflects the fact that the cattle pens of the Imperial
College of Tropical Agriculture were not far away. All specimens
of Tabanidae and Muscidae from this site were females. On the
other hand, the records of Syrphidae included males as well as fe-
males, suggesting that they were taken on flowers. There were many
cattle near the nesting site at Cali, again possibly serving as a source
of Muscidae, although no tabanids were found to be used as prey at
this site or at any of the localities in Argentina.

We observed several females hunting beeflies at Santa Maria,
Catamarca, Argentina (RWM, note no. C38). These wasps flew
about flowers growing in the sand, apparently hunting flies visually.
Having located a beefly, the wasp would hover some 30-40 cm be-
hind it, about 15 cm above the sand, and follow the fly until it
landed on a flower. Then it would dash suddenly at the fly, some-
times producing an audible clash. However, although more than 20
attempts to take beeflies were observed, in every case the fly darted
almost straight up at the last moment and escaped from the wasp.
The infrequent use of beeflies as prey (Table 2) may reflect their
lack of success with flies of this group. At Santa Maria, Rubrica
females were also seen to dash after Microbembex wasps visiting the
flowers, again unsuccessfully.

The way in which the prey is carried to the nest is similar to that
of other bembicine wasps. It is held firmly by the middle legs,
ventral side up and with the head forward. Since the wasp must dig
through the small pile of soil over the entrance, as much as 20-60
seconds may be required to gain entry (Fig. 2). As the wasp enters,
the prey slides backward and the grasp is shifted to the hind legs,
whereupon wasp and prey disappear down the burrow. However,
flies are occasionally dropped or become stuck in the entrance. In
such cases the wasp enters the nest, turns around inside, and comes
out and drags the prey in with her mandibles. On leaving, the wasp
makes a quick closure, then flies about briefly and returns one or more
times to scrape soil over the entrance.

Both at St. Augustine, Trinidad, and Cali, Colombia, we noted
instances in which females carrying prey were pounced upon by
other females. This usually resulted in the females struggling on
the ground and ended by one escaping and diving precipitously into
her nest with the prey.

342

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Table 2. Dipterous records for Rubrica surinamensis

Species of fly

Number of specimens

Trinidad

Colombia Argentina

STRATIOMYIDAE

H edriodiscus chloraspis Wiedemann

1 2

H. pulcher Wiedemann

1 64

Hermetia illucens L.

6

2

Hoplitimyia fasciata F.

4 8

H. subalba Walker

5

Labostigmina inermis Wiedemann

15

Stratiomys connexa van der Wulp

4

TABANIDAE*'

Esenbeckia prasiniventris Macquart

1

Leucotabanus exaestuans L.

1

Tabanus nebulosus DeGeer

2

T. claripennis Bigot

4

T. lineola var. carneus Bellardi

2

T. colombensis Macquart

4

BOMBYLIIDAE

Villa sp.

4

SYRPHIDAE

Eristalis erraticus Curran

3

E. sp. nr. fasciatus Wiedemann

4

E. tenax L.

I

E. testaceicornh Macquart

1

E. vinetorum F.

1

E. sp.

8

Quichuana aurata Walker

3

Volucella obesa F.

9

7

MUSCIDAE

Morellia scapulata Bigot

1

Musca dome stic a L.

8

1

Stomoxys calcitrans L.

5

3

SARCOPHAGIDAE

Oxysarcodexia sp.

1

Sarcophaga spp.

1

1

CALLIPHORIDAE

Cochliomyia macellaria F.

4

Phaenicia sp.

2

TACHINIDAE

Cylindromyia sp.

1

Protogoniops sp.

2

Ptilodexia sp.

9

* Several of these records of Tabanidae were published by Bequaert
(1944). His names are here updated according to Fairchild and Aitken
(I960).

1974] Evans , Matthews & Callan — Rubrica surinamensis 343

Use of Unusual Prey

Despite the many records of R . surinamensis taking flies (there
are numerous published records as well as over 200 new records
summarized in Table 2), we are now able to report 4 records of the
use of non-dipterous prey, 2 from Trinidad and 2 from Argentina.
We assume that these were in the nature of “mistakes/’ that is, that
the wasp struck them in the course of fly-hunting and for some reason
failed to discriminate them from their more usual prey. Since some
of the Lepidoptera were found in cells only as wings, it is obvious
that the bodies had been accepted as food by the larvae.

On 2 December 1941, at St. Augustine, Trinidad, EMC ob-
served a female flying toward her nest carrying a dragonfly, Perithe-
mis moona Kirby (Odonata, Libellulidae) . This is a yellow-winged
species with a body length of about 25 mm. As the prey was held
well back, probably by the middle legs, it extended some distance
behind the abdomen of the wasp. The wasp was caught with her
prey, which was dead; the nest was unfortunately not located. On
11 March 1947, at St. Augustine, E. A. Fitzpatrick took a female
that was carrying a small skipper butterfly, Panoquina sp. (Lepi-
doptera, Hesperiidae) . The prey was dead when caught.

We found the wings of Lepidoptera in two nests at Las Termas,
Santiago del Estero, Argentina (HEE, note nos. C63, 64). One
nest contained the wings of a skipper ( Monca sp., Hesperiidae)
along with 3 flies and a half-grown larva. The other nest, only 2
m away, contained the wings of two moths (both probably Loxostege
sp., Pyralididae) . This cell contained many flies and a nearly full-
grown larva.

Cell Cleaning

As noted in the introduction, two observers in Argentina have
seen females flying from their nests carrying the remains of flies or
bits of debris. However, this has not been reported by other workers.
Although our studies of the aggregations in Trinidad and Colombia
were fairly detailed, we did not observe this behavior, and the
numerous nests excavated at Cali usually contained many fly re-
mains with the larva or cocoon.

However, our experience at Yacochuya, near Cafayate, Argentina,
was quite different. When we discovered this small aggregation on
the morning of 2 February 1972, a number of females were seen to
fly from their nests and drop small objects some distance away.

344

Psyche

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Studies over the next few days revealed that most females cleaned
the cells of uneaten prey or pieces of prey each morning before they
began provisioning. Wasps backed out of their burrows, holding the
debris in their mandibles, then flew off 2-6 m, ascending to a height
of 1-2 m, and dropped it on the ground. Many females flew over a
nearby hedge and dropped their load on the other side. It appeared
that smaller pieces of prey were dropped closer to the nest than
larger ones (sometimes whole, uneaten flies), although we have no
quantitative data to support this. Cell cleaning usually required only
5-8 minutes, during which time 3-6 loads were carried from the nest.
The following notes on one nest (HEE, C55-2) may serve as an
example.

This female first entered her nest at 1106 hours, clearing the
entrance as she entered. After a minute or two she backed out with
a large, whole fly which she carried on the wing a distance of 4 m
and dropped. She returned to the nest quickly, this time remaining
inside only 10 seconds and emerging with a fly which she carried
6 m and dropped from a height of 1.5 m. On the next trip she re-
mained in the nest 10 seconds and emerged with a small particle
which she dropped 3 m away from a height of 1.5 m. On the fourth
and final trip she remained in the nest 8 seconds and emerged with
a very small particle which she dropped quite close to the nest. She
then returned to the nest, closed the entrance (having left it open
while cleaning), and flew off at 1114. She returned with her first
fresh prey at 1 1 1 7.

We did not observe cell cleaning at Santa Maria or Las Termas,
Argentina, but our observations were not prolonged in either site,
and were mostly made in the afternoon. Most of the nests we ex-
cavated at these sites contained little debris, strongly suggesting that
cell cleaning had occurred earlier in the day. We are under the
impression that cell cleaning is a common feature in populations in
western Argentina, as it evidently is in the Buenos Aires region
(Brethes, 1902; Llano, 1959).

Final Closure

Several final nest closures were observed at Cali, Colombia. In
each case the female emerged from the entrance repeatedly while
scraping soil behind her; she often moved as much as 4 cm away,
scraping soil into the hole, then either backed in or turned around
and walked to the burrow, turning around in the entrance and back-
ing in. In no case were females seen to take flight and land in the

1974] Evans, Matthews J & Callan — Rubrica surinamensis 345

entrance, then turn around and back in, as occurs commonly in other
sand wasps. Much use is made of the mandibles in biting soil around
the entrance, the wasp often producing a loud buzz in the process.
When the burrow is nearly full, it can be seen that the soil is
packed in with rapid blows of the tip of the abdomen, which is
turned downward and forward and moved in a somewhat circular
pattern. Finally, small pebbles may be dragged backward over the
entrance.

Nest Associates

The fly Raviniopsis spinosa (Hall) (formerly in the genus Pachy-
graphomyia ) (Sarcophagidae) , is a constant nest associate of Rubrica
surinamensis in Trinidad, where it was first recorded by Vesey-
Fitzgerald (1940). It has been noticed by EMC many times in
Trinidad, and also on the Paria Peninsula of Venezuela (Callan,
1954). The fly was also common at Cali, Colombia^ usually to be
seen perched near a nest entrance on the alert for a wasp returning
with prey. One was seen to dash quickly upon a fly just being
brought into a nest, presumably larvipositing upon it. Six of 37 cells
dug out at Cali contained maggots, 2-4 per cell. In every case the
wasp larva appeared healthy, and several had attained full size.

None of the maggots found in cells at Cali were reared through
to adulthood, but at St. Augustine, Trinidad, on 14 March 1943,
EMC found a nest containing a maggot along with a large wasp
larva. The maggot was fed on dead houseflies and pupated on 16
March, producing an adult R. spinosa on 28 March. On another
occasion, at St. Augustine, a female R. spinosa was captured and 21
minute first instar larvae were squeezed from her abdomen. Some
of these larvae were placed on a large Rubrica larva, but they made
no attempt to attack it. Five of the maggots were placed separately
in small glass vials. Two or three dead houseflies were put in each
vial and others were subsequently added. Two of these larvae eventu-
ally pupated, producing adult R. spinosa 12-14 days later.

Other species of Sarcophagidae evidently also act as inquilines in
Rubrica nests. One of several flies taken from nest entrances at Cali
was identified as Sarcophaga sp. Three flies reared from nests at
St. Augustine on 2 December 1941 proved to be Sarcodexia sterno-
dontis Townsend. It should be noted that Sarcodexia, Sarcophaga,
and Raviniopsis are all members of the subfamily Sarcophaginae,
although it is Miltogramminae that are usually found associated with
the nests of Hymenoptera.

34 6

Psyche

[June

Curiously, we found none of these Sarcophaginae in any of the
study areas in Argentina. One fly noted trailing a Rubrica that was
digging in a road near Yacochuya proved to belong to the genus
Senotainia , of the Miltogramminae. One of 14 cells excavated at this
locality contained 2 small maggots in debris surrounding a cocoon,
but we did not rear these successfully. Brethes and Llano made no
note of inquilinous flies in eastern Argentina, and Brethes noted that
the cells he excavated contained little in the way of debris, doubtless
the result of cell cleaning.

A minute fly of the family Chloropidae, Liohippelates pusio
(Loew), was also recorded by Vesey-Fitzgerald (1940) in asso-
ciation with R. surinamensis in Trinidad. This fly has also been
observed at St. Augustine in the immediate neighborhood of nests
and also entering open burrows in the absence of the wasp. This
fly is no doubt also a nest scavenger, and its larvae probably feed on
the uneaten remains of prey left by the wasp larva.

Structure of Cocoon

Ten cocoons of R. surinamensis were collected at Cali, Colombia,
2 more near Cafayate, Argentina. The cocoons from the two lo-
calities are very similar, and the following notes pertain to all 12.
The cocoons are ovoid, more pointed at the posterior end, and have a
hard wall made up of soil particles and salivary secretions, as usual
in Bembicini. They vary in length from 25 to 28 mm (mean 26.7
mm) and in maximum width from 8.5 to 10 mm (mean 9.6 mm).
The pores are more or less evenly distributed around the widest part
and number from 4 to 6 (actually 5 of the cocoons we collected have
4, 4 have 5, and 3 have 6). Each pore is complex, being darkened
and strongly elevated, some of them rising as much as 0.4 mm above
the cocoon surface; each is somewhat bilobed, each lobe having
several small protuberances, each of which terminates in a minute
opening.

Brethes (1902) reported only 2 pores per cocoon, an unusually low
number for so large a cocoon, causing the species to occupy an anoma-
lous position in the diagrammatic representation of cocoon size versus
number of pores presented by Evans (1966, Fig. 209). Even 4-6 is
a low figure for a cocoon of this size, but the fact that each pore has
numerous small apertures may compensate for this.

Structure of the Mature Larva

There is no detailed published description of the larva of this

1974] Evans j Matthews t & Callan — Rubrica surinamensis 347

Figs. 4-9. Structures of full-grown larva of Rubrica surinamensis. (4)
Head, anterior view. (5) Antennal orbit and papilla. (6) Labium, oral
surface. (7) Body, lateral view. (8) Spiracle of most anterior pair. (9)
Labrum (left) and epipharynx (right).

348

Psyche

[June

species, and since Rubrica is a somewhat unusual genus with respect
to adult structure and behavior, it seems worth presenting a descrip-
tion of larval characters. Our description is based on 15 full-grown
larvae from Colombia and Argentina. We noted no variation worthy
of note and no differences between larvae from the two widely sep-
arate areas.

Length 30 mm ; maximum width 8 mm. Body robust, more tapered
anteriorly than posteriorly; terminal segment blunt, supra- anal lobe
surpassing subanal; pleural lobes well developed; middle body seg-
ments weakly divided into two annulets dorsally; each segment except
last two with a transverse ventral swelling (Fig. 7). Spiracular atria
about .20 mm in diameter except second pair only .16 mm in diame-
ter; external opening of atrium small, about .08 mm in diameter
(second pair .07 mm), peritreme very narrow; walls of atrium lined
with many rings of minute spinules; opening into subatrium large,
armed with a strong circlet of spines (Fig. 8). Integument smooth,
visibly setose only under high power; each segment dorsally with a
few minute setae, longest measuring only .06 mm; parts of venter
with sparse, weak spinules.

Head 1.9 mm wide, height (exclusive of labrum) 1.9 mm (Fig. 4).
Parietal bands absent; antennal orbits very small, only .10 mm in
diameter, papillae short, about .04 mm long, only slightly longer
than basal width (Fig. 5). Front deeply impressed just mesad of
each antennal orbit. Head with scattered minute punctures, some of
them giving rise to minute setae, especially on clypeus. Labrum
.70 mm wide, emarginate medially, disc with more than 40 rather
strong setae as well as a number of non-setigerous punctures, apical
margin strongly bristly, paralleled by a row of about 40 barrel-
shaped sensilla.; epipharynx strongly spinulose, but with a bare cen-
tral band which bears six sensory pores on each side of the: midline,
also with several additional pores above the central band (Fig. 9).
Mandibles .95 mm long, .5 mm wide at the base, tridentate and
with a weak fourth tooth between the apical two; sides with several
minute setae. Hypopharynx and mesal margins of maxillae densely
spinulose; maxillae with several lateral setae; maxillary palpi and
galeae subequal in length, the latter more slender, somewhat horn-
shaped. Oral surface of prementum mostly papillose, with a few
spinules laterally and basally; spinnerets barely exceeding labial
palpi (Fig. 6).

The larva of Rubrica differs from that of nearly all other Bem-
bicini in lacking all evidence of parietal bands and in having the
spinules on the prementum confined to the sides and base. The short

1974] Evans , Matthews J & Callan — Rubrica surinamensis 349

antennal papilla and almost total lack of body setae and spinules
suggest Steniolia. On the whole, larval characters suggest that
Rubrica may not be as closely related to Bembix and to Stictia as
might be assumed on the basis of adult structure.

Discussion

The somewhat isolated position of Rubrica among the genera of
Bembicini is suggested not only by larval characters but by a number
of behavioral peculiarities, of which the following five are especially
noteworthy.

( 1 ) Females make a small pile of earth and pebbles over the nest
entrance, to which they return and use as a perch from time to time
when not engaged in other activities. Members of other genera that
spend inactive periods away from the nest tend to return occasionally,
but usually hover briefly over the entrance (e.g. Steniolia).

(2) Although R. surinamensis is a predator on flies, there are
several records of females taking Lepidoptera (Hesperiidae, Pyra-
lididae) and one record of a dragonfly being used as prey (Libel-
lulidae). Evidently the females do not discriminate as sharply as
do most other Bembicini. For example, the many hundreds of records
for North American Bembix include no non-dipterous prey (Evans,
I957> 1966). On the other hand, Janvier (1928) found that the
fly predator Zyzzyx chilensis (Eschscholz) occasionally takes a skipper
(Hesperiidae). Manfredo A. Fritz has informed us (in litt.) that
he has taken skippers from the nests of the fly predator Trichostictia
guttata (Taschenberg) in Argentina. It should be noted that Lepi-
doptera form the normal prey of members of the genera Edit ha and
Stictiella. Use of Odonata as prey is more unusual, but two Aus-
tralian species are now known to prey on damselflies, at least one of
them also on flies (Evans and Matthews, 1973). Evidently there is
enough similarity in the flight characteristics of Lepidoptera and
Odonata to those of Diptera that the switch-over is not difficult.

Lin (1971) recently discovered that some members of an aggre-
gation of 2000 females of Stictia Carolina Fabricius in Oklahoma
preyed not only upon skippers but upon small brown cicadas (Ho-
moptera), although flies constitute the usual prey of this species.
He postulated that competition for food in this dense population
caused females to accept unusual prey.

(3) Cell cleaning by flying from the nest and dropping the debris
some distance away is well marked in Argentinian populations, but
it has not been observed in northern South America. It is perhaps

350

Psyche

[June

no coincidence that dipterous nest scavengers appear much more
prevalent in Trinidad, Venezuela, and Colombia than they are in
Argentina. We assume that cell cleaning has evolved in response
to these nest associates, which probably often cause the female to
bring in additional flies and thus prolong the nesting cycle. Cell
cleaning is known in several species of Bembix (Evans, 1966, pp. 358,
451), although none of these wasps are known to carry the debris as
far as Rubrica carries it.

(4) The mature larva constructs 4-6 pores in the walls of the
cocoon, but these pores are complex, each consisting of a bilobed
elevation containing several minute perforations. The construction
of these pores has not been observed in Rubrica , and the function of
the pores has not been clearly demonstrated in any bembicine wasps.

(5) Females spend the night away from the nest, and they have
been reported to cluster on vegetation, although we have not ob-
served this. Clustering is known in several genera of Bembicini,
especially Steniolia and Zyzzyx. Species of Stictia and Bicyrtes also
spend the night on vegetation, although in all known species of
Bembix the female sleeps in the nest, and in Microbembex the female
sleeps in a short burrow apart from the brood nest.

Thus on the whole behavioral and larval features present a some-
what ambiguous picture as to the relationships of Rubrica to other
genera, although they do emphasize that it is a distinctive taxon not
closely related to Bembix. Of particular interest is the intraspecific
variation in the behavior of R. surinamensis. The occasional use of
non-dipterous prey suggests a loosening of the stereotypy of hunting
behavior, in the direction of the broad-spectrum prey selection of
some species of Glenostictia or of the total lack of specificity of all
species of Microbembex. Cell cleaning in R. surinamensis appears
to be an example of geographic variation in behavior, populations in
the southern parts of the range apparently having evolved a mechan-
ism for reducing the success of nest scavengers that northern popu-
lations have not evolved (at least so far as presently known). Nest
depth and number of cells are probably related to hardness of the
substrate, shallow nests with several cells being an adaptive response
to very hard soil, reducing the time and wear involved in nest con-
struction. All of these points are deserving of further study by per-
sons with aggregations of the species at their disposal. Rubrica
surinamensis is evidently one of the most successful of South Ameri-
can solitary wasps, and a more thorough knowledge of its biology
may teach us much about the origin and adaptiveness of behavioral
attributes.

1974] Evans , Matthews & Callan — Rubrica surinamensis 351

Acknowledgments

We are indebted to various specialists of the Systematic Ento-
mology Laboratory, U.S. Department of Agriculture, for identifying
the Diptera: R. Gagne, L. Knutson, C. W. Sabrosky, A. Stone, and
W. W. Wirth. Dr. H. de Souza Lopes, of Rio de Janeiro, kindly
identified the Raviniopsis spinosa. J. C. Bequaert identified some of
the Trinidad Tabanidae, and the late R. C. Shannon made pre-
liminary identifications of many of the Trinidad Diptera. The
dragonfly (Odonata) was identified by D. C. Geijskes, the Hes-
periidae by J. M. Burns. Evans and Matthews would like to ex-
press their indebtedness to Drs. William and Mary Jane Eberhard,
who served as their hosts in Colombia, and to Dr. A. Willink, who
assisted in many ways during their research in Argentina. The
wasps, prey, and associates collected during these studies have been
deposited at the Museum of Comparative Zoology, Harvard Uni-
versity. Evans and Matthews’ studies were supported by the Na-
tional Science Foundation, U.S.A., grant GB 8746.

Literature Cited

Belt, T.

1874. The Naturalist in Nicaragua. John Murray, London. 403 pp.
Bequaert, J. C.

1944. Further studies of the Tabanidae of Trinidad, B.W.I. Psyche
51: 12-21.

Bodkin, G. E.

1917. “Cowfly tigers”, an account of the hymenopterous family Bem-
becidae in British Guiana. Jour. Bd. Agri. Brit. Guiana 10: 119-
125.

Brethes, J.

1902. Notes biologiques sur trois Hymenopteres de Buenos Aires. Rev.
Mus. La Plata 10: 193-205.

Callan, E. McC.

1945. A wasp preying on house-flies and stable-flies. Nature 155: 146.
1950. Observations on tropical wasps in Trinidad. Proc. 8th Internat.

Congr. Ent. Stockholm pp. 204-206.

1954. Observations on Vespoidea and Sphecoidea from the Paria
Peninsula and Patos Island, Venezuela. Bol. Ent. Venez. 9: 13-27.
Evans, H. E.

1957. Studies on the Comparative Ethology of Digger Wasps of the
Genus Bembix. Comstock Publ. Assoc., Ithaca, N.Y. 248 pp.

1966. The Comparative Ethology and Evolution of the Sand Wasps.
Harvard Univ. Press, Cambridge, Mass. 526 pp.

Evans, H. E., and R. W. Matthews

1973. Systematics and nesting behavior of Australian sand wasps.
Mem. Amer. Ent. Inst. 20: 1-387.

352

Psyche

[June

Fairchild, G. B., and T. H. G. Aitken

1960. Additions to the Tabanidae (Diptera) of Trinidad, B.W.I. Ann.
Ent. Soc. Amer. 53: 1-8.

Janvier, H.

1928. Recherches biologiques sur les predateurs du Chili. Ann. Sci.
Nat., Zool. (10)11 : 67-207.

Lin, C.S.

1971. Bionomics of Stictia Carolina at Lake Texoma, with notes on
some neotropical species (Hymenoptera : Sphecidae). Texas Jour.
Sci. 23: 275-286.

Llano, R. J.

1959. Observaciones biologicas de insectos bonaerenses. Supl. Rev.
Educ. Prov. Buenos Aires, La Plata, Argentina. 136 pp.
Vesey-Fitzgerald, D.

1940. Notes on Bembicidae and allied wasps from Trinidad (Hym. :

Bembicidae and Stizidae). Proc. R. Ent. Soc. London 15(A):
37-39.

A NEW SPECIES OF OPITHES FROM MEXICO
WITH A KEY TO THE SPECIES
(COLEOPTERA: STAPHYLINIDAE ) *

By Ian Moore
Staff Research Associate
Division of Biological Control
University of California, Riverside

The generic name Ophites was validated by Erichson ( 1839, P* 29)
by inclusion in a key to the genera of the tribe Paederini. In the
second half of the same work (1840, p. 627) Erichson described in
detail the genus and three included species from Colombia. Sharp
(1876) added a new species from Brazil and Lynch-Arribalzaga
(1884) added another from Argentina. In 1901 Fauvel described a
sixth species from Colombia. In 1952 Blackwelder called attention
to the fact the name Ophites was preoccupied by Wagler, 1830, and
proposed the substitute name Opithes.

In 1904 Fauvel described from Brazil two genera, Mimophites
and Bolbophites , which closely resemble Opithes. Members of each
of these genera have the slender neck and narrow pronotum of
Opithes. Blackwelder in 1944 placed the former next to Ophites
( — Opithes) and the latter far removed in the subtribe Echiasteres.
Seevers (1965) reviewed Mimophites and Bolbophites. He stated
“Mimophites appears to belong to Casey’s (1905) subtribe Stilici, a
group included in Blackwelder’s (1944) Lathrobii.” In Opithes
the first antennal segment is as long as the next six combined, more
than one-third of the length of the entire antenna; the gular sutures
are narrowly but distinctly separate; the neck is about one-fourth
the length of the head and the last segment the maxillary palpus is
not subulate but is almost as wide at base as the apex of the third
segment and is conical. Mimophites differs from Opithes in that the
first antennal segment is short, no longer than the next two com-
bined, and the antennae are not anteriorly flexile; the neck is short;
the gular sutures are united and the last segment of the maxillary
palpus is subulate and quite slender. Bolbophites differs from
Opithes in that the first antennal segment is no longer than the next
three together; the gular sutures are united and the last segment of
the maxillary palpus is subulate.

*Manuscript received by the editor June 10, 1974.

353

354

Psyche

[June

Bolbophites is placed in the subtribe Echiasteres because the pro-
sternum is produced posteriorly and thence expanded laterally to the
hypomera, a character which is usually concealed by the anterior
coxae and consequently difficult to see. It is readily distinguished by
the pronotum and elytra being provided with large, conspicuous
knobby protrusions. Mimophites includes several species. Borgmeier
(1949) reviewed the genus and provided a key to the species. Fauvel
(1904) included two species in Bolbophites at the time he described
the genus. No species have been added since.

Blackwelder (1939) included Ophites { — Opithes) in a key to
the genera of the subfamily Paederinae where he indicated its close
relationship to Homaeotarsus by the presence of geniculate, anteriorly
flexile antennae. Specimens of Mimophites and Bolbophites ap-
parently were not available to Blackwelder as he did not include
them in his key.

In this work I describe a new species of Opithes from Mexico and
for the first time present a key to the species of the genus.

Nothing has been recorded concerning the habits of the various
species nor of the habitats in which they are found. Unfortunately,
Mr. Crandall cannot recall the circumstances under which he cap-
tured the specimen described here from Mexico.

1.

2.

5-

6.

Key to the species of Opithes Blackwelder

Head and pronotum ferrugineus. 2

Head and pronotum black or piceus 3

Surface of head anterior to eyes with two foveae, length 12mm.;

Mexico. c ran dalli new species

Surface of head anterior to eyes with three foveae, length 10 mm. ;

Argentina. fauveli Lynch-Arribalzaga

Head and pronotum opaque. 4

Head and pronotum shining. 5

Abdomen densely opaque; Brazil. stilicoides Sharp

Abdomen semi-shining; Columbia. velitarsis Erichson

Tibiae in part pale. 6

Tibiae entirely black; Columbia. versatilis Erichson

Elytra black; Columbia. raphidioides Erichson

Elytra aeneus; Columbia bugnioni Fauvel

Opithes crandalli new species
Description of holotype.

Color . — • Head, pronotum and first four and one-half abdominal

1974]

Moore — New Species of Opithes

355

segments bright ferrugineus with the head slightly darker. Elytra
deep black with the humeri and base piceus and the apices narrowly
flavate. Apical two-thirds of fifth tergite black. Antennae and mouth
parts ferrugineus. Legs pale flavate with the apices of the femora
and the bases of the tibiae piceus, tarsi ferrugineus. Beneath largely
ferrugineus with the mesasternum and metasternum piceus, apical
half of fifth and most of sixth sternites black, the narrow apex of the
latter flavate.

Head. — Head three-fifths as wide as long, widest across the eyes,
the posterior margins of which are a little less than half the distance
from the apex to the base; subparallel in anterior half, thence,
rapidly narrowed to a cylindrical neck which is about one-fifth the
length and one-fourth the width of the head. Anterior margin of
head straight. Supra-antennal ridge prominent. Eye prominent, one-
fifth the length of head. Antenna three-fifths longer than head,
densely pubescent from the third segment, first segment as long as
next six together, second segment twice as long as wide, narrower
than apex of first, third through fifth segments each about two and
one-half times as long as wide, about as wide as second, sixth through
tenth segments each slightly shorter and just perceptibly wider than
the preceding so that the tenth is only about one-fifth longer than
wide, eleventh about as long as wide, pointed in apical third. Upper
surface of head shining, with barely perceptible reticulate ground
sculpture, finely, evenly, sparsely punctured throughout with a fine
short pubescence interspersed with less numerous longer setae; at the
base of each antennal ridge with a prominent fovea between which
is a slight cntral impression, otherwise evenly convex. Each side of
head behind the eyes with two large umbilicate punctures, the first
on the upper surface one-third of the length of the eye from it, the
other on the lower surface about the length of the eye from it.
Under surface of the head impunctate except for about two dozen
scattered fine punctures bearing fine long setae. Gular sutures widely
separated in front, rapidly converging and thence very narrowly but
distinctly separate to the neck where they diverge slightly. Maxil-
lary palpus four-segmented, the first segment short, second long and
slender, slightly arcuate, widest at apex, third slightly longer than
second, hardly wider, widest at apex, almost straight, fourth segment
semi-membranous, almost as wide at base as apex of third, almost as
long as wide, conical with apex rather abruptly acuminate and mem-
branous. Labial palpus three-segmented, first segment about twice
as long as wide, second segment about as wide but half again as long
as first, third segment about as long as first, acicular. Mandibles

356

Psyche

[June

Figure 1. Opithes crandalli Moore, dorsal view.

1974]

Moore — New Species of Opiihes

357

long, slender, falcate, each with two large internal teeth, those of
the left of nearly equal size, those of the right with the proximal
tooth smallest.

Thorax. — Pronotum four-fifths as long as head, a little less than
half as wide as long, widest near the middle, rapidly constricted to
the front where it is about twice as wide as neck, rounded at the
middle and narrowed posteriorly for a short distance, then straight
and nearly parallel to the narrowly rounded basal angles ; base
straight, about one-fifth narrower than widest point; surface evenly
convex, highly polished, impunctate on disc with large scattered
umbilicate punctures along the base and for a short distance forward
along each side. Superior lateral line short, about half the length of
the pronotum, evanescent at each end. Prosternum longitudinally
carinate with a few fine scattered setae. Lateral prosternal carina
obliterated. Mesosternum narrow, not produced between the coxae.
Metasternum long, broad, with an impressed longitudinal central
line, very finely, very sparsely pubescent. Elytra hardly wider than
head, conjointly almost one and one-half times as long as wide,
humerus obtusely angulate with a minute tooth at the angle, sides
nearly straight and nearly parallel, outer apical angles rounded,
apices oblique, inner apical angles obtuse, sutures elevated, disc with
six or seven irregular rows of crowded coarse punctures, interspaces
shining. Legs very long and slender, anterior tarsus not dilated,
without dense spatulate setae beneath. First segment of posterior
tarsus longer than last segment, segments two, three and four de-
creasing in length.

Abdomen. — Parallel, finely, sparsely, punctured, the punctures set
with fine black setae with a few scattered larger setae particularly
along the posterior margin. Last two segments somewhat more
densely punctured than the others. Apical margin of sixth tergite
about one-third of distance on each side produced in a large blunt
tooth, the margin between the teeth broadly evenly arcuately emargi-
nate. Vestiture beneath similar to above. Apex of sixth sternite
arcuate with a very slight emargination in the central eight.

Length. — 12 mm.

Sex unknown but probably a female because of the condition of
the sixth sternite, other species in the male have a deep incision in
the apex of that sclerite.

Mexico, Sinaloa, Mazatlan, December 1966, R. H. Crandall, Jr.,
collector.

Disposition of type. — California Academy of Sciences, San Fran-
cisco.

Figures 2-6. Opithes crandalli Moore. 2, left mandible; 3. right man-
dible; 4, posterior tarsus; 5, sixth tergite ; 6, fourth segment of maxillary
palpus.

1974]

Moore — New Species of Opithes

359

Notes. — This species is very similar in color to fauveli Lynch-
Arribalzaga from Argentina but is without the central fovea on the
disc of the head and has the abdomen more regularly marked with-
out splotches of black.

This striking species is named for its collector, R. H. Crandall,
Jr., who with his father has added many fine specimens to our col-
lection.

Literature Cited

Blackwelder, R. E.

1939. A generic revision of the staphylinid beetles of the tribe Pae-
derini. Proc. U. S. Nat. Mus. 87: 93-125.

1944. Checklist of the coleopterous insects of Mexico, Central America,
The West Indies, and South America, part 1. U. S. Nat. Mus.
Bull. 185: 1-188.

1952. The generic names of the beetle family Staphylinidae with an
essay on genotypy. U. S. Nat. Mus. Bull. 200: i-iv, 1-483.

Borgmeier, T.

1949. Neue Gatungen und Arten termitophiler Staphyliniden aus Bra-
zilien, nebst einem Katalog aller bischer aus der neotropischen
Region beschriebenen Arten (Col. Staph.). Revista Ent., 21:
625-676, illus.

Erichson, W. F.

1839. Genera et species staphylinorum coleopterorum familaea. (part
1), PP- 1-400. Berlin.

1840. Genera et species staphylinorum coleopterorum familaea. (part 2),
pp. 401-954. Berlin.

Fauvel, C. A.

1901. Voyage ide M. de Dr. Ed. Gugnion au Venezuela, en Colombie
et aux Antilles. Staphylinides. Revue d’Entomologie, 20: 251-252.

1904. Staphylinides myrmecdphiles du Bresil Revue d’Entomologie, 32:
276-283, illus.

Lynch-Arribalzaga, F.

1884. Los Estafilinos de Buenos Aires. Bol. Acad. Nat. Sci. Cordova,
7: 5-392.

Seevers, C. H.

1965. The systematics, evolution and zoogeography of staphilinid
beetles associated with army ants (Coleoptera: Staphylinidae).

Fieldiana: Zoology 47: 138-351, figs. 14-37.

Sharp, D.

1876. Contributions to an insect fauna of the Amazon Valley. Trans.
Ent. Soc. London, 1876: 27-424.

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PSYCHE

A JOURNAL OF ENTOMOLOGY

Vol. 8 1 September-December, 1974 No. 3-4

CONTENTS

Host-plant Utilization by Pieris napi Populations in California (Lepi-

doptera: Pieridae). A. M. Shapiro 361

New Protonemura (s.l.) from Nepal (Plecoptera: Nemouridae).

P. P. Harper 367

The Eyeless Catopocerus Beetles (Leiodidae) of Eastern North Amer-
ica. S. B, Peck 377

Post-immobilization Wrapping of Prey by Lycosid Spiders of the

Herbaceous Stratum. J. S. Rovner and S. J. Knost 398

Erratum. M. Moglich and B. H oil dob l er 415

Pteralia of the Paleozoic Insect Orders Palaeodictyoptera, Megasecop-

tera, and Diaphanopterodea (Paleoptera) . J. Kukalova-Peck 416

Systematics of the Trapdoor Spider Genus Aliatypus (Araneae: An-

trodiaetidae). F. A. Coyle 431

A Review of the Agyrtes (Silphidae) of North America. S. B, Peck 501
Application of the Transmission Electron Microscope to the Examina-
tion of Spider Exuviae and Silk. R, F. Foelix 507

Plastral Respiratory Devices in Adult Cryphocricos (Naucoridae:

Heteroptera) . M. C. Parsons and R. J. Hewson 510

Home Ranges of Male Cerceris simplex macrostricta (Hymenoptera,

Sphecidae). J. Alcock and G. Gamboa 528

Additional Notes on Stephanitis takeya in New England (Heterop-
tera: Tingidae). N. S, Bailey 534

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Officers for 1974-1975

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Secretary .

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PSYCHE

Vol. 8 1 September-December, 1974 No. 3-4

HOST-PLANT UTILIZATION BY PIERIS NAPI
POPULATIONS IN CALIFORNIA
(LEPIDOPTERA: PIERIDAE)

By Arthur M. Shapiro1
Department of Zoology, University of California,

Davis, Calif. 95616

Oligophagous or polyphagous insects frequently exhibit ecologi-
cally or geographically complex patterns of host-plant utilization.
Such patterns have recently been documented for the butterflies
Colias alexandra Edwards (Pieridae) (Ellis, 1974) and Euphydryas
editha Boisduval (Nymphalidae) (White and Singer, 1974). This
paper reports a similar situation among California populations of
the Gray- Veined White, Pieris napi Linnaeus (Pieridae) and notes
its potential significance in interspecific competition.

Pieris napi is a circumpolar species; in its extensive boreal and
temperate range it has been recorded on a great variety df plants of
the family Cruciferae. In California it is widely distributed in
foothill and lower montane (Yellow Pine-Incense Cedar-Douglas
Fir) environments in the Coast Ranges south to San Luis Obispo
County and in the Sierra Nevada at least as far south as Yosemite,
but its host preferences have been very poorly documented. In their
survey of Yosemite butterflies, Garth and Tilden (1963) listed in
an Appendix “some plants on which Yosemite butterflies feed as
larvae.” Unfortunately these records, which are mostly not at-
tributed, do not appear to be limited to Yosemite populations. Garth
and Tilden list four Crucifer genera as hosts of P. napi: Barbarea
(winter cress, yellow rocket), Brassica (mustard), Raphanus (rad-
ish), and Dentaria (milkmaids, crinkleroot, toothwort). Of these,
all but Raphanus are known hosts of P. napi in the northeastern
United States. Tilden (1965) recorded P. napi in the “San Fran-
cisco Bay area” on Dentaria only.

This research was financed in part by grant D-804 from the Committee
on Research of the Academic Senate, U.C. Davis.

Manuscript received by the editor August 16, 1974.

36l

362

Psyche

[September-December

Figure 1. Locations of California Pieris napi populations.

1974]

Shapiro — Pieris napi Populations

363

During 1974 a number of Sierran and Coast Range populations
of P. napi were sampled for photoperiodism studies, resulting in the
incidental accumulation of data on host selection. The locations of
these populations are shown in figure 1. The pattern of host selection
proved to be of considerable interest, as reported below.

San Andreas Reservoir , San Mateo County. — This is a coastal,
facultatively bivoltine population from a gully (elevation about 200
feet) subject to summer fog. On 10 April 1974, 58 ova of P. napi
were found by searching 86 plants of Barbarea verna (Mill.) Asch.,
and three females were observed ovipositing on this plant. No ova
were found on 25 plants of Dentaria californica Nutt, growing
nearby in the same canyon. Ova on B. verna were placed on leaves
(mostly on lower surfaces), stems, and pedicels. Most of the Bar-
barea plants had only cauline leaves and were in the early stages of
flowering. Dentaria were in the advanced stages of flowering and
had some green siliques.

Mix and Gates Canyons , Solano County. — This is a strictly uni-
voltine population from summer-arid canyons of the east slope of the
Vaca Mountains, central Inner Coast Ranges, open to the Sacra-
mento Valley (elevations 500 to 2000 feet). On 4 April 1974,
39 ova of P. napi and 21 of Anthocaris sara Reakirt were found on
66 plants of Barbarea verna; the sara ova were mostly in the in-
florescence and on the upper surfaces of leaves, while the napi ova
were scattered as described for San Mateo County, but seldom on
the upper surfaces of leaves. No Pierid ova of any kind were found
on 40 Dentaria californica near the Barbarea. On 19 May 1974,
14 larvae of P. napi were collected on B. verna. They were feeding
on leaves, and the green siliques had not been damaged although
most of the plants were nearly or quite defoliated; only one napi
larva was found feeding on siliques, and this was on a plant whose
leaves and petioles had been entirely consumed. Seven larvae of
A. sara were found on five plants of Sisymbrium officinale (L.)
Scop, (hedge mustard) and two on B. verna. All of these were
feeding on green siliques only, on plants whose leaves were undam-
aged. Four of 20 Dentaria examined had been defoliated, appar-
ently by Pierid larvae, but no larvae were collected.

Near Washington, Nevada County. — Located in the South Yuba
River canyon at about 2600 feet on the west slope of the Sierra, this
is also a univoltine population. On 3 May 1974, 78 ova of P. napi
were collected by the author and S. R. Sims from about two dozen
immature Arabis glabra (L.) Bernh. (tower mustard). The ova

364

Psyche

[September-December

had been laid on both upper and lower surfaces of rosette leaves,
lower surfaces of the (oppressed ) cauline leaves (often near the
point of attachment, as was also true on Barbarea verna ), and on
stems. One larva (3rd instar) was found, and two ovipositing
females were observed. On 22 May, 16 additional ova and 5 larvae
were found at the same spot on the same plants, which were now
mature and beginning to set fruit. Abundant evidence of larval
feeding was found on the rosettes and lower cauline leaves, but the
flowers and siliques were undamaged. Fourteen plants of flowering
Barbarea orthoceras Ledeb. were found in a grassy meadow 0.7 mile
up the canyon, within the area where adult P. napi had been seen,
but no ova, larvae, or feeding damage could be located on them.

Lang Crossing , Nevada County. — This population is at about 4500
feet in the South Yuba River canyon, at or near the upper altidudinal
limit of Sierran P. napi; it is also univoltine. On 22 May 1974,
3 ova of P. napi were found in a stand of 30 immature Arabis glabra
and none on 20 flowering Barbarea orthoceras growing immediately
adjacent to them. On 2 June the same stand was again searched,
producing 11 ova of P. napi , 20 of P. rapae Linnaeus, 3 of i. sara
and 2 of Anthocaris lanceolata Lucas from A. glabra, and 8 of
A. sara from B. orthoceras. An additional 34 ova of P. napi were
collected from A. glabra elsewhere in the vicinity. Four larvae of
P. napi were found feeding on rosette leaves of A. glabra, and two
of A. sara on inflorescences and siliques of B. orthoceras. On 26 June
four mature A. sara and two immature A. lanceolata larvae were
found on siliques of A. glabra and one mature A. sara larva on
siliques of B. orthoceras. Many A. glabra plants showed feeding
damage to the rosettes, but only seven P. rapae larvae were found;
probably most napi had already pupated. Pieris rapae, which is
multivoltine, was flying in abundance at Lang Crossing on 19 July.

Despite the large number of Pierid species and individuals, the
visible impact of feeding on the Crucifers at Lang Crossing was
quite small. In particular, Arabis glabra plants often produced
several hundred siliques on leafy, two- to four-foot stems.

Discussion

There are ten species of obligate Crucifer-feeding Pierids in
California, occurring in various combinations at different localities.
Some of these are spring-univoltine, a few are spring-bivoltine, and
some are multivoltine. How they partition the available Crucifers
among themselves may shed valuable light on the broad problems

1974]

Shapiro — Pieris napi Populations

365

of niche differentiation, competitive exclusion, and species packing.
Emmel (1974) reported three species of inflorescence feeders ( Pieris
protodice Boisduval and LeConte, A. sara and A. lanceolata ) on
Arabis glabra in Riverside County, 2 June 1973. Shapiro (1974)
described five- and six-species Pierid assemblages in two high Sierran
localities and concluded that competition is reduced by behavioral
mechanisms (habitat selection) .

Pieris napi occurs with up to six other Crucifer-feeding Pierids
at the four localities described above. Based on adult collections, the
Crucifer-feeding Pierid faunas of the four localities are (inflorescence
feeders are marked “I”; others primarily leaf feeders) :

San Andreas Reservoir: P. napi, P. rapae , A. sara (I), Euchloe
ausonides Lucas ( I ) .

Mix and Gates Canyons: P. napi , P. rapae, P. sisymbrii Boisduval
( I ? ) , A . sara ( I ) , E. ausonides ( I ) .

Washington: P. napi, P. rapae, A. sara (I), A. lanceolata (I),
E. ausonides (I) .

Lang Crossing: P. napi , P. rapae, P. sisymbrii (I?), A. sara (I),
A. lanceolata (I), E. ausonides (I), E. hyantis Edwards (I).

Not all of these breed in the same microhabitats. At Lang Cross-
ing, for example, P. sisymbrii and E. hyantis are found only in
exposed rocky situations and appear to breed only on Streptanthus,
and are thus not in competition with the woodland species. In
examining the fauna of particular plants in particular habitats, the
division of Pierids into a leaf-feeding and an inflorescence/silique
feeding guild seems paramount. At Mix and Gates Canyons, where
Barb area is obviously in “short supply” and frequently completely
defoliated, at least one species from each guild (P. napi, A. sara)
can occur on this plant. At Lang Crossing the combined visible
impact of four species — two of each guild — on A rabis glabra is
so small that it is tempting to speculate that the populations are
regulated by other factors below the level at which interspecific
competition would be significant.

Despite the potential for interspecific competition, at each of the
study areas one plant received the bulk of the attention from Pierids
while another appeared largely or wholly unutilized. It remains to
be seen whether this reflects nutritional or toxicological unsuitability
of certain Crucifer species. (In unpublished laboratory studies,
Shapiro and F. Slansky ( pers . comm.) have found variation in the
suitability of native and weedy Crucifers as hosts of Pieris rapae

366

Psyche

[September-December

and other species.) It is also possible that host selection by ovi-
positing females is closely tied to host plant phenology, and that
flowering condition of the plants, and possible correlated changes in
mustard oil concentrations, may determine the patterns observed.

References

Ellis, S. L.

1974. Field observations on Colias alexandra Edwards (Pieridae).
J. Lepid. Soc. 28: 114-125.

Emmel, J. F.

1974. in R. L. Langston. Zone 1. Season Summary, Lepid. Soc. News,
25 June, p. 3.

Garth, J. S. and J. W. Tilden

1963. Yosemite Butterflies. J. Res. Lepid. 2: 1-96.

Shapiro, A. M.

1974. Ecological and behavioral aspects of coexistence in six Crucifer-
feeding Pierid butterflies in the central Sierra Nevada. Amer.
Midi. Naturalist, in press.

Tilden, J. W.

1965. Butterflies of the San Francisco Bay Region. Berkeley and Los
Angeles: University of California Press. 88 pp.

White, R. R. and M. C. Singer

1974. Geographical distribution of host plant choice in Euphydryas
editha (Nymphalidae) . J. Lepid. Soc. 28: 103-106.

NEW PROTON EMURA (S. L.) FROM NEPAL
(PLECOPTERA; NEMOURIDAE )*

By P. P. Harper
Departement des Sciences biologiques
Universite de Montreal
C. P. 6128, Montreal, Que.

Described below are the stoneflies of the genus Protonemura
(Nemouridae) collected in 1967 by the Canadian Nepal Expedition
organized by the Entomological Research Institute of the Canada
Department of Agriculture in Ottawa. The genus Protonemura is
understood here in its widest sense (sensu lato) as used by Kimmins
(1946, 1950) and Aubert (1967) ; the genus is obviously in need of
a revision as it contains a wide assemblage of forms some of which
bear larval gills and some which do not. The species considered here
are apparently gill-less as far as can be ascertained from the adults.

The material examined comes from six localities: Godavari at
altitudes of 50°° ft and 6000 ft, Bhurunche at an altitude of 8500
9500 ft, and four localities known only from their geographical co-
ordinates and altitudes: 27°56'N, 85°oo'E at 9900 ft and 10100 ft,
27°57/N, 84°59'E at 10100 ft, 27°58'N, 85°oo' E at moo ft, and
28°oo'N, 85°oo'E at 9900 ft. Most of the specimens were collected
in Malaise traps and therefore the collection dates indicate the period
between visits to the traps.

All the specimens are preserved in alcohol and are deposited in the
Canadian National Collection of Insects in Ottawa. The drawings
have been prepared from cleared specimens. Only the species repre-
sented by male specimens have been named, the others have been
designated by letters to prevent useless future synonymy.

Protonemura mira, n. sp.

Figures 1-4

Length of body 11 ( c? )_I4 (?) mm, to tips of wings 16 ( c? )-
19 ( $ ) mm.

General coloration dark reddish brown; antennae black; anterior
and lateral margins of pronotum light ; wings uniformly brown ; legs
brown, tarsi and distal end of femora darkened ; metafemora also
with a subapical dark ring. Abdomen mostly membranous except for
the genital segments and a pair of sclerotized tergal plates on each
segment in the male. Genitalia reddish brown.

* Manuscript received by the editor November 18, 1974.

367

368

Psyche

[September-December

Figures 1-4: Protonemura mira Harper, n. sp. 1. Terminal segments of

male with paraprocts slightly pulled out (dorsal). 2. The same (ventral).
3. Epiproct (lateral). 4. Terminal segments of female (ventral).

Male genitalia: median lobe of sternum IX long and narrow,
ventral vesicle short. Paraproct: median lamella small, slightly
curved, extending just beyond sternum IX; subanal plate wide, its
process forming two dark heavy hooks ; subanal vesicle short ; external
appendage finger-like, heavily sclerotized and gradually curved.
Cerci long and membranous. Tergum IX with a few spinules on
its posterior margin. Epiproct long and slender with an expanded
tip bearing a double row of ventral spinules.

Female genitalia: subgenital plate sclerotized, wide, with rounded
corners, but not produced posteriorly. Vaginal structures as in Fig-
ure 4, somewhat reminiscent of Protonemura A of Aubert (1967?
Fig. 99), but the subgenital plates are different.

1974]

Harper — New Protonemura

369

Holotype: cT , Godavari, 6000 ft, 17-20. VII. 1967 (CNC type
no. 13430). Allotype: 9, and paratype: 9> same data.

Diagnosis: P. mira clearly belongs to the P. indie a Kimmins group
as defined by Aubert (1967) ; the shape of the process of the para-
procts of the male and the subgenital plate as well as the vaginal
structures of the female will easily separate mira from the other
known species of the group.

Protonemura godavariensis, n. sp.

Figures 5-8

Length of body 10-13 mm, to tips of wings 13-15 mm.

General coloration dark brown. Wings uniformly brown. Legs
brownish, femora with two subapical dark bands. Abdomen brown-
ish, genitalia dark brown.

Figures 5-8: Protonemura godavariensis Harper, n. sp. 5. Terminal seg-
ments of male (dorsal). 6. The same (ventral). 7. Epiproct (lateral).
8. Terminal segments of female (ventral).

370

Psyche

[September-December

Male genitalia: median lobe of sternum IX narrow and of mod-
erate length; ventral vesicle reaching beyond middle of sternum.
Paraproct: median lamella short and pointed, heavily sclerotized;
subanal plate wide, triangular, with broadly rounded angles; distal
process forming a long curved trough-like appendage; external ap-
pendage finger-like, heavily sclerotized externally and bearing a
subapical lateral tooth; the external appendage extends just beyond
the subanal plate. Cerci long and membranous. Terga IX and X
bearing small fields of spinules. Epiproct long and slender, its tip
expanded laterally and bearing ventrally a short double row of
heavy teeth.

Female genitalia: subgenital plate well sclerotized, prolonged pos-
teriorly into a trapezoidal lobe which is broadly and regularly
emarginate. Vaginal structures as in Figure 8.

Holotype: cf, Godavari, 5000 ft, 20-23.VII.1967 (CNC type no.
1 3431); allotype: $, same data; paratypes: cf, $, same locality,
20.VII.-4.VIII. 1967.

Diagnosis: as the preceding species, P. godavariensis belongs to
the P. indica group. The trough-like process of the subanal lobe
of the male as well as the vaginal structures of the female are good
diagnostic characters. The species is very close to P . assami Aubert
and may eventually prove to be the same.

Protonemura funicula, n. sp.

Figures 9-1 1

Length o'f body, 11-12 mm, to tips of wings, 18-19 mm.

General coloration brown ; head and disk of pronotum dark brown.
Wings clear, veins of the front wings marginated with brown. Legs
banded : femora with a median and distal dark band, tibiae narrowly
darkened at base and darkened again distally, tarsi dark. Abdomen
mostly membranous except the genital segments; in the male a pair
of small ventral sclerites and a pair of larger dorsal sclerites are
borne at the base of each abdominal segment.

Male genitalia: sternum IX long with a short rounded median
lobe; ventral vesicle large reaching three quarters of the length of
the sternum. Paraproct: median lamella narrow; subanal plate
partly membranous, irregularly triangular, prolonged distally into
a heavily sclerotized process which is slightly recurved dorsally; this
process is pointed apically and bears a sharp subterminal spine; ex-

1974]

Harper — New Protonemura

371

Figures 9-11: Protonemura funicula Harper, n. sp. 9. Epiproct (lateral).

10. Terminal segments of male (dorsal). 11. The same (ventral).

ternal appendage moderately sclerotized, finger-like, and reaching out
as far as the process. Cerci long and membranous. Terga IX and
X with small median fields of spinules. Epiproct narrow and flat,
asymmetrical distally and prolonged by a very long whip-like process.

Female: unknown.

Holotype: cT , 27°57'N, 84°59'E, 10100 ft, 26-31.V.1967 (CNC
type no. 13432) ; paratypes: cf , 27°56'N, 85°oo'E, 10100 ft, 23-29.
V.1967; cf1, 27°58'N, 85°oo'E, moo ft, 25.V.1967; 3 cT , 28°oo/N,
85°oo'E, 9900 ft, 27.V.-22.VI.1967.

Diagnosis: P. funicula can be placed in the P. filigera (Kimmins)
group of Aubert ( 1 967 ) . The shape of the subanal lobes and the
particularly long appendage of the epiproct are distinctive.

372

Psyche

[September-December

Protonemura adunca, n. sp.

Figures 12-14

Length of body 10-11 mm, to tips of wings, 13-14 mm.

General coloration dark brown; antennae and disk of pronotum
nearly black. Wings amber, veins darker. Legs particularly the hind
pair banded; femora with a median and a distal band. Abdomen
membranous except the genital segments; in the male there is also
a pair of small sclerites at the base of each tergum.

Male genitalia: sternum IX broadly pentagonal, prolonged into a
long and tapering median lobe; this median lobe bears distally an
apical expansion which is outturned at a right angle from the lobe
and forms a beak-like extension ending in two sharp points. Para-
proct: median lamella narrow and rounded apically; subanal plate
sclerotized at base, divided distally into three lobes, the inner lobe
short and rounded, the intermediate lobe membranous, and the outer
lobe long and digitiform, fused laterally with the external appendage;
external appendage fused with subanal lobe for most of its length,
terminating distally in a beak-like point. Cerci long, membranous,
and expanded slightly at tips. Tergum IX emarginate centrally, the
emargination bearing short spinules. Tergum X excavated anteriorly,
and prolonged into two short median lobes covered with spinules.
Epiproct cylindrical, asymmetrical beyond middle, and terminating in
a short curved lobe; a short irregular group of spinules at base of
epiproct.

Female: unknown.

Holotype: cf , 27°57'N, 84°59'E, 10100 ft, 26-31. V.1967, (CNC
type no. 13433); paratypes: cf , same data; cf, 27°58'N, 85°oo'E,
moo ft, 18.V.1967; 4cf, 28°oo'N, 85°oo'E, 9900 ft, 21.V.-22.VI.
1967; 2 cf j Bhurunche, 8500-9500 ft, 23.V.1967.

Diagnosis: P. adunca probably fits into the P. filigera group on the
basis of its epiproct. The shape of the distal end of the median lobe
of sternum IX is quite distinctive and unique.

Protonemura mastigophora, n. sp.

Figures 15-17

Length of body 10 mm, to tips of wings, 13 mm.

General coloration dark brown. Head nearly black (antennae
missing). Wings brownish with some darker clouds. Legs banded,
particularly the hind pair; femora with a dark distal band and a
paler median one. Abdomen membranous except for the genital seg-
ments.

1974]

/ larper — New Protonemura

373

Figures 12-14: Protonemura adunca Harper, n. sp. 12. Terminal seg-

ments of male (dorsal). 13. The same (ventral). Figures 15-17: Proto-
nemura mastigophora Harper, n. sp. 15. Epiproct (lateral). 16. Terminal
segments of male (dorsal). 17. The same (ventral).

374

Psyche

[September-December

Male genitalia: sternum IX long and pentagonal; median lobe
short and inconspicuous; ventral vesicle large. Paraproct: median
lamella long, thin, and extending backwards as far as the subanal
plate; subanal plate terminating in two finger-like lobes, a shorter
median lobe, and a larger lateral lobe; external appendage fused to
lateral lobe of subanal plate and extending three quarters of its
length. Tergum IX produced posteriorly into two asymmetrical lobes
separated by a large median excavation ; each lobe bears a distal patch
of spines. Tergum X normal. Epiproct short and stocky, remark-
ably asymmetrical, and bearing a terminal whip-like appendage.

Female: unknown.

Holotype: cf, 27°56'N, 85°oo'E, ioioo ft, 23-29.V.1967 (CNC
type no. 13493)-

Diagnosis : another member of the P. filigera group, P. mastigo-
phora is close to the species filigera (Kimmins 1946), parafiligera
Aubert 1967 and metafiligera Aubert 1967. It can be best dis-
tinguished from the others by the shape and the sclerotizations of
its epiproct; these are not as well produced in P. filigera the species
it resembles most; Kimmins’ (1946) drawing shows no detail of
the inner structure of the epiproct, but a specimen in the United
States National Museum (det. R. W. Baumann) does not possess
the structures described here for mastigophora.

Protonemura A
Figure 18

In general structure and coloration, this female agrees with P.
funicula and P. adunca; it could be the female of either.

Female genitalia: the subgenital plate is very slightly trilobed.
Vaginal structures as in Figure 18.

Material examined: ? , 27°56'N, 85°OOrE, 9900 ft, 23-29.V.1967 ;
6$, 27°57'N, 84°59'E, ioioo ft, 19-31.V.1967 ; 5$, 27°58'N,
moo ft, 8-24.VI.1967 ; 4?, 28°oo'N, 85°oo/E, 9900 ft,
14.V.-22.VI.1967.

Protonemura B
Figure 19

Total length 10 mm, to tips of wings, 13 mm.

General coloration brown. Wings uniformly amber. Legs banded
in the usual manner.

1974]

Harper — New Protonemura

375

Figures 18-22: Terminal segments of unassociated females (ventral).
18. Protonemura A. 19. Protonemura B. 20. Protonemura C. 21. Proto-
nemura D. 22. Protonemura E.

Female genitalia: subgenital plate small, and rounded posteriorly.
Sternum VII sclerotized medially. Vaginal structures indistinct.
Material examined: $ , 28°0O/N, 85°oo'E, 9900 ft, 27.V.-22.VI.

1967.

Protonemura C
Figure 20

Total length 11-12 mm, to tips of wings, 15-16 mm.

General coloration brown. Wings yellowish-brown, veins dark
brown, marginated with brown. Legs brown with a subterminal
white band.

Female genitalia: subgenital plate short, slightly bilobed. Sternum
VII heavily sclerotized bearing an important posterior nipple-like
protuberance. Vaginal structures indistinct.

Material examined: 22$, 27°58/N, 85°oo'E, moo ft, 2-24.VI.
1967.

376

Psyche

[September-December

Protonemura D
Figure 21

Total length 15 mm, to tips of wings, 18 mm.

General coloration dark brown. Wings blackish. Pro- and meso-
femora light brown with an apical darkening, meta-femora brown
with a subapical light ring.

Female genitalia: sternum VII lightly sclerotized medially. Sub-
genital plate forming posteriorly two pairs of lobes separated by a
wide emargination. Vaginal structures very characteristic, as in
Figure 21.

Material examined: 2?, Godavari, 6000 ft, 14-20.VII.1967.

Remarks: This species evidently belongs to the P. indica group; it
can be separated from P. indica Kimmins 1946 and P. quadridentata
Kimmins 1950 (see Figures 18, 29 & 30 in Aubert 1967) the species
it resembles most by the shape of its vaginal structures and by its
medially pigmented (and sclerotized) seventh sternum.

Protonemura E
Figure 22

Total length 13 mm, to tips of wings, 16 mm.

General coloration similar to that of Protonemura D.

Female genitalia: subgenital plate produced into two rounded pos-
terolateral lobes. Vaginal structures very distinctive, as in Figure 22.

Material examined: $, Godavari, 6000 ft, 17-20.VII.1967.

Acknowle d gm en ts

I thank Mr. J. E. H. Martin of the Canada Department of Agri-
culture in Ottawa for .allowing me to study this material and Dr.
R. W. Baumann of the United States National Museum for the
loan of material and the comparison of specimens.

References

Aubert, J.

1967. Les Nemouridae de l’Assam (Plecopteres) . Mitt. Schweiz.
Entomol. Ges. 39: 209-253.

Kimmins, D. E.

1946. New species of Himalayan Plecoptera. Ann. Mag. Nat. Hist.
(11) 13: 721-740.

1950. Some Assamese Plecoptera, with descriptions of new species of
Nemouridae. Ibid. (12)3: 194-209.

THE EYELESS CATOPOCERUS BEETLES (LEIODIDAE)
OF EASTERN NORTH AMRICA*

By Stewart B. Peck
Department of Biology, Carleton University
Ottawa, Ontario, Canada KiS 5B6

The Leiodid beetle genus Catopocerus Motschoulsky 1869 is
composed exclusively of small (1.8-4. 5 mm body length), eyeless,
wingless, partially depigmented inhabitants of moist forest soil and
duff, and occasionally of caves. The genus, the only member of tbe
subfamily Catopocerinae, is known to occur only in North America.
The distribution of the genus is principally the unglaciated mountain
forests of the eastern and western portions of the continent.

Hatch ( 1957) has reviewed the nine described species from western
North America. These range from the San Francisco area northward
in coastal forests to Sitka, Alaska. Undescribed species occur in this
region and also in southern California, Arizona, and Colorado (per-
sonal data). The known and new western species will be covered in
a future paper which will also include data from an assemblage of
over 2000 specimens collected throughout Oregon by Ellen Benedict
of Portland State University.

This paper reports on the two described eastern North American
species, and describes three new eastern species.

The genus possesses a character common to the family; the eighth
antennal segment is smaller than either the seventh or ninth segments
(except in C. pusio Horn of California in which the seventh and
eighth segments are equally small). They may be distinguished from
other leiodids by a combination of the 'following characters: eyeless-
ness, winglessness, oval shape, dorsal-ventral flattening, open procoxal
cavities, five segmented abdomen, and separated metacoxae (Peck,
1973:50).

Methods

In the field, series of specimens were taken from moist forest litter,
soil, and well-rotted logs. The debris was sifted in the field through
a one-half inch mesh screen to remove large objects. The sifted debris
was carried and stored in large plastic bags until it could be processed
in “Berlese” funnels. When processed, three liters of litter were
placed in each funnel (45 cm high, 30 cm across the top, with the

* Manuscript received by the editor December 17, 1974.

377

378

Psyche

[September-December

screen io cm from the top) on a double layer of cheesecloth sup-
ported by the screen. A 40 watt bulb was used over the litter. Two
thin wood strips held the top o'f the funnel above the rim of the
bottom and provided ventilation and for the escape of excess moisture.
Beetles stopped falling into the collecting bottle at the bottom of the
funnel in less than 12 hours. During periods of maximum operation,
up to 24 funnels were used, each holding three liter samples of sifted
litter for twelve hours. Thus, up to 144 liters of sifted litter could
be processed each day.

From 1967 to 1974, in search of Catopocerus and other leiodid
beetles, I have extracted the fauna in the above method from 4361 kg
of sifted forest litter from the eastern United States. Most of the
beetles extracted have been deposited with the CNCI and FMNH,
and the arthropod residues are in the FMNH.

Barber’s Fluid (Valentine, 1942) was used as a preservative under
the funnels. Species determination can be reliably made only by
examination of the male genitalia. This examination is facilitated bv
the Barber’s Fluid which does not harden the tissues of the beetles
as does alcohol. The few specimens found under rocks were collected
into Barber’s Fluid. Some specimens were collected in caves at rotted
pig liver bait, and in carrion baited pitfall traps (Peck, 1973).
Specimens were cleaned of adhering debris by an ultrasonic cleaner.
Dry material was relaxed in boiling water. Dissections were made
with minuten needles, in alcohol. Genitalia were observed in alcohol,
as temporary glycerine mounts, and dry. Specimens were mounted on
points using an alcohol soluble glue. Illustrations were prepared
from temporary glycerine mounts.

Measurements are given in millimeters, and were made using a
calibrated ocular micrometer disc. The following abbreviations are
used in the paper: HW, head width; PW, pronotal width; PL,
pronotal length ; EL, elytral length ; and EW, elytral width. Lengths
were measured along the midline. Elytral lengths are from the apex
of the scutellum. Widths are maximum widths. Rounded surfaces
are measured as chords of arcs.

The following abbreviations are used to indicate the sources of the
material examined: AMNH, American Museum of Natural History;
ANSP, Academy Natural Sciences Philadelphia (Horn colln., now
in MCZ) ; CAS, California Academy of Sciences; CM, Carnegie
Museum; CNCI, Canadian National Collection of Insects; FMNH,
Field Museum of Natural History; INHS, Illinois Natural History
Survey; MCZ, Museum Comparative Zoology, Harvard University;
SBP, Stewart B. Peck; SVAM, St. Vincent Archabbey Museum

1974]

Peck — Catopocerus

379

Collection, Latrobe, Penn.; UK, University of Kansas, Entomology
Collection; USNM, United States National Museum.

Bionomics

The general collector may occasionally encounter Catopocerus
under large rocks and logs in mountain forests. Large series are
most usually taken in the spring and fall and only by extraction from
soil and litter with the Tullgren modification of a Berlese funnel.
Some species have been found in caves in the eastern interior low
plateaus. Here they are usually under rocks in organic soil near cave
entrances. The use of carrion bait has been useful in attracting the
beetles in caves.

Food probably consists of organic debris and associated yeasts and
fungi. Several species of Catopocerus have been repeatedly taken on
subterranean fungi in Oregon and Washington (Fogel and Peck, in
MS) and the eastern species may have similar habits. A laboratory
colony of the beetles reproduced and developed on bakers yeast on
moist soil. C. hamiltoni was taken congregated around a dead beetle
larva, and the many beetles taken on carrion bait in caves were un-
doubtedly attracted to the carrion for the purposes of feeding and/or
oviposition.

Only one certain case of sympathy is known. Thirty-four C. ulkei
and 1 1 C. appalachianus were taken together in io kg of litter near
Whitmer, W. Va.

Reproduction may occur throughout the year in caves, but in forest
litter is probably most active in the spring. Larvae and teneral adults
have been found most commonly in early spring in litter. One species
has been reared from eggs. At I5°C, the egg stage lasts about 17
days, the larval stage about 29 days, and the pupal stage 20 days.

Three species have been found in caves in Alabama and Tennessee.
However, this association with caves is only facultative, for these soil-
inhabiting (edaphobitic) beetles probably accidentally enter caves,
but once there can survive and reproduce.

One of these species, C. appalachianus is found at higher elevations
in the Appalachians near the soil surface under rocks and in litter.
If the preferred environment of the beetles is a montane type of
stable, cool, and moist environment, this will be found at lower
elevations deeper in the soil. Collecting of surface soil and litter has
never taken the beetles at the lower elevations of the Cumberland
Plateau or Highland Rim of Alabama or Tennessee. Thus, in the
cave region, the beetles’ normal soil habitat must be too deep to be

[September-December

1974]

Peck — Catopocerus

381

normally sampled, and this deep soil habitat will bring the beetles
into frequent contact with rock crevices filled and partly filled with
soil, which may open into caves with their cool and moist conditions.
Hence, in the southern Appalachians, caves can be a more convenient
sampling site for soil inhabitants which are usually hidden under a
concealing thickness of soil. Other such montane soil inhabitants that
are infrequently found at lower elevations except in caves are Anil-
linus carabids (Barr, 1969) and Arianops pselaphids (Barr, 1974).
Collecting in prolonged wet weather or during winter months might
find all of these beetles closer to the surface in the Cumberland
Plateau or Highland Rim regions of Alabama and Tennessee.

Artificial key to the Catopocerus

1 a. Body length 4mm or over C. hamiltoni

ib. Body length 3mm or under 2

2a. Hind tibia with setal bearing excavation on dorsal posterior sur-
face (fig. 2) C. alabamae , n. sp.

2b. Hind tibia with setae only, no excavation on dorsal posterior

surface 3

3a. Third antennal segment clearly longer than second, five clearly
larger than four and six; northeastern Alabama C. jonesi, n. sp.
3b. Third antennal segment as long as or only slightly longer than
the second, five the same size as four and six; widely distributed

4

4a. Tip of aedeagus curved downward, forming 8o° to iOO° angle

with ventral surface of aedeagus (Fig. 4) C. ulkei

4b. Tip of aedeagus only slightly curved downward, angle with ven-
tral surface of aedeagus 1350 or greater (Fig. 6)

C. appalachianus , n. sp.

Catopocerus hamiltoni (Horn)

Fig. 3 ; Map 2

Pinodytes hamiltoni Horn, 1892:45. Lectotype here designated as female
in MCZ, Horn colln. (no. 3027), seen. Type locality: “vicinity of Allegheny
City”, Pennsylvania.

Figures 1-11. 1, habitus of Catopocerus appalachianus. 2, metatibia of

C. alabamae. 3, aedeagus of C. hamiltoni. 4, side of aedeagus of C. ulkei.
5, ventral tip of aedeagus of C. ulkei. 6, side of aedeagus of C. appala-
chianus. 7, ventral tip of aedeagus of C. appalachianus. 8, side of aedeagus
of C. jonesi. 9, ventral view of aedeagus of C. jonesi. 10, side of aedeagus
of C. alabamae. 11, ventral view of aedeagus of C. alabamae, tip incom-
plete because of damage in dissection. Scale line for figs. 3-11 only.

3^2

Psyche

[September -December

Diagnosis. The large size, the third antennal segment which is
i.'5 times or more the length of the second segment, and the shape
of the aedeagus serve to distinguish this species.

Redescription. Measurement of lectotype: HW, 0.85. PW, 1.63.
PL, 1.23. EW, 1.63. EL, 2.74.

Length 4. 1-4.7 mm. Width 1.66-1.73 mm. Color uniform dark
reddish-brown, shining. Shape oblong, moderately elongate, not very
convex.

Head without trace of eyes; sparsely punctate. Antennae well
supplied with long hairs throughout length, last three segments with
dense vestiture; segment III 1.5 X or more the length of II; III
slightly wider than II; IV, V, VI as wide as II, subglobose, slightly
longer than wide; VII, IX, X cup shaped; VII longer than but nar-
rower than IX and X; VIII shorter than but slightly wider than
VI ; XI conical.

Pronotum widest at middle, sides evenly arcuate; narrower at
front than back ; hind angles rectangular ; front angles rounded ;
front margin evenly emarginate to receive head; hind margin weakly
convex ; disc finely but not closely punctate ; covered with fine strigae.

Elytra fused ; at base slightly wider than base of pronotum ; widest
at middle; sides slightly arcuate; narrowing in apical third; each
with seven striae of close punctures, less distinct at sides and apex,
intervals flat, sparsely punctate.

Protibia gradually widening to apex; setose on apical half of inner
margin. Mesotibia gradually widening to apex; sharply outward
flaring in outer margin at apex; spinose on outer concave margin;
inner margin sinuous; setose on apical half of inner margin. Meta-
tibia gradually widening to apex; inner and outer margin slightly
sinuous; setae on apical half of inner margin. Penultimate abdominal
segment, 'fourth, with transverse depression on either side.

Male protarsomeres not dilated. Abdominal segment five truncate.
Aedeagus poorly sclerotized ; thin and straight in lateral view ; wide
and straight in dorsal view; tip dorsoventrally flattened, slightly up-
turned at edge, smoothly rounded in dorsal view, 14 setae along
edge; parameres separated, longest on ventral surface but not reach-
ing tip of aedeagus, partially covering anterior ventral surface and
dorsal median surface of aedeagus (fig. 3).

Female abdominal segment five rounded.

Note on lectotype. The specimen is damaged. It possesses only
two prothoracic and one metathoracic legs. One antenna is missing
its terminal segments.

1974]

Peck — Catopocerus

383

Distribution. Southwestern Pennsylvania (map 2). The type
locality, Allegheny City, lying on the north shore of the confluence
of the Allegheny and Monongahela Rivers, was incorporated into
the city of Pittsburgh near the beginning of this century.

Material examined. Pennsylvania. Allegheny, 1 female, lectotype
3027 (ANSP) ; 2, (ANSP, CM). Beatty, 1 female (USNM).
Charleroi, 2 (MCZ, UK). Jeannette, vi. 1901, 2 (CM). Pitts-
burgh, ix, 2 (CM). St. Vincent, 3 (Ulke colln. in CM; MCZ).
Washington Co., 1 female (Fall colln. in MCZ). State label only,
1 paratype no. 3027 (ANSP); 1 (UK).

Biology. The available information on the species has been pre-
sented by Hamilton (1897). He found the first specimen in Decem-
ber, 1872, “about a foot under ground beneath a large impacted
boulder in a wild mountainous place.” Several individuals were seen
but only one was collected because the pale color and remarkable
swiftness led Hamilton to believe they were young roaches. This
and a second specimen found dead by Hamilton the following June
on a wooded hillside were those which Horn had for description.
Other specimens were taken later by other collectors in winter months
under bark, under stones, and by sifting decaying leaves. A group of
twelve was taken in late November under a log in a wooded ravine
around a large dead elaterid beetle larva upon which they were prob-
ably feeding. I have not seen specimens taken since the turn of the
century.

Catopocerus ulkei Brown
Fig. 4, 5 ; Map 1

Catopocerus ulkei Brown 1933 ; 215. Holotype female in CNC, seen.
Type locality; District of Columbia.

Pinodytes cryptophagoides Mannerheim (in part), Horn, 1880: 249;
1892: 46; Hamilton, 1894: 16. Misidentified.

Diagnosis. The species is separated from others by its small size,
the subequal second and third antennal segments, and the flattened,
downward curved tip of the aedeagus.

Redescription. Length 1.4-2. 2 mm. Width 0.71-0.94 mm. Color
uniform dark reddish-brown, shining. Shape oblong.

Head with trace of eyes as depigmented spot on side of head in
some pale specimens ; vertex finely punctulate, finely striolate. Anten-
nae well supplied with hairs throughout length; segments VII, IX,
X supplied with dense vestiture of long and short hairs; I stout; II
and III equally long; III narrower at base than II; IV, V, VI equal,
globose; VII as long as broad; VIII smaller than VII, larger than

384 Psyche [September-December

Range

Mean

Standard Deviation

males

females

males

females males

females

HW

0.44-0.51

0.44-0.54

0.49

0.50

0.024

0.034

PW

0.81-0.91

0.80-0.91

0.85

0.88

0.034

0.054

PL

0.54-0.63

0.54-0.64

0.59

0.60

0.30

0.047

EL

1.08-1.20

1.10-1.28

1.15

1.12

0.052

0.071

EW

0.80-0.92

0.84-0.99

0.88

0.90

0.034

0.045

PW/PL

1.41-1.58

1.41-1.54

1.48

1.46

0.05

0.05

EL/EW

1.28-1.36

1.29-1.38

1.32

1.34

0.03

0.03

PW/HW

1.72-1.86

1.70-1.86

1.77

1.77

0.05

0.04

EW/PL

1.42-1.56

1.45-1.60

1.50

1.51

0.05

0.06

Table

1. Variation

in a sample of

: a population

of Catopo

cerus

ulkei

topotypes from

the District

of Columbia.

10 males

and 10

females. Measurements in millimeters.

VI; IX smaller than X, larger than VII; IX and X broader than
long; XI conical.

Pronotum widest just behind middle; sides evenly arcuate; nar-
rower at front than back; hind angles slightly obtuse; front angles
rounded ; front margin slightly concave ; hind margin straight ; disc
finely and sparsely punctulate; finely striolate.

Elytra fused ; at base slightly wider than base of pronotum ; widest
at middle ; sides slightly curved ; narrowing in apical third ; surface
finely punctate, some punctures vaguely indicating striae; finely
striolate; apical declivity smooth.

Protibia widening to apex in distal 2/3; setose on apical half of
inner margin; mesotibia gradually widening to apex in apical 2/3;
inner margin setose in apical third, faintly sinuous; outer margin
spinose; outer apical angle truncate. Metatibia gradually widening
to apex; outer margin smooth; inner margin smooth to serate; outer
apical angle slightly truncate.

Male first four protarsomeres dilated and supplied with long
lateral hairs; first three mesotarsomeres feebly dilated. Aedeagus in
lateral view arcuate; ventral surface of anterior end projecting for-
ward and curved downward ; tip dorsoventrally flattened and turned
downward forming 8o°-ioo° angle with ventral surface of aedeagus;
parameres long and thin, exceeding tip of aedeagus, hinged at aedea-
gus base to ventrally deflected elongate basal pieces (figs. 4, 5).

1974]

Peck — Catopocerus

385

Female tarsomeres not expanded ; abdominal segments as in male.

Variation. Table 1 presents meristic variation for the topotypic
population. All populations examined which are assigned to this
species have aedeagi of the males identical to aedeagi of the topotypic
population but for two exceptions. The two samples from south-
western North Carolina are 220 air miles to the southwest of the
next nearest locality from which males have been collected. These
southwestern collections differ in the angle of deflection of the aedea-
gal tip, A distinct acute angle of about 8o° is formed between the
ventral surface of the aedeagus and the tip of the aedeagus. All
other populations possess aedeagi in which the angle is usually slightly
obtuse. Only rarely is it a right angle.

The variation of measurements for a sample of four males and
eight females from Joanna Bald, Cherokee County, North Carolina,
when compared to that in table 1, showed that the size of the south-
ern populations is larger, but there is a great deal of overlap within
the range of the measurements. Variation of the ratios, when com-
pared to the topotypic population shows less difference, and that which
does exist is contained within an overlap of one standard deviation
from each mean. This numerical data demonstrates little or no
difference from the topotypic C. ulkei. Since there is a large gap in
knowledge of the range of the species between western Virginia and
southwestern North Carolina, I am unwilling to interpret the taxo-
nomic significance of the greater curvature of the tip of the aedeagus
in the southwestern North Carolina populations. It may be a distinct
subspecies if its distribution is geographically separated from typical
C. ulkei. Or the population may represent one end or segment of a
cline of gradual deflection of the aedeagal tip. Further collecting
may tell.

Distribution. The species is known to occur in southwestern and
southeastern Pennsylvania, the District of Columbia, Maryland,
North Carolina, Virginia, and West Virginia (map 1). Future col-
lecting might show it to be continuously distributed along the Pied-
mont and eastern edge of the Appalachians to the southern known
locality in Cherokee County, North Carolina.

Material examined and habitat data. District of Columbia: no
other data, 32. Washington, D. C., 1 ; 1.5, 1; 5.3, 1; Hubbard &
Schwarz. Maryland: Garrett Co.; Deer Park, 4.7, 1, Hubbard &
Schwarz; 2 mi. E. Keysers Ridge, 2500' elev., 18.vi.1968, S. Peck,
3 in 220 lbs. log-stump litter. North Carolina: Cherokee-Graham
County; Andrews, Joanna Bald, 4700' elev., 26.vii.1967, S. Peck &
A. Fiske, 13 in 89 lbs rotted chestnut log; Joanna Bald, 5.viii.i96o,

386

Psyche

[September-December

Map 1. East central United States and Catopocerus distributions. Closed
dots, C. ulkei. Open dot, C. alabamae. Closed triangles, C. jonesi.

T. C. Barr, i male. Clay-Macon County; Tusquitee Bald, q.viii.
i960, T. C. Barr & M. C. Bowling, 1 male. Pennsylvania: x, 1.
Allegheny County; Pittsburgh, x, 3. St. Vincent, 9. Cambria
County; Nicktown, M. Wirtner, 3. Philadelphia County; 28. vi.
1 9 1 1 , G. M. Greene, 1. Westmoreland County; Chestnut Ridge,
E. of Youngstown, 27.vi.1961, W. Suter, 1 in litter under Rhodo-
dendron; 11.vii.1961, W. Suter, J. Wagner, D. Reichle, 1 in forest
litter. Jeannette, vii, 2; ix, 3; x, 1. Virginia. Caroll County;
Groundhog Mt., 2900' elev., Blueridge Parkway, 23.vii.1967, S.
Peck & A. Fiske, 1 female in 46 lbs. rotted stump and log litter in
Rhododendron thicket. Giles County; Mountain Lake, 1-5.vii.1970,
W. B. Muchmore, 8. Madison County; Shenandoah National Park.
Dark Hollow Falls, 2.W.1967, S. Peck, 1 under rocks. Lewis Mt.,
2000' elev., 10.vi.1967, S. Peck, 104 in rotted logs. Page County;
Shenandoah National Park. Hawksbill Mt., 17.x. 1954, D. G. Kis-
singer, 1. Stony Man Overlook, 2.W.1967, S. Peck, 3 under rocks
and 6 in litter. West Virginia. Greenbrier County; outside Ar-
buckle Cave, 14.vii.1970, W. B. Muchmore, 1 in litter. Mercer
County; 5.5 mi. N. Princeton, Brush Creek Falls, 13.V.1971, W.
Shear, 1 in cliffside litter. Pocahontas County; W. side Cranberry
Glades, base of mountain, streamside, under rock, 28.vi.1967, T. C.

1974]

Peck — Catopocerus

387

Barr, 1 female. Hills Creek Falls, 6.V.1970, W. Shear, 1; ig.vi.
1971, Shear and Platnick, 2. Randolf County; Bickle Knob, 3800'
elev., E. of Elkins, ig.vi. 1968, S. Peck, 34 in 179 lbs. of log litter.
5 mi. S. Whitmer, 18.vii.1971, S. Peck, 3000' elev., 34 in litter.
(AMNH, CAS, CM, CNC, FMNH, INHS, SBP, SVAM, UK,
USNM).

Catopocerus appalachianus New Species
Fig. 6, 7 ; Map 2

Holotype male (MCZ no. 32228) from Balsam Gap, 5500' elev.,
Mt. Mitchell, Yancey Co., North Carolina, 9.VK1967, S. Peck.
Paratypes: all material listed in section on material examined.

Diagnosis. Most similar to C.~ ulkei~ but differing distinctly from
it and all other species in the shape of the aedeagus. Usually differ-
ing from C. ulkei in its larger size and greater EW/PL ratio.

Description. Length 2.14-2.86 mm. Width 0.83-1.10 mm. Color
uniform dark reddish-brown, shining. Shape oblong.

Head with faint trace of eyes as depigmented spots on side of head
in pale specimens ; vertex finely punctulate, finely striolate. Antennae
with long hairs throughout length; segments VII, IX, X with dense
vestiture of long and short hairs; segment I stout; II stouter and
only slightly shorter than III; IV, V, VI subglobose and subequal;
VII as long as broad, widest at apical third; VIII as long as and
wider than VI, smaller than VII and IX ; IX and X of equal length,
X wider; XI conical.

Pronotum widest at basal third; sides evenly arcuate; narrower at
front than back ; hind angles slightly obtuse ; front angles rounded ;
front margin slightly concave; hind margin straight; disc finely and
sparsely punctulate; finely striolate.

Elytra fused ; at base slightly wider than base of pronotum ; widest
near middle, sides slightly curved ; narrowing in apical third ; surface
finely punctate, punctures indicating striae; finely striolate; apical
declivity smooth.

Protibia widening to apex in distal 2/3; setose on apical half of
inner margin ; a few spines on apical half of outer margin. Mesotibia
straight along outer margin ; expanding gradually along inner margin
to apex from proximal third; inner margin setose in apical third,
faintly sinuous; outer margin spinose; outer apical angle rounded.
Metatibia gradually widening to apex; outer margin smooth; outer
apical angle slightly truncate, possessing two large spines; dorsal
surface of apical end with patch of setae; slight downward bow in
middle in lateral view.

388

Psyche

[Septembe

r-December

Range

Mean

Standard

. Deviation

males

females

males

females

males

females

HW

0.49-0.70

0.48-0.63

0.60

0.55

0.070

0.054

PW

0.81-1.15

0.84-1.08

1.01

0.95

0.097

0.080

PL

0.58-0.80

0.55-0.75

0.70

0.68

0.072

0.064

EL

1.15-1.55

1.19-1.54

1.40

1.34

0.131

0.107

EW

0.84-1.14

0.89-1.10

1.00

0.98

0.094

0.074

PW/PL

1.36-1.55

1.43-1.60

1.44

1.49

0.05

0.04

EL/EW

1.34-1.44

1.32-1.42

1.36

1.38

0.04

0.04

PW/HW

1.58-1.82

1.69-1.80

1.66

1.74

0.06

0.04

EW/PL

1.90-2.09

1.98-2.24

1.99

2.10

0.06

0.07

Table

2. Variation

in a sample of a population

of Catopc

icerus

appalachianus from Mt. Mitchell, North Carolina. 15 females
and 15 males. Measurements in millimeters.

Male first four protarsomeres expanded and with long hairs; first
three mesotarsomeres feebly dilated ; metatibia with inner edge weakly
to strongly serrated. Aedeagus in lateral view robust and arcuate;
ventral surface of anterior end projecting forward; from base nar-
rowing slightly for 2/3 length, narrowing markedly in distal third,
tip slightly downcurved ; in ventral view sides expanding slightly for
2/3 length, then evenly converging to tip; tip flattened; parameres
gradually tapering to thin point which extends beyond aedeagus tip;
basal piece of parameres pointing forwards along sides of aedeagus
(figs. 6, 7).

Female tarsomeres not expanded; abdominal segments as in male;
inner margin of metatibia smooth.

Variation. Table 2 presents meristic data on the topotypic popu-
lation. In some specimens the outer margin of the elytra at the an-
terior angles bears a few coarse, low teeth. The males of the species
variably possess a saw-toothed inner margin of the hind tibia, ranging
from absent to highly developed.

The variation in size and proportions within this species overlap
in some measured populations with C. ulkei size and proportions.
Hence, care should be used in separating the species using external
criteria. A sample of four males of C. appalachianus from Whitetop
Mt., Virginia is composed of noticeably smaller individuals. Their

1974]

Peck — Catopocerus

389

PL mean corresponds to that of the topotypic population of C. ulkei.
The Whitetop Mt. sample EW/PL mean also corresponds to that
of topotypic C. ulkei and not the topotypic C. appalachianus sample.
This size overlap demonstrates the clear value of using the distinct
and less variable features of the aedeagus. Because of these size
similarities some female specimens may be difficult to indentify. Lo-
cality records for these two species in their zone of overlap may be
treated with certainty only when males are present.

Etymology. The name refers to the Appalachian region in which
the beetle is found.

Distribution. The species has roughly a linear range of 480 miles
along the Appalachian Mts. from Madison County, Alabama north-
eastward to Pocahontas County, West Virginia (Map 2). It is
known from a few intermediate areas in Virginia, Tennessee, and
especially from North Carolina in the vicinity of Mt. Mitchell. One
collection has been made in the Cumberland Plateau of Kentucky
and one in the Shawnee Hills of Southern Illinois.

Material examined and habitat data. Alabama. Madison County:
Barclay Cave, 26.vii.1965, S. & J. Peck, 1 in trap; 18.viii.1965, S.
Sc J. Peck, 1 in trap; 20.viii.1965, S. Sc J. Peck, 1 in trap; i.ix.1965,
S. Sc J. Peck, 2 in trap; 2.vii.i967, S. Peck and A. Fiske, 1 at car-
rion bait; 24.viii.1971, S. Peck, 1 at bait; io.v.72, S. Peck, 4 at bait;
13.vii.1973, S. Peck, 1 at bait. New Hope, Cave Spring Cave, 23.
viii. 1971, S. Peck, 3 at bait in side chamber 400' in cave. Ellis Cave,
5 .vii. 1 967 , S. Peck and A. Fiske, 2 at carrion bait; ix.1968, S. Peck,
8 at carrion bait; 2 reared from eggs laid in culture 3.viii.i969, S.
Peck. Illinois. Union County; Pine Hills Field Station, 15.V.
1967, J. M. Campbell, 1 at malt trap site #9. Kentucky. Harlan
County; Big Black Mt., 4000' elev., 11.vii.1968, S. Peck, 1 in 302
lbs. log and stump litter. Jackson County; The Rises, S. Fork Sta-
tion Camp Creek, 16.ii.1967, T. Marsh, W. Andrews, 1 in log
berlese. North Carolina. Avery County: Beech Mt., 15.vii.1960,
Barr and Bowling, 10; Grandfather Mt., 22.viii.1960, Barr and
Bowling, 1 male. Blowing Rock to Linville, 3000-4000', 7UX.1930,
P. Darlington, 1. Buncombe County: Asheville, A. P. Jacobs, 1.
Blue Ridge Parkway, 5 mi. W. Craggy Gardens, 3500' elev., 22.
vii. 1 967, S. Peck and A. Fiske, 6 in 63 lbs. Chestnut stump litter;
Craggy Dome, 22.vii.1960, Barr and Bowling, 9. Haywood County;
Richland Balsam, 1. viii. i960, Barr and Bowling, 3. Henderson
County; 0.3 mi. S.W. of Bat Cave, 1000' elev., 22.vii.1967, S. Peck
and A. Fiske, 2 in 39 lbs. of log and leaf litter. Madison County;
Rich Mt., 3000' elev., 25.vii.1967, S. Peck and A. Fiske, 4 in 85 lbs.

390

Psyche

[September-December

Map 2. East central United States and Catopocerus distributions. Open
dots, C. hamiltoni. Closed dots, C. appalachianus.

of rotted stump. Mitchell County; Blue Ridge Parkway, mile 326.8,
3200' elev., 22.vii.1967, S. Peck and A. Fiske, 2 in 64 lbs. of litter
at base of large dead pine tree: Roan Mt., 5000' elev., 25.vii.1967,
S. Peck and A. Fiske, 2 in 72 lbs. of litter along rotted beach log.
Round Knob, 27.6, Hubbard and Schwarz, 1. Yancey County;
Hamrick, 3000-3100' elev., 30.xii.1946, Hairston, 1 in leaf litter:
Mt. Mitchell, Balsam Gap, 5300' elev., 9.VL1967, S. Peck, 9 in
birch-beech litter: Mt. Mitchell, 5500' elev., 9.VL1967, S. Peck, 10
in birch-beech litter and 1 in basal tree hole: Mt. Mitchell, 4000'-
6000' elev., June 1939, E. D. Quirsfeld, 10 : Mt. Mitchell, 26 with
no other data: Valley of Black Mts., July 28, 1906, W. Beuten-
muller, 3; August 19, 1906, W. Beutenmuller, 1. Black Mts., vi-
vii.1902, Van Dyke colln., 14. Tennessee. Coffee County. Beech
Grove, Burke Cave, 24.V.73, S. & J. Peck, 1 male, 1 female, at
carrion bait. Great Smoky Mt. National Park; Chimneys, 3000'
elev., ix. 1 9. 1 944, A. Nicolay, 2 females. Newfound Gap, 5000-
5200', August 31, 1930, P. Darlington, 1 female. Clingmans Dome,
7.VH.1960, T. C. Barr. Unicoi County; Unaka Mtn., 5.vii. 1 953 ,
H. & A. Howden, 1. Virginia. Grayson County; Whitetop Mtn.,
4000' elev. 24.vii.1967, S. Peck and A. Fiske, 10 in 77 lbs. of log

1974]

Peck — Catopocerus

39i

and stump litter; Whitetop Mtn., 13.vii.1960, T. C. Barr, M. C.
Bowling, 4. West Virginia. Mercer County; Athens, 6.vi.i97i,
W. Shear, litter, 1. Randolf County; 5 mi S. Whitmer, i8.vii. 1971,
S. Peck, 3000', 11 in litter. Pendleton County; Spruce Knob, 4000'
elev., 26.viii.1964, S. Peck & T. 'C. Barr, 2 under rocks and 2 in
litter. (AMNH, CAS, CNC, FMNH, MCZ, SBP, USNM).

The Illinois collection, separated from the rest of the species’
range, is also of special interest because of the unusual nature of the
habitat. The collector, Dr. J. M. Campbell, has supplied me with
more detailed data on the locality, and I quote from a letter; “Malt
trap no. 9 was set in a small stand of Pin, us echinata on a very dry
site on the S. W. slope near the top of a ridge. There was very little
litter on the soil ; what little litter there was consisted of pine needles.
This entire area of S. Illinois is characterized by having very small
stands of pine scattered on the S. W. slopes of dry, rocky ridges. It
is possible that any beetles found in this site live in the more mesic
ravines which are nearby.”

This species may also exist in Missouri. One female labeled “St.
Louis, Mo., iv.28” from the Liebeck collection is in the Fall collec-
tion in the MCZ. The measurements of the specimen fall into the
range of variation of C. ulkei, but the Illinois record of C. appa-
lachianus suggests that it might more likely occur in Missouri if we
accept the accuracy of the specimen label

Life Cycle Data. Eight adult C. appalachianus , captured Septem-
ber 1, 1968 in Ellis Cave, Madison Co., Alabama, were kept alive
in laboratory culture at I5°C in plastic dishes of moist soil from
Ellis Cave. All adults had died by May 1969. Two beetles becom-
ing adults in May 1969 died in June and July 1970, suggesting that
full adult life may be about one year. Copulation occurred in cul-
ture. A total of 37 eggs were laid on the soil surface, with soil
crumbs stuck to their sides. The number of females producing these
eggs is not known. The month of egg laying and number of eggs
were as follows: October, 10; November, 5; December, 3; January,
2; March, 14; April, 3. Reproduction may thus occur throughout
the year, especially under favorable temperature conditions such as in
caves or deep rock crevices. The eggs were white and oblong, a
sample of six measured .35-.60 mm (mean .48 mm) by .5 5~-75 mm
(mean .63 mm).

A sample of 8 eggs hatched 13 to 20 days (mean 17 days) after
being laid. A sample of 7 larvae had a total life span in this stage
of from 26 to 30 days (mean 29 days). The first instar lasted 8
days for 1 larva. The total number of instars was not determined

392

Psyche

[September-December

but may be three. Approximately the first 20 days of larval life were
spent feeding on the soil surface or in shallow burrows at the edge
of the yeast. Approximately the last ten days (6 to n recorded) of
larval life were spent in deepened vertical burrows which were made
into pupation chambers. The larvae in these cells became more plump
and swollen. Pupation lasted from 16 to 22 days (mean of 20 in a
sample of 7). After ecdysis the adults remained in their cells for
about a week before emerging. When the adults became reproduc-
tively active is not known.

Some of the larvae and pupae were preserved for a study of im-
mature characteristics. They are in the MCZ beetle larva collections
and will be described in a later paper.

Catopocerus jonesi New Species
Figs. 8, 9 ; Map 1

Holotype male (MCZ no. 32229) from Eudy Cave, 1 mi. S.
Oleander, Marshall County, Alabama, 23.vi.1942, W. B. Jones leg.
Paratypes, in SBP collection: Alabama. Morgan County, Shine
Cave (in Newsome Sinks) 7.vii. 1957, L. Varnedoe, 1 female; Van-
dever Cave, 3 mi. S.S.W. Laceys Spring, 22.V.1972, S. & J. Peck,
1 male and 1 female in rotten sticks at back of cave. Jackson
County; House of Happiness Cave, 4 mi. S.W. Scottsboro, 23.V.
1972, S. & J. Peck, 2 males and 1 female at carrion bait near en-
trance.

Diagnosis. The species can be distinguished by the third antennal
segment being longer than the second, and the fifth being longer
than the fourth and sixth, and by the conspicuous toothed ventral
projection of the aedeagus.

Description. Holotype measurements, HW, 0.62. PW, 1.12.
PL, 0.80, EL, 1.56. EW, 1. 1 7.

Length 2.54-2.65 mm. Width 1. 15- 1.23 mm. Color uniform
dark reddish-brown, shining. Shape oblong, sides parallel.

Head with no indications of eyes; clypeus slightly truncate; frons
punctate; vertex punctate and finely striolate. Antennae well sup-
plied with long hairs; segments IX, X, XI, possessing dense mat of
long and short hairs; segment I robust; II cylindrical, shorter than
III by 1/4; III conical, as broad as II; IV, VI longer than broad,
shorter than V; VII distinctly longer than broad, slightly wider than
VIII and longer than VIII by a third; IX slightly narrower than X,
a third wider than VIII ; XI conical at tip.

Pronotum widest at middle; sides evenly arcuate; narrower at
front than back; hind angles slightly obtuse; front angles rounded;

1974]

Peck — Catopocerus

393

front margin slightly concave; hind margin slightly concave; disc
finely punctulate ; finely striolate.

Elytra fused; at base slightly wider than base of pronotum; widest
near middle; anterior margins subparallel; surface punctate, punc-
tures indicating striae; finely striolate; sides and apex smooth. Pro-
tibia straight on outer margin ; sinuous on inner margin ; widening
to apex in apical 2/3; setose on apical half of inner margin; spinose
along outer margin ; two large spines at outer apical angle ; two
shorter spines on inner margin at origin of tarsus; comb of short
spines below origin of tarsus. Mesotibia straight and spinose along
outer margin; curved and setose along inner margin; expanding to
apex in apical 2/3; outer apical angle sharp, possessing two spines;
one large spine along ventral edge of apical margin. Metatibia
stout, gradually widening to apex ; outer and inner margin smooth ;
curved slightly so that apex is deflected outward ; straight in lateral
view; one large spine at outer apical angle and one in middle of
apical margin; dense patch of setae along upper and inner margin
of apex. Fifth abdominal segment evenly rounded.

Male first three protarsomeres widely expanded and supplied with
a dense mat of hairs; first three mesotarsomeres less expanded and
hairy. Aedeagus in lateral view bent near base, basal anterior pro-
jection deflected downward, body of aedeagus slightly sinuous and
tapering, ventral surface projecting posteriorly to form shelf under
genital orifice, dorsal surface curving downward slightly; in ventral
view robust, sides parallel, dorsal surface tapering only at apex and
smooth in outline, ventral surface with apical tooth-like projection;
parameres gradually tapering to point, terminating close to aedeagal
apex, basal pieces thin (figs. 8, 9).

Female tarsomeres not expanded.

Variation. No noteworthy variation is evident in the seven known
specimens.

Etymology. I am pleased to name this species for Dr. Walter B.
Jones, retired State Geologist of Alabama, in recognition of his as-
sistance to me and to other students of Alabama cave fauna.

Distribution. Known only from Morgan, Marshall and Jackson
Counties in northeastern Alabama (map 1 ).

Catopocerus alabamae New Species
Figs. 10, 1 1 ; Map 1

Known only from Holotype male (MCZ no. 32230) from Cave
Spring Cave, Chapman Mtn., 1 mi. N.E. Huntsville, Madison Co.,

394

Psyche

[September-December

Alabama, found in the stomach of a plethodontid salamander Eurycea
Iucifuga, captured 8.ix.i965, S. Peck leg.

Diagnosis. Most similar to C. jonesi but differing from it and
all other species in the shape of the aedeagus, the very widely ex-
panded male tarsal segments, and the distinct setal bearing excavation
on the inner and upper margin of the hind tibia.

Description. Measurements of the Holotype. HW, .70. PW,
1.25. EL, 1.78. EW, 1.30. Color uniform dark reddish brown,
shining. Shape rectangular, rounded at front and back.

Head with no traces of eyes; vertex finely punctulate; frons
coarsely punctate. Antennae with hairs on all segments; segments
IX, X, XI also supplied with dense cover of short hairs at their
apical margins; segment I stout; II shorter than III, III slightly
conical; IV, VI larger than broad, shorter than V; VII larger than
wide, larger than VIII, smaller than IX; IX longer than wide; X
as long as wide; XI conical at tip.

Pronotum widest at middle; sides evenly arcuate; narrower at
front than back ; hind angles slightly obtuse ; front angles rounded ;
front margin slightly concave; hind margin slightly concave; disc
finely puntulate, coarsely set with punctures.

Elytra fused ; at base slightly wider than base of pronotum ; widest
near middle; surface finely punctulate, with larger punctures indi-
cating 5 distinct and a sixth indistinct stria. Protibia widened at
apex, protarsi conspicuously expanded and setose. Mesotibia widened
at apex, mesotarsi conspicuously expanded and setose. Metatibia with
setae bearing excavation in posterior third of dorsal and inner sur-
face. Metatarsi not expanded. Fifth abdominal segment evenly
rounded.

Aedeagus in lateral view (fig. 10) wide, without ventral projec-
tion, expanded in posterior half. In ventral view (fig. 11) wide and
slightly expanded toward posterior. Parameres gradually tapering,
terminating close to aedeagal apex, basal pieces thin.

Notes on holotype. The tip and left side of the aedeagus were
damaged in dissection. The illustrations are reconstructed except for
the dorsal tip whose full size and shape are not known.

Etymology. The name refers to the state of Alabama.

Distribution and habitat notes. The species is probably a deep
soil inhabitant, limited to northeastern Alabama. The type locality
is a small cave under the sandstone cap of Chapman Mountain. A
seep spring in the cave gives the cave its name. I have visited the
cave many times since 1965 in unsuccessful attempts at obtaining
more specimens by searching by hand under rocks and litter and by

1974]

Peck — Catopocerus

395

using bait. The salamander which obtained the beetle as food un-
doubtedly did so from a small and/or inaccessible population in or
near the cave. Catopocerus are not commonly eaten by terrestrial
cave salamanders. I have examined the digestive tract contents of
433 salamanders from caves within the range of Catopocerus in the
southeastern U. S. (Peck, 1974: Richardson and Peck, in MS) and
no other Catopocerus have been found.

Evolution and Distribution

We may assume that the genus had a past transcontinental range.
The present division of the generic range into the ecologically iso-
lated eastern and western forested regions may date from the Mio-
cene when drying conditions caused a retreat of the Neotropical
flora from western North America. This range division was probably
widened with the later Pliocene arid trend associated with the north-
ward invasion of the Madro-Tertiary geoflora which gave rise to
the recent xerophytic vegetation (Axelrod, 1958).

Since this division, there is little that can be said of speciation and
distributional events for the eastern species. A more rich history is
evident in the western species, but will be discussed in a later paper.
The three eastern species with limited distributions ( C . hamiltoni,
C. alabamae, and C. jonesi ) probably experienced a contraction to
their present ranges, from a wider ancient range, in connection with
Pleistocene climatic events. None of these species is closely related
to the other.

C. applachianus and C. ulkei, however are similar enough to have
been derived from a single common ancestral species which may have
inhabited the Appalachians from at least Pennsylvania to North
Carolina. A division of the range into two main units during an
early interglacial may have allowed C. ulkei to diverge in the north
from Virginia perhaps to Pennsylvania, and allowed C. appalachianus
to diverge in the south in North Carolina. C. ulkei may have then
dispersed southward during the Illinoisan glaciation to southwestern
North Carolina. These populations may have become isolated and
somewhat divergent during the Sangamon interglacial. Other overlap
of the ranges of the two species may have occurred by dispersal in the
Illinoisan, as well as the Wisconsinan. However, judging from the
lack of other geographic variation, most of the present distributions
were probably gained in the Wisconsinan. This was the time when
the larger and perhaps more vagile C. appalachianus moved north
into Virginia and West Virginia. The populations of this species in

396

Psyche

[September-December

the Cumberland Plateau of Kentucky and Alabama, the Highland
Rim of Tennessee, and the Shawnee Hills of southern Illinois are
probably not remnants of an ancient distribution but also date from
the Wisconsinan glaciation when more wet and cool montane-type
climatic conditions prevailed in the Interior Basins, favoring the
distribution and dispersal of “montane” beetles.

The comparatively large ranges and low levels of geographic vari-
ability in C. ulkei and C. appalachianus are remarkable in the light
of an assumed low dispersal potential resulting from the beetles’
winglessness, eyelessness, and soil habitation. Other ecologically
similar beetles, for instance Anillinus and Arianops (Barr, 1969;
1974) in the eastern United States have much smaller ranges and
are more highly speciated. In contrast then, it must be that the
vagility of some Catopocerus is higher than would seem likely at
first, because of an incompletely understood ability to withstand
various mechanisms of long distance dispersal, similar to that of some
European soil Colydiid beetles (Peck, 1972).

Acknowledgements

I thank the following persons and their institutions for allowing
examination of specimens: M. G. Emsley, ANSP; L. H. Herman,
AMNH ; H. B. Leech, CAS; J. M. Campbell, CNCI; H. Dybas
and R. Wenzel, FMNH ; M. W. Sanderson, INHS; P. J. Darling-
ton and J. F. Lawrence, MCZ; J. M. Kingsolver, USNM; Fr.
Jerome Rupprecht, SVAM ; G. E. Wallace, CM; and G. Byers,
UK. Thomas C. Barr, University of Kentucky, and Henry F.
Howden have provided specimens from their personal collections.
Collections made in regions administered by the National Park Ser-
vice were permitted by Ernest G. Whanger, Blue Ridge Parkway,
and V. R. Bender, Great Smoky Mountains National Park. A debt
of gratitude is due to many persons who assisted me in collecting,
especially James H. Peck in 1965, Alan Fiske in 1967, and Jarmila
Kukalova-Peck. Dr. and Mrs. Walter B. Jones of Huntsville, Ala-
bama, are thanked for their hospitality in providing a base for field
work conducted intermittently in and near Alabama over seven years.
My wife, Jarmila, prepared the final illustrations. My specimens
were collected during field work supported by National Science
Foundation Grants GB 3167 and GB 7346 (Professor Reed C.
Rollins, principal investigator, Biological Laboratories, Harvard Uni-
versity) and by Canadian National Research Council operating
grants. John Lawrence and Ernst Mayr kindly read an early ver-
sion of the manuscript.

1974]

Peck — Catopocerus

397

Literature Cited

Axelrod, D.

1958. Evolution of the Madro-Tertiary gloflora. Bot. Rev., 24: 433-
509.

Barr. T. C., Jr.

1969. Evolution of the (Coleoptera) Carabidae in the southern Appala-
chians, 67-92 in P. Holt ed., The distributional history of the
biota of the southern Appalachians, part I: Invertebrates. Re-
search Div. Mono. 1, Virginia Polytech. Inst., Blacksburg, Va.
295 pp.

1974. The eyeless beetles of the genus Arianops Brendel (Coleoptera,
Pselaphidae) . Amer. Mus. Nat. Hist. Bull. 154(1): 1-51.

Brown, W. J.

1933. Two undescribed species of the old family Silphidae with notes
on some characters that have been used to divide the group.
Can. Ent., 65: 213-215.

Hamilton, J.

1894. Catalogue of the Coleoptera of Alaska with the synonymy and
distribution. Trans. Amer. Ent. Soc., 21: 1-38.

1897. Pinodytes hamiltoni and Anthicus formicarius. Ent. News, 8:
34-35.

Hatch, M. H,

1957. The beetles of the Pacific Northwest; part II: Staphyliniformia.
University of Washington Press, Publications in Biology, vol. 16,
Seattle, 384 pp.

Horn, G. H.

1880. Synopsis of the Silphidae of the United States with reference
to the genera of other countries. Trans. Amer. Ent. Soc., 8:
219-322.

1892. Random studies in North American Coleoptera. Trans. Amer.
Ent. Soc., 14: 40-47.

Peck, S. B.

1972. The eyeless European soil Colydiid, Anommatus duodecimstriatus ,
in North America (Coleoptera; Colydiidae). Coleop. Bull., 26:
19-20.

1973. A systematic revision and the evolutionary biology of the

Ptomaphagus ( Adelops ) beetles of North America (Cole-

optera, Leiodidae; Catopinae), with emphasis on cave-inhabiting
species. Bull. Mus. Comp. Zool., 145(2): 29-162.

1974. The food of the salamanders Eurycea lucifuga and Plethodon
glutinosus in caves. Nat. Speleol. Soc. Bull., 36(4):1-10.

Valentine, J. M.

1942. On the preparation and preservation of insects, with particular
reference to Coleoptera. Smithsonian Misc. Coll., 103(6): 1-16.

POST-IMMOBILIZATION WRAPPING OF PREY
BY LYCOSID SPIDERS OF THE
HERBACEOUS STRATUM1

By Jerome S. Rovner and Susan J. Knost
Department of Zoology, Ohio University, Athens, Ohio 45701

The use of silk for wrapping prey is generally associated with
those spiders that construct trapping webs; nevertheless, there are
indications in the literature that wandering spiders sometimes use
silk for this purpose. In some of the latter cases the silk is applied
after the prey has been subdued by biting, a use corresponding to
that of the diguetid and linyphiid web-weavers studied by Eberhard
(1967). Post-immobilization wrapping in the latter two families
was suggested to prevent the prey from falling out of the web during
subsequent attacks (ibid.), and in araneid spiders to have the same
and additional roles, depending on species and prey size (Robinson
et al., 1969). The function of this behavior in the non-web-weaving
ctenids (Melchers, 1963), theraphosids (Eberhard, 1967), and
lycosids (Rovner, 1971) remained unknown and was the subject of
the present study. Our findings suggested that post-immobilization
wrapping by wandering spiders serves the same general function that
it probably does in web-weavers — to prevent prey from dropping
from the spider’s elevated location down to the ground whenever it
is released from the chelicerae during feeding, grooming, or subse-
quent capture attempts.

Methods

We observed individuals of Lycosa rabida Walckenaer (females
— 12-19 mm), Lyoosa punctulata Hentz (females = 13-15 mm),
and Schizocosa crassipes (Walckenaer) (females — 8-10 mm) for
the presence of and nature of prey wrapping. Whereas S. crassipes
was collected during spring on forest leaf litter, both species of
Lycosa were found in grassy fields, L. rabida during the summer
and L. punctulata during early fall. The spiders were collected in
Athens Co., Ohio, USA.

Spiders were housed individually in plastic cages (70 X 125 X
70 mm high) with water available ad lib., and given mealworms
(larvae of Tenebrio molitor) for maintenance feedings. Laboratory

^This study was supported in part by National Science Foundation Grant
GB 35369 to J. S. Rovner.

Manuscript received by the editor September 20, 1974.

398

1974]

Rovner Knost — Wrapping of Prey

399

temperatures averaged 25.8 zb 2. 7° (SD) over the entire investi-
gation.

Initial observations were made on spiders in boxes identical to
their bousing cages. Each individual had been food-deprived for up
to 7 days. In this phase of our study we used the following types
of prey: terrestrial isopods ( Armadillidium sp.), 6-8 mm; Japanese
beetles ( Popillia japonica)} 10-12 mm; mealworm larvae, 18-20 mm;
small grasshoppers (Cyrtacanthacridinae) , 6-8 mm; small ground
crickets (Nemobiinae) , 10-13 mm; and vestigial-winged fruit flies
( Drosophila ?nelanogaster) , 2 mm. We offered three prey at time
— O, and additional single prey at + 10 and +20 min (all of the
same species) in most of these tests. (For fruit flies, groups of eight
were added at each of the three times.) Observation periods were
at least 30 min in duration. The spider then was returned to its
home cage with its captured prey.

To determine the functions of the various spinnerets in wrapping,
we sealed pairs of spinnerets with paraffin while female L. rabida
were under C02 anesthesia. Two females underwent sealing of the
anterior spinnerets; two had the anterior and median spinnerets
sealed; and two, the median and posterior spinnerets. These spiders
were subsequently observed and filmed (Bolex Macrozoom Super
8 mm camera, Model 160) during prey-wrapping. We also exam-
ined the silk on the prey and substratum, after chasing away the
female from her just-wrapped prey.

To study the preference of L. rabida (and, to some extent, L.
punctulata) for the ground vs. the herbaceous stratum, we con-
structed artificial “field habitats” in three terraria (0.2 X 0.4 X 0.2
m high). Cardboard “foliage” was fixed in a plaster base which
provided the “ground surface” on 24 of the terrarium bottom. The
remainder of the glass bottom was left uncovered to provide an
alternative ground surface in case the plaster had a repellent effect
on the spiders (which turned out not to be the case). Vertical and
sloping surfaces projecting into space were provided by the artificial
foliage (Fig. 1). Spiders could climb to a height (limited by a glass
lid) of 0.19 m.

Three female L. rabida (tagged with non-toxic enamel) were
housed in each terrarium. (Three individuals were cannibalized in
the course of this experiment and were replaced with equivalently
tagged substitutes.) To prevent possible conditioning effects, we fed
the spiders on the ground as often as on the foliage. (Feeding was
done at times other than the daily observation periods.) We observed
the nine spiders for 2-hr blocks of time each day for 10 days during

400

Psyche

[September-December

Fig. 1. Female Lycosa rabida (with prey) on artificial foliage.

the afternoon (on most days, 1300-1500), and recorded the duration
of time each spider spent on the ground vs. the foliage. At ^-hr
intervals we also measured the height of those spiders that were on
the foliage. After completing this phase of our study, we observed
and filmed prey capture and wrapping by spiders on the artificial
foliage. Analysis of these and other films was aided by use of a
Bell & Howell Super 8 mm Multi-motion projector.

Field observations were made on L. rabida during daylight hours
(usually 1300-1500), and during both day (1300-1500) and night
(2030-2200) conditions on L. punctulata. The spiders were detected
at a distance and their location within the habitat noted. By ap-
proaching carefully, we were able to film undisturbed L. rabida in
the field.

Prey-wrapping Behavior

Whereas individuals of S. crassipes did not use silk after prey
capture, those of L. rabida and, to a lesser extent, L. punctulata
did so. In the latter two species, silk was not used to immobilize
the prey, since wrapping began after the prey had ceased most of its
struggling, usually several min after capture (Table 1).

1974]

Rovner & Knost — JV rapping of Prey

401

Table I. Bouts of prey-wrapping by female Lycosa rabida {*) and Lycosa
punctulata (f) when offered multiple prey during a 30-min period. A
total of 12 spiders was tested on each type of prey. (Sample sizes in
parentheses.)

Prey

Bouts/

captures

Mean

revolutions/

bout

Mean

duration

(sec)

Mean onset
after bite
(min)

^Mealworms

16/40

4.5(15)

36(13)

11.4(8)

^Grasshoppers

28/27

4.4(27)

34(22)

2.7(9)

^Crickets

50/40

3.2(48)

26(39)

1.1(8)

fCrickets

6/32

2.2 ( 6)

16 ( 6)

9.3(3)

When beginning a bout of wrapping, the spider spread and wiggled
its spinnerets, flexed its abdomen ventrad, and then pressed its anterior
spinnerets to the substratum. (Sometimes additional attachment
disks were placed on the substratum in this manner within an arc
of 6o° or less.) Then the spider began to pivot above the prey.
In about 10% of the cases the spider pivoted ^4 revolution (or
sometimes revolution), fixed an attachment disk to the substratum,
and then began to turn in the opposite direction. However, in most
cases the spider pivoted in only one direction, while remaining cen-
tered above the prey. During the first revolution, the silk was
attached to the substratum at 2-5 points, the number perhaps de-
pending on the prey’s bulk. Fewer attachment disks were placed
during the next turn or two, and usually only one or none during
later revolutions.

During the first two or three turns, the spider usually held the
prey in its chelicerae (Fig. 2). Consequently, as the spider revolved,
the prey animals beneath it pivoted around with the spider. (Very
large animals, while not held in the chelicerae during wrapping, and
therefore not having their weight supported by the spider, were
contacted by the palps, chelicerae, and sternum of the spider pivoting
above.) If, as was the case with most prey, the spider held the prey
in its chelicerae during early revolutions, it then released the prey
during a subsequent turn and continued to pivot for one or more
revolutions (Fig. 3). Throughout the process, the palps were used
to contact or manipulate the prey. In L. rabida 80% of the wrapping
bouts involved four or fewer revolutions (mean = 3.3 ± 2.1 SD;
range = 1-14; n — 106). Some views of one such bout are sketched
in Fig. 4.

All three pairs of spinnerets were used during prey-wrapping.
Data obtained from experimental animals indicated the same roles

402

Psyche

[September-December

Fig. 2. Female Lycosa rabida placing an attachment disk on the sub-
stratum during the early phase of prey-wrapping. The multiple prey are
held in her chelicerae and palps.

Fig. 3. Female Lycosa rabida during the later phase of prey-wrapping.
The prey have just been released from her cheliceral grasp. (Her palps
continue to contact the prey.) Swathing silk issues from her posterior
spinnerets.

1974]

Rovner & Knost — W rapping of Prey

403

for the spinnerets in lycosids as in other spiders that wrap prey.
When the anterior spinnerets were sealed, no attachment disks were
produced. The prey was wrapped but not fixed to the substratum.
The same was true when the anterior and median spinnerets were
sealed. When the median and posterior spinnerets were sealed,
attachment disks were fixed to the substratum and dragline laid
around the prey, but no swathing silk. In this case wrapping was
not effective. Multiple prey were not tied together; indeed, the prey
items were often pushed apart by the legs of the pivoting spider.

As the result of normal wrapping behavior, silk was placed around
the bodies of the prey animals and attached to the substratum at
intervals. Although the pivoting spider had pulled the threads taut,
the silk was not dense enough to produce a very tightly wrapped
package, as is produced by orb-weavers, for example. Indeed the
wrapping was so sparse that it would be unnoticed by the casual
observer. Nevertheless, the prey group did form a more compact
mass than it had prior to wrapping. When the spider resumed feed-
ing, it lifted the prey away from the substratum a. short distance
(rather than lean down to feed) ; however, the lines running to the
substratum generally remained intact due to their elasticity.

After prey-wrapping, the spider sometimes groomed its chelicerae
and palps, while remaining directly above the prey, and then resumed
feeding. Additional prey that approached within reach were often
captured; and one or more bouts of wrapping followed. However,
regardless of whether further captures were made after the first bout
of wrapping, subsequent bouts on the previously wrapped prey
occurred later in the observation period in many cases.

Table II. Number of Lycosa rabida making single vs. multiple captures
when offered multiple prey during a 30-min period (second figure) and the
number of those captures in which wrapping occurred (first figure). Twelve
spiders of each sex were tested on each type of prey.

Males Females

Prey

Single

capture

Multiple

capture

Single

capture

Multiple

capture

Isopods

— / 0

— /o

1/ 4

— / 0

Beetles

0/ 1

— /O'

2/ 6

— / 0

Mealworms

0/ 9

— /o

1/ 3

8/ 9

Grasshoppers

2/ 7

2/2

1/ 1

11/11

Crickets

0/ 4

2/3

— / 0

12/12

Totals

2/21

4/5

5/14

31/32

404

Psyche

[September-December

D

H

1974]

Rovner & Knost — Wrapping of Prey

405

L. rabida wrapped various types of prey, especially in situations
in which multiple prey were captured (Table II). While only 19%
of the males captured multiple prey, 68% of the females did so.
Considering both sexes together, wrapping occurred in 20% of the
cases in which only single captures were made, and in 95% of the
cases in which two or more prey were captured. Such correlations
meant that females wrapped prey much more often than males,
performing at least one bout of wrapping in 78% of the observation
periods in which they captured prey (single or multiple), while males
did so only 23% of the time. None of our spider species wrapped
fruit flies.

As shown in Table I, individuals of L. punctulata wrapped prey
much less often than L. rabida and did so less intensively (fewer
turns and shorter bout duration). Furthermore, it was only four
of the twelve females that did so; none of the twelve males wrapped
prey.

Stratum Selection

Field observations. — We watched penultimate and adult L. rabida
on sunny, calm days (early July) in a grassy field near the edge of a
woods. Undisturbed individuals rested motionless on the leaves and
stems of grasses and other vegetation. The spiders typically were on
horizontal or sloping surfaces (rather than vertical ones). Those
individuals we happened to observe were on the upper leaves of
relatively low vegetation or part-way up the stems and leaves of
taller vegetation.

In response to mild disturbance, the spiders walked or climbed a
short distance and then resumed their motionless stance. Strong
disturbance (as that which resulted from our walking near them
during our collecting efforts) caused the spiders to run and climb

Fig. 4. [Opposite page]. Sketches, based on a Super 8 mm film,
showing dorsal views of prey-wrapping by a female Lycosa rabida. The
several items of the multiple prey-capture are drawn as an homogenous
mass (stippled) for convenience, and should not be regarded as a wrapped
bundle. Lighting conditions permitted seeing the silk in views B, E, and

G. Times were rounded to the nearest sec. A. Placing 1st attachment disk
(Time=0). B. One-quarter into the 1st revolution ( + 4 sec). C. Near end
of 1st revolution (+ 7 sec). D. One-third into 2nd revolution (+9 sec).
E. Near end of 2nd revolution (+ 12 sec). Prey were released from the
chelicerae at the beginning of the next revolution. F. One-third into 3rd
revolution ( + 14 sec). G. Beginning of 4th revolution ( + 18 sec).

H. End of 4th and final revolution (+21 sec).

406

Psyche

[September-December

swiftly through the foliage, sometimes breaking into a series of leaps
and dashes that carried them long distances along the top of low-
growing vegetation in a few seconds. Thus, these animals usually
remained in the herbaceous stratum; only occasionally did an in-
dividual plunge downward and hide near the ground.

Penultimate L. punctulata were seen in the fall in the same fields,
and typically traveled through the herbaceous stratum when dis-
turbed. At night these spiders were seen resting on or moving
slowly upon the vegetation or, much less frequently, upon the ground.

Laboratory studies. — The results of our study of diurnal stratum
selection by undisturbed female L. rabida in an artificial habitat are
summarized in Fig. 5. (Temperature was 25-26° and relative hu-
midity > 60% throughout the 10 days.) During the 20-hr survey,
the spiders were on the cardboard foliage 73% of the time. Most
of that time the animals rested motionless on one of the “leaves” at
an average height of 0.12 m, i.e., more than half-way up the “plant.”
The long axis of their bodies was usually in a plane ranging from
nearly horizontal to a slope of about 6o° from the horizontal. In
the latter situation the spider faced up- or downward. In those
cases in which the spider rested vertically on a “stem” the spider

INDIVIDUAL

Fig. 5. Stratum selection by female Lycosa rabida in an artificial
habitat. During 20 hr, spread over 10 days, we recorded the proportion
of time spent on cardboard foliage (solid bar) vs. on the ground. At half-
hr intervals within observation periods, we also measured the height of
those spiders that were on the foliage (hatched bar).

1974]

Kovner & Knost — Wrapping of Prey

407

typically faced downward. Individuals rested more often in an
upright position on the upper surface of the leaf (illumination —
approx. 400 lux) than they did in an inverted position beneath the
leaf (illumination = approx. 130 lux).

We considered each spider’s behavior within the 2-hr periods of
observation and noted that each remained within the “herbaceous
stratum” during the entire 2 hr in 5.3 (range = 4-7) of the ten
periods. Each spider remained on the ground throughout the 2-hr
period in only 1 .6 ( range = 0-3 ) of the ten periods. The spider
moved from the ground to the foliage or vice versa in the remaining
3.1 (range = 2-4) of the ten periods.

Some observations made on several female L. punctulata in the
artificial habitat indicated that these spiders also tended to spend
their time resting on the foliage. However, we did not quantify
this part of our study.

After completing the 1 0-day survey of stratum selection by L.
rabida, we observed and filmed prey-wrapping by these spiders on
the artificial foliage. The behavior resembled that performed on the
floors of the housing containers, as we described above. Wrapping

Fig. 6. Female Lycosa rabida wrapping prey on artificial foliage. This
leaf sloped at an angle of about 60° to the horizontal.

408

Psyche

[September-December

usually was performed at the capture site and therefore was carried
out in all planes, including the vertical. The spiders behaved on
sloping leaves and vertical stems in a manner similar to that seen on
the horizontal cage floors; however, pivoting was slower on the
narrow surfaces, apparently due to the spider’s having to seek foot-
holds as it revolved “above” the prey (Fig. 6).

In some cases, we intentionally disturbed the spider before or
after prey-wrapping. In the former situation, the prey sometimes
was released from the chelicerae and dropped to a lower leaf or to
the ground. On the other hand, if wrapping already had occurred,
they prey did not 'fall from the capture site when released from the
chelicerae, even on a vertical surface. When multiple prey were
wrapped on a sloping surface, they : ( i ) remained together rather
than roll apart and (2) hung at the wrapping site rather than drop
to the ground, when the spider was chased away.

A further bit of evidence for the adaptedness of L. rabida to the
herbaceous stratum was provided later by the construction of a
“shelter web” (about 50 mm3) and, within it, an egg sac (infertile)
by a female on a nearly horizontal leaf 0.13 m above the ground in
the artificial habitat.

Discussion

Preference for herbaceous stratum. — S. crassipes is well-known
as an inhabitant o'f woodland leaf-litter (Kaston^ 1948; Fitch, 1963) ;
indeed, lycosids are generally regarded as ground-dwellers (Lowrie,
1968). However, L. rabida and L. punctulata can be collected by
sweeping grass and shrubs (Kaston, 1948). Kuenzler (1958) also
found L. rabida on the lower trunk and branches of trees in open
woodland. Whitcomb et al. (1963) describe finding L. rabida in
cotton fields on the ground during the day, but at night “. . . half-
way up the cotton stalks feeding on bollworm or cabbage looper
moths . . .” as well as on noctuid moths. At night, Eason and
Whitcomb (1965) collected 80% of their specimens of L. rabida
2/3 m or more above the ground in tall grass and bushes.

According to Eason and Whitcomb (1965), " L . punctulata is
mostly captured on or near the ground, since it has less tendency to
climb into bushes than does L. rabida ” While finding both species
in grassland, Fitch (1963) reported that L. punctulata preferred
“. . . relatively open or barren situations as compared to L. rabida ,
which prefers a tail-grass habitat.” Kaston (1948) stated simply
that both species have the same habits.

1974]

Kovner & Knost — Wrapping of Prey

409

Our field and laboratory observations are in agreement with the
general idea expressed above — L. rabida and L. punctulata are
adapted for spending a significant part of their life in the herbaceous
stratum, rather than being, like most lycosids, ground-dwellers.
(Obviously, stratum selection may vary during the life of an in-
dividual under different conditions, both external (temperature,
humidity, wind, etc.) and internal (hunger, sexual state, etc.)). A
difference between the species in regard to habitat or stratum selec-
tion, suggested by some authors, was not examined by us, but might
be revealed by quantitative field and laboratory studies. The rela-
tively longer legs of L. rabida suggests a greater adaptation to the
herbaceous stratum than that of L. punctulata, as does the stronger
tendency of L. rabida to wrap prey.

While these lycosids are often seen running about, perhaps mi-
grating from one habitat to another in response to micro-climatic
conditions, or perhaps merely reacting to the collector’s movements,
most of their time is probably spent at rest on the foliage. This
seemed to be the case in the field, and was certainly the situation in
the artificial habitat. As has been noted elsewhere (Edgar, 1969),
lycosid spiders generally remain motionless and wait for their prey
to come within reach. Like the web-weaving spiders, the wandering
spiders of the herbaceous stratum are resting on a medium that pro-
vides effective transmission of vibrational cues from the prey.

Function of prey-wrapping. — We hypothesize that post-immo-
bilization prey-wrapping in L. rabida and L. punctulata is a be-
havioral adaptation for life in the herbaceous stratum. It reduces
the possibility of losing the prey items when the spider releases them
from its cheliceral grasp between bouts of feeding. The spider may
release its grasp to make an additional capture, to groom, to drink,
or to locate another point on the prey’s body for further feeding.
Also, when startled by another animal or by shaking of the sub-
stratum due to wind, the spider’s momentary release of the prey
would often result in its loss. No such problem exists for ground-
dwelling forms.

Our hypothesis is supported by the absence of wrapping after the
capture of small prey, which are easily held in the chelicerae and
ingested in a relatively short time. It appears, then, that post-immo-
bilization wrapping serves the same primary function in wandering
spiders as it does in web-weavers — to prevent the prey from dropping
to the ground from the spider’s elevated location. Also, it is im-
portant when multiple prey are captured by a wandering spider,

4io

Psyche

[September-December

since the number of prey that the spider can hold in its chelicerae
during feeding is limited.

As to the functions of the components of prey-wrapping, we sug-
gest the following. The swathing silk serves to enclose the prey, a
role that is not only important for holding multiple prey together,
but which is also of value in holding together the pieces that often
result from the process of ingesting a single prey item. This latter
function may be one reason for the occurrence of additional bouts
of wrapping during the course o'f feeding. The dragline may be
entangled in the swathing band and serve to attach the prey package
to the substratum via the attachment disks, although it seemed that
the swathing band itself was connected to the attachment disks.
Since attachment to the substratum seems to be of prime value in
preventing loss of acquired food material, the re-anchoring that
occurs in additional wrapping bouts during the course of feeding
probably makes up for the breakage of lines that accompanies the
physical and chemical processing of the prey by the feeding spider,
and thus may be another reason for extra bouts of wrapping.

The temporal pattern is one of providing multiple attachment
points, along with swathing, during the early phase of each bout
(while the prey is held in the chelicerae) and then devoting most
of the remaining time to swathing (after the prey has been released
from the chelicerae). The early phase (attachment) may be im-
portant for effective wrapping, since it enables the spider to release
the prey and pivot around it, pulling the swathing lines taut.

Role of aciniform and other silk glands. — Comstock (1912) and
others have described the aciniform glands of web-weaving spiders
as the source of the swathing silk used for prey-wrapping, as well as
the fine fibers of egg sacs. These glands are present in lycosids, but
their function had remained unknown (Richter and Van der Kraan,
1970). On the basis of histological data, Richter (1970a) suggested
roles for the aciniform glands in lycosid spiders - — - in producing silk
involved in molting, egg sac construction, and sperm web construction.

Our data indicate that the aciniform glands also have one of the
same functions in lycosids as they do in the web-weavers — produc-
tion of swathing silk for wrapping prey. When the spigots for these
glands were closed experimentally (by sealing the median and
posterior spinnerets), wrapping behavior was ineffective. Such spiders
pivoted, fixed attachment disks, and laid a dragline; however, no
broad bands of silk were placed on the prey. Indeed, the several
items of multiple prey were often pushed apart by the legs of the
pivoting spider.

1974]

Rovner & Knost — W rapping of Prey

411

Our sealing of the anterior spinnerets (site of spigots for the
piriform glands) prevented attachment disk production, as was well-
known in the literature. Such spiders could wrap the prey but not
attach it to the substratum. This was also the case with spiders that
had their anterior and median spinnerets sealed, with the further
limitation of being incapable of producing draglines. In summary,
for complete wrapping of the prey, these lycosids are using all three
pairs of spinnerets and employing silk from three types of silk glands
(the third being the ampullaceal glands — for production of drag-
lines— whose spigots are on the anterior and median spinnerets).
This agrees with observations made by Melchers (1961) on prey-
wrapping by normal individuals of Cupiennius salei , a large ctenid
spider.

Since the wrapping of prey by lycosids (and at least some other
wandering spiders) probably correlates with a. preference for the
herbaceous (or higher) stratum, we hypothesize that the size of the
aciniform glands, source of swathing silk, would also. This expecta-
tion is based on Richter’s (1970^) demonstration of such a corre-
lation between ampullaceal gland size (dragline silk) and preferred
vegetation structure in Pardosa spp. Indeed, we should expect lyco-
sids with larger ampullaceal glands to have larger aciniform glands,
based on our hypothesis and on Richter’s findings.

Stimuli releasing prey-wrapping. — The correlation between the
frequencies of prey-wrapping and multiple prey-capture suggests that
the latter event provides a stimulus situation to release prey-wrap-
ping. The value of wrapping for retaining multiple prey has been
discussed above. However, wrapping certainly occurs after single prey
are captured, if these prey are about as large or larger than the
spider’s body size. It seems, then, that the stimulus for post-immo-
bilization wrapping is that of ,a relatively large volume or mass of
prey material beneath the spider, not the absolute number of prey
captured. We suspect volume rather than weight to be the stimulus,
since very large prey are wrapped without being held in the cheli-
cerae following a capture in which the bite was maintained without
the spider supporting the weight of the prey resting on the sub-
stratum.

Prey-wrapping was initiated after the prey had ceased most or all
of its struggling. Groups of small crickets or grasshoppers, each
member of which succumbed quickly to the spider’s bite, were
wrapped within a minute or two after capture. Mealworms, which
performed active twisting of the free end of their body for a long
time, were not wrapped until about 10 min after capture. Very

412

Psyche

[September-December

large, single prey were wrapped only after a lengthy period, i hr or
more, had elapsed. It appears that the absence of vigorous movements
by the prey is a necessary part of the stimulus situation for the onset
of wrapping.

Post-immobilization wrapping in other wandering spiders. — Our
two species of Lycosa performed post-immobilization wrapping,
whereas Schizocosa crassipes did not. We modified our original
speculation that this behavior might be characteristic of the Lycosidae
to the notion that it might be at least genus-specific. Even this is
no longer tenable, in light of our preliminary observations on other
species as well as data obtained by R. J. McKay (personal commu-
nication). We now regard post-immobilization prey-wrapping as a
behavioral adaptation that may be included in the repertoire of
various unrelated wandering spiders that have specialized for spend-
ing part or most of their time some distance above the ground on
vegetation.

Post-immobilization wrapping occurs after the prey has stopped
struggling, occupies a relatively brief amount of time during the
lengthy feeding process, and is used for prey only when the total
volume exceeds some threshold value. These are the probable rea-
sons why this behavior has not yet been reported in many species in
which it is likely to occur. Authors specifically studying feeding
behavior have seen post-immobilization wrapping in two other fam-
ilies of wandering spiders — ctenids (Melchers, 1961, 1963) and
theraphosids (Eberhard, 1967), although neither of these authors
determined the function.

A personal communication from E.-A. Seyfarth, that wrapping
occurred in Cupiennius salei , called our attention to the work of
Melchers (1961, 1963) on this ctenid species. (Comparison with our
species was facilitated by our being able to view Melchers’ (1961)
film of this behavior.) Post-immobilization wrapping by C. salei is
very similar to what occurs in our Lycosa spp. as to the sequence of
events and movements, stimuli releasing the behavior, etc. However,
two differences are obvious : ( 1 ) C. salei waits for prey on a vertical
surface. If it tumbles to the ground during prey capture, it carries
the prey back up to an elevated point on the vertical surface — the
glass wall of the terrarium in Melchers’ study. Individuals of our
species never did the latter, but readily stayed on the ground to feed
if the capture ended there. (2) The density and breadth of the
swathing bands are much greater in C. salei than in our Lycosa spp.
Points 1 and 2 are probably interrelated and suggest that this ctenid

1974] Rovner & Knost — Wrapping of Prey 413

spider is even more specialized for living on vegetation than are our
Lycosa spp.

Scopula hairs, which enable spiders to climb smooth surfaces, have
been studied in lycosids, theraphosids, and other wandering spiders
(Foelix and Chu-Wang, 1974). Since these structures are an im-
portant adaptation for many of the spiders which spend their life on
vegetation, they may serve as a morphological indicator of behavioral
correlates o'f this life style, including post-immobilization prey-wrap-
ping. Indeed, a scopula is present in those species in which this
behavior has been seen. On the other hand, since scopula hairs on
the tarsus and metatarsus are probably important for the capture
and manipulation of prey (Rovner and Knost, in preparation), their
presence may not always be indicative of habitat preference and
related behaviors.

Experimentally modified male L. rabida that are unable to achieve
palpal insertions “tie-down” females during attempted copulation
(Rovner, 1971), a behavior resembling post-immobilization wrapping
of large prey. Male thomisid spiders perform a similar behavior as
part of their normal pre-copulatory activity. (In neither case does
the wrapping have any restraining effect on the female, who easily
tears the threads and departs when the copulation is completed.)
Perhaps the pre-copulatory wrapping in thomisids is a ritualized
behavior derived from a present or past use of silk for wrapping
prey in this family.

Eberhard (1967) associated post-immobilization wrapping with
certain primitive types of aerial web-building, similar to that ex-
hibited by the present-day diguetid spiders. The behavior shown
by our Lycosa spp. of the herbaceous stratum, and by other wander-
ing spiders, suggests that the evolution of post-immobilization wrap-
ping may not be as closely associated with the building of aerial webs
as Eberhard had hypothesized.

Summary

Post-immobilization wrapping of large, single prey and groups of
smaller prey occurs in Lycosa rabida and, to a lesser extent, Lycosa
punctulata. Observation of these spiders in both the natural habitat
and an artificial habitat indicated their preference for the herbaceous
stratum. Post-immobilization wrapping is probably an adaptation for
life in this stratum, since it reduces the possibility of losing prey when
it is released from the cheliceral grasp. The swathing bands, at-
tachment disks, and draglines involved in wrapping are produced

414

Psyche

[September-December

with silk from the aciniform, piriform, and ampullaceal glands re-
spectively. Post-immobilization prey-wrapping, well-known in web-
weavers and now known in ctenids, lycosids, and theraphosids, is
likely to be found in other wandering spiders that live on foliage.

Literature Cited

Comstock, J. H.

1912. The Spider Book. Garden City, N. Y. : Doubleday, Page. 725

pp.

Eason, R. and W. H. Whitcomb

1965. Life history of the dotted wolf spider, Lycosa punctulata Hentz
(Araneida: Lycosidae). Proc. Arkansas Acad. Sci. 19: 11-20.
Eberhard, W.

1967. Attack behavior of diguetid spiders and the origin of prey
wrapping in spiders. Psyche 74: 173-181.

Edgar, W. D.

1969. Prey and predators of the wolf spider Lycosa lugubris. J. Zool.,
Lond. 159: 405-411.

Fitch, H. S.

1963. Spiders of the University of Kansas Natural History Reserva-
tion and Rockefeller Experimental Tract. Misc. Publ., Mus. Nat.
Hist., Univ. Kansas 33: 1-202.

Foelix, R. F. and I. W. Chu-Wang

1974. Scopula hairs in spiders. Proc. Sixth Int. Congr. Arach. In
press.

Kaston, B. J.

1948. Spiders of Connecticut. Bull. State Geol. Natur. Hist. Surv.,
Hartford 70: 1-874.

Kuenzler, E. J.

1958. Niche relations of three species of lycosid spiders. Ecology 39:
494-500.

Lowrie, D. C.

1968. The spiders of the herbaceous stratum of the Jackson Hole
region of Wyoming. Northwest Sci. 42: 89-100.

Melchers, M.

1961. Cupiennius salei (Ctenidae) Einspinnen der Beute und Nahrung-
saufnahme. Encycl. Cinemat., E 421.

1963. Zur Biologie und zum Verhalten von Cupiennius salei (Keys-
erling), einer amerikanischen Ctenide. Zool. Jahrb. Abt. System.
91: 1-90.

Richter, C. J. J.

1970a. Morphology and function of the spinning apparatus of the wolf
spider Pardosa amentata (Cl.) (Araneae, Lycosidae). Z. Morph.
Tiere 68: 37-68.

1970#. Relation between habitat structure and development of the
glandulae ampullaceae in eight wolf spider species ( Pardosa ,
Araneae, Lycosidae). Oecologia (Berl.) 5: 185-199.

Richter, C. J. J. and C. Van Der Kraan

1970. Silk production in adult males of the wolf spider Pardosa amen-
tata (Cl.) (Araneae, Lycosidae). Neth. J. Zool. 20: 392-400.

1974]

Kovner & Knost — Wrapping of Prey

415

Robinson, M. H., H. Mirick, and O. Turner

1969. The predatory behavior of some araneid spiders and the origin
of immobilization wrapping. Psyche 76: 487-501.

Rovner, J. S.

1971. Mechanisms controlling copulatory behavior in wolf spiders
(Araneae: Lycosidae). Psyche 78: 150-165.

Rovner, J. S. and S. J. Knost

Prey capture by lycosid spiders. In preparation.

Whitcomb, W. H., H. Exline, and R. C. Hunter

1963. Spiders of the Arkansas cotton field. Ann. Ent. Soc. Amer. 56:
653-660.

ERRATUM. — In our paper on “Social 'Carrying Behavior and
Division of Labor During Nest Moving in Ants”, Psyche , Vol. 81,
No. 2, pages 219-236, the last line of the caption of Fig. 10 should
read: Dotted Line: Formica sanguinea; Solid Line: Camponotus
sericeus. — Michael Moglich and Bert Holldobler (Biological Dept.,
MCZ-Laboratories, Harvard University).

PTERALIA OF THE PALEOZOIC INSECT ORDERS
PALAEODICTYOPTERA, MEGASECOPTERA AND
DIAPHANOPTERODEA ( PALEOPTERA) *

By Jarmila Kukalova-Peck
Department of Geology, Carleton University
Ottawa, Ontario, Canada

For an understanding of insect evolution the structure of the wing
base is of major significance. However, the fossil record of pteralia
in extinct orders is extremely scanty. This paper is concerned with
the wing bases of certain Paleozoic Paleoptera, namely, Palaeodictyop-
tera, Megasecoptera and Diaphanopterodea from the Upper Carboni-
ferous (Namurian) of Czechoslovakia, the Upper Carboniferous
(Stephanian) of France, and the Lower Permian of Czechoslovakia
and Kansas. Independently of the Neoptera, the Diaphanopterodea
acquired the ability to flex the wings backwards over the abdomen.
In this respect, the order is of special interest, and an attempt is
made here to compare the pteralia of the Diaphanopterodea with
those of extant Ephemeroptera.

Our present knowledge of the wing base in Paleozoic Paleoptera
is restricted to the axillary plate of several palaeodictyopteran adults
(Kukalova, i960, 1969-70), and palaeodictyopteran nymphs (Woot-
ton, 1972; Sharov, 1971) [See figures 1, 2 and 4]. Recently, the
wing base has been described in the Diaphanopterodea, Family El-
moidae (Kukalova-Peck, 1974) (fig. 8). In the present paper, the
axillary plates in Martynoviidae and Asthenohymenidae of the Order
Diaphanopterodea are included, and for the first time the axillary
sclerites in Megasecoptera are described.

The interpretation and terminology of the pteralia in extinct Pale-
optera are necessarily dependent upon the detailed functional mor-
phology of extant Ephemeroptera and Odonata. At the same time,
the wing base structures found in extinct orders provide an evolu-
tionary view and might be helpful in unraveling the enigmatic archi-

*This research has been aided in part by a Publication Grant from
Carleton University and in part by a National Science Foundation Grant,
GB 39720, F. M. Carpenter, Principal Investigator, Harvard University.
I am deeply indebted to Dr. Carpenter for his assistance in finding speci-
mens and for placing them at my disposal at the Museum of Comparative
Zoology. I also wish to express my sincere gratitude to Dr. R. J. Wootton
(University of Exeter, England) and Dr. P. A. Adams (California State
University at Fullerton) for their valuable critical comments.

Manuscript received by the editor December 15, 1974.

416

1974]

Kukalova-Peck — Pteralia

417

Fig. 1. Partially fused axillary plate of Boltopruvostia nigra (Palaeo-
dictyoptera, Homoiopteridae) . Enlarged base of fore wing. Redrawn
from Kukalova, 1960. API — anal plate; bg — basal groove; CuPl -—
cubital plate; MP1 — median plate; ScRPl — subcosto-radial plate. Upper
Namurian, Czechoslavakia.

tecture of extant Paleoptera. In the Odonata, the thorax and pteralia
have been discussed in detail by Tannert (1958), Neville (i960),
and Hatch (1966), but interpretation of these structures is still
subject to debate. The axillary plate is considered either as the fused
radio-anal plate, or as a compound structure, with the radio-anal
plate, axillary sclerites, and subalare incorporated. However, the
wings of the Odonata are highly specialized and possess many unique
features and therefore are less suited for comparison with Paleozoic
orders than the more primitive wings of the Ephemeroptera. The
pteralia in Ephemeroptera have been recently studied by Brodskyi
(1970), Matsuda (1970), and Tsui and Peters (1972) (fig. 5),
but the axillary plate was not described in functional terms; it was
mostly referred to as “median plate”. In this paper, the terminology
is derived from Tannert’s (1958) account on the Odonata and the
respective axillary plates are called by the terms of incorporated basal
plates of the veins. Since the fossil specimens do not contribute to
our knowledge of whether the other pteralia are fused with the
axillary plate, this problem should be considered as fully open to
future emendations based upon a detailed study of functional mor-
phology.

418

Psyche

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Fig. 2. Palaeodictyopteran nymphal wing pad with partially separate
basal plates. Idoptilus onisciformis (Graphiptilidae). After Wootton,
1972, F — -deep furrow along the thorax; PI — basal plates. Westphalian,
England.

Palaeodictyoptera and Ephemeroptera

In the most primitive, yet unknown, Paleoptera all main veins
probably originated at the wing base from the separate basal plates.
In the process of evolution, the basal plates gradually became fused.
The intermediate stage of partial fusion is documented in the primi-
tive palaeodictyopteron, Boltopruv ostia nigra (Namurian 'C of Czech-
oslovakia, fig. i ) . In this species the subcostal and radial plates are
fully grown together (forming the subcosto-radial plate), the median
plate is separate, and the cubital plate is fused with the small anal
plate (forming the cubito-anal plate). The partially separated basal
plates are also indicated in the wing pad of the nymph Idoptilus
h onisciformis (Wootton 1972, Westphanian of England) (fig. 2).

Within the evolutionary history of Palaeodictyoptera, full fusion
of the plates was acquired in the more specialized forms. Basal plates
fused together into a single subcosto-anal plate but still retaining an
indication of individual outlines, have been found in Moravia con-
vergens (Lower Permian of Czechoslovakia, fig. 3). In Palaeodicty-
optera from the Upper Carboniferous of France (fig. 4), the sub-
costo-anal basal plate is fully fused and its subcostal part becomes
reduced in size.

In the primitive Ephemeroptera (Siphlonuridae, fig. 6), the costal
brace in the fore wing starts at the anteroproximal angle of the
axillary plate ; in the hind wing, it sometimes starts from a transverse
elevation on the axillary plate, forming a convex hook directed against
the stem of M. The costal brace is, in all probability, homologous
to the postcostal vein of Palaeodictyoptera (fig. 4B, pv), plus the
cross vein between Sc and R. The base of ephemeropteran wings is
weakly sclerotized and the stems of the main veins adjacent to the
axillary plate are mostly not discernible. However, if the vein stems

1974]

Kukalova-Peck — Pteralia

419

in primitive mayflies are thoroughly examined, their connection with
the axillary plate can be traced. This is true in many Siphlonuridae,
especially in the hind wings (fig. 6B). In the fore wing of Siphlonu-
rus inirus (fig. 6A), the axillary plate is domed in the middle. At
the anteroproximal angle, there is a short groove, which is also present
in Palaeodictyoptera (fig. 1, 3). Expanded stems of Sc and R (which
might in this Siphlonurus be represented by the fused R + M + Cu
stems) meet at a suture and clearly penetrate the axillary plate. In
the hind wing of the same species, the groove at the anteroproximal
angle is reduced to a small pit. R is located at the anterior part of
the dome and is closely followed by the short stem of M, from which
MA and MP diverge almost immediately. The short stem of Cu

basal groove; pa — precostal area; ScAPl — subcosto-anal plate; tf —
transverse furrow. Lower Permian, Czechoslovakia.

420

Psyche

[September-December

meets MP and instantly splits into CuA and CuP. As described
above, all main veins, with the exception of the anal veins, start in
the hind wing directly and separately from the domed part of the
axillary plate, as in more advanced Palaeodictyoptera (fig. 4).

The main difference between the axillary region of the Palaeo-
dictyoptera and that of the Ephemeroptera is in the attachment of
the anal veins. In the Palaeodictyoptera, the anal veins probably
always originated directly from the subcosto-anal plate (figs. 3, 4),
or the anal plate was closely attached to the cubital plate (fig. 1).
The anal area in Palaeodictyoptera is often crossed by diverse convex
ridges, which are sometimes V-shaped ( Homaloneura , Kukalova
1969), and by transverse grooves (figs. 3, 4A), or the anal veins
start from a cuticular thickening ( Dunbaria , Kukalova-Peck 1971).
As pointed out by Wootton (1974, personal communication), these
structures are in the place of the anal brace of Ephemeroptera. In
the primitive mayflies, the anal veins start at the anal brace, which is
basally attached to a small plate, separated from the large axillary
plate. It seems possible that while the axillary plate originated by
fusion of the subcosto-cubital plates, the anal plate stayed detached.
This condition is partially reminiscent of Diaphanopterodea and
Megasecoptera, as will be shown below. The anal brace in Ephem-
eroptera is followed, both anteriorly and posteriorly, by two concave
furrows. By location and, very probably, by function for flight, the
furrows may be compared with the concave transverse furrow, cross-
ing the anal area in almost all Palaeodictyoptera (figs. 3, 4A).

The weak sclerotization of the axillary plate in Ephemeroptera and
the vanishing of the adjacent stems are probably the changes which

Fig. 4. Fused subcosto-anal plates of Palaeodictyoptera. A. Dictyop-
tilus sepultus (Eugereonidae) , enlarged base of the fore wing. B. Steno-
dictya parisiana ( Dictyoneuridae) , enlarged base of the hind wing. After
Kukalova, 1970. pa — precostal area; pv — postcostal vein; ScAPl — sub-
costo-anal plate; tf — transverse furrow. Upper Carboniferous, France.

1974]

Kukalova-Peck — Pteralia

421

occurred later in evolution. This conclusion is supported by the
morphology of primitive Permian mayfly families, the Protereismati-
dae and the Misthodotidae. In both, the stems of main veins were
strong, reaching the wing base, and the anal brace was less pro-
nounced than in extant mayflies (fig. 7). It should be noted that the
adult life span in Permian Ephemeroptera was probably longer than
in Recent species; the presence of large and apparently fully func-
tional mouthparts in the adult Permian Misthodotidae (Tschernova,
1965) is very suggestive that these particular mayflies, at least, were
active feeders. Flight was presumably important in obtaining food,
as well as in courtship activities.

According to Brodskyi (1970), the second axillary sclerite in
Recent Ephemeroptera is firmly fused with the axillary plate, al-
though this opinion is not shared by some other specialists; the sche-
matic figure of the axillary region in extant mayflies (Leptophle-
biidae) in the interpretation of Tsui and Peters (1972) is shown in
figure 5. In the fossil material described, no remnants of the axillary
region have been mentioned so far.

Fig. 5. Axillary region of fore wing of ephemeropteron, Aprionyx
tricuspidatus (Leptophlebiidae, adult male). After Tsui and Peters, 1972.
AxPl — axillary plate; Ax 1, 2, 3 — axillary sclerites; HP — humeral
plate; T — tegula. Recent.

Diaphanopterodea

The Diaphanopterodea have mouthparts, genitalia and wing vena-
tion pattern very similar to those of Palaeodictyoptera, from the
primitive forms of which they were probably derived (Carpenter,

422

Psyche

[September-December

Fig. 6. Subcosto-cubital plate of ephemeropteron, Siphlonurus mirus
(Siphlonuridae) . A. Fore wing; B. Hind wing. Original. Anb — anal
brace; API — anal plate; bg — basal groove; Cb — costal brace; ScCuPl
— ^subcosto-cubital plate; tf — transverse furrow. Recent.

1974]

Kukalova-Peck — P ter alia

423

1963). In comparison with the average size of Palaeodictyoptera,
they are much smaller and they developed the ability to flex the wings
backwards over the abdomen. The axillary plate of the wing base
and the basal fold were recently described by Kukalova-Peck (1974)
in Permodiapha carpenteri of the family Elmoidae (fig. 8A). Here,
the axillary plate is compared with that of the related family Marty-
noviidae as preserved in Martynovia protohymenoides (specimen
No. 4600, Museum of Comparative Zoology, Harvard University)
from the Lower Permian of Kansas (fig. 8B).

In both forms, the general patterns of the wing base structure are
similar. The larger part of the axillary plate in Martynovia is
formed by the fused medio-cubital plate, or perhaps by the cubital
plate itself. Different is the development of the anal plate, which is
distinctly outlined and two-lobed, located anteriorly to the basal fold.
The anal plate is firmly connected by the anterior margin with the
axillary plate. The basal fold encircles the axillary plate anteriorly
and distally and separates it from the sclerotized region, extending
between CuP and the posterior wing margin (fig. 8B, S). The
axillary plate in both Martynovia and Permodiapha is obliquely
crossed by a convex strut, which perhaps participated in the folding
of the wings. The incompletely preserved axillary sclerite contacts

,-C

Fig. 7. Fore wing base of Permian mayfly, Protereisma permianum
(Protereismatidae). After Carpenter, 1933. Lower Permian, Kansas.

424

Psyche

[September-December

Fig. 8. Medio-cubital plate of Diaphanopterodea. A. Permodiapha car-
penteri (Elmoidae), hind wing. After Kukalova-Peck, 1974. B. Marty-
novia protohymenoides (Martynoviidae) , fore wing. Original. API —
anal plate; Ax — axillary sclerite; B — basal fold; MCuPl — medio-
cubital plate; S — sclerotized region. Lower Permian, Czechoslovakia and
Kansas.

1974]

Kukalova-Peck — Pteralia

425

Fig. 9. Incomplete wing base of Asthenohymen dunbari (Diaphanop-
terodea, Asthenohymenidae) . Fore wing. Original. B — basal fold; S —
sclerotized region. Lower Permian, Kansas.

with the proximal margin of the axillary plate and with R. It is in
the position of 2 Ax in Recent Ephemeroptera ( sensu Tsui and Peters
1972, fig. 5).

In Asthenohymenidae, the incompletely preserved wing base, in-
cluding only the basal fold and part of the sclerotized region, is
known. The structure of the wing base, as preserved in Asthenohy-
men dunbari (specimen No. 3835, Museum of Comparative Zoology,
fig. 9) seems to be basically similar to that of Martynovia and
P ermodiapha.

The fused axillary plate of Diaphanopterodea is homologous only
to the posterior part of the subcosto-anal plate of Palaeodictyoptera.
The subcostal, radial, and possibly also the median basal plates be-
came completely reduced, perhaps in connection with narrowing of
the wing base, coalescence of veins and with acquisition of wing
folding ability.

Megasecoptera

The Megasecoptera and Palaeodictyoptera have very similar mouth-
parts, genitalia and wing venation pattern, and undoubtedly developed
from common ancestral stock. The Megasecoptera differed, accord-
ing to our previous understanding, only in the shape of the wings
and in the more pronounced crowding of Sc and R towards the
costal margin. However, the study of the wing base has revealed a

426 Psyche [September-December

— anal plate; Axe — convex axillary sclerite; Axe — elongated axillary
sclerite; np — narrow plate; ScPl — subcostal plate. Lower Permian,
Kansas.

major difference in the pteralia, not only with respect to Palaeodicty-
optera, but also to all other paleopterous orders.

The well preserved wing bases with axillary sclerites were found
in specimens of two comparatively small species: Protohymen permi-
anus (specimen No. 3060) and Protohymen readi (holotype No.
3258), in the Museum of Comparative Zoology. Since the structures
in both specimens are similar, only the wing bases of Protohymen
permtanus, which is better preserved, are described (fig. 10).

1974]

Kukctlova-Peck — Pteralia

427

Towards the base, the wing narrows markedly in the posterior
part, while the anterior margin remains almost straight. Shortly
before the base, the posterior margin curves posteriorly to form a
prolonged axillary lobe parallel with the tergal margin. The serrated
costa is attached at the very base to a long narrow plate on the lateral
margin of the tergum. This plate is connected anteriorly with a
prominent, darkly colored lobe. The subcosta is convexly bent near
the base. Both costa and subcosta terminate on a triangular, highly
convex basal plate, which is posteriorly connected to R + M and
proximally to the small, convex axillary sclerite. This is associated
with the tergal margin, the basal plate, the costa, the narrow plate
on the tergum, and the anterior arm of the large elongated sclerite.
The cubitus is concavely bent towards the very end of R shortly
before the base, so that it creates the mirror image of the subcosta.
The anal vein originates from a small anal plate, which is divided
by a perpendicular suture into two small equal sections. The anal
plate is directed obliquely and posteriorly against the posterior arm
of a large, anteroposteriorly elongated sclerite. This closely contacts
the tergum along almost its entire proximal margin. The elongated
sclerite is highly convex posteriorly from the level of the end of the
anal plate. In the anterior, flatter part, the axillary sclerite divides
into two arms. The proximal arm is longer and slightly bent to at-
tach to R + M and to the small axillary sclerite; the distal arm is
shorter and directed to connect the curved end of the anal plate.
This sclerite is markedly larger in the hind wing.

The megasecopteran pteralia, as described above (figs. 10, 11), are
so highly specialized that homology at our present level of knowledge
is nearly impossible. The narrow plate laterally on tergum (np)
and the elongate sclerite (Axe) as well seem to be compound struc-
tures in which several elements might be incorporated. The small
convex sclerite (Axe) is approximately in the position of iAx of
Ephemeroptera ( sensu Tsui and Peters 1972; fig. 5). The little
basal plate (ScPl) at the beginning of Sc is perhaps homologous to
the subcostal basal plate of hypothetical primitive Paleoptera. The
highly specialized character of megasecopteran pteralia shows that the
construction of the wing attachment in Paleoptera was even more
varied and diverse than it has been assumed. However, it should be
taken into consideration that the pteralia of Protohymen may not be
typical for the whole order Megasecoptera because of its relatively
small size. In small insects, reduction and other modification of
axillary sclerites sometimes occur.

428

Psyche

[September-December

Fig. 11. Reconstruction of Protohymen (Megasecoptera, Protohymenidae) .
All structures shown are preserved in specimens of Protohymen permianus
and P. readi, except for the meso- and metathoracic legs, which are re-
stored on the basis of the prothoracic legs. Original. Lower Permian,
Kansas.

1974]

Kukalova-Peck — Pteralia

429

Summary

In the several paleopterous orders, the “axillary plates” are by no
means to be considered as fully homologous structures. They are
formed by fusion of various basal plates, which can eventually vary
in the degree of fusion and in the individual size within a single
order (for instance in Palaeodictyoptera, and in Diaphanopterodea).

There is little doubt that in the hypothetical primitive ancestor of
Paleoptera the main veins originated from the separate basal plates.
From this condition, the axillary plate was derived by fusion of the
following plates : in Palaeodictyoptera, of subcosto-anal plates ; in
Ephemeroptera, of subcosto-cubital plates; in Odonata, of radio-
anal plates ; in Diaphanopterodea, of medio-cubital plates ; in Megase-
coptera, only small subcostal plate remained.

In spite of the variability mentioned above, the general plan of the
axillary region and adjacent part of the wings is shared by all paleop-
terous orders, and certain structures, functionally significant for the
flight, repeat many times throughout Paleoptera. For instance, sep-
arated or semi-separated anal plates occur in Magasecoptera, Dia-
phanopterodea, and Ephemeroptera. A deeply concave furrow cross-
ing transversely the anal area is found in almost all Palaeodictyop-
tera and in the Ephemeroptera. Transverse reinforcement of the
proximal half of the wing formed by coalesced veins and supporting
cross veins, cuticular thickenings and pigmented stripes is present in
Odonata and in “odonatoid” (dragonfly-convergent) Palaeodictyop-
tera, etc. These structures will not be understood until more study
of the functional morphology has been done, particularly on extant
mayflies, dragonflies, and on wings in general.

References

Brodskyi, A. K.

1970. Organisation of the Flight System of the Mayfly Ephemera
vulgata L. (Ephemeroptera). Ent. Rev. Wash., 49: 184-188.
Carpenter, F. M.

1933. The Lower Permian Insects of Kansas. Part 6. Delopteridae,
Protelytroptera, Plectoptera and a New Collection of Protodonata.
Odonata, Megasecoptera, Homoptera, and Psocoptera. Proc.
Amer. Acad. Arts & Sci., 68 (11): 411-503.

1963. Studies on Carboniferous Insects from Commentry, France.
Part V. The Genus Diafhanoptera and the Order Diaphanop-
terodea. Psyche, 70(4): 240-256.

Hatch, G.

1966. Structures and mechanics of the dragonfly pterothorax. Ann.
Ent. Soc. Amer., 50: 702-714.

430

Psyche

[September-December

Kukalova, J.

1960. New Palaeodictyoptera of the Carboniferous and Permian of
Czechoslovakia. Sbornik UUG, 25: 236-251.

1969- Revisional Study of the Order Palaeodictyoptera in the Upper

1970. Carboniferous Shales of Commentry, France. Part 1. Psyche,
76(2): 163-215; Part 2. Psyche, 76(4): 439-486; Part 3. Psyche,
77(1) : 1-44.

Kukalova-Peck, J.

1971. The Structure of Dunbaria (Palaeodictyoptera). Psyche, 78(4):
306-318.

1974. Wing-folding in the Paleozoic Insect Order Diaphanopterodea
(Paleoptera) , with a Description of New Representatives of the
Family Elmoidae. Psyche, 81 (2): 315-333.

Matsuda, R.

1970. Morphology and Evolution of the Insect Thorax. Mem. Ent. Soc.
Can. 76: 3-431.

Neville, A. C.

1960. Aspects of Flight Mechanics in Anisopterous Dragonflies. J. Exp.
Biol., 37: 631-656.

Sharov, A. G.

1971. Morphological Features and the Way of Life of Palaeodictyop-
tera. Dokl. 24. tschtenii pam. N.A. Cholodkovskogo. Akad. Nauk
SSSR, Moscow: 49-63.

Tannert, W.

1958. Die Fluegelgelenkung bei Odonaten. Deutsche Ent. Zeitschr.,
(N.F.), 5 (5) : 394-455.

Tschernova, O. M.

1965. Some Fossil Mayflies (Ephemeroptera, Misthodotidae) found in
Permian Deposits in the Ural Mountains. Ent. Rev. Wash., 44:
202-207.

Tsui, T. P. and W. L. Peters

1972. The Comparative Morphology of the Thorax of Selected Genera
of the Leptophlebiidae (Ephemeroptera). J. Zool., Lond., 168:
309-367.

WOOTTON, R. J.

1972. Nymphs of Palaeodictyoptera (Insecta) from the Westphalian
of England. Palaeontology, 15: 662-675.

SYSTEMATICS OF THE TRAPDOOR SPIDER GENUS
A LI A TY PUS (ARANEAE: ANTRODIAETIDAE)*

By Frederick A. Coyle

Department of Biology, Western Carolina University,
Cullowhee, North Carolina, 28723

Introduction

Aliatypus species are all rather stocky mygalomorph spiders (Figs.
45-49) which construct a burrow with a trapdoor entrance from
which they capture prey. In general morphology and behavior,
Aliatypus bears striking resemblance to the distantly related trapdoor
spider family Ctenizidae, but this similarity is clearly the result of
convergent, or at least parallel, evolution; Aliatypus is an atypoid
mygalomorph taxon most closely related to Antrodiaetus, Atypoides,
and the Mecicobothriidae. Aliatypus species appear to be restricted
to California and Arizona (Maps 1-4) where they live in ravine
banks, road banks, or other slopes in habitats ranging from hot, dry
sagebrush scrub communities to wet coast redwood forests and cool
California red hr mountain forests. They are among the most
abundant trapdoor spiders in California.

Aliatypus has been badly neglected; only one species has been
described (Banks, 1896; Smith, 1908) and little natural history
information has been published (Smith, 1908; Gertsch, 1949; Coyle,
1971). During the last seven years a concerted collecting effort,
largely by Wendell Icenogle and myself, has increased the availability
of adult specimens from a dozen to 330 and has thereby made pos-
sible this revision. My chief goal in this study has been to define
accurately the species limits by means of an analysis of variation.
The methods employed are essentially those of my earlier studies
(Coyle 1968, 1971) and are summarized in the Methods section of
this paper. Discussions of variation patterns are included in order
to improve our understanding of geographic variation in mygalo-
morph spiders and guide future research on Aliatypus. The consid-
erable amount of behavioral and ecological data which has been
collected will be published separately in a paper on Aliatypus natural
history.

* Manuscript received by the editor January 1, 1975.

431

432

Psyche

[September-December

I hope that these studies of Aliatypus will stimulate others to
investigate these fascinating spiders. Considerably more collecting
is necessary before Aliatypus systematics can be confidently under-
stood. As pointed out in the discussions of variation, small sample
sizes and sizeable geographic gaps from which no samples are avail-
able have greatly limited the strength of some of my conclusions.
The variation discussions, locality records, and distribution Maps 2-4
(Map 4 marks the distribution of unidentifiable Aliatypus speci-
mens.) should help direct future collecting efforts.

Acknowledgements

Willis Gertsch first recognized the potential diversity within
Aliatypus and suggested that I revise the genus. Wendell Icenogle
deserves tremendous credit for the hours he labored collecting about
55 percent of the specimens upon which this study is based. He was
the first collector of five new species. Without his field work this
study would be very incomplete. Jim Horton, my department head
during most of this research, provided needed space and release time.
I thank the following individuals and institutions for loans of
Aliatypus specimens: William Azevedo, Michael Bentzien, Patrick
Craig, Willis Gertsch (American Museum of Natural History),
Wendell Icenogle, Herbert Levi (Museum of Comparative Zoology),
Patrick Marer, Robert Schick (California Academy of Sciences),
and Mel Thompson. This research and its publication have been
supported by a grant (GB-34128) from the National Science
Foundation.

Evolution

PHYLOGENY

Aliatypus Antrodiaetus, Atypoides , the Mecicobothriidae, and
the Atypidae form a distinct monophyletic taxon (Coyle, 1971).
Aliatypus is probably an old group, so that the details of its relation-
ship to these other atypoid mygalomorph taxa are not now clear.
The question of whether Aliatypus is more closely related to Antro-
diaetus and Atypoides or to the mecicobothriids was discussed earlier
(Coyle, 1971), but will remain unresolved until after the completion
of a careful comparative study of all atypoid mygalomorphs.

In Figure 1 I have presented a hypothetical phylogeny of the
genus Aliatypus. This speculation is based upon a comparison of
character states in living Aliatypus species and related genera. It is

1974]

Coyle — Genus Aliatypus

433

Figure 1: Suggested phylogeny of Aliatypus species. 1. Character

states of hypothetical ancestral stock: No ICS keel or OCS keel. Seminal
receptacles with moderately long, sinuous, non-tapering stalks and medium
sized bulbs. Posterior sigilla small and well separated. Legs of moderate
length. Thoracic groove a deep pit. Leg I setation as in majority of
species. Moderately lar,ge body. 2. Seminal receptacle stalks become
short and straight. Legs become proportionately shorter. 3. Posterior
sigilla enlarge. Thoracic groove lost. Leg I setation changes. 4. Pos-
terior sigilla enlarge. Legs become proportionately shorter. Become adapted
to dryer habitats. 5. Become adapted to more humid and cooler habitats.
6. ICS keel develops. 7. Seminal receptacle stalks become tapered. 8. Con-
ductor tip changes form. Body size reduced. 9. Seminal receptacle stalks
become less elongate and less sinuous. Body size reduced. 10. Body size
reduced.

meant to be a useful working hypothesis, subject to revision. Char-
acters which were relied upon most heavily are palpus form, seminal
receptacle form, and posterior sigilla size and placement. The actual
direction of evolution in some characters may well be the reverse of
those suggested. It is certain that Aliatypus contains two distinct
groups of closely related species — A. calif ornicus, A. janus , A. iso-
latus, A. aquilonius , and A. gnomus on the one hand and A. tro-
phonius, A. erebus , A. plutonis, and A. torridus on the other — and
two distinct species, A. gulosus and A. thompsoni , each rather dis-
tantly related to all the others.

434

Psyche

[September-December

GEOGRAPHIC VARIATION AND SPECIATION

As is the case in Antrodiaetus (Coyle, 1971), Atypoides (Coyle,
1968 ), and probably most other burrowing mygalomorph genera
(See, for example, Main, 1957 ; Loksa, 1966; and Forster and
Wilton, 1968.), there is considerable geographic variation in some
species of Aliatypus. Detailed descriptions of these geographic vari-
ation patterns can be found in the Taxonomy section. My purpose
here is to consider some of the causes of these patterns.

The environmental tolerance ranges and dispersal ability of a
species are key factors in determining what environmental conditions
constitute barriers which can fragment and isolate its populations so
that genetic divergence can take place. Little pertaining specifically
to Aliatypus dispersal can be added to my earler discussion of dis-
persal ability in antrodiaetids (Coyle, 1971). In summary, the prob-
ability of successful colonization of distant localities by long distance
aerial dispersal is extremely low ; aquatic rafting, short distance
spiderling dispersal, and male wandering are probably the only im-
portant means of dispersal under natural conditions. As in Antro-
diaetus and Atypoides, environments with very low humidity, such
as deserts or semiarid grasslands, are the outstanding barriers to
dispersal and thus gene flow in Aliatypus. However, some species of
Aliatypus, notably A. plutonis and A. torridus, are less well restricted
by dry barriers than are most other antrodiaetids.

The following discussions, although partly speculation, should, like
any working hypotheses, help direct further research. Frequent refer-
ral to Map 1 will help to understand them.

The Central Valley of California, a semiarid grassland in its
recent natural state, appears to be a strong barrier to gene flow
between coastal and Sierran populations of both Aliatypus calif ornicus
and Aliatypus erebus. The genetic discontinuity between these popu-
lations may even be great enough to merit calling them incipient
species. A similar situation exists in Atypoides riversi (Coyle, 1968
and 1971). Apparently, during Pleistocene glacial periods when the
climate was wetter and cooler, dispersal of these species occurred
across favorable wooded parts of the Central Valley. The recent
discovery of isolated A. calif ornicus and A. riversi populations in the
Sutter Buttes of the Central Valley indicates that this was once part
of such a corridor allowing gene flow across the valley. Similar dis-
persals across the Central Valley during Pleistocene glacial periods
are also indicated by distribution patterns of the salamander genera
Ensatina and Taricha, which, like Aliatypus, require rather mesic
habitats (Stebbins, 1949; Riemer, 1958). I suspect that the antro-

1974]

Coyle — Genus Aliatypus

435

Map 1. Known distribution ranges of Aliatypus species in relation to
present major arid habitat barriers which formed during the retreat of
the Wisconsin glaciation.

diaetid trans-valley connections existed during the most recent (Wis-
consin) glacial period and consequently became severed as recently
as 13,000 years ago. Perhaps the A. calif ornicus population at Mari-
posa, which is phenotypically more distinct from the coastal popula-
tion than is the north Sierran population, was last connected with
the coastal population during an earlier glacial period ; or perhaps its
trans-valley connections were simply severed earlier during the retreat
of the last (Wisconsin) glacial period than were those of the north
Sierran populations. Perhaps continued expansion of habitat barriers
during the present post-glacial period has tended to restrict gene flow
between the morphologically divergent northern populations of A.
erebus and its south Sierran populations.

There is much geographic variation in Aliatypus janus, but the
samples are so small and scattered that it is difficult to recognize
important barriers to gene flow. The northernmost samples probably

436

Psyche

[September-December

represent the kind of semi-isolated, genetically divergent, peripheral
populations found in many species. Aliatypus thornpsoni variant
populations are also at the periphery of the species range, where
suitable habitats are probably uncommon and semi-isolated. The
Tehachapi Mountains apparently provide (or recently provided) an
east- west corridor of favorable habitats for the dispersal of A. janus
and A. thornpsoni between the southern end of the Sierra Nevada
Mountains and the coastal mountain ranges. The morphologically
divergent nature of the A. thornpsoni samples at Tehachapi and in
the southern end of the Sierra Nevada Mountains indicates that this
corridor is not currently supporting much dispersal.

The phenotypic differences between the two known Aliatypus
isolatus populations in Arizona are almost certainly caused by the
disruption of gene flow after the recent (Wisconsin) glacial period
as desert and grassland barriers expanded all over the Southwest to
isolate various mesic mountain habitats. Pollen analyses (Martin
and Mehringer, 1965) demonstrate that during the Wisconsin glacial
period (which apparently lasted in that area until about 13,000 years
ago), woodland and forest habitats favorable for A. isolatus extended
continuously throughout western and northern Arizona. Thus the
similar geographic variation patterns in A. isolatus and Antrodiaetus
apachecus (Coyle, 1971) probably have a common cause.

The extreme similarity of allopatric Aliatypus janus and Aliatypus
isolatus , when viewed with Southwest pollen analyses (Martin and
Mehringer, 1965) in mind, leads to the conclusion that these two
species were formed when a recent interglacial expansion of the
Sonoran and Great Basin Deserts severed a previously widespread
ancestral population. Convincing evidence that 17,000 to 23,000
years ago (during the Wisconsin glacial period) woodland extended
continuously from current A . isolatus localities to the present range
of A. janus , strongly indicates that these sister species may be only
15,000 years or so old. Indeed, it is possible that genetic divergence
has not even progressed far enough for the development of repro-
ductive isolating mechanisms.

There are three other pairs of closely related Aliatypus species —
A. janus and A. aquilonius , A. calif ornicus and A . gnornuSj and A.
erebus and A. trophonius — which are not as similar as A. janus
is to A. isolatus. It is possible that each of these pairs originated
from a trio of ancestral species fragmented by arid barriers, such as
the present Central Valley, during an earlier Pleistocene interglacial.
Interestingly, each of these pairs consists of a large and a small species.

1974]

Coyle — Genus A liatypus

437

Methods

COLLECTING METHODS

Because of their covert behavior, A liatypus spiders are rarely col-
lected except when one concentrates on special collecting strategies.
Banks and slopes of promising habitats (to be described thoroughly
in a paper on A liatypus natural history) are best searched in daylight
by carefully examining suitable microhabitat surfaces for the out-
line of a closed trapdoor. However, whenever the trapdoors are
sealed, such as during dry periods, they may become covered with
loose soil particles or other debris; carefully shaving away the top
layer of soil may then be the only way to locate burrows. Night
collecting is usually less satisfactory than daytime collecting since
even unsealed Aliatypus doors are only cracked open at night and
not easily located with artificial light. At night it is sometimes pos-
sible to trap active spiders at their burrow entrances by thrusting a
knife blade into the soil and across the burrow lumen just below the
spider, but frequently the soil is too hard. More information can
be gained by careful excavation of the burrow in daylight. An army
trench shovel, a small, chisel-head, rock hammer, a large pocket knife,
and pruning shears are all useful for excavating in the often hard
and root-bound soil. Penultimate males, easily recognized by their
swollen pedipalpal tarsi, will often molt to adulthood if kept in a
cool, humid, and dark environment. Just before and during the
mating season (usually during the wet fall and winter months)
recently matured males may be found in their burrows prior to
abandoning them. Wandering males are best collected at night by
hand or with pitfall traps in dense burrow aggregations during opti-
mum mating weather.

ANALYSIS OF VARIATION

I have examined the taxonomy of Aliatypus by means of an analy-
sis of variation nearly identical to the analysis employed in my re-
visions of Antrodiaetus (Coyle, 1971) and Atypoides (Coyle, 1968).
Such an analysis largely overcomes difficulties posed by the relatively
simple reproductive anatomy, by the instar heterogeneity of adult
female samples, and by heightened geographic variation, difficulties
which appear to be common to most mygalomorph spider taxa. The
material analyzed consists of 252 adult females and 78 adult males.
The sample size for each species is indicated in Tables 1 and 2.

Initially, variation in a large number of qualitative and quantita-
tive characters was briefly surveyed, and from these characters the
diagnostically most promising were selected and their variation studied

438

Psyche

[September-December

in depth. Variation of the quantitative characters (measurements,
meristic characters, and ratios formed from these) was analyzed with
the aid of an IBM 360 Model 30 computer. A Fortran IV program
directed the computer to calculate the mean and standard deviation
of each character for each local population sample of each sex and
for certain groupings of local samples into larger infraspecific samples
or species samples. The computer then compared these samples pair-
wise in any desired combination, giving for each character for each
comparison a value of the distinctness of the two samples. This
“distance” value equals the difference between the means of the two
samples divided by the sum of their standard deviations.

This variation analysis was performed with the following number
of characters: 23 measurements, one meristic character, and 39 ratios
for males; 20 measurements, six meristic characters, and 50 ratios
for females. The measurements and meristic characters were defined
so as to be clearly delimited. Their abbreviations and definitions are
as follows (see Figs. 2-7) :

Figures 2-7: Measurements used in Aliatypus revision. See text for

definitions. Figures 8-9: Macrosetae types. 8: ensiform. 9: attenuate.

CL Maximum length of carapace measured as distance (along

median longitudinal axis) between lines tangent to anterior-
most and posteriormost edges of carapace, with lateral
border of carapace in horizontal plane.

PCL Length of pars cephalica measured as distance from anterior
edge of thoracic groove along median longitudinal line.

CW Maximum width of carapace along line perpendicular to
median longitudinal axis.

1974]

IFL

ITL

IML

ITarL

IVFL,

PFL

PPL

PTL

PTX

PTT

PED

PCA

SL

SW

PSS

PSL

OQW

ALS

ALD

AMS

Coyle — Genus A liaty pus 439

Length of femur I taken as length of straight line connect-
ing the proximal and distal points of articulation. All leg
and pedipalp segment length measurements were made in
side view along retrolateral surface of appendages after
removing them from spider.

Length of tibia I taken as length of straight line connect-
ing proximal and distal points of articulation.

Length of metatarsus I taken as length of straight line con-
necting proximal point of articulation with distalmost point
of segment.

Length of tarsus I taken as length of straight line connect-
ing most proximal exposed point of tarsus with distalmost
point of dorsal surface.

IVTL, IVML, IVTarL Leg IV segment lengths meas-
sured in same manner as corresponding leg I segments.
Length of pedipalpal femur measured same as IFL.

Length of pedipalpal patella measured as straight line dis-
tance from proximal to distal end along dorsal surface.
Length of pedipalpal tibia measured same as ITL.

Distance from proximal point of articulation on tibia to
point where PTT line intersects PTL line.

Maximum diameter, taken perpendicular to line defining
PTL, of pedipalpal tibia in lateral view.

Straight line distance from base of embolus to tip of con-
ductor.

Maximum distance from PED line to outer edge of OCS
along line perpendicular to PED line.

Maximum length of sternum on line parallel to median
longitudinal axis. Anterior border of sternum is its pointed
anterior extension lateral to labium.

Maximum width of sternum perpendicular to line defining

SL.

Minimum distanoe between posterior sigilla.

Maximum diameter of right posterior sigillum.

Maximum width of eye group (ocular quadrangle) on line
perpendicular to median longitudinal axis of carapace. All
eye measurements are made in dorsal view with lateral
border of carapace horizontal.

Minimum distance between anterior lateral eyes.

Maximum diameter of left anterior lateral eye.

Minimum distance between pupils (light colored saucer-
shaped central area of eye) of anterior median eyes.

[September-December

440 Psyche

AMD Transverse diameter of left anterior median eye pupil.

EGS Number of epiandrous gland spigots. These are located
just anterior to genital opening on abdomen of adult males.
CTP Number of cheliceral teeth in prolateral macrotooth row on
left chelicera.

CTR Number of cheliceral teeth in retrolateral row of smaller
macroteeth on left chelicera.

CMT Number of cheliceral microteeth between these two rows
on left chelicera.

PTSP Number of ensiform macrosetae on prolateral surface of
tarsus of female pedipalp.

PTSR Number of ensiform macrosetae on retrolateral surface of
tarsus of female pedipalp.

IMS Number of ensiform macrosetae on metatarsus of leg I of
female.

All measurements and counts were performed by myself with the
same Wild M-5 stereomicroscope with 20 X eyepieces and an eye-
piece micrometer scale. The measurements are accurate to one mi-
crometer unit for each of the three different powers of magnification
used. One micrometer unit had the following values for the follow-
ing characters: 0.0770 mm for CL; 0.0385 mm for PCL, CW, SL,
SW and all leg and pedipalp segment lengths; and 0.0092 mm for
PTT, PED, PCA, PSS, PSL, and all eye measurements.

A female specimen was included in a population sample only if it
was reproductively active (with maturing eggs in abdomen or brood
in burrow) or had a longer carapace than the smallest reproductively
active female in that sample. Many first adult instar females, a few
older adult instar females, and rarely a large immature female make
up the portion of a sample which is not reproductively active.

MORPHOLOGICAL TERMINOLOGY

Setae. Postocular setae form a longitudinal row or longitudinal
cluster along the median longitudinal axis of the pars cephalica just
behind the eye group. A macroseta is a very large seta. Called spines
by many authors, these macrosetae are attached to the exoskeleton
proper by means of an obvious socket which allows for some move-
ment. An ensiform macroseta is one which tapers rather abruptly at
its terminal end and is therefore rigid for its entire length ( Fig. 8 ) .
An attenuate macroseta tapers gradually and is therefore very slender
distally (Fig. 9) .

1974]

Coyle — Genus A liatypus

441

Palpus. The conductor of the Aliatypus palpus (Fig. 97) consists
of an inner conductor sclerite (ICS) and an outer conductor sclerite
(OCS) which lies outside and partly cradles the ICS and the em-
bolus. The ICS base is forked, with the more well developed of the
two branches being called the proximal branch.

Female genitalia. In Aliatypus the bursa copulatrix , which opens
just anterior and ventral to the uterus opening in the epigastric fur-
row, is bilobed and weakly sclerotized. The four seminal receptacles
(Fig. 163) are functionally paired so that the two on the right side
open close to one another into the right lobe of the bursa copulatrix
and the other two open together into the left lobe. Each seminal
receptacle is weakly sclerotized and consists of a narrow stalk and a
distal expanded bulb. The seminal receptacle is either homogeneously
sclerotized or the bulb is slightly less sclerotized than the stalk.
Aliatypus stalks are usually sinuous, frequently even highly looped or
coiled. These loops and coils are not confined to a single plane and
are often irregular so that the degree of looping or coiling is very
difficult to quantify. It is, however, possible to make a rough quanti-
tative comparison by counting the number of bends per stalk, as is
done in Figure 163, when the stalk is treated as a two-dimensional
structure.

Abdominal tergites. The anterior portion of the abdominal dorsum
of Aliatypus is provided with one or more segmentally arranged,
rather heavily sclerotized patches which are presumably vestigial ter-
gites (Figs.45-49) . These tergites are numbered from anterior to
posterior, tergite I , ter git e II , and ter git e III. Tergite II is always
present in both sexes and is always larger than tergites I or III.

METHODS OF PRESENTATION

Type specimens. The holotypes of all species described in this
paper are deposited in the Museum of Comparative Zoology. All
paratypes are from the type locality and are labeled as paratypes.
The paratypes for each species are deposited in about equal numbers
in the Museum of Comparative Zoology and the American Museum
of Natural History. Quantitative character values are given for each
holotype in Table 3.

Key. Whenever quantitative characters are used in the key, the
known range of values is used. Proceed cautiously when these
ranges are based upon very small samples. Sample size for each spe-
cies is given in Tables 1 and 2.

Diagnoses. Each diagnosis lists, in the approximate order of their
usefulness, those characters most useful in identifying a given species.

442

Psyche

[September-December

Proceed cautiously when using diagnoses based upon very small
samples. In Tables i and 2 the diagnostically most useful character
values are circled for each species or group of species.

Descriptions. The quantitative character values recorded in Tables
i and 2 are an essential part of each species description. Each descrip-
tion is a composite of all adult specimens at hand. Only characters
of at least some diagnostic value are included. Colors are described
from fully sclerotized specimens dead for two months to six years
and immersed in alcohol under strong fluorescent light.

Illustrations. Illustrations were carefully drawn in pencil on
translucent paper over a squared grid template with the aid of a
squared grid reticle in the eyepiece of the Wild M-5 stereomicro-
scope. The penciled drawings were then traced in ink on heavier
paper. Nearly all figures of seminal receptacles are drawn from re-
productively active females.

Variation discussions. For each species, variation of all 63 male
quantitative characters, all 76 female quantitative characters, and a
number of qualitative characters was examined, and all characters
which show marked variation are discussed. The sizes of all popula-
tion samples discussed can be found in the modified Dice-Leraas dia-
grams or in the records section.

Records. Only specimens which I have examined are listed.
Within each county citation, all records from a given locality are
separated from those of other localities by a dash. Collection dates
are listed only for males. When a male symbol is surrounded by
parentheses, it means that the specimen was collected when immature
on the date given and matured later in captivity. When no male or
female symbol follows a record, this means that only immatures were
collected.

Taxonomy

AL1ATYPUS Smith, 1908

Aliatypus Smith, 1908, Ann. Ent. Soc. Amer., 1(4): 231. Type species by
monotypy Atypoidcs californica Banks, 1896, Jour. New York Ent.
Soc., 4(4): 88 — Bonnet, 1955, Bibliographia Araneorum, 2:225. —
Coyle, 1971, Bull. Mus. Comp. Zool., 141 (6): 372.

Descriptive diagnosis. Carapace: Figs. 45-53. Thoracic groove a
deep pit which varies greatly in shape (from transverse to slightly
longitudinal; borders rounded or angular); may be absent or re-
duced to a shallow depression. A large seta on ocular prominence

1974]

Coyle — Genus Aliatypus

443

between anterior median eyes. Chelicerae: Figs. 39-40, 45-49. Fe-
male with row of 6 to 13 macroteeth on prolateral edge; another
row of 2 to 7 smaller macroteeth on retrolateral side of closed fang;
and 4 to 68 microteeth between these rows. Male also with retro-
lateral row of cheliceral teeth. Anterior dorsal outline of chelicerae
evenly rounded in both sexes. Female with rastellum. Pedipalps:
Figs. 78-91, 96-120. Femur, patella, and tibia of male very elongate.
Male tibia swollen at least ventrally near distal end. Embolus very
long and slender. Outer (convex) edge of OCS folded over to
cradle embolus. ICS extends to but not beyond OCS tip, and is
intimately combined distally with OCS. Ridge runs most of length
of ICS. Legs: Figs. 92-95. One to 4 (rarely more than 1) tricho-
bothria dorsally near distal end of metatarsus IV. Male tibia and
metatarsus I each with macrosetae distributed in scattered but con-
sistent pattern ventrally over most of length. Spinnerets : Figs. 43-44.
Three pairs (AL, PM, PL); all functional (with spigots). AL
spinnerets 2-segmented with at least several spigots clustered at tip
of distal segment. PM spinnerets unsegmented. PL spinnerets 3-
segmented ; distal segment shorter than other two together. Geni-
talia: Figs. 12 1- 1 94. Bursa copulatrix bilobed and usually very
weakly sclerotized. Seminal receptacles paired, each pair opening
into one lobe of bursa copulatrix. Stalks weakly sclerotized and
usually relatively elongate and sinuous. Bulbs as sclerotized or
slightly less sclerotized than stalks. Behavior: Burrow entrance a
trapdoor. Egg sac pendulous and occludes burrow lumen.

The following features of Antrodiaetus and Atypoides readily dis-
tinguish them from Aliatypus: Thoracic groove very narrow and
longitudinal. No row of large cheliceral teeth on retrolateral side of
closed fang. Male pedipalp, especially patella, relatively short. ICS
tip distinctly separated from OCS tip. Metatarsus IV with 5 to 21
trichobothria dorsally near distal end. AL spinnerets absent or un-
segmented with at most one spinneret apically. Burrow entrance a
collar or turret. Egg sac attached to one side of wall and does not
occlude burrow lumen.

The following features of the Mecicobothriidae readily distinguish
it from Aliatypus: Thoracic groove very narrow and longitudinal.
No single large seta between anterior median eyes. No row of large
cheliceral teeth on retrolateral side of closed fang. No rastellum.
Male pedipalp, especially patella, relatively short. Metatarsus IV
with more than 4 trichobothria dorsally near distal end. PL spin-
nerets very elongate; distal segment as long or longer than two basal
segments combined. Sheet web retreat without trapdoor.

444

Psyche

[September-December

Discussion. As I have said earlier in this paper and before (Coyle,
I97I), Aliatypus may be as closely related to the Mecicobothriidae
as to the other antrodiaetid genera, Antrodiaetus and Atypoides.
However, it is best to retain Aliatypus within the family Antrodiaeti-
dae until a careful comparative study of all atypoid mygalomorph
taxa demonstrates otherwise.

As discussed earlier, A. calif or nicus , A. ]anus, A. isolatus, A. aqui-
lonius, and A. gnomus form a group of closely related species, A.
trophonius , A. erehus, A. plutonis, and A. torridus form another
group of closely related species, and A. gulosus is quite distinct from
all other Aliatypus species. I will not formally designate species
groups, however, because the placement of A. thompsoni, which ap-
pears to be intermediate in its relationship to the two groups of spe-
cies, would be rather arbitrary.

Key to Species of Aliatypus
Males

1. Palpus unique (Fig. no); sperm reservoir looped very loosely,

and embolus base close to ICS base. Pedipapal tibia (Fig. 85)

banana shaped and elongate; PTX/PTL = 0.37-0.45

gulosus

Palpus otherwise (Figs. 96-109, m-120); sperm reservoir
much more tightly coiled, and embolus base distant from
ICS base. Pedipalpal tibia (Figs. 78-84, 86-91) not banana
shaped; PTX/PTL = 0.64-0.82. 2

2. PSL/PSS = 1.28-2.38. CL/IML = 1.02-1.24. Tibia and

metatarsus I (Fig. 93) with short ensiform macrosetae and
strongly appressed background setae. Thoracic groove nearly

always absent or shallow. thompsoni

PSL/PSS = 0.14-1.17. CL/IML = 1.37-2.07. Tibia and
metatarsus I (Figs. 92, 94-95) with more elongate macrosetae
and suberect or erect background setae. Thoracic groove a
deep pit. 3

3. Weakly sclerotized, finger-like extension at tip of conductor

(Fig. 106). PTL/PPL = 1.03-1.06 and CL/PSS = 4.80-

5.3 1 . aquilonius

No such extension at tip of conductor (Figs. 96-105, 1 07-1 20).
PTL/PPL = 1.10-1.46 or CL/PSS = 5.50-10.34. 4

4. ICS forms at least a weak keel distally (Figs. 96-101) 5

Conductor either without a keel (Figs. 102-113, 119-120), or,

if keel present, then it is an extension of the OCS and is
closer to the conductor tip (Figs. 114-118). 6

1974]

Coyle — Genus Aliatypus

445

5. PFL = 3-07-3.54 mm. CL/PPL = 1.66-1.67. ICS keel weak

and conductor tip quite slender (Fig. 101). gnomus

PFL = 4.08-6.73 mm. CL/PPL = 1.33-1.61. ICS keel
stronger and conductor tip not as slender (Figs. 96-100). ....
calif ornicus

6. Distal one-fourth of conductor slender and tapers evenly (Figs.

102-105, 107-109), and PTX/PTL = 0.73-0.82. 7

Either the distal one-fourth of conductor at least moderately
broad and keel-like and tapers suddenly to point at end (Figs.
114-118), or PTX/PTL = 0.64-0.68 8

7. CL/ALS = 7.70-8.08. Known only from Arizona. .... isolatus
CL/ALS — 8.70-1 1. 1 1. Known only from California. . janus

8. PTX/PTL = 0.64-0.68. PTL/PPL = 1.31-1.32. Distal half

of pedipalpal tibia venter not much more swollen than proxi-
mal half (Fig. 91). Palpus as in Figs. 119 and 120. torridus
PTX/PTL = 0.72-0.78. PTL/PPL = 1. 02-1. 22. Distal
half of pedipalpal tibia venter much more swollen than proxi-
mal half (Figs. 87-90). Palpus as in Figs. 114-118. 9

9. CL/IFL = 1.03-1.08. plutonis

CL/IFL = 1.25-1.33 10

10. CL/PCA = 6.64-6.96. CL/AMD = 21.7-22.5. CL = 2.9-
3.7 mm. Palpus as in Fig. 116. trophonius

CL/PCA = 9.17-9.56. CL/AMD = 25.2-34.8. CL = 4.4-
6.6 mm. Palpus as in Figs. 114-115 erebus

Females

1. Seminal receptacle stalks straight and short (Figs. 151-154).

gulosus

Seminal receptacle stalks sinuous (Figs. 1 21-150, 1 55-194). .. 2

2. Thoracic groove absent or only a shallow depression (Figs. 52-

53). IMS/PSS = 33.3-116.6. thompsoni

Thoracic groove a deep pit (Figs. 47, 49). IMS/PSS = 6.1-
34-7 3

3. CL/IVTL p 2.29-2.86. SW/PSL = 6.19-20.00. PSL/PSS

— 0.13-0.67. 4

CL/IVTL = 2.88-3.52. SW/PSL = 3.81-6.56. PSL/PSS
= 0.55-2.00 8

4. AMD/AMS = 1.36-1.71. Seminal receptacle stalks weakly

sinuous (Figs. 149- 150). gnomus

AMD/AMS — 0.39-1.06. Seminal receptacle stalks strongly
sinuous (Figs. 121-148). 5

4+6

Psyche

[September-December

5. Known only from Arizona. isolatin'

Known only from California. 6

6. IFL/IVFL — 0.96-1.02. ITarL — 0.46-0.80. aquilonius

IFL/IVFL = 1. 02- 1. 1 3. ITarL = 0.92-1.38. 7

7. Seminal receptacle stalks about equal diameter throughout

length (Figs. 121-131) californicus

Seminal receptacle stalks usually much thicker basally than at
distal end (Figs. 132- 143). janus

8. IVFL/IVML =1.02-1.08 plutonis

IVFL/IVML = 1. 1 1-1.23 9

9. IFL = 1.84-2.84. CL/PTSR := 0.98-1.31. Seminal recep-

tacle stalks proportionately long and with 3-5 bends; bulbs

small to medium sized (Figs. 172- 173). trophonius

IFL = 3.27-5.38. CL/PTSR — 1.39-3.03. Seminal recep-
tacle stalks proportionately shorter, with 1-3 bends; bulbs
medium sized to very large (Figs. 174-187, 192-194). .... 10

IO. CMT = 4-1 1. CL/OQW = 3.43-3.99 torridus

CMT = 11-68. CL/OQW = 4.05-4.93 erebus

Aliatypus californicus (Banks)

Figures 10-17, 39, 45-4$, 54, 65, 78-79, 92, 96-100, 121-131. Map 2.

Atypoides calif ornica Banks, 1896, Jour. New York Ent. Soc., 4(4) : 88.
Holotype a penultimate male from Black Mountain, California, 23
October, in Museum of Comparative Zoology; examined.

Aliatypus californicus : Smith, 1908, Ann. Ent. Soc. Amer., 1 (4): 232. —

Comstock, 1912, The Spider Book, p. 251. — Gertsch, 1949, American
Spiders, p. 132. — Bonnet, 1955, Bibliographia Araneorum, 2: 225. —
Coyle, 1971, Bull. Mus. Comp. Zool., 141 (6): 372. — Kaston, 1972,
How to Know the Spiders, p. 60.

Comments on the type locality and previous descriptions. Banks
(1896) wrote only “Black Mtn., Calif.” for the type locality. Other
species described in the same paper were collected by the same col-
lector (R. W. Doane) from the Palo Alto area. This implies that
the type locality of A. californicus is Black Mountain in northwest-
ern Santa Clara County. Smith (1908), who knew Doane well,
states confidently that the type was collected on Monte Bello Ridge
of this mountain. The following species description is based partly
upon a male and a female collected on Montebello Road, presumably
very close to the type locality.

Banks’ (1896) description is based upon an immature specimen,
is brief, and does not include characteristics which distinguish A.
californicus from some other Aliatypus species. Smith’s (1908) de-

1974]

Coyle — Genus Aliatypus

447

scription is more thorough, includes illustrations, and describes a
crucial diagnostic feature, the ICS keel. This shows up in Fig. 4a
of Smith’s Plate XV. Although Smith correctly states that the
pedipalpal tibia is longer than the patella, he erred in drawing it
shorter (Smith’s Fig. 3, Plate XV). The specimens upon which
Smith based his description were placed in the Stanford University
collection but have since been lost. Having closely examined the type
specimen and Smith’s description, I am confident that the material
upon which the following description is based is conspecific with that
described by Banks and Smith.

Diagnosis. Males: The presence of a keel on the ICS together
with the shape of the conductor tip (Figs. 96-100) distinguish this
species from all others. Closely related A. janus differs from A.
calif ornicus in the ratio IML/ITarL (Table 1). A. isolatus, also
closely related, is distinguished by the ratios IML/ITarL, PPL/
PFL, and PTL/PPL (Table 1). Any of the other species can be
separated from A. calif ornicus with the appropriate ratio selected
from the following: CL/IML, CL/PSL, and PTL/PPL (Table
1). Females: A. calif ornicus females are difficult to distinguish from
those of A. janus and A. isolatus. A. calif ornicus seminal receptacle
stalks are of about equal diameter throughout (Figs. 121-131), un-
like those of A. janus (Figs. 132- 143) and A. isolatus (Figs. 144-
146) which are much narrower distally than at the basal end. In
A. isolatus the postocular row of cephalic setae extends to a point
one-half or more of the distance from the anterior edge of the cara-
pace to the thoracic groove; in A. calif ornicus it extends to a point
less than one-half the distance to the thoracic groove. A. aquilonius
and A. gnomus, both closely related to A. calif ornicus, can be dis-
tinguished from the latter by their small body size (Table 2; espe-
cially ITarL). A. aquilonius has distinctively smaller CL/PTSR
and IFL/IVFL values (Table 2) and seminal receptacles with more
swollen stalk bases (Figs. 147-148) than A. calif ornicus (Figs. 12 1-
1 3 1 ) . A. gnomus has a distinctively larger AMD/AMS value
(Table 2) and shorter, less sinuous seminal receptacles (Figs. 149-
150) than A. calif ornicus (Figs. 121-131). All other species can be
separated from A. calif ornicus by either seminal receptacle form or
appropriate ratios chosen from the following (Table 2): CL/IFL,
CL/ITL, CL/IML, SW/PSL, and PSL/PSS.

Description. See Tables 1-3.

Male: Carapace. Figs. 45-46. Thoracic groove a deep pit; usually
rounded anteriorly, elongate, and tapering posteriorly ; sometimes
nearly circular or triangular. Postocular setae form a narrow longi-

448

Psyche

[September-December

tudinal row. Sternum. Fig. 54. Posterior sigilla small to medium
sized and well separated. Pedipalps. Figs. 78-79, 96-100. Tibia
strongly swollen ventrally near distal end. Embolus base well sep-
arated from ICS base. ICS ridge distally develops into thin keel
which then disappears so that conductor tip is pointed. Inner (con-
cave) edge of OCS nearly smooth to rough. Leg I. Fig. 92. Tibia
and metatarsus with ventral, suberect, mostly attenuate macrosetae.
Rest of metatarsus setae mostly long, slender, and suberect. Abdo-
men. Figs. 45-46. Tergites I and III reduced to small patches or
spots at bases of macrosetae. Coloration. Pars thoracica light yellow
to pale yellow-brown. Pars cephalica darker ; light brown to medium
brown; darkest along margin and median longitudinal line. Cheli-
cerae like pars cephalica. Pedipalpal femur and patella dorsally a
darker orange-brown or red-brown.

Females: Carapace. Figs. 47-48. Thoracic groove a relatively
small deep pit of varying shape; usually rounded anteriorly and
tapered posteriorly; sometimes circular, elongate-oval, transverse-
oval, or triangular. Postocular setae form narrow row which ex-
tends back to a point Y to almost V2 of distance from anterior edge
of carapace to thoracic groove. Sternum. Fig. 65. Posterior sigilla
small to medium sized and well separated. All or nearly all per-
ipheral sternal setae slender; a few may be moderately stout. Long-
est setae scattered all over sternum, but more abundant anteriorly.
Chelicerae. Figs. 39, 47-48. Genitalia. Figs. 121-131. Seminal re-
ceptacle stalks weakly sclerotized, nearly constant diameter through-
out length, long, and with 3 to 9 bends (usually 6 to 9 bends).
Bulbs relatively small to medium sized, slightly less sclerotized than
stalks. Coloration. Pars thoracica light yellow to light grey-yellow.
Pars cephalica darker; often especially dark around margin and
median longitudinal line; darker parts medium brown to darker red-
brown ; lighter parts light yellow-brown to orange-brown. Chelicerae
match darker parts of pars cephalica.

Variation. Males: The coastal population samples average con-
siderably larger in body part dimensions than the Sierran population
samples, with the sharpest discontinuity being in PED, PTT, and
ITarL (Fig. 10). Somewhat discontinuous geographic variation is
found in the ratios CL/PPL (Fig. 11), CL/PTX (Fig. 12), and
PPL/PFL. This and weaker variation in other characters show a
recurrent pattern: some phenotypic discontinuity between the rather
homogeneous coastal populations on the one hand and the Sierran
populations on the other; considerable discontinuity between the two
Sierran populations, with the coastal populations more similar to the

1974]

Coyle — Genus Aliatypus

449

Mariposa
Coloma ♦
Aukum
coastal

1—3

1 ^

Mariposa 1 — 3

Coloma* | 4

Aukum i 1

coastal i_. — j|_ j 7

10

1.2 1.3 1.4 1.5 1.6 1.7

ITarL

1.9

1.5 1.6 1.7 1.8 1.9 2.0

12 CL/PTX

Mariposa
Coloma *
Aukum
coastal

-f-4

1 3

11

1.3

1.4 1.5 1.6

CL/PPL

A. janus
Mariposa
Coloma *
Aukum
coastal

— R

25

— 1—3

13

2.120

2.30

2.40 2.50

2.60

CL/ITL

A. janus

25

Mariposa

Coloma*

Aukum

coastal

A. janus
Mariposa
Coloma ♦
Aukum
coastal

25

4-3

- "|" 3 35

14

1.30 1.40 1.50

CL/IFL

1.00 1.05 1.10 1.15

15 IFL/EFL

Figures 10-17: Geographic variation of Aliatypus calif ornicus. Map of

sample localities and modified Dice-Leraas diagrams. (Horizontal line
represents the observed range, vertical line the mean, open rectangle the
standard deviation, and number to right of range line the sample size.)
10-12: males. 10: ITarL (in mm) variation. 11: CL/PPL variation.
12: CL/PTX variation. 13-17: females. 13-15: A comparison of coastaL
A. calif ornicus, Sierran A. calif ornicus, and A. janus. 13: CL/ITL varia-
tion. 14: CL/IFL variation. 15: IFL/IVFL variation. 16: CL/IVTL

variation. 17: IVFL/IVTarL variation.

450

Psyche

[September-December

northern Sierran (Coloma-Aukum) population than to the Mariposa
population.

Noteworthy geographic variation also occurs in thoracic groove
shape, pedipalpal femur shape, and conductor form. Almost all
coastal specimens have an elongate thoracic pit which narrows pos-
teriorly (Fig. 45). The Coloma-Aukum specimens have roughly
circular or slightly transverse pits. The Mariposa specimens have
slightly elongate pits which narrow posteriorly and have a roughly
triangular shape. Coastal specimens have rather strongly bowed
pedipalpal femurs (Fig. 79), Mariposa pedipalpal femurs are less
strongly bowed, and Coloma-Aukum ones are nearly straight (Fig.
78). Most of the Sierran specimens have less well developed ICS
keels and relatively wider conductor tips (Figs. 98-100) than do
coastal specimens (Figs. 96-97). However, there is considerable
variation among Sierran populations, with Coloma males having a
rather well developed keel and narrow conductor tip (Fig. 98), and
Mariposa males having smaller keels and proportionately wider con-
ductor tips (Figs. 99-100).

Females: There is a moderate amount of geographic variation
among the coastal populations. Although some individual population
samples (especially the Mt. Diablo and Soquel area samples) are
quite different from another sample in a few ratios (Figs. 16-17),
no population sample is distinct from the rest of the entire coastal
sample in any character. The three northern Sierran (Coloma-
Aukum) specimens are quite similar to the coastal populations in all
characters with the exception of seminal receptacle form. Variation
in the number of stalk bends in the northern Sierran receptacles
(Figs. 1 27- 1 29) spans the gap between the more sinuous coastal
stalks (Figs. 1 21-125) and the less sinuous Mariposa stalks (Figs.
130-131). Northern Sierran females tend to have proportionately
large receptacle bulbs (Figs. 128-129). The Mariposa sample
differs rather strongly from the coastal samples in four characters

(Figs. 13-15) : IFL/IVFL, IVFL/IVTL, CL/IFL, and CL/ITL.
It likewise differs markedly from the northern Sierran sample in
these plus a fifth, CL/IVTL (Fig. 16).

In summary, male and female variation patterns indicate that there
is little, if any, gene flow between coastal and Sierran populations
across the unfavorable Central Valley, that there is little gene flow
between the northern and more southern Sierran populations, and
that the coastal population is genetically more similar to the northern
Sierran populations than to the Mariposa population. I do not feel
that the variation discontinuities between coastal populations and the

1974J

Coyle — Genus Aliatypus

451

Coloma-Aukum population are great enough to indicate that repro-
ductive isolation would develop if the populations were brought to-
gether. The recent discovery in the Sutter Buttes area of the Central
Valley of a moderately large immature female A. calif ornicus (Fig.
126) supports this conclusion by suggesting that gene flow could
have occurred rather recently across the Central Valley.

The female sample from Mariposa is quite divergent from all other
A. calif ornicus samples. However, without an analysis of more and
larger Sierran samples, I am reluctant to conclude that this Mari-
posa population is a different species. It is possible but unlikely that
this Mariposa female sample is conspecific with A. janus. The com-
puter analysis shows that this sample is more similar to A. janus
than to the rest of the entire A. calif ornicus sample. If this popula-
tion is conspecific with A. janus, its variation patterns shown in
Figures 13-15 could be the result of character displacement with the
sympatric A. calif ornicus population. However, the collection of
three A. calif ornicus males in the same location with the Mariposa
females argues strongly against this possibility.

Distribution. Mountains and foothills of the San Francisco Bay
region, western foothills of the Sierra Nevada Mountains, and at
least one area in the Central Valley (Map 2).

Records. California, coastal populations: Contra Costa Co.:
0.5 mi. E of South Gate of Mt. Diablo St. Pk., 1300 ft., 3$. —
Orinda Village. San Mateo Co.: Butano State Park, 9 . — Huddart
Park, 9 • Santa Clara Co.: Montebello Rd., 4 mi. W of junc. with
Stevens Canyon Rd., 2300 ft., 10 Oct. 1971, cT, 9* — Mt. Loma
Prieta, 9 mi. W of Morgan Hill, 1800 ft., 10 Oct. 1970, cf, 4$-

— Marsh Rd., 0.5 mi. S of Calaveras Reservoir, 900 ft., 7 Oct.
1970, 2 cf j 3 ? . — Alum Rock Park, 600 ft., 11 Oct. 1970, cf , 9 ;
23 Oct. 1970, cf, 39 ; 79- Santa Cruz Co.: 4.5 mi. N of Soquel
Center, 300 ft., 12 Oct. 1971, cf, 49 ; 29- — Bates Creek, 3 mi.
NE of Soquel, 200 ft., 3 9 • — Henry Cowell St. Pk., Redwood
Loop Nature Trail, 250 ft., 9 • — 1.7 mi. W of Felton on Felton
Empire Rd., 1000 ft., 9* — 4-3 mi • W of Felton on Ice Cream
Grade Rd., 1600 ft. sierran populations: El Dorado Co.: Omo
Ranch Rd., 1.5 mi. NE of Aukum, 2200 ft., 9 Nov. 1972, cf , 29 •

— 0.5 mi. SE of Coloma, 750 ft., 8 Nov. 1972, 3 cf » 9 • Mariposa
Co.: 0.5 mi. NW of Mariposa on rt. 49, 2000 ft., 14 Oct. 1969,
(cf ), 9 ; 8 Oct. 1971, 2 cf , 39 ; 29* central valley popula-
tion: Sutter Co.: Moore Canyon of Sutter Buttes, 4.5 mi. NW of
Sutter, 200 ft.

452

Psyche

[September-December

Aliatypus janus new species
Figures 18-24, 55, 66, 83-84, 102-105, 132-143. Map 2.

Type specimens and etymology. Holotype male from 5 mi. south
of Hume Lake, Fresno Co., California, 16 October 1973 (W. R.
Icenogle). One male and three female paratypes. Janus was the
Latin god of gates and doors.

Diagnosis. Males: A. janus is very similar to A. isolatus; CL/
ALS (Table 1) is useful in separating these two allopatric species.
A. janus is distinct from closely related A. calif ornicus in the absence
of an ICS keel (Figs. 102-105) and in IML/ITarL (Table 1).
A. janus is distinct from A. aquilonius in palpus tip form (Figs. 102-
105), SW/PSS, CL/PSS, and CL/ALS (Table 1). A. janus
differs from A. gnomus in CL/ALS, CL/AMD, and body size
(Table 1). Sympatric A. erebus differs from A. janus in CL/
PTT, PSL/PSS, CL/IFL (Table 1), and palpus form (Figs.
102-105). A thompsoni, also sympatric with A. janus, is distinct
in PSL/PSS, CL/IML (Table 1), thoracic groove form, and
leg I setation. Other species can be separated from A. janus with
an appropriate ratio from the following (Table 1): CL/ALS,
PTX/PTL, and PTT/PTL. Females: The only really helpful
character to use in separating A. janus from closely related A.
isolatus is SL/SW (Table 2). Only seminal receptacle form ap-
pears useful in separating A. janus from closely related A. cali-
f ornicus (See A. calif ornicus diagnosis). A. janus has a larger
IFL/IVFL, longer ITarL, and more IMS (Table 2) than does
A. aquilonius. A. janus is distinct from A. gnomus in AMD/
AMS, IFL/IVFL, body size (Table 2), and seminal receptacle
form (Figs. 132- 143). A. janus is distinct from sympatric A.
erebus in seminal receptacle form (Figs. 132-143), CL/PSL, SW/
PSL, and IFL/IVFL (Table 2). A. janus is distinct from sym-
patric A. thompsoni in thoracic groove form, seminal receptacle
form (Figs. 132-143), CL/PSS, and PSL/PSS (Table 2). The
remaining Aliatypus species can be separated from A. janus by either
CL/IFL or IFL/IVFL (Table 2), and by seminal receptacle form
(Figs. 1 32- 1 43).

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep pit; usually roughly
triangular or T-shaped. Postocular setae form moderately long nar-
row row. Sternum. Fig. 55. Posterior sigilla small to medium sized
and well separated. Pedipalps. Figs. 83-84, 102-105. Tibia markedly
swollen ventrally near distal end. Embolus base distant from ICS

1974]

Coyle — Genus Aliatypus

453

base. Conductor tapers evenly to narrow tip which is sharp or an-
gularly truncate; tip may be bent or straight. Inner (concave) edge
of OCS smooth to somewhat rough. Leg. I. A few to all ventral
macrosetae on tibia and metatarsus are ensiform ; background setae
long, slender, not appressed, and very densely distributed. Abdomen.
Tergites I and III reduced to small patches or spots at bases of
macrosetae. Coloration. Pars thoracica pale yellow to light brown.
Pars cephalica markedly darker; medium to dark red-brown or
chestnut. Chelicerae usually slightly darker red-brown than pars
cephalica. Dorsal surface of pedipalpal patella and tibia same as
chelicerae or darker.

Females: Carapace. Thoracic groove a deep pit; usually roughly
triangular or T-shaped; rarely a transverse furrow, transversely
oval, or circular. Postocular setae form a very narrow longitudinal
row. Sternum. Fig. 66. Posterior sigilla medium sized and well
separated. Great majority of peripheral sternal setae slender; usually
a few to many stout ones on anterior-lateral margins. Longest setae
scattered over most of sternum, but slightly more abundant anteriorly.
Genitalia. Figs. 1 32-1 43. Stalks of seminal receptacles very weakly
to moderately heavily sclerotized ; almost always much thicker at
base than at distal end; weakly to strongly sinuous (3-9 bends).
Bulbs small to medium sized ; less sclerotized than stalks. Coloration.
Pars thoracica pale yellow to rather dark yellow-gray. Pars cephalica
slightly to much darker; pale brown to chestnut; median longitudinal
band and posterior border darker than rest. Chelicerae orange-
brown, red-brown, or chestnut; slightly darker than dark part of
pars cephalica.

Variation. Males: Although the male population samples are very
small, they should provide at least an indication of the geographic
variation patterns within this species. The strongest geographic
variation occurs in pedipalpal tibia shape and conductor tip form.
As Figures 18, 21, 83, and 84 indicate, the Yosemite and Briceburg
specimens have markedly more elongate and slender pedipalpal tibiae
than do the other samples. However, because of a relatively short
pedipalpal patella, the Hume Lake and Sequoia specimens exhibit a
PTL/PPL ratio similar to the Yosemite and Briceburg specimens
and distinct from the other samples (Fig. 19). Although conductor
tip shape varies considerably, the variation is rather continuous and
clinal (Figs. 102-105). The Yosemite conductor tips (Fig. 103) are
rounded in ventral view and strongly bent dorsad. The Briceburg
conductor tip is very similar except that it is not as strongly bent
dorsad. The Benton Station conductor tip is like the Yosemite tips.

454

Psyche

[September-December

Yosemite
Briceburg
Benton Sta.
Hume L
Pinehurst
Sequoia
Glenville
Squaw Flat

18

1.00 1.10 1.20 1.30 1.40 1.50

CL/PTL

Yosemite
Briceburg
Benton Sta.
Hume L
Pinehurst
Woodlake
Glenville
Squaw Flat

20

2.10

IML/lTarL

Yosemite
Briceburg
Mammoth L
Hume L
Pinehurst
Woodlake
Sequoia
Glenville

22

6.00 700

CL/PSS

Yosemite
Briceburg
Mammoth L
Hume L
Pinehurst
Woodlake
Sequoia
Glenville

23

4.00

5.00 6.00

CL/HTarL

Yosemite
Briceburg
Mammoth L.
Hume L
Pinehurst
Woodlake
Sequoia
Glenville

600 700 8.00 9.00 10.00 11.00

24 sw/psl

Yosemite

.

Briceburg

.

Benton Sta.

.

Hume L

. .

Pinehurst • •

Woodlake •

Sequoia

.

Glenville •

Squaw Flat •

19 1,0

" 1.20 1.30

PTL/PPL

Yosemite ••

Briceburg •

Benton Sta.

•

Hume L

••

Pinehurst

•

Woodlake

•

Sequoia

•

Glenville

•

Squaw Flat

•

21 0.18 0.20

0.22 0.24 0.26 0.28 0.30
PTT/ PTL

Figures 18-24: Geographic variation of Aliatypus janus. Modified Dice-

Leraas diagrams and map of sample localities. 18-21: males. 18: CL/PTL
variation. 19: PTL/PPL variation. 20: IML/ITarL variation. 21: PTT/
PTL variation. 22-24: females. 22: CL/PSS variation. 23: CL/IVTarL
variation. 24: SW/PSL variation.

1974]

Coyle — Genus A liatypus

455

The Hume Lake conductor tips (Fig. 102) are blunt or slightly
angularly truncate, and bend dorsad almost as strongly as the
Yosemite conductor tips. The Pinehurst conductor tips (Fig. 1O4)
are slightly broader and thicker, are angularly truncate, and bend
only very slightly dorsad. The Woodlake, Glenville (Fig. 105), and
Squaw Flat conductor tips are similar to those at Pinehurst, but are
more angularly truncate and completely unbent.

Rather strong variation occurs in the ratio IML/ITarL (Fig. 20),
with Yosemite and Briceburg specimens distinct from Hume Lake
and Pinehurst specimens. Tibia and metatarsus I setation varies
geographically with Yosemite, Briceburg, Benton Station and Hume
Lake specimens having 10 to 50 percent of the ventral macrosetae
ensiform and relatively long densely distributed background setae.
All other specimens have Jo to 100 percent of the ventral macrosetae
ensiform, and shorter, less erect, less densely distributed background
setae. The Yosemite and Briceburg specimens differ from the rest in
that the pars thoracica is not markedly lighter than the pars cephalica.
The Yosemite and Benton Station specimens differ from the rest in
that the postocular setae extend back to a point at one-half or more
of the distance from the anterior edge of the cephalothorax to the
thoracic groove.

Fe?nales: The female population samples also show a considerable
amount of geographic variation, which is not surprising for such a
widespread species. The chief variation pattern is that most samples
(Hume Lake, Pinehurst, Woodlake, Sequoia, and Glenville) are
quite similar, but that the northern samples (Yosemite, Briceburg,
and Mammoth Lakes) do not form as homogeneous a grouping and
each differs almost distinctively from most other samples in a few
characters. The very small sample sizes limit the strength of any
conclusions.

Discontinuous geographic variation in quantitative characters in-
volves those ratios incorporating PSS and PSL (Figs. 22 and 24).
In these ratios the Mammoth Lakes population, with its smaller,
more widely spaced posterior sigilla, is quite divergent. CL/IVTarL
(Fig. 23) exhibits rather strong geographic variation with the
Yosemite specimen very divergent. Other quantitative characters
exhibit less geographic variation.

Variation in seminal receptacle form is illustrated by Figures 132-
143. All samples except those from the Yosemite-Briceburg area
have very similar seminal receptacles. In both the Yosemite and
Briceburg specimens, the stalk base is not much thicker than the
distal end, a condition similar to that of A. calif ornicus. Additional

456

Psyche

[September-December

noteworthy variation among female samples is as follows: the Mam-
moth Lakes, Pinehurst, Woodlake, and Glenville specimens are
lighter colored than most specimens from other localities; the Wood-
lake and Glenville specimens have zero to two stout peripheral sternal
setae (nearly all other specimens have many more) and fewer post-
ocular setae than nearly all other specimens.

Variation in both sexes indicates, not surprisingly, that gene flow
is restricted in portions of this species’ geographic range. Gene flow
between the more northern populations (Yosemite, Briceburg, Ben-
ton Station, and Mammoth Lakes) and those to the south may be
especially restricted. Possible competition or hybridization with A.
calif ornicus in the area of sympatry around Mariposa may have an
important effect upon the genetics of these northern populations. The
Yosemite and Briceburg specimens are especially divergent and may
eventually prove to represent a distinct species. Obviously, more and
larger samples must be collected in order to obtain an accurate picture
of geographic variation and of factors affecting gene flow within
this species.

Distribution. Central and southern Sierra Nevada Mountains and
part of the Coast Range Mountains north of Los Angeles (Map a).

Records. California. Fresno Go.: Loop road 5 mi. S of Hume
Lake, 6000 ft., 16 Oct. 1973, 2cf, 9$. — Hwy. 245, 1 mi. E of
Pinehurst, 4300 ft., 18 Oct. 1973, 2cf, 3$. Kern Co.: Glenville,
15 Nov. 1969, cf- — 10 mi. SW of Glenville, 2$. Madera Co.:
7 mi. W of Mammoth Lakes, 8700 ft. 2?. Mariposa Co.: near
Briceburg, 1300 ft., cf. — 4 mi. N of Briceburg on Hwy. 140,
1300 ft., $. — Yosemite Nat’l Pk., 3.4 mi. E of Yosemite Cr.
bridge on Hwy. 120, about 8000 ft., 10 Aug. 1972, 2cf • — Yosem-
ite Nat’l Pk., 20 mi. E of Crane Flat on Hwy. 120, about 8000 ft.,
?. Mono Co.: Benton Station, 26 Oct. 1941, cf. Tulare Co.:
Sequoia Nat’l Pk., Congress trail near General Sherman Tree, 6800
ft., 13-14 Aug. 1972, cf, 5$. — 14 mi. N of Woodlake on Hwy.
69, 1200 ft., 10 Nov. 1972, cf, 3$. Ventura Co.: 5.5 mi. S of
Squaw Flat, 29 Nov. 1970, cf •

Aliatypus isolatus new species
Figures 56, 67, 80, 107-109, 144-146. Map 3.

Type specimens and etymology. Holotype male from Cave Springs
Campground, about 9 mi. north of Sedona in Oak Creek Canyon,
Coconino Co., Arizona, 19 August 1972 (F. A. Coyle). One male
and four female paratypes. The specific name is a Latin adjective
meaning isolated.

1974]

Coyle — Genus Aliatypus

457

Diagnosis. The geographic range of this species (Map 3) is well
separated from those of all other Aliatypus species. Males: Because
of its proportionately long metatarsus I, short tarsus I, and short
pedipalpal patella, A. isolatus is best distinguished from most of its
congeners by the ratios IML/ITarL, PPL/PFL, and PTL/PPL
(Table 1). The palpus structure (Figs. 107-109) of A. isolatus is
distinctly different from that of most species. CL/ALS (Table 1 ) is
the best character for distinguishing A. isolatus from closely related
A. janus. Females: A. isolatus is extremely similar to A. janus and
A. calif ornicus ; refer to diagnoses of these species. A. isolatus differs
from similar A. aquilonius in having a transverse thoracic pit rather
than a rounded or elongate one, and in its larger body size (especially
longer tarsi; Table 2). A. isolatus can be separated from A. gwomus
by seminal receptacle form (Figs. 144-146), AMD/AMS and
CL/AMD (Table 2), and body size (Table 2). CL/IVTL,
CL/IFL' and PSL/PSS (Table 2) clearly distinguish A. isolatus
from all other species.

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep transverse pit or groove.
Postocular setae form a relatively long narrow row. Sternum.
Fig. 56. Posterior sigilla small and far apart. Pedipalp. Figs. 80,
107-109. Tibia swollen ventrally near distal end. Embolus base
distant from ICS base. Conductor tapers rather evenly to narrow,
sharp, angularly truncate tip which is bent. Inner (concave) edge
of OCS smooth. Leg I. Tibia and metatarsus with all or nearly all
ventral macrosetae attenuate; background setae elongate, slender,
densely distributed, and not appressed. Abdomen. Tergite I small.
Tergite III absent. Coloration. Pars cephalica and chelicerae light
brown to dark brown ; much darker than light grey-yellow pars
thoracica. Pedipalps darker than pars cephalica; medium orange-
brown to dark red-brown.

Females: Carapace. Thoracic groove a deep transverse pit or
groove. Postocular setae row extends to point at least one-half of
distance from anterior edge of carapace to thoracic groove. Sternum.
Fig. 67. Posterior sigilla small and far apart. Peripheral setae
slender. Longest setae scattered all over sternum. Genitalia. Figs.
144-146. Seminal receptacles weakly sclerotized. Base of stalk rela-
tively thick and nearly straight. Stalk becomes much narrower
distally; long; 4-7 bends; often very irregularly looped. Bulbs small.
Coloration. Pars thoracica pale yellow to pale yellow-brown. Pars
cephalica darker; light yellow-brown to medium brown. Chelicerae
slightly darker than pars cephalica.

458

Psyche

[September -December

V aricition. Males: A comparison of the two small samples of two
males each indicates rather strong geographic variation. The Oak
Creek Canyon sample has a markedly larger body size ( CL = 4.8
mm, 4.9 mm), a proportionately longer pedipalpal patella (CL/
PPL = 1.53, 1.64), and darker coloration than the Santa Catalina
Mountain sample (CL = 3.5 mm, 3.8 mm; CL/PPL = 1 .75,
1.77)* The conductor tip is proportionately a bit narrower and is
bent more strongly (Figs. 107-109) in the Oak Creek Canyon males.
Females: No marked geographic variation occurs in any of the ratio
characters. The Oak Creek Canyon sample averages larger in body
size, but the ranges of the two samples overlap. Seminal receptacle
stalks are more irregularly sinuous in the Santa Catalina Mountain
sample than in the Oak Creek Canyon sample (Figs. 144-146). The
two samples show broadly overlapping color variation.

Distribution. Arizona ( Map 3 ) .

Records. Arizona. Coconino Co.: 0.2 mi. S of Manzanita Camp-
grd. in Oak Creek Canyon about 6 mi. N of Sedona, 4400 ft., 9 •
— Cave Springs Campgrd. in Oak Creek Canyon about 9 mi. N of
Sedona, 4900 ft., 19 Aug. 1972, 2 cf, 4 $. Pima Co.: Molino
Basin Campgrd. in Santa Catalina Mtns., 4500 ft. — 1.5 mi. below
Bear Cr. Picnic Area along Hwy. to Mt. Lemon, 54°° ft., 27 March
1970, 2 (d*), 5 $. — Bear Cr. Picnic Area in Santa Catalina
Mtns., 5800 ft. — General Hitchcock Picnic Area in Santa Cata-
lina Mtns., 6000 ft.

Aliatypus aquilonius new species
Figures 57, 68, 81, 106, 147-148. Map 2.

Type specimens and etymology . Holotype male from Grizzly
Creek Redwoods State Park, Humboldt Co., California, 8 August
1972 (F. A. Coyle). One male and 14 female paratypes. The
specific name is a Latin adjective meaning northern.

Diagnosis. Males: The weakly sclerotized, finger-like extension
at the tip of the palpus (Fig. 106) is distinctive. The pedipalpal
patella is proportionately long (Fig. 81), so that appropriate ratios
from among the following distinguish A. aquilonius from any
other species (Table 1): PTL/PPL, PPL/PFL, CL/PPL, CL/
PSS, SW/PSS. A. aquilonius males are markedly smaller than
those of coastal A. calif ornicus and some other species (Table 1).
Females: The following characters best distinguish A. aquilonius
from similar species: smaller AMD/AMS (Table 2) and more
strongly coiled seminal receptacle stalks (Figs. 147-148) than in

1974]

Coyle — Genus Aliatypus

459

A. gnomus; smaller CL/PTSR and IFL/IVFL (Table 2) than in
A. calif ornicus ; smaller IFL/IVFL and fewer IMS (Table 2) than
in A. janus ; fewer IMS and smaller CL/ITL (Table 2) than in
A. isolatus. Because of its small widely spaced posterior sigilla
(Fig. 68) and proportionately long tibia IV, A. aquilonius is easily
separated from the rest of the species by the following characters
(Table 2): CL/IVTL, CL/PSS, CL/PSL, SW/PSS, SW/PSL,
and PSL/PSS.

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep longitudinal groove or
a deep rounded pit. Postocular setae form a relatively short and
narrow longitudinal band. Sternum. Fig. 57. Posterior sigilla faint,
small, and far apart. Pedipalp. Figs. 81, 106. Distal half of tibia
ventrally moderately swollen. Embolus base distant from ICS base.
Palpus tipped with weakly sclerotized finger-like extension. Inner
(concave) edge of OCS smooth. Leg I. Tibia and metatarsus with
most of ventral macrosetae attenuate ; background setae long, slender,
erect, and rather sparsely distributed. Abdomen. Tergites I and III
reduced to small patches or spots at bases of macrosetae. Coloration.
Cephalothorax and chelicerae nearly homogeneous light yellow-brown.
Pedipalps slightly darker.

Females: Carapace. Thoracic groove a deep pit; circular, irregular,
or longitudinal. Postocular setae few; form a moderately long,
roughly single row. Sternum. Fig. 68. Posterior sigilla small and
far apart. Peripheral setae slender. Longest setae absent from large
central area. Genitalia. Figs. 147-148. Seminal receptacles very
weakly sclerotized. Base of stalk relatively thick, elongate, and
nearly straight. Stalk long, becomes much narrower distally, 3-6
bends. Bulbs very small. Coloration. Cephalothorax homogeneous
light yellow-brown. Chelicerae darker light brown to medium brown.

Variation. There is very little variation among the four males.
Females: The Grizzly Creek sample (n — 12) has a markedly larger
mean body size (CL m 4.60 zb .74 mm; range = 3-4 mm-5.8 mm)
than the Redway sample (n == 14) (CL =fi 3.40 zb .41 mm; range
— 2.8 mm-4.1 mm). Generally, the larger the specimen the more
heavily sclerotized and elongate the posterior sigilla; the posterior
sigilla are round in the smallest specimens and twice as long as wide
in the largest specimens. There is continuous variation in the degree
of coiling of seminal receptacles; all specimens fall between, or are
similar to, the conditions illustrated by Figures 147 and 148.

Distribution. Humboldt Co. in northwestern California (Map 2).

460

Psyche

[September-December

Records. California. Humboldt Co.: Grizzly Creek Redwoods
St. Pk., 400 ft., 8 Aug. 1972, 2 ( cT ) , 12 $. — 1.4 mi. W of Red-
way on road to Briceland, 400 ft., 7 Aug. 1972, ( cf ), 14? . — 2 mi.
W of Briceland, 400 ft., 15 Sept. 1971, cf, ?.

Aliatypus gnomus new species
Figures 49, 58, 69, 82, 101, 149-150. Map 2.

Type specimens and etymology. Holotype male from Henry
Cowell Redwoods State Park, Santa Cruz Co., California, 3 August
1972 (F. A. Coyle). One male and four female paratypes. The
specific name is a Latin noun meaning dwarf.

Diagnosis. Males: At least one of the following ratios (Table 1)
will separate this species from any one of the other species: CL/PPL,
CL/PSL, PSL/PSS, and PTL/PPL. The palpus, with an ICS
keel and a slender conductor tip (Fig. 101), is quite distinct from
that of all species. The ICS keel is narrower and the conductor tip
more slender than in closely related A. calif ornicus. These palpus
features, CL/PPL, CL/PSL, and body size (Table 1) are the best
characters for separating A. gnomus from A. calif ornicus. In all of
these characters, A. gnomus is more distinct from coastal A. cali-
f ornicus populations than from the Sierran A. californicus popula-
tions. Females: A. gnomus has distinctively large, close set AME’s,
so that one ratio, AMD/AMS (Table 2), clearly separates this
species from all other species. Also, the weakly sinuous seminal
receptacles (Figs. 149- 150) are distinctive. A. gnomus is distinctively
smaller (Table 2) than most other species.

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep rounded pit. Post-
ocular setae form rather short narrow row which is broadest an-
teriorly. Sternum. Fig. 58. Posterior sigilla small and far apart.
Pedipalp. Figs. 82, 101. Distal half of tibia ventrally swollen.
Embolus base distant from ICS base. ICS with a thin narrow keel
distallv. Conductor tip slender and tapers evenly to fine point. Inner
(concave) edge of OCS smooth. Leg I. Tibia and metatarsus with
most of ventral macrosetae attenuate; background setae long, slender,
moderately densely distributed and more or less appressed on tibia but
suberect on metatarsus. Abdomen. Tergites I, II, and III all well
developed; II largest. Coloration. Pars cephalica and chelicerae pale
brown. Pars thoracica pale yellow. Pedipalps darker than pars
cephalica.

1974]

Coyle — Genus Aliatypus

461

Females: Carapace. Fig. 49. Thoracic groove a deep circular or
longitudinally oval pit. Three or four large postocular setae form
single, moderately long, row; sometimes additional tiny setae. AME’s
relatively large and separated by less than diameter. Sternum.
Fig. 69. Posterior sigilla small and far apart. Peripheral setae
slender. Longest setae less common centrally than toward the pe-
riphery. Genitalia. Figs. 149- 150. Seminal receptacle stalks ex-
tremely weakly sclerotized, weakly sinuous (1-4 weak bends), and
same diameter throughout length. Bulbs almost transparent and
proportionately small to medium sized. Coloration. Pars thoracica
pale yellow. Pars cephalica and chelicerae a darker pale brown.

Distribution. Known only from the type locality (Map 2).

Record. California. Santa Cruz Co.: Henry Cowell Redwood
St. Pk., along Hwy. 9, 3.3 mi. S of Felton center, 400 ft., 3 Aug.
1972, 2 ( cf ) , 4 ? •

Aliatypus gulosus new species
Figures 25-26, 59, 70, 85, no, 151-154. Map 2.

Type specimens and etymology. Holotype male from Salt Creek,
1.5 miles north of Dana Point, Orange Co., California, 6 December
1968 (W. R. Icenogle). Three males and 17 female paratypes. The
species name is a Latin adjective meaning gluttonous.

Diagnosis. Males: The palpus (Fig. no) of A. gulosus is dis-
tinctively different from that of all other species. The very loosely
looped sperm reservoir, the closeness of the embolus base to the ICS
base, the jagged, serrate inner (concave) edge of the OCS, and the
evenly tapered conductor tip are some of the more distinctive palpus
features. The distinctive, long banana shape of the pedipalpal tibia
(Fig. 85) is expressed quantitatively by the excellent diagnostic
ratios (Table 1) PTX/PTL and PTL/PPL. Females: The semi-
nal receptacles (Figs. 1 51-154) of i. gulosus are distinctively dif-
ferent from those of all other species. The stalks are short and
straight and the bulbs are relatively large. The appropriate ratio
selected from among the following will clearly distinguish A. gulosus
from any other species (Table 2) : CL/IFL, CL/IML, CL/PSS,
SW/PSS, SW/PSL, PSL/PSS, and IMS/PSS. Any of the last
five ratios distinguish A. gulosus , with its small, widely spaced pos-
terior sigilla, from the other southern California species (A. thomp-
soni, A. plutonis, and A. torridus) .

Description. See Tables 1-3.

462

Psyche

[September-December

Chetsworth
Crescenta Park
Sierra Madre
Eaton Can.

Salt Cr

Chatsworth .

Crescenta Park •

Sierra Madre •

Eaton Can. — f— 1 7

Salt Cr —

25

1.60 1.70 1.80

cl/ivfl

26

1.70 1.80 1.90

CL/IFL

Figures 25-26: Geographic variation of Aliatypus gulosus females.

Modified Dice-Leraas diagrams. 25: CL/IVFL variation. 26: CL/IFL

variation.

Males: Carapace. Thoracic groove a deep, roughly circular pit.
Postocular setae form a roughly triangular grouping. Sternum.
Fig. 59- Posterior sigilla small and well separated. Pedipalps.
Figs. 85, 1 10. Tibia banana shaped. Sperm reservoir large and
loosely looped. Embolus base very close to ICS base. Distal half of
conductor tapers evenly to tip. Inner (concave) edge of OCS with
minute jagged serrations. Leg. I. Very similar to A. plutonis leg I
in proportions and setation. Most ventral macrosetae on tibia and
metatarsus are ensiform. Abdomen. Tergites I, II, and III all well
developed; II largest and I smallest; sometimes II and III are fused
together or nearly so. Coloration. Carapace light yellow-brown to
medium red-brown; margins of pars cephalica often slightly darker.
Chelicerae like lightest parts of carapace or slightly darker. Pedi-
palps dorsally like lightest parts of carapace or lighter.

Females: Carapace. Thoracic groove a deep transverse pit; slightly
to much wider than long. Postocular setae form a roughly double
row. Sternum. Fig. 70. Posterior sigilla rather small and far apart.
Usually, all or most peripheral sternal setae slender; stout setae most
likely found near anterior-lateral margins of sternum. Longest setae
scattered rather evenly over sternum. Genitalia. Figs. 151-154*
Seminal receptacles very weakly sclerotized. Stalks short and straight.
Bulbs relatively large. Coloration. Pars thoracica pale yellow to
medium brown. Pars cephalica centrally about same color; darker
(light brown to chestnut brown) around margin. Chelicerae like
darker portion of pars cephalica.

Variation. While the total male sample (one male from Eaton
Canyon and five from Salt Creek) is remarkably homogeneous in all
characters, the female samples exhibit patterns of geographic variation
which indicate some limitation to gene flow between the Salt Creek
and the Los Angeles area populations. Females: Three characters
exhibit marked geographic variation. The Salt Creek and Eaton
Canyon samples differ somewhat markedly in two ratios, CL/IFL
and CL/IVFL (Figs. 25-26). Also, Salt Creek specimens possess

1974]

Coyle — Genus A liatypus

463

fewer stout peripheral sternal setae than most Los Angeles area
specimens; a few of the latter possess many stout setae scattered along
the entire sternal margin. The Chatsworth specimen is markedly
variant. All its peripheral sternal setae are stout, the very long
sternal setae are almost completely limited to the anterior half of the
sternum, the sternum is exceptionally wide (SL/SW == 1*02), and
femur I is longer than femur IV (IFL/IVFL — 1.05). Some
minor and non-geographic variation occurs in the relative positions
of the seminal receptacles (Figs. 1 51-154).

Distribution. The Los Angeles Basin of southern California
(Map 2) .

Records. California. Los Angeles Co.: Eaton Canyon Park,
3 Jan. 1965, cT i 7$- —Sierra Madre, Bailey Canyon, $. —
Crescenta Valley Park, 9- — Chatsworth, . Orange Co.: Salt
Creek, 1.5 mi. N of Dana Point, 60 ft., 6 Dec. 1968, 2cf, 4$ ;
5 Sept. 1969, cf ; 12 Nov. 1969, <A ; 13 ? .

Aliatypus thompsoni new species
Figures 27-33, 40-44, 50-53, 60, 71-73, 86, 93, m-113, 155-171-

Map 4.

Type specimens and etymology. Holotype male from Chatsworth,
Los Angeles Co., California, 25 November 1967 (M. E. Thompson).
Three male and 15 female paratypes. This species is named after
Mel Thompson, who has collected most of the male specimens studied.

Diagnosis. Males: A. thompsoni is distinct from all other species
in two ratio characters (Table 1), PSL/PSS and CL/IML. The
posterior sigilla are large, faint, and closely spaced (Fig. 60) and
metatarsus I is relatively long ( Fig. 93 ) . The relatively short ventral
macrosetae, all of which are ensiform, and the appressed background
setae of tibia and metatarsus I (Fig. 93) are also distinctive. The
thoracic groove is nearly always absent or shallow (Figs. 50-51).
Females: Unlike all other species, A. thompsoni females either have
no thoracic groove or only a shallow vestige (Figs. 52-53). The
small PSS (Table 2; Figs. 71-73) distinguishes this species from all
others except A. tropkonius. The large numbers of PTSR and IMS
(Table 2) distinguish this species from many others. An appropriate
ratio from among the following will distinguish A. tho?npsoni from
any other species: CL/PSS, SW/PSS, and IMS/PSS (Table 2).
A. thompsoni seminal receptacles (Figs. 1 55-1 7 1 ) , with their long,
many-looped, non-tapered stalks and relatively small to medium sized
bulbs, are diagnostically useful.

Description. See Tables 1-3.

464

Psyche

[September-December

Males: Carapace. Figs. 50-51. Thoracic groove absent, a slight
double longitudinal depression, or a shallow double longitudinal pit.
Postocular setae form a short narrow longitudinal row. Sternum.
Fig. 60. Posterior sigilla faint, large, and close to one another.
Pedipalps. Figs. 86, m-113. Tibia strongly swollen ventrally near
distal end. Embolus base distant from ICS base. Conductor tip
shaped like knife blade tip. Inner (concave) edge of OCS smooth
to moderately rough. Leg I. Fig. 93. Tibia and metatarsus with
ventral, erect to suberect, relatively short, ensiform macrosetae ;
nearly all rest of setae appressed. Abdomen. Tergites I and III
much smaller than tergite II; usually reduced to small patches or
spots at bases of macrosetae. Coloration. Pars thoracica pale yellow
to pale yellow-brown. Pars cephalica darker; pale brown to medium
brown. Chelicerae dorsally similar to or slightly darker than pars
cephalica. Pedipalps dorsally vary from pars thoracica color to
cheliceral color; sometimes with orange or red tint.

Females: Carapace. Figs. 52-53. Thoracic groove absent or vesti-
gial; if latter, it is usually a shallow depression or rarely a shallow
rounded pit. 2-5 postocular setae form a very narrow longitudinal
row. Sternum. Figs. 71-73. Posterior sigilla faint, often irregularly
shaped, large, and very close together. Peripheral sternal setae all
slender ; sometimes a few to many anterior-lateral ones stout. Longest
setae scattered over most of sternum. Chelicerae. Fig. 40. Genitalia.
Figs. 1 55-171. Seminal receptacles extremely weakly sclerotized.
Stalks very long, nearly same diameter throughout length and
strongly looped (4-11 bends), sometimes irregularly. Bulbs small to
medium sized. Coloration. Pars thoracica pale yellow to yellow.
Pars cephalica darker (at least around margins and median longi-
tudinal line) ; light yellow to light brown. Chelicerae darker than
pars cephalica; light brown or light orange-brown to medium red-
brown.

Variation. Males: Several male characters show strong geographic
variation. Two variation patterns are common : often the Sierra
Madre and Henninger Flats samples are similar to one another and
divergent from the rest; sometimes the Tehachapi Mountain sample
is divergent.

The Henninger Flats males are smaller than any others and the
Tehachapi Mountain sample exhibits the largest body size average,
but body size values in the other samples fill the gap between these
two extremes (Fig. 27). The sternum is markedly broader (Fig. 30)
and the pars cephalica is distinctly more elongate (Figs. 28, 50-51)
in the Sierra Madre and Henninger Flats samples than in all others.

1974]

Coyle — Genus Aliatypus

465

Tehachapi • 4

Placerita 3

Chatsworth 1 4

Pac. Palisades •

Las Barras Can. • 2

Henninger F 1 4

Sierra Madre •

Tehachapi
Placerita
Chatsworth
Pac. Palisades
Las Barras Can.
Henninger F
Sierra Madre

27

3.5 4.0 4.5 5.0

CL

28

1.30 1.40 1.50 1.60 1.70

cl/pcl

Tehachapi
Placerita
Chatsworth
Pac. Palisades
Las Barras Can.
Henninger F
Sierra Madre

Tehachapi
Placerita
Chatsworth
Pac. Palisades
Las Barras Can.

Henninger F 1 4

Sierra Madre •

29

1.20 1.30 1.40 1 50

CL/ITL

30

780 1.90 1 ZOO 1 zio ' Z20>

CL/SW

A. janus
Mariposa
A. thompsoni

erf— 1 — 66

A. janus
Mariposa
Kernville
Tehachapi
Squaw Rat
Santa Ynez
Placenta
Chatsworth
Limekiln Can.
Pac. Palisades
Baldwin Hills
Las Barras Can.
Henninger F
Eaton Can.
Sierra Madre

-^-11
fT't 1 . — 15

E=j^3 12

8

31

1.20 1.30 1.40 1.50 1.60 . 1.70

cl/pcl

Kernville
Tehachapi
Squaw Flat
Santa Ynez
Placerita
Chatsworth
Limekiln Can.
Pac. Palisades
Baldwin Hills
Las Barras Can.
Henninger F
Eaton Can.
Sierra Madre

6

I 3

■12

33

8.0 12.0 16.0 20.0
CL/PSL

~~ 160 1.80 2.00 2.20
cl/sw

Figures 27-32: Geographic variation of Aliatypus thompsoni. Motdified

Dice-Leraas diagrams and map of sample localities. 27-30: males. 27: CL
(in mm) variation. 28: CL/PCL variation. 29: CL/ITL variation. 30:
CL/SW variation. 31-32: females. 31: CL/PCL variation compared with
that of A. janus and the enigatic Mariposa population. 32: CL/SW
variation. Figure 33: Modified Dice-Leraas diagram showing CL/PSL

variation of females of Aliatypus thompsoni, A. janus, and the enigmatic
Mariposa population.

466

Psyche

[September-December

Thoracic groove form is correlated with pars cephalica shape. Those
specimens (Sierra Madre and Henninger Flats) with an exception-
ally elongate pars cephalica lack any vestige of a thoracic groove
(Fig. 51). Specimens with a more normal pars cephalica possess at
least a vestigial thoracic groove in the form of a shallow depression
(Fig. 50). Tehachapi Mountain specimens have slightly to markedly
deeper thoracic grooves than other specimens.

Tehachapi Mountain males have two or three trichobothria dor-
sally near the distal end of metatarsus IV. All other A. thompsoni
males as well as all other Aliatypus males have only one tricho-
bothrium in that position. Variation in palpus conductor tip form
is illustrated by Figures 111 to 113. The Tehachapi Mountain con-
ductor tips (Fig. hi) are slightly but consistently different from
those in all other samples. Tehachapi Mountain males tend to have
a darker pars cephalica and chelicerae, and redder pedipalps than do
other specimens. The Las Barras Canyon specimens have noticeably
more elongate legs (Fig. 29) than most others.

Fe?nales: A few female characters exhibit strong geographic vari-
ation. These are characters which also vary strongly in the male
sample, and the geographic patterns of these variations are like those
in the male sample.

The pars cephalica is markedly more elongate in the Sierra Madre,
Eaton Canyon, and Henninger Flats samples than in all other sam-
ples (Figs. 31, 52-53). These same three southeastern samples have
on the average a considerably broader sternum than most other sam-
ples (Figs. 32, 71-73). The thoracic groove is completely absent in
these three samples with an elongate pars cephalica (Fig. 53), and is
only a faint depression (slightly more heavily sclerotized than its
immediate surroundings) (Fig. 52) in all other samples except the
Tehachapi Mountain and Kernville samples. These latter specimens
have a slightly deeper depression or a shallow pit.

The Tehachapi Mountain and Kernville specimens have two to
four trichobothria dorsally near the distal end of metatarsus IV. All
other females of A. tko?npsoni and all other Aliatypus species have
only one such trichobothrium. Variation in seminal receptacle form
is illustrated by Figures 155 to 171. The Tehachapi Mountain and
Kernville specimens have shorter receptacle stalks and fewer loops
per stalk than other specimens elsewhere. The centrally located popu-
lations (Baldwin Hills, Pacific Palisades, Las Barras Canyon, Lime-
kiln Canyon, Chatsworth, and Placerita) all have a dense cluster of
short stout setae at the anterior median edge of the carapace, while

467

1974] Coyle — Genus A liaty pus

all other populations have at most only a few very small or moder-
ately stout setae here.

The similar geographic variation patterns for both sexes of A.
thompsoni indicate two areas of reduced gene flow; one between the
northeastern populations (Tehachapi Mountain and Kernville) and
the rest of the species, and the other between the southeastern popu-
lations (Sierra Madre, Eaton Canyon, and Henninger Flats) and
the rest of the species. In the former area it is likely that there is a
paucity of suitable habitats. It is not as apparent why there might
be reduced gene flow in the latter area. One female that was col-
lected with the Tehachapi Mountain population looks suspiciously
like a product of interbreeding between A. thompsoni and A. torridus
or A. plutonis. If hybridization has occurred in this area, it could
be responsible for some of the variant nature of the Tehachapi and
Kernville populations.

A possible central Sierran population of A. tho?npsoni. Seven
A liaty pus females with only a faint depression for a thoracic groove
were collected at Mariposa, California (0.5 mi. north of town limits
on Hwy. 49). These are similar to A. thompsoni in all characters
execpt posterior sigilla size and separation and PTSR. The posterior
sigilla of the Mariposa sample are markedly smaller (Fig. 33) and
farther apart than in A. thompsoni. The PTSR of the Mariposa
sample ranges from four to six with a mean of 4.3, while that of
A. thompsoni ranges from five to eight with a mean of 6.2. Except
for CL/PC L (Fig. 31) and the condition of the thoracic groove,
this sample is even more similar to the sympatric species, A. janus.
It is less similar to sympatric A. calif ornicus and very unlike sym-
patric A. ere bus.

There are a number of possible explanations for this situation.
Perhaps this Mariposa sample is conspecific with A. thompsoni. Per-
haps it is a reproductively isolated northern derivative of A. thomp-
soni. Perhaps it is a variant population of A. janus which has
undergone a drastic shift in pars cephalica and thoracic groove struc-
ture. Perhaps it is a product of hybridization between A. thompsoni
and A. janus. Intensive field work in the Mariposa area is required
to solve this problem.

Distribution. Foothills of the Los Angeles area north into the
Santa Ynez Mountains in the west and the southern end of the
Sierra Nevada Mountains in the east (Map 4).

Records. California. Kern Co.: Piute Mtns. S of Kernville, $.
— along Water Canyon Rd., 4500-5500 ft., Tehachapi Mtns. S of

468

Psyche

[September-December

Tehachapi, 7 Sept. 1967, (d) ; 10 Oct. 1968, 2 d, 2 9 ; 16 Jan.
T97°) d ; 9 • Los Angeles Co.: Sierra Madre, 900 ft., 4 Feb. 1973,
d ; 9 • —Eaton Canyon Park near Altadena, 8 9- — Henninger
Flats near Altadena, 15 Nov. 1968, 2 d , 2 9 ; 10 Jan. 1968, d ;
16 Jan. 1970, d ; 10 9- —Las Barras Canyon, 2 mi. SE of
Tujunga, 1500 ft., 12 Oct. 1972, 2 d, 9* — Baldwin Hills, 2 9.
— Pacific Palisades, Feb. 1945, d, 9- — Chatsworth, 1000-1200
ft., 2 Oct. 1966, d ; 9 Oct. 1966, d, 2 9 ; 28 Oct. 1967,
d , 9 ; 25 Nov. 1967, d, 3 9 ; 9 9- — Limekiln Canyon, 2.5
mi. NW of Granada Hills, 1300 ft., 3 9- — Placerita Canyon St.
Pk., 31 Oct. 1968, 3 cf, 5 9 ; 6 9 . Santa Barbara Co.: Santa
Ynez Mtns., Stagecoach Rd., 200 yds. W of junc. with Hwy. 154,
2200 ft., 5 9* — Santa Ynez Mtns., Paradise Rd. 2.1 mi. E of
junc. with Hwy. 154, 9 • Ventura Co.: 5 mi. S of Squaw Flat, 2 9 •

Aliatypus trophonius new species
Figures 61, 74, 87, 95, 116, 172-173- Map 4.

Type specimens and etymology. Holotype male from 4.5 mi.
north of Soquel, Santa Cruz Co., California, 13 October 1971 (W.
R. Icenogle). One male and nine female paratypes. Trophonius was
a Boetian oracular god who snatched inquirers underground to give
them revelations.

Diagnosis. Males: The palpus of this species (Fig. 116) is quite
different from that of all other species, except A. erebus and A.
plutonis. The OCS is quite broad to near the tip where it suddenly
tapers to a fine point. Because of proportionately short appendage
articles, a strongly swollen pedipalpal tibia (Fig. 87), and relatively
large close-set anterior median eyes, any one of the following ratios
(Table 1) will separate A. trophonius from nearly all other species:

CL/IFL, CL/ITL, CL/IML, CL/PPL, PTT/PTL, CL/PTT,
CL/AMD. A. trophonius is small, so that many dimensions, espe-
cially PFL and PTL (Table 1) are useful diagnostically. The most
similar species, A. erebus , can be separated from A. trophonius best
with the following ratios (Table 1) CL/PCA, CL/ITarL, and
CL/AMD. Also, A. trophonius has a proportionately shorter prox-
imal branch of the ICS base (Fig. 116) and a relatively broader
conductor just proximal to the conductor tip than does A. erebus.
Females: A. trophonius has relatively short leg I articles and rela-
tively large, close spaced, posterior sigilla, so that it can be separated
from any species by using the appropriate ratio from among the
following (Table 2) : IFL/IVFL, CL/IFL, CL/ITL, CL/IML,

1974]

Coyle — Genus Aliatypus

469

SW/PSS, SW/PSL, and PSL/PSS. Small A. trophonius is easily
separated from large species by IFL, ITL, and I ML (Table 2).
Closely related A. erebus can be distinguished from A. trophonius
by the ratios (Table 2) CL/IMS, CL/AMD, and CL/PTSR, by
body size (CL and other measurements in Table 2), and by the
proportionately longer A. trophonius seminal receptacle stalks with
more bends and relatively smaller bulbs (Figs. 172- 173). A. tro-
phonius seminal receptacles are also markedly different from those of
several other species.

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep circular to elongate-
oval pit. Postocular setae form roughly triangular grouping. Ster-
num. Fig. 61. Posterior sigilla medium sized and rather well sep-
arated. Pedipalps. Figs. 87, 116. Articles relatively short. Tibia
greatly swollen ventrally, especially near distal end. Embolus base
well separated from ICS base. OCS broad to near its tip where it
suddenly tapers to fine point. Inner (concave) edge of OCS smooth.
Leg I. Fig. 95. Majority of ventral macrosetae on tibia and meta-
tarsus are ensiform ; background setae fairly sparse, long and slender,
more erect on metatarsus. Abdomen. Tergites I and III reduced to
spots at bases of macrosetae. Coloration. Pars thoracica pale yellow.
Pars cephalica darker; light brown to medium brown around mar-
gins and along median longitudinal line, lighter elsewhere. Chelicerae
and pedipalps dorsally like pars cephalica.

Females: Carapace. Thoracic groove a deep pit; roughly circular
or slightly transverse. Postocular setae form longitudinal triangular
band. Sternum. Fig. 74. Posterior sigilla moderately large and
somewhat elongate. Peripheral sternal setae slender. Longest setae
scattered rather evenly over sternum. Genitalia. Figs. 1 72-1 73.
Stalks of seminal receptacles very weakly sclerotized, only slightly
more sclerotized than bulbs. Stalks moderately long and slender;
about same diameter throughout length; with 3 to 5 bends. Bulbs
proportionately small to medium sized. Coloration. Pars thoracica
pale yellow to pale yellow-brown. Pars cephalica darker ; light brown
to medium red-brown around margins and along median longitudinal
line, lighter elsewhere. Chelicerae like darkest part of pars cephalica.

Distribution. Known only from the low foothills of the Coast
Range just west and south of San Francisco Bay (Map 4).

Records. California. San Francisco Co.: San Francisco, 9 •
Santa Cruz Co.: 4.5 mi. N of Soquel center on Soquel-San Jose Rd.,
300 ft., 13 Oct. 1971, 2 cf , 9 9 .

470

Psyche

[September-December

Aliatypus erebus new species
Figures 34 -38, 62, 75, 88, 114-115, 174-187. Map 4.

Type specimens and etymology. Holotype male from Fossil Ridge
on south side of Mt. Diablo, Alameda Co., California, 24 November
to 5 December, 1970 (W. E. Azevedo). Three male and two fe-
male paratypes. Erebus was the Latin god of darkness.

Diagnosis. Males: The ratios CL/PED and CL/PCA (Table 1)
will separate A. erebus males from those of the closely related species,
A. trophonius, A. plutonis, and A. torridus. For other characters
useful in separating A. erebus from each of these species, see their
diagnoses. A. erebus ’ conductor form (Figs. 114-115) and strongly
swollen pedipalpal tibia (Fig. 88) are distinct from that of most
other species. The ratios PTT/PTL, CL/ITarL, CL/IFL, CL/
PED, and CL/PSL (Table 1) best distinguish A. erebus from the
less closely related species. Females: A. erebus’ seminal receptacles,
with large bulbs and short stalks with normally only two (at most
three) bends (Figs. 174-187), are distinct from those of all species
except A. torridus. CL/PSL, SW/PSL, or PSL/PSS (Table 2)
will separate A. erebus specimens nicely from distantly related spe-
cies. CL/I MS, IMS/PSS, CL/PTSR, PTSR, or IMS (Table 2)
distinguish A. erebus from A. thompsoni. CL/IMS, CL/AMD,
CL/PTSR, CL, and IVTarL (Table 2) distinguish A. erebus from
closely related A. trophonius. IVFL/IVML and CMT (Table 2)
best distinguish A. erebus from closely related A. plutonis. CMT,
CL/OQW, CL/CMT, and IMS/CMT (Table 2) are the most
helpful characters when separating A. erebus specimens from those
of very similar A . torridus.

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep circular or elongate
oval pit; somewhat narrowed posteriorly. Postocular setae numerous
and form a roughly triangular grouping. Sternum. Fig. 62. Pos-
terior sigilla large and well separated. Pedipalps. Figs. 88, 114-115.
Pedipalpal tibia robust; greatly swollen ventrally near distal end.
Embolus base well separated from ICS base. Proximal branch of
ICS base elongate. OCS rather broad to near its end where it
quickly tapers to a fine point. Inner (concave) edge of OCS smooth
to very rough. Leg I. Ventral tibia and metatarsal macrosetae
attenuate and ensiform ; background setae rather elongate and not
closely appressed. Abdomen. Tergites I and III reduced to patches
or spots at bases of macrosetae. Coloration. Pars thoracica grey-
yellow. Pars cephalica markedly darker; medium brown to dark

1974]

Coyle — Genus Aliatypus

471

red-brown. Chelicerae match pars cephalica. Pedipalps dorsally like
or slightly lighter than pars cephalica.

Females: Carapace. Thoracic groove a deep pit; elongate or trans-
verse; rounded or triangular. Postocular setae numerous; form an
elongate, roughly triangular grouping. Sternum. Fig. 75. Posterior
sigilla large and rather well separated. Stout setae usually distributed
around entire periphery of sternum; always common along anterior
lateral margins. Most of central setae extremely elongate. Genitalia.
Figs. 174-187. Seminal receptacles very weakly sclerotized. Stalks
rather short; with 1 to 3 strong bends; same diameter throughout
length. Bulbs large to very large. Coloration. Pars thoracica grey-
yellow to pale brown. Pars cephalica darker; pale brown to dark
chestnut brown ; darkest along margins and median longitudinal line.
Chelicerae light brown to dark chestnut brown.

Variation. Males: The two male samples, which are separated by
the Central Valley, differ considerably in body size and EGS num-
ber but are very similar in most ratio characters. The male from
Tuolumne Co. is markedly larger (CL = 6.6 mm), has a propor-
tionately longer pedipalpal patella (CL/PPL = 1.57; PTL/PPL
— 1.02) and metatarsus I (CL/IML = 1.76; IML/ITarL =
1.96), and proportionately closer posterior sigilla (SW/PSS — 5.41)
than does the Mt. Diablo sample (CL = 4.4 mm-4.9 mm; CL/
PPL = 1.94-2.03: PTL/PPL = 1. 13-1.16; CL/IML = 1.94-
2.07; IML/ITarL = 1.66-1.71; SW/PSS = 3.30-4.35). The

only marked difference in palpus form is in conductor tip shape

(Figs. 114-115).

Females: The four population samples from the southern half of
the Sierra Nevada Mountains (Sonora, Mariposa, Shaver Lake, and
Pinehurst-Miramonte) are similar in all characters. Each of the other
three samples (Mt. Diablo, Wilbur Springs, and Nevada City) are
divergent from this homogeneous south Sierra grouping in some
characters. Of these divergent samples, the Mt. Diablo sample is
less divergent than either the Wilbur Springs or Nevada City sam-
ples.

The Mt. Diablo sample is divergent from the south Sierra grouping
in body size (Fig. 34), IFL/IVFL, and IFL/ITarL, and is some-
what divergent from all samples in CL/CMT (Fig. 35) and IMS/
CMT. The Wilbur Springs sample is divergent from the south
Sierra grouping in body size (Fig. 34), CL/SL (Fig. 38), CL/SW,
ITL/IML, and IMS/PSS (Fig. 36), and is divergent from all
samples in CL/SL (Fig. 38), CL/SW, and IMS/PSS (Fig. 36).
T^? Nevada City sample is divergent from the south Sierra group-

472

Psyche

[September-December

Mt. Diablo
Wilbur Spgs. •
Nevada City
Sonora
Mariposa
Shaver L.
Pinehurst <
Miramonte

34

6.0

Mt. Diablo

Wilbur Springs
Nevada City

Sonora

Mariposa

Shaver L.
Pinehurst *
Miramonte

35

0.10

•2

1 3

1 4

0.20 0.30 0.40 0.50 0.60

CL/CM T

Mt.' Diablo • 2

Wilbur Spgs. • •

Nevada City • •

Sonora } 3

Mariposa i |; 8

Shaver L. 1 3

Pinehurst + i 4

Miramonte

36

10 12 14 16 18 20 22 24

IMS/PSS

Mt. Diablo
Wilbur Spgs.
Nevada City
Sonora
Mariposa
Shaver L.
Pinehurst +
Miramonte

37

+3

"iso 1 ioo ' 200

N FL/'EZ’TL

Mt. Diablo
Wilbur Spgs.
Nevada City
Sonora
Mariposa
Shaver L.
Pinehurst +
Miramonte

38

-H3

1.50 1.60

CL/SL

1.80

Figures 34-38: Geographic variation of Aliatypus erebus females. Modi-

fied Dice-Leraas diagrams and map of sample localities. 34: CL (in mm)
variation. 35: CL/CMT variation. 36: IMS/PSS variation. 37: IVFL
IVTL variation. 38: CL/SL variation.

ing in IVFL/IVTL (Fig. 37), IFL/IVFL, IVTL/IVML, and
ITL/IML, and is divergent from all samples in IVFL/IVTL
(Fig. 4) and ITL/IML. Mt. Diablo, Sonora, Mariposa, Pinehurst,
and (two of three) Shaver Lake specimens have stout setae dis-
tributed all around the margins of the sternum. The Wilbur Springs,
Nevada City, Miramonte, and (one of three) Shaver Lake specimens
have only a few stout marginal setae, and these only at the anterior
lateral sternal margins. In addition, the Wilbur Springs specimens
are unique in not having short stout setae scattered oyer the periphery
of the central region of the sternum. The range of Color variation is
considerable, with the Sonora specimens darkest, the Pinehurst and

1974]

Coyle — Genus Aliatypus

473

Miramonte specimens a bit lighter, and all other specimens noticeably
lighter than these. Variation in seminal receptacle form (Figs. 174-
187) is continuous with no divergent populations. Bulb size and
stalk diameter seem to be the most variable aspects of seminal re-
ceptacle form.

More and larger samples are needed before firm conclusions can
be made about the genetic relationships of these populations. I feel
that the best working hypothesis suggested by this analysis of varia-
tion is that the Mt. Diablo, Wilbur Springs, and Nevada City popu-
lations are, because of distance and ecological barriers to gene flow,
markedly different genetically from the south Sierra populations, but
that this isolation is either incomplete or has not been of long
enough duration for reproductive isolating mechanisms to develop.
A thorough search for more populations is especially needed in central
and northern California in the eastern part of the Coast Range, the
Central Valley, and the western foothills of the Sierra Nevada
Mountains.

Distribution. Eastern edge of the Coast Range in central Cali-
fornia and western slope of the Sierra Nevada Mountains (Map 4).

Records. California. Butte Co.: 16 mi. E of Oroville on Lump-
kin Rd., 1300 ft., $. Colusa Co.: 3 mi. SE of Wilbur Spgs. on
Bear Cr. Rd., 1250 ft., 2 9- Contra Costa Co.: south side of Mt.
Diablo, Fossil Ridge, 31 Oct.-6 Nov. 1970, cf ; 24 N0V.-5 Dec.
1970, 2 cf \ 5 Dec. 1970-20 Feb. 1971, cf. —0.5 mi. E o'f South
Gate of Mt. Diablo St. Pk., 1300 ft'., 2$. Fresno Co.: Shaver
Lake, 2$. — Hwy. 168, 1.5 mi. S of Dinkey Cr. Rd., near Shaver
Lake, 5400 ft., 9 • — Hwy. 245, 1 mi. E of Pinehurst, 4500 ft..
3 9- — Mt. Miramonte, 9* Mariposa Co.: 0.5 mi. NW of Mari-
posa along Hwy. 49, 2000 ft., 89- Nevada Co.: Hwy. 49, 10.7
mi. S of Grass Valley, 1300 ft. — Hwy. 20, 2 mi. NE of Nevada
City, 3000 ft., 2 9- Tuolumne Co.: Draper Mine Rd., 6 mi. E of
Sonora, 2700 ft., 39- — probably near Sonora, summer 1968, cf-

Aliatypus plutonis new species
Figures 63, 76, 89-90, 94, 117-118, 188-191. Map 4.

Type specimens and etymology. Holotype male from University of
California at Riverside campus, Riverside Co., California, 31 Octo-
ber 1968 (W. R. Icenogle). One male and four female paratypes.
The specific name is the genitive of Pluto, the Latin god of the
nether world.

Diagnosis. Males: A. plutonis can be distinguished best from
closely related and sympatric A. torridus by pedipalp and palpus

474

Psyche

[September-December

characters. The pedipalpal patella of A. plutonis is distinctively more
elongate (Figs. 89-90; PPL/PFL and PTL/PPL in Table 1), its
pedipalpal tibia has a distinctively different shape (Figs. 89-90;
PTX/PTL in Table 1), and its OCS is drawn out into a broad
thin lateral keel just below the tip (Figs. 117-118). A. plutonis
can be distinguished from closely related but allopatric A. erehus by
the following characters (Table 1) : CL/PCA, CL/PED, CL/IFL,
and the broad, thin OCS keel (Figs. 117-118). The following ratios
will separate A. plutonis specimens from those of the other species:
PSL/PSS, CL/PSL, CL/ITL, and CL/PPL (Table 1). Females:
A. plutonis females are distinct from those of all other species by
virtue of their low IVFL/IVML value (Table 2). This is the best
character to use in separating A. plutonis from its closest relatives,
A. erehus and A . torridus. The seminal receptacle stalks of A. plu-
tonis are more elongate and more slender (Figs. 188-191) than those
of A. torridus or A. erehus. SW/PSS and PSL/PSS (Table 2) are
useful in separating A. plutonis from most of the other species.

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep pit, longer than wide;
anterior border rounded ; narrow posteriorly. Postocular setae usually
form a double row anteriorly. Sternum. Fig. 63. Posterior sigilla
large and moderately well separated. Pedipalps. Figs. 89-90, 117-
1 1 8. Tibia strongly swollen ventrally near distal end; slightly
swollen more proximally. Embolus base distant from ICS base.
Conductor tip very sharp. Thin lateral keel-like extension of con-
ductor (OCS) just proximal to tip so that conductor is markedly
narrower just proximal of this keel. Inner (concave) edge of OCS
smooth to slightly rough. Leg I. Fig. 94. Tibia and metatarsus
with ventral, erect, elongate, attenuate and ensiform macrosetae;
background setae elongate and not closely appressed. Abdomen.
Tergites I and III reduced to small patches or spots at bases of
macrosetae. Coloration. Pars thoracica grey-yellow to light brown.
Pars cephalica darker, at least along margins and median longitudinal
line; pale brown to medium brown. Chelicerae match lighter or
darker portion of pars cephalica. Pedipalps dorsally like pars thora-
cica.

Female: Carapace. Thoracic groove a large deep pit; usually
roughly triangular with front wall straight or procurved. Postocular
setae distribution variable; single or roughly double longitudinal
row or long narrow triangular grouping. Sternum. Fig. 76. Pos-
terior sigilla large and moderately well separated. Stout setae dis-
tributed around entire periphery of sternum. Most of central setae

1974]

Coyle — Genus Aliatypus

475

extremely elongate. Genitalia. Figs. 188-191. Seminal receptacles
very weakly sclerotized. Stalks rather short, with a few strong loops
(3-5 bends), moderately thick, and same diameter throughout length.
Bulbs rather large. Coloration. Pars thoracica grey-yellow. Pars
cephalica darker; light brown to medium brown; darkest along mar-
gins and median longitudinal line. Chelicerae match darkest parts
of pars cephalica.

Variation. Males: Both Riverside males have a more slender pedi-
palpal tibia (Fig. 90) than all other specimens, which are all similar
to Figure 89. The palpus form of most specimens is like Figure 118
or intermediate between this and the Palomar Mountain specimen
(Fig. 1 17). Females: As illustrated (Figs. 188-191), there is a small
amount of largely intrapopulation variation in bulb size and stalk
diameter.

Distribution. Southwestern California south of the San Bernardino
Mountains (Map 4).

Records. California. Riverside Co.: U. of Calif, at Riverside
campus, 1250 ft., 27 Oct. 1967, cf ; 31 Oct. 1968, cf ; 4? • — S of
Banning on Hwy. 243, 5.4 mi. S of junc. with I-iO, 3300 ft., 21
Aug. 1968, (2cf ), $ ; 3 ? • San Diego Co.: Hwy. 395, 4 mi. E of
Fallbrook, 800 ft.; 20 Sept. 1971, cf. — Palomar Mtn., Nate
Harrison Grade Rd., 2350 ft., 6 Jan. 1972, cf.

Aliatypus torridus new species
Figures 64, 77, 91, 1 19-120, 192-194. Map. 4.

Type specimens and etymology. Holotype male from Mountain
Center, Riverside Co., California, 3 October 1968 (W. R. Ice-
nogle). One male and four female paratypes. The specific name is a
Latin adjective meaning dry and hot.

Diagnosis. Males: The pedipalpal tibia (Fig. 91) of this species
has a distinctive shape and the pedipalpal patella is relatively short
so that PTX/PTL and PTL/PPL (Table 1) together distinguish
A. torridus from all other species. For other characters which also
distinguish A. torridus from closely related A. trophonius , A. erebus,
and A. plutonis, see these species’ diagnoses. Females: A. torridus
is difficult to distinguish from A. erebus; CMT number, CL/OQW,
CL/CMT, and IMS/CMT (Table 2) are the most helpful diag-
nostic characters. A. torridus is distinguished from closely related
and sympatric A. plutonis by IVFL/IVML, PSL/PSS (Table 2),
and its shorter, thicker seminal receptacle stalks (Figs. 192- 194).
Among the following characters can be found at least one that will

476

Psyche

[September-December

distinguish A. torridus from any one of the other Aliatypus species
(Table 2) : PSL/PSS, CL/PSL, CL/IFL, and seminal receptacle
form (Figs. 192- 194).

Description. See Tables 1-3.

Males: Carapace. Thoracic groove a deep pit with front wall
broad and procurved ; transverse to slightly longer than wide ; nar-
rowed posteriorly. Postocular setae grouped in form of narrow
triangle. Sternum. Fig. 64. Posterior sigilla rather large and well
separated. Pedipalp. Figs. 91, 1 19-120. Tibia swollen ventrally
over most of its length. Embolus base distant from ICS base. Inner
(concave) edge of OCS smooth to slightly rough. Leg I: Tibia and
metatarsus setation very similar to that of A. plutonis. Abdomen.
Tergites I and III reduced to small spots at bases of macrosetae.
Coloration. Pars thoracica grey-yellow to pale brown. Pars cephalica
darker; light brown to medium brown; darkest along margin and
median longitudinal line. Chelicerae match either lighter or darker
portion of pars cephalica. Pedipalps dorsally like pars thoracica.

Females: Carapace. Thoracic groove a large, deep, roughly tri-
angular pit with transverse anterior wall ; usually wider than long.
Postocular setae form a long slender triangular grouping. Sternum.
Fig. 77. Posterior sigilla large and well separated. Stout setae dis-
tributed around entire periphery of sternum. Most of central setae
extremely elongate. Genitalia. Figs. 192- 194. Seminal receptacles
weakly to very weakly sclerotized. Stalks short, with 1 to 3 bends,
thick, and same diameter throughout length. Bulbs rather large.
Coloration. Pars thoracica pale yellow to pale brown. Pars cephalica
darker; grey-yellow to medium brown; darkest along margins and
median longitudinal line. Chelicerae medium brown.

Variation. Males: Considering the large geographic distance sep-
arating the two population samples, there is surprisingly little differ-
ence in pedipalp and palpus form (Figs. 1 19-120). Moderately
strong differences between these two small samples show up only in
the following ratios: CL/ALS (El Paso Mountain sample with
higher mean), CL/ITL (Mountain Center mean higher), and
IFL/ITarL (El Paso Mountain mean higher).

Females: Seminal receptacle form is remarkably similar throughout
all three population samples (Figs. 192- 194). Noteworthy geographic
variation occurs in five ratio characters. The Yucaipa sample
(n = 2) has markedly smaller mean values of IMS/CTP and
CL/CTP than both the Mountain Center (n = 4) and El Paso
Mountain (n = 2) samples. In three other characters, CL/OQW,

1974]

Coyle — Genus Aliatypus

477

CL/AMD, and IFL/ITL, the Mountain Center and El Paso
samples are markedly different from one another and the Yucaipa
sample is intermediate.

Distribution . Interior southern California from the San Jacinto
Mountains north to the southern Sierra Nevada Mountains (Map
4).

Records. California. Kern Co.: NE edge of El Paso Mtns.,
i mi. W of Hwy. 395, 3800 ft., 3 Jan. 1969, 2 c? ; 2$. — ■ NE of
El Paso Mtns., spring 1963, c?* Riverside Co.: Mountain Center,
300 yds. W of junc. of Hwys. 243 and 74, 4400 ft., 3 Oct. 1968,
2c? ; 4$ . San Bernardino Co.: Yucaipa, 2800 ft., 2? .

Literature Cited

Banks, N.

1896. New Californian spiders. J. New York Ent. Soc., 4(4) : 88-91.
Coyle, F. A.

1968. The mygalomorph spider genus Atypoidcs (Araneae: Antro-

diaetidae). Psyche, 75: 157-194.

1971. Systematics and natural history of the mygalomorph spider
genus Antrodiaetus and related genera (Araneae: Antrodiaeti-
dae). Bull. Mus. Comp. Zool., 141 (6): 269-402.

Forster, R. R., and Wilton, C. L.

1968. The spiders of New Zealand. Part 2. Otago Mus. Bull. No. 2:
1-180.

Gertsch, W. J.

1949. American Spiders. Princeton, D. Van Nostrand Co., 285 pp.
Loksa, I.

1966. Nemesia pannonica O. Herman (Araneae: Ctenizidae). Ann.

Univ. Sci. Budapest, Sect. Biol., 8: 155-171.

Main, B. Y.

1957. Biology of aganippine trapdoor spiders (Mygalomorphae : Cteni-
zidae). Australian J. Zool., 5(4): 402-473.

Martin, P. S. and Mehringer, P. J., Jr.

1965. Pleistocene pollen analysis and biogeography of the Southwest.
Pp. 433-451. In H. E. Wright and D. G. Frey (eds.), The
Quaternary of the United States. Princeton, Princeton Univ.
Press.

Riemer, W. J.

1958. Variation and systematic relationships within the salamander
genus Taricha. Univ. of California Publ. Zool., 56: 301-390.

Smith, C. P.

1908. A preliminary study of the Araneae Theraphosae of California.
Ann. Ent. Soc. America, 1 (4): 207-249.

Stebbins, R. C.

1949. Speciation in salamanders of the plethodontid genus Ensatina.
Univ. of California Publ. Zool., 48: 377-526.

are defined in Methods section.

478

Psyche

[September-December

=3 C O

CL 03 *r—

CD +->

03 CM

T d

03 O

• H r— r— \ r—

I LO
CM CO
O'* o

o

CO O* CO
LO O

I CM I
Or— |\

CO CM CO

• — CD CU
< 0)5-
C jQ
M— 03 -O
O CC 03

00 CO

CO
I <3-
r— cn
CO CO

CM CO

O

CM CM r— r—

O'*

cm co

CM -H CM

CM CM CM CM

CO CO
CO -+i

CO -n r—

O CO
CM CM

O CO
CM CM

CM CM CM CM

> — CO

S- 4-> 03
03 CD
(JOE
•r- C
-»-> 03*0

LO CO
CM CM

I CO
LO CO
CO CO

CO -fl CO

I co 10

o co

cm o co

co co co co

<J CO
CD LO

O *r-
*r- (J C
4-> CD (D
CO CL E
•r- CO CD

I CO
LO 03
CM CM

CM -f| CM
I CO I CM

00 o ^

CM CM CM CM

<3- *3- CO 1

3.956

californicus 4.08-6.73 2.34-4.31 2.65-4.92 1.92-3.77 0.62-1.07 0.81-1.36 0.36-0.75

1974]

Coyle — Genus Aliatypus

479

VO CO
CO LO

d d

o cr>

d d

vo o

CO

d o

co co

d d

co co
o o

I ID
CO
*3- <3-

CO^t

LO

d d

LO LO

d d

cr> cm
o r—

o

I CO
LO CO

CO LO

d o

LO LO

d d

1 LO

cr> lo
cx* o

CT> O
d -H

cr>
o o

CO CO

d d

LO LO

d d

LO LO

o d

cn o
d -h

co co

d d

C\J 41

i

o cr>

lo co

LO

co co

f'oj \

I LO

od o
lo co

LO O
CM CO

CO o
r^s cm

CO -H
I LO
LO ^
I— CO

CO CO

i

.168

Table 1 (continued)

480

Psyche

[September-December

cal ifornicus 0.10-0.21 0.12-0.18 6-34 1.03-1.15 1.56-1.78 1.57-1.73 2.69-3.22

1974]

Coyle — Genus Aliatypus

481

D CO

r> r—

(co 'N

•3- 1

CO

vo 00

LO O

'co~~"\

CO

O

f oo\

CO LO

10 0

VO

CO

- -H
1

0 CT»
sJ LO

1

co

LO LO

T in
co co

T LO

LO LO
VO LO

T cr>
LO 00

C\J co

T CO
lo

CM CO

CM

1 0

CM LO

CM
1 VO

0
lo cr>

r~ " -H
1 cr»
00

co *3*

1

1 — *3*

vo

1 n- r-

■-

[cm CM

*- ■-

I CM
cn co
o «—

o

1

co o

r— r— O 1

o

I CO
CM CO

I — o

O -H
I 00
O CM

CO
O'! I—

•— o

O -H
I r—

10

o o

00 00 00 00

00 00

LO CO

o o

Table 1 (continued)

482

Psyche

[September-December

PSL/PSS PPL/PFL PTL/PPL PTX/PTL PTT/PTL IML/ITarL SW/PSS

1974]

Coyle — Genus Aliatypus

483

0.655

Table 2. Measurement (in mm), meristic character, and diagnostic ratio values for adult females of Aliatypus
species. For each species or group of species, the most useful diagnostic characters are circled. Range, mean,
and standard deviation given for measurements and ratios. Range and mean given for meristic characters. Number of
females containing maturing eggs or with brood is in parentheses after N. Character abbreviations are defined in
Methods section.

484.

Psyche

[September-December

CT> r—

— +1 ■ — -H i — -H

I <y> I CO I r—

ID CD

in

1 o 1 in
r-* co
in in ov

r— +»

00 o

O -fl
1 in
in «—
*3- m

1 in
in
00 o

73

CVJ *3-

ov o

- r- O r— OO

\ody

(p 0 J 00

0- Vpoj

0 r—

O r—

0 0

. / 0
.229

.00

.318

.84

.363

cn

•3- LO

00 co

cvj inis

*5f r- r—

in 1 —

in co *5}- m

^•1— CO r—

0

CD CO

in co

0
0 in
CO 1—

in
00 ov
co 0

ov co

. — CVJ

CO +1
I CVJ

co
r-*. co

CVJ -fl »— CVJ ±

1 1 01 1 o

o co 00 CVJ r — o

inoi co 1 — in cvj

CVJ -H

I cd
CO

00 o

Or- r— .

CO

CO -H

in cvj
cvj -H

co -h
1 in
o cd
co

CVJ -H r—

1 cvj 1 in 1
o 00 00 r— cvj

o in co *3* m o

co o

r— CVJ r-

CVJ -fi
I CVJ
CD

CVJ -fi
I CD
CO co

r— CVJ f— I— r— .

^ CO

OV *5j- CO CVJ

inN in

in -f<
1 in
cvj in

r— 5*

00 *3-

m -f| cvj -h in -h *=i* -h
10 1 in 1 cvj 1 in

m co co *5i-oo »— in

1 — o coco co in o

in -H
1 co

(D

in -n ±

l CD 10

*5i- *3-

CO CO CVJ o

in -fi
1 co
00 r—
in in

-h in -H

I Ol I ID

in cd ino

co cvj

cvj co in *5i* in

73

in «5i-

r—

co
cn o
in ^1-

co
o o

o CVJ

O CD
CO CO

LO -fi (O -H

1 in 1

r— in
^1- in co

cd

CVJ co

in +1 <0 -fi in -n

1 01 1 1 — 1 in

• — cvj in 01 in

in r— 10

in o

CO «5f

I CD

00 in
co

cvj »—

r- 1 —

f— CVJ

3.656±

californicus 3.61-5.23 2.07-3.07 3.11-4.46 1.15-1.53 3.31-5.23 2.69-4.15 0.79-1.49

1974]

Coyle — Genus Aliatypus

4^5

r— CO r—

4! r— -H

in isA\
cj co 10 10

d* d- o

mo r— r—

r- 41 O r-

<y\ cr> co co

n in in

00 00

r— O r-

in CO

CO <N1 CJ

CJ IO CJ CO

in 41 d- 4i

r— 1 in
co d* »—
d* in d

CO CJ CO

CO 41
I CO
10 cn

C\J

IO CsJ
CsJ CO CsJ CO

in
co 00
cj in

co co

in
in 41

lo in
CO 41

in o

r—

in 41

O CJ
CJ CJ

1 in
in

d o

m

cn o cj

o in »—

41 »— 41

1 — 1 d

ci* r—

co o co

cj co cj co

G> t—

I o

o 00
00 00

I CO I 00

O 00 O CJ

O r— O CJ

co 41
cj

ro 41 d 41

1 1 co

cl- O'* NfO

ino in co

CJ 41
1 m
in

1 00 1 o

d cr» d • —

o cr> co 01

co co co co

co -h
d- 00

00 co

CJ 41
1 d
CJ CJ

o in

m r—

CJ 41
1 d

cr> cj

CJ 41 • —

cj

o

1 — in in ci-

in co in co

O CO
co d

CJ CJ

o m

CJ CJ

Table 2 (continued)

486

Psyche

[September-December

O CO
CM •
I C\J

CO CO

co M-

LO LO

1

co co

r— CO C\J
I I

CO CO CO <4

co 10
1

CM CM

LO CO

co co

*4 CO
CO CO

I— CO
I

co

00 o

1

co

>4

co co

cr> 00

CTl CO

co c-

co cm 1—

o +1

I CO
CM LO

OO OO OO OO

OO O O

00 00

10

CO «4

CO O

d -H
I 10

CO CM
I— CM

C TV
CM CM
CM O

CM O

d -h

o

cti

>4 o

CM
co co

CO o

O -H o 41
I CTl l .—
1— CO CM CM

O +
I CTl
LO CTl
CM CM

O O

00
CC CO
CO o

co
co co
co o

CO
CO CM
CM O

i — 1 — co co

COCO CM CO COCO CO CM

coo *3- o coo coo

CO
CO CM
CO o

o •

CO
CO CM
CO O

I CM
«4 o
CM CO

o 4<
I CO
LO 00
CM CM

1 1 — 1 r-. 1 ■ — 1 co

CM CTl LOO CM CO CTl CO

CM CM CM CO CM CM CM CO

O O OO

o o o o

LO I —

d d

I 00
1 — l-~
LO CO

d d

O 41

1 co

CO o

co 10

.— 41 041 041 r— 41

L CM I 00 I LO I CTl
1 — CTl CO CM LO LO CTl CTl

ION LON LOCO r-'.CTl

I CO

co r-.
00 cn

00 CTl

d d

o o 00

00 00

I co

00 r—

CO CTl

I CM

co r-s
00 00

1—41 1 — 41 ■ H CM-jH

1 cm 1 co 1 10

CM CO CTl LO 00 *4 COLO

•— <4 o co cn.— co r-~

.— .— o

I—.— OO OO

cal i form cus 4-9 4-7 12-20 1.28-1.53 2.16-2.61 2.60-3.22

487

1974]

Coyle — Genus A liatypus

LO

CO r— r— .

00
OJ CO
ID 1—

O
CO -fl

CO LO
OJ CXI

OJ LO
OJ OJ

OJ -fl OJ -H
I CO 1 LO
IS CO O VO

^ LO LO VO

OJ

1 cr>
1— co

LO LO
OJ OJ )

00 -h

I CX»
VO CO

00 I—

r— r— VO I— VO r—

VO O'* O OJ O'* OJ

OJ OJ CO CO OJ CO

O OJ
CO CO

VO VO 00 00 o

^ LO Cj*

roo oj] lo o

i LO VO/ CO

I— 00

I

LO VO

00 OJ o

I

LO VO VO VO

O-* r— CO

OJ
VO vo

o

^ VO

vo cr*

I

LO LO

o

vo vo

Table 2 (continued)

488

Psyche

[September-December

*d"
r— CsJ
CM o

VO CVJ

1 CM r— CO /00 CM

r— 41
I CO
CO CO

I CsJ

o

r— CM

OO r— O

* T &

CM IO

r- -H
I to
CM VO
O O

I LO

r— CM

o o

o> cn
o o

•— 41 r- 41

I CM I <J\

co o co cn

O'* O CTV CX*

Or- OO

O 4i
I CM
O LO

o 41
I LO
O CM
CM CO

*d- O
O 4i

O *
1 CO

r- O

o o o o

00
LO CM
CO r-

00 o\

r— r— (OO CO
CO I — O CO

I o\
co

O'* LO

CO 41
I VO

co r^

LO O'*

1 co

cm

co LO

cn f

CM /O
1— LO *

VO CM
1 — *td-

O O

1— 41
1

cr*

cm co

o o

O 41
VO co

r- CO 1

vo

0 r- NO

00
CT> O

vo

nJTN

LO
O LO

CM
O CM

vo

00
CO CO

Z~^\

fo ^

sj «d- co lo

0 41 VO 41

< — «d-

10 41

CO LO
LO 41

cn 1
CM

, LO

vo 41

cn vo
LO 41

cm
co 41

LO vo
vo 4l

cm

LO 41

co csj
-H

0 CO O O

00

CM «d*

cm

LO O

lo O'*

LO CM

CT\

cn

0 CO 1— I—

0 *d* lo

CO co
co

0 oi
co co

co

^CM CMj

vo vo

O') «d-

LO LO

co *d-

CT) CM

\cmco/

CM r—
\^d~ LO^

CO LO

co d

VO CTt
\cp coj

CTv I—

I VO
cn o
co

vo cm

r— CO

I LO
. — CM
VO CO

SL/SW SW/PSS SW/PSL PSL/PSS AMD/AMS IMS/CMT IMS/PSS

489

1974]

Coyle — Genus Aluttypus

californious janus isolatus aquilonius gnomus gulosus thompsoni tvophonius erebus plutonis torridus

Psyche

[September-December

490

00 l~-~ <3- I-', o

O co o 10 o co

*3- CM CO ro C\l CM

s 00 in ^ lo

CM N I — CO 03 r-.

I— 00 CM CXI 1 — O

. — O CO CO LO

o in cn m in n

1 — O CM r— O O

M- M- in cc M-

1 — CO CM 1 — 1 —

CVJ

I— O O O O CM

•3- CO CO 00 ■— o

n in <f o co co lo co

Ml CM CO Ml CM CM r— Ml

O 00 Ml 1 — CD O') CO CD

O CD LO CO <31 in cf I —

CO 00 CM O O O CM CM

CM *3- CM <3- CD CT> r^.

Ml Ml 1 — CO CM 1 — r—

o>

OO.— OOO Or—

CM CO (O) O') CO (\J N CM N O CM >3 CO >3 CO LO r— CD 1m 1m O') LO

CD CD Im COr— CO Ml |M Ml !m O 1 — CDLnCO CO LO LO O LO CM t— r—

<3 CM CO COCMCMr— COCMCMCMr— OOCMCMOO 1— O O O O CM

LO Ml (MLOCMLOLOCDCMCOi — COCOOOOOCDCOLOLOCO r^.
D^r— CO r^COOlr— COCOCDCOCOCOLOOCO^l-CMCDLOCMr— r—

CO CM CM CM I — I — I — CM I — I — I — OOOCMi — OOOOOO O CM

Ml CO CO Ml r —

CM LO Ml O O CO

Ml- CM CO LO CO CO

N N W CO to
00 o 1 — co 00

r— Ml- CO CO CM O

co o o o r — co

CM N CO O CM Ml

1 — O CM CM O O

O) CD LO LO CO

O Ml" CM r- 1—

CM

1— O O O O CM

LO Ml- COCOCOi — LOOCOi — Ml" Ml" r — LO Ml" CM CO CO) CD LO Ml" CO

tM 1— O N O Mf CD 1 — LO LO CO CO CO CO 1 — COD^i — CD ML CM 1 — 1 —

CO CM CO COCMCMr— Ml- CM CO r- OOOCMr— OOOOOO O CM

Cm Ml- CM CO ID [X N CO r — COCOCOCmCOCMCMCOOCMCM i —

1— I'm LO CDCDCDOOCOCMCOLOI'^COrMCOCOt— COMlCMr— r—

COr— CM CMr— r— r— COr— CMr— OOOr— r— OOOOOO O

CD CO CD CM r-M 1m CO LO 1m 1 — CDCOCOCDCOMlCDOOCMOCD CM

1 — co mi- r— cnoocMr — cmcomicococococooimmIcmo r —

CO rr CM COr— CMr— COCMCMr— OOOr— r— OOOOOO Or—

cm 00 lo o co I'm 1 — 01 — r^OLor^Mj-or^LOCMCMLOLO co

oicdo coocMLncoor-r''.cncMLOcooor'.CMCMCocMr— r—

LO

Ml- W Mf MtCOCOr— LOCOCOCMOr— OCMCMOOr— OOO O CM

LO r — LO LO O
LO r — CO r— , — LO

LO CO Mf LO CO CO

CO CM CM r— Ml- Mf

IM Ml CT) CO 03 01

r— LO CM CO CM O

CO 1 — 1 — r — CO CO

CM LO CO CO 00 CO

r— O OO CM O O

CM r— CD 00 CM

t CO CM t — r —

CD

r— O O O O CM

r— CO O CO o 00

LO CO r O C OOO

LO CO Ml Ml CM CM I —

00 D 01 CO cl N r- O Cm
COr^COCMCMCOCOCM r—

COCMOOr— OOO O

_I _J _J X
Li_ CL. I— t—
O- Cl. CL Cl.

O <C

LU O _l 3
cl a. 00 00

o

00

00

CL

I 3 C/3 O CO

CO O' — I — I 2l

CL O C «=C <C

O 00

2: o

<C LlJ

1974]

Coyle — Genus A liatypus

49

Maps 2-3. Distribution of A liatypus species.

492

Psyche

[September-December

Map 4. Distribution of Aliatypus species. (Unidentifiable specimens
are immatures and females which cannot be assigned to any recognized
species.) Figures 39-40: Cheliceral teeth of Aliatypus females. (Ventral
view of left chelicera.) 39: A . calif ornicus, Soquel. 40: A. thompsoni,

paratype. Figures 41-42: Eyes of Aliatypus thompsoni (Dorsal view with
lateral border of carapace horizontal.) 41: male, holotype. 42: female,

paratype. Figures 43-44: Spinnerets of Aliatypus thompsoni. (Ventral
vieAV.) 43: female, paratype. 44: male, paratype.

1974]

Coyle — Genus Aliatypus

493

Figures 45-53: Whole body and carapace views of Aliatypus. 45-48:

A. calif ornicus. 45-46: male, Calaveras Reservoir. 47-48: female, Soquel.
49: A. gnomus female, paratype. 50-53: A. thompsoni . 50-51: males. 50:
Placerita Canyon. 51: Henninger Flats. 52-53: females 52: Placerita

Canyon. 53: Henninger Flats.

494 Psyche [September-December

Figures 54-77: Sternum, labium, and sigilla of Aliatypus species. 54-64:

males. 54: A. calif ornicus, Alum Rock Park. 55: A. janus, holotype. 56:
A. isolatus , paratype. 57: A. aquilonius, holotype. 58: A. gnomus, holo-
type. 59: A. gulosus, holotype. 60: A. thompsoni , paratype. 61: A. tro-
phonius, holotype. 62: A. erebus, holotype. 63: A. plutonis, Banning. 64:
A. torridus, holotype. 65-77: females. 65: A. calif ornicus, Montebello Rd.
66: A. janus, paratype. 67: A. isolatus, paratype. 68: A. aquilonius, para-
type. 69: A . gnomus, paratype. 70: A. gulosus, paratype. 71-73: A.
thompsoni . 71: paratype. 72-73: Eaton Canyon. 74: A. trophonius , para-
type. 75: A. erebus, Sonora. 76: A. plutonis, Banning. 77: A. torridus,
paratype.

1974]

Coyle — Genus Aliatypus

495

Figures 78-91: Aliatypus male pedipalps. (Retrolateral view of left
pedipalps.) 78-79: A. calif ornicus. 78: Coloma. 79: Montebello Rd. 80:
A. isolatus, holotype. 81: A. aquilonius, holotype. 82: A. gnomus, holotype.
83-84: A. janus. 83: Yosemite Nat’l Park. 84: Glenville. 85: A. gulosus,
holotype. 86: A . thompsoni, holotype. 87: A. trophonius, holotype. 88:
A. erebus, holotype. 89-90: A. plutonis. 89: Banning. 90: holotype. 91:
A. torridus, holotype. Figures 92-95: Tibia and metatarsus of leg I of
Aliatypus males. (Retrolateral view of left leg.) 92: A. calif ornicus,
Alum Rock Park. 93: A. thompsoni, holotype. 94: A. plutonis, holotype.
95: A. trophonius , holotype.

Figures 96-109: Aliatypus palpi. (Prolateral view of left palpus and

view of tip after palpus rotated 90° on longitudinal axis of distal part of
conductor.) 96-100: A. calif ornicus. 96: Montebello Rd. 97: Alum Rock
Park. 98: Coloma. 99-100: Mariposa. 101: A. gnomus, holotype. 102-105:
A. janus. 102: holotype. 103: Yosemite Nat’l Park. 104: Pinehurst. 105:
Glenville. 106: A. aquilonius , Briceland. 107-109: A. isolatus. 107: holo-
type. 108-109: Santa Catalina Mtns.

1974]

Coyle — Genus Aliatypus

497

Figures 110-120: Aliatypus palpi. (Same views as previous plate.)

110: A. gulosus, holotype. 111-113: A . thompsoni. Ill: Tehachapi Mtns.
112: holotype. 113: Henninger Flats. 114-115: A. erebus. 114: holotype.
115: Sonora. 116: A. trophonius, holotype. 117-118: A. plutonis. 117:
Palomar Mtn. 118: Banning. 119-120: A. torridus. 119: El Paso Mtns,
120: holotype.

498

Psyche

[September-December

121 122 123

Figures 121-148: Aliatypus seminal receptacles. (Dorsal view.) 121-131:

A . calif ornicus. 121: Montebello Rd. 122: Bates Creek. 123: Henry Cowell
Redwoods State Park. 124: Alum Rock Park. 125: Mt. Diablo State Park.
126: Sutter Buttes (immature). 127: Coloma. 128-129: Aukum. 130-13 1 :
Mariposa. 132-143: A. janus. 132: Briceburg. 133: Yosemite Nat’l Park.

134-135: Mammoth Lakes. 136-137: paratypes. 138-139: Pinehurst. 140-

141: Sequoia. 142: Woodlake. 143: Glenville. 144-146: A. isolatus. 144:

paratype. 145-146: Santa Catalina Mtns. 147-148: A. aquilonius. 147:

paratype. 148: Redway.

499

1974] Coyle — -Genus Aliatypus

Figures 149-171: Aliatyfus seminal receptacles. (Dorsal view.) 149-150:
A. gnomus paratypes. 151-154: A. gulosus. 151: Eaton Canyon. 152:

Sierra Madre. 153-154: paratypes. 155-171: A. thompsoni. 155: Kern-

ville. 156-157: Tehachapi Mtns. 158: Santa Ynez. 159: Squaw Flats.

160-161: Placerita Canyon. 162: Baldwin Hills. 163: Limekiln Canyon.
164-166: paratypes. 167: I. as Barras Canyon. 168: Hcnninger Flats. 169-
170: Eaton Canyon. 171: Sierra Madre.

Psyche [September-December

Figures 172-194: Aliatypus seminal receptacles. (Dorsal view.) 172-173:

A. trophonius paratypes. 174-187: A. erebtis. 174-175: paratypes. 176-177:
Wilbur Springs. 178-179: Nevada City. 180-181: Sonora. 182-183: Mari-
posa. 184-185: Shaver Lake. 186: Pinehurst. 187: Miramonte. 188-191:
At plutonis . 188-189: paratypes. 190-191: Banning. 192-194: A . torridus.

192: El Paso Mtns. 193: Yucaipa. 194: paratype.

A REVIEW OF THE AGYRTES (SILPHIDAE)
OF NORTH AMERICA*

By Stewart B. Peck

Department of Biology, Carleton University,

Ottawa, Ontario, Canada

Dr. John Lawrence and Dr. Roy Crowson pointed out to me that
the type specimen of Lendomus politus Casey (placed in Myceto-
phagidae by Casey) seemed to be a silphid in the genus Agyrtes.
This led me to study the Casey type and to review the species of
Agyrtes of North America as the first part in a projected series of
reviews of the silphid genera of North America.

The genus Agyrtes is Holarctic. Seven species are known and a
key to them is given by Hlisnikovsky (1964). Of these, two species
occur in and are limited to North America. The habits of the North
American species are unknown, but the European species occur in
forests under bark, in moss, and at rotting mushrooms.

Agyrtes longulus (LeConte)

Figures 1, 2, 8, 9, 10.

N ecrophilus longulus LeConte, 1859: 282. Lectotype here designated as
female in MCZ, LeConte colln. (no. 3148), bearing gold disc, seen.
Type locality: Table Mountain, below San Francisco, California, by
original designation.

Agyrtes longulus (LeC.). Horn, 1880:246. Hatch, 1957:8.

Lendomus politus Casey, 1924: 184. New Synonym. Holotype male in
USNM, Casey colln. (no. 37497), seen. Type locality; Anticosti Island,
Quebec, Canada. No other type data.

Diagnosis. Antennal club composed of four segments, first three
of club with rings of dense pubescense on inside apical margin, third
antennal segment appreciably longer than the second. Prothorax
conspicuously wider than long, less narrowed towards the front,
posterior angles distinct. Aedeagus in side view with pronounced
bend (fig. 10), paramere an elongated spatula (fig. 9).

Distribution. The species ranges from southern California north-
wards through the coastal mountains to British Columbia and south-
ern Alaska, and inland to Idaho. In addition to the type I have seen
the following material :

* Manuscript received by the editor December 17 , 1974.

501

502

Psyche

[September-December

1974]

Peck — A gyrtes

503

Canada. British Columbia. Creston. i5-xi-3i, G. Stace Smith
(abbreviated subsequently as GSS), 1 (CNCI) ; 29-0-32, GSS, 1
(CAS): n-xi-32, GSS, 1 under wood at 2000' (CAS); io-xi-33,
GSS, 1 on ground, 2000' (MCZ) ; 29-xi-33, GSS, 1 in flood debris
at 1750' (MCZ); i-xi-45, GSS, 1 (MCZ) ; 2i-xi~50, GSS, 3 on
snow (CAS); 24-xi-50, GSS, 1 on snow (CAS); 27-xi-50, GSS,
23 on snow (CAS); io-xii-50, GSS, 1 on snow (CAS); 1952,
GSS, 4 (FMNH) ; 26-iii-56, GSS, 1 (CNCI); i8-iv-56, GSS, 1
(CNCI). Queen Charlotte Islands. Massett, 10 (USNM) ; no
locality, 5 (CAS, CNCI, USNM). Salmon Arm, n-iv-28, HB
Leech, 1 (CNCI); i2-iv-35, H. Leech, 1 (MCZ); 2i-xii-35, SH
& O Leech, 1 on snow (CAS). Terrace, viii-1927, ME Hippsley, 1
(MCZ) ; ME Clark, no other data, 1 (MCZ). Vancouver, 28-i-3i,
HB Leech, 2 (CAS, CNCI); 26-iii-33, HB Leech, 1 flying, (Fall
colln. MCZ); no other data, 1 (LeConte colln., MCZ).

United States. Alaska. Sitka, 5 (USNM). California, Alameda
County, Berkeley, 22-ii-20, JO Martin, 3 (CAS). Glenn County,
Elk Creek, 17-1V-13, R Hopping, 1 on Pinus ponder osa (CAS).
Marin County, Mill Valley, 22-ii-26, Van Duzee, 1 (CAS) ; 8-X-51,
RE Leech, 1 at light (CAS); 9-L56, HB Leech, 1 (CAS). Mt.
Tamalpais, 23-V-09, Van Dyke, 1 (CAS). Taylorville, 28-XH-19,
JO Martin, 1 (CAS). Monterrey County, Carmel, 27-xii-i4, 1
(CAS). Plumas County, 2 (USNM). San Francisco County, San
Francisco, 8-i-n, Van Dyke, 1 (CAS). Santa Clara County, no
other data, 1 (CAS). Sonoma County, Eldridge, 17-xii-io, Van
Dyke, 2 (CAS) ; southern Sonoma County, 9-xii-iO, Van Dyke, 2
(CAS); 11-xii-io, 1 (CAS); 14-xii-io, Van Dyke, 3 (CAS). 29-
i-11, Van Dyke, 3 (CAS). Oregon. No other data, 1 (USNM).
Clackamas County, Colton, iv-35, ES Ross, 1 (CAS) ; Yamhill
County, 2-iii-34, ES Ross, 1 (CAS). Washington. King County,
Seattle, 3-V-14, 2 (Fall colln., MCZ) ; 28-H-98, 1 (CAS) ; no other
data, 3 (CAS). Thurston County, Olympia, no other data, 1 (Fall
colln., MCZ).

Hatch (1957) also mentions that the species occurs in northern
Idaho.

Biology. Virtually nothing is known of the habits of the species
or of its life cycle. The seasonality data show that it is most active
in the winter months. The species has been taken most often on
snow but also in flood debris, and under chips of live oak. It is
presumably associated with forested habitats.

Notes on synonymy. The type of Lendomus politus Casey was
examined and dissected. It was found to be identical, within reason-

504

Psyche

[September-December

able limits of variation in the shape of the aedeagus, with Agyrtes
longulus. Since a number of beetles in eastern Canada have been
introduced from Europe (Lindroth, 1957) there was the possibility
that the Agyrtes of Casey might be a species introduced from Europe.
It was compared with the European A. castaneus (F.) and the
aedeagus drawings of Hlisnikovsky (1964) and was found to be
distinctly different.

Since Casey’s specimen is the only Agyrtes known from eastern
North America, I think that an error was made in the labeling of
Casey’s Agyrtes , and we may continue to believe that the genus
Agyrtes and the species A. longulus occur only in the coastal and
interior mountains of western North America.

Agyrtes similis Fall
Figures 3, 4, 5, 6, 7.

Agyrtes similis Fall, 1937; 29. Holotype male in MCZ, Fall colln. (no.

24015), seen. Type locality: Pasadena, California.

Diagnosis. Antennal club composed of five segments, first four of
club with rings of dense pubescense on inside apical margins, third
antennal segment only a little longer than the second. Prothorax
wider than long but more elongate and more gradually narrowed
toward the front, posterior angles rounded. Aedeagus in side view
relatively straight (fig. 7), paramere a rounded spatula (fig. 6).

Figures 5-10. 5. A. similis aedeagus, dorsum, Pasadena, California.

i. A. similis paramere, setae omitted. 7. A . similis aedeagus, lateral view.
8. A. longulus aedeagus, dorsum, Mill Valley, Marin Co., California. 9.
A. longulus paramere, setae omitted. 10. A . longulus aedeagus, lateral
view.

1974]

Peck — A gyrtes

505

Distribution. The species is known only from the coastal ranges
of central and southern California. In addition to the type I have
seen the following specimens :

California. Alameda County, Alameda, i-iv-90, 1 (Koebele colln.,
CAS). Los Angeles County. Pasadena, Jan., 1; Feb., 1; Mar., 1;
(Fenyes Colln., CAS). Pomona, mountains, 15-XU96, 1 (Fall
colln., MCZ) . “Cala,” no other data, 2 (Horn colln. in MCZ).

Biology. Nothing is known of the habits or life cycle of this
beetle. The collection records show that it is active only in the
winter months. It is presumably associated with forested habitats.

Notes. There is a remarkable similarity in the dorsal view of the
aedeagus and of the paramere of A gyrtes bicolor Lap. of central
Europe (as illustrated by Hlisnikovsky, 1964) and of A. similis, but
other characters in the key of Hlisnikovsky seem to separate the
species. The aedeagus of A. bicolor in side view is not illustrated.
I have not studied material of A. bicolor, although I have seen the
Hlisnikovsky collection in Prague.

Acknowledgements

I wish to thank the following for having made available for study
material in the collections under their care: M. J. Campbell, Cana-
dian National Collection of Insects, Ottawa (CNCI) ; H. Dybas,
Field Museum of Natural History, Chicago (FMNH) ; J. M.
Kingsolver, U. S. National Museum, Systematic Entomology Lab-
oratory (USNM). J. F. Lawrence, Museum Comparative Zoology,
Harvard University (MCZ) ; H. B. Leech, California Academy of
Sciences, San Francisco (CAS). H. F. Howden and J. F. Lawrence
are thanked for reviewing the manuscript. L. E. C. Ling of Carle-
ton University took the SEM photographs.

Literature Cited

Casey, T. L.

1924. Memoirs on the Coleoptera, XI, 346 pp.

Fall, H. C.

1937. Miscellaneous notes and descriptions (Coleoptera). Can. Ent.,
69: 29.

Hatch, M. H.

1957. The beetles of the pacific northwest. Part II: Staphyliniformia.
Seattle: Univ. Wash. Pub. Biol., 16, 384 pp.

Hlisnikovsky, J.

1964. Zur Kenntnis der Gatung Agyrtes Lap. (Coleoptera, Silphidae,
Agyrtini) . Reichenbachia, 2: 275-278.

506

Psyche

[September-December

Horn, G. H.

1880. Synopsis of the Silphidae of the United States, Trans. Amer.
Ent. Soc., 8: 219-322.

LeConte, J. L.

1859. Additions to the Coleopterous fauna of northern California and
Oregon. Proc. Acad. Nat. Sci. Philadelphia, 281-293.

Lindroth, C. H.

1957. The faunal connections between Europe and North America.
New York, Wiley and Sons, 344 pp.

APPLICATION OF THE TRANSMISSION ELECTRON
MICROSCOPE TO THE EXAMINATION OF
SPIDER EXUVIAE AND SILK*

By Rainer F. Foelix

Lehrstuhl fur Zellphysiologie, Ruhr-Universitat Bochum
D 4630 Bochum, West Germany

In recent years the scanning electron microscope (SEM) has been
used fruitfully to study the fine structure of many arthropods,
including spiders. The advantage of the SEM lies in the simple
method of preparation — no embedding and sectioning has to be
done — and its large depth of focus, which yields almost three-
dimensional views of surface structures. Drawbacks of the SEM
are a limited resolution (1 50-200 A) and, as well, lack of availability
to most researchers. The transmission electron microscope (TEM)
has a much better resolution ( < 10A) than the SEM, but requires
objects of less than o. 1-0.2 /x thickness to allow penetration of the
electron beam.

It was found that by simply mounting parts of spider exuviae
(preferably of early instars) on Formvar-coated copper grids, details
of hairs and claws can be seen with the TEM. If a leg tip of an
exuvia is placed on a water droplet on the Formvar membrane, it will
firmly adhere to the membrane after the water has evaporated. The
preparation can then be viewed immediately in the TEM. Better
stability and more contrast may be achieved by shadowing the speci-
men at an angle of 40-50° with a metal (e.g. copper). The very
delicate scopula hairs (Fig. 1-4) are especially well-suited for exam-
ination with the TEM, but other hairs and bristles can also be easily
surveyed and classified. For instance, the open tip of chemosensitive
hairs (Foelix, 1970) is convincingly demonstrated with the TEM

(Fig. 5).

Spider silk, especially the extremely fine threads of cribellate silk
(Fig. 6), has been studied with the TEM before (Lehmensick and
Kullman, 1957; Friedrich and Langer, 1969). It should be empha-
sized here that shadowing with copper yields much more stable prep-
arations. Ecribellate silk is often too thick (1-2 /x) to be examined
with the TEM, but accurate measurements of the diameter of various
threads can be performed (Fig. 7). The substructure of the sticky

"* Manuscript received by the editor December 10, 1974.

507

508

Psyche

[September-December

1974]

Foelix — Spider Silk

509

spiral of orb weavers, however, can be revealed : The glue substance
partially disintegrates under the electron beam and the two axial
threads become clearly visible (Fig. 8).

It is hoped that this simple technique will be usefully applied to
morphological as well as taxonomical problems.

References Cited

Foelix, R. F.

1970. Chemosensitive hairs in spiders. J. Morph. 132: 313-334.
Friedrich, V. L. and R. M. Langer

1969. Fine structure of cribellate spider silk. Amer. Zool. 9: 91-96.
Lehmensick, R. and E. Kullmann

1957. Feinbau der Fiiden einiger Spinnen. Zool. Anz. Suppl. 19: 123-
129.

Explanation of Figures on Opposite Page

Fig. 1. Leg tip of Philodromus aureolus (nymph 2) as seen with the
SEM. Scopula hairs (sc) surround the combed main claw. 1400 X. Inset:
Scopula hairs bear fine extensions with terminal ‘end-feef. 4200 X- (Cour-
tesy of Dr. W. Gnatzy). Fig. 2. The ‘end-feet’ of a scopula hair of
Philodromus seen with the TEM. 7000 X. Fig. 3. Leg tip of Alopccosa
accentuata (nymph 2) showing one main claw (M), chemosensitive hairs
(c), mechanosensitive hairs (m) and a scopulate hair (sc). TEM, 630 X-
Fig. 4. Scopulate hair of Micrommata virescens. Note the small end-feet.
TEM, 4200 X. Fig- 5. Tip of a chemosensitive hair of Philodromus. Note
the terminal pore opening (arrow). TEM, 12600 X- Fig. 6. Cribellate
silk from Amaurobius ferox. The diameter of single threads is less than
200 Little knobs become accentuated by the metal shadowing. TEM,
28000 X- Fig. 7. Ecribellate silk from the silken cell of a Heliophanus.
A mesh-work of fine fibers (0.1 n) underlies several stronger fibers (0.5 n).
TEM, 2800 X, unshadowed. Fig. 8. Sticky spiral thread from Zygiella
x-notata. The peripheral glue substance has precipitated around the double
axial threads (arrow). TEM, 630 X-

PLASTRAL RESPIRATORY DEVICES IN ADULT
CRYPHOCRICOS ( NAUCORIDAE : HETEROPTERA) *

By Margaret C. Parsons and Rosemary J. Hewson

Department of Zoology
University of Toronto
Toronto, Ontario M5S 1A1

Introduction

Adult Cryphocricos live in very rapid, well-aerated streams, under
stones and gravel at depths of from 6 to 24 inches (Fritz Plaumann,
personal communication ; John Polhemus, personal communication ;
Randall Schuh, personal communication). Under these conditions it
would be difficult for these insects to come to the water surface for
atmospheric air, as do “slow-water” Naucoridae and most other
aquatic Heteroptera, and neither Polhemus (personal communication)
nor Schuh (personal communication) have observed them to do so.

Even if it were possible for Cryphocricos to obtain atmospheric air,
the insects lack a means of storing large amounts of it for underwater
respiration. Aquatic Heteroptera with “air-bubble” respiration carry
much atmospheric air under their long forewings and trapped by long
hairs on the ventral surfaces of their bodies. Long-winged Crypho-
cricos, however, are very rare. Most individuals lack hindwings and
have forewings which reach only to the third abdominal segment.
The reduced subalar space on the pterothorax is too small to hold
any appreciable amount of air (Parsons 1974). Thorpe (1950)
concluded, from examination of a dried specimen of Cryphocricos
sp., that it is “a bubble-carrier rather than a true plastron insect”
and has “a substantial air film”. Polhemus, however, examined C.
hungerfordi in the field and observed no such air layer except on the
six pairs of small ventral abdominal sense organs (Polhemus, per-
sonal communication). Parsons (1974) observed a very thin air
layer on the ventral surfaces of the thorax and abdomen in formalin-
preserved C. barozzii, but this layer is visible only under the stereo-
scopic microscope and is usually hidden by the debris which covers
the insect.

Young’s (1944) suggestion that Cheirochela (Naucoridae) sur-
vives by cutaneous respiration appears highly dubious, and it is un-
likely that Cryphocricos uses this method to any great extent. The

*Manuscript received by the editor February 6, 1975

510

1974]

Parsons & Hewson — Cryphocricos

51 1

exoskeleton is quite thick, and the relatively large size of the insect
( C . hungerfordi is 8. 5-9.5 mm long; C. barozzii is 10-13 mm long)
produces an unfavorable ratio of volume to surface area.

Among the aquatic Heteroptera, true plastral respiration has thus
far been experimentally demonstrated only in Aphelocheirus (Thorpe
and Crisp 1947a, 1947b; Thorpe 1950). Whether this insect is a
naucorid (Usinger 1956) or represents a separate family, the Aphelo-
cheiridae (Parsons 1969) is debatable. Aphelocheirus possesses a
very thin, permanent air layer, or “plastron”, which is retained by
short, densely-packed hairs on most of the dorsal and ventral surfaces
of the body. The plastron is exposed to the water and acts as a gill,
obtaining enough dissolved oxygen from the water to make the in-
sect independent of atmospheric oxygen (Thorpe and Crisp 1947a;
Thorpe 1950). Cryphocricos and Aphelocheirus share such morpho-
logical similarities as reduced forewings, vestigial hindwings, lack of
a large air store, and paired abdominal pressure receptors of a type
not yet reported in any other Heteroptera ( Aphelocheirus , Thorpe
and Crisp 1947c, Larsen 1955; Cryphocricos , Thorpe 1950, Parsons
1974). These similarities suggest that Cryphocricos , like Aphelo-
cheirus, has a plastral type of respiration. Hinton (1969a, p. 198)
has remarked that “Por ejemplo, entre insectos adultos hay por lo
menos cuatro origenes independientes de respiracion de plastron den-
tro de la familia Naucoridae: Aphelocheirus , Cryphocricos , Heleo-
coris, e Idiocarus” . He included a photomicrograph of thoracic
plastral hairs in Heleocoris but did not further elaborate on his
statement.

Although the presence of plastral respiration would be best dem-
onstrated by both morphological and physiological studies, live speci-
mens of Cryphocricos were not available for experimentation. Our
investigation, like Hinton’s (1969b) study on aquatic beetles, was
therefore confined to morphology. Its purposes were ( 1 ) to discover
whether the exposed surfaces of the body possess hydrofuge devices
which might retain a thin plastral air layer, and (2) briefly to com-
pare any such structures with those of other insects in which plastral
respiration is believed to occur.

Materials and Methods

Species used :

Cryphocricos barozzii Signoret (preserved in formalin; collected
in Brazil by F. Plaumann; two specimens)

C. hungerfordi Usinger (preserved in alcohol; collected in Mexico
by J. T. Polhemus ; one specimen )

512

Psyche

[September-December

Aphelocheirus aestivalis Fabricius (preserved in Bourn’s fluid;

collected in England by D. T. Crisp; two specimens)

The specimens were first dissected in 80% alcohol under a stereo-
scopic microscope. They were cut midsagittally with a razor blade
and the ventral and dorsal surfaces of one or both halves were sep-
arated along the lateral edge. On one specimen of Aphelocheirus a
small section of the metathoracic episternum was removed so that the
cut edge could be examined. A major obstacle was the presence of
large amounts of silt and other debris on the external surfaces of
all three species. Treatment with weak solutions of either potassium
hydroxide or Sparkleen detergent failed to remove it. Somewhat
better results were obtained by scraping small areas with watch-
maker’s forceps.

The specimens were then mounted with double-sided tape on
aluminum stubs, freeze-dried, and coated, in an Edwards vacuum
evaporator, Model 4, with a layer of gold which was approximately
200-400 A thick. They were examined in a Cambridge Stereoscan,
Model 2A, and photographed with a Singer camera, using Ilford
120 roll film.

Observations

Ventral Surface
( 1 ) Plastral Hairs

In both species of Cryphocricos most of the ventral surface is
covered with densely-packed hairs (Fig. 3). They occur on the pro-

Fig. 1. Cryphocricos barozzii; ventral view, legs removed. Stippling
on fourth through sixth abdominal sternites indicates area of microtrichia
(Fig. 11) ; stippling on ventral abdominal paratergites, left side of figure,
indicates area of short leaf-like setae (Figs. 6-8). Longer leaf-like setae
occur on the six pairs of abdominal sense organs. Most of the Ventral
surface is covered with plastral hairs (Figs. 3, 4).

Abbreviations: EM, epimeron ; ES, episternum; FW, forewing; LS,

longitudinal sulcus; MCI, metacoxal indentation on abdomen; PT, ventral
abdominal paratergite; SO, abdominal sense organ; SP, abdominal spiracle;
ST, sternite. Arabic numerals indicate abdominal segments; Roman nu-
merals indicate thoracic segments.

Fig. 2. Cryphocricos barozzii; dorsal view, right forewing removed,
revealing smooth subalar surfaces on right side (even stippling). Other
surfaces are roughened. For distribution of plastral hairs, see text.

Abbreviations: TI, TII, Till, thoracic terga ; Tl, T2, T3, abdominal
terga.

2 mm

1974]

Parsons & Plewson — Cryphocricos

513

Z

LlI

(0

_J

S07
PT 8

5H

Psyche

[September-December

thoracic epimeron and sternum (Fig. i, EM I, ST I), the meso-
thoracic epimeron, episternum, and sternum (EM II, ES II, ST II),
the metathoracic episternum (ES III), and all seven ventral ab-
dominal paratergites (PT 2-PT 8). They also cover all of the third
and seventh abdominal sternites and all but the anteromedial por-
tions of the fourth through sixth sternites (ST 4-ST 6).

The close spacing of the hairs makes it difficult to see their bases
and to measure their length. Their visible portions are at least 6 /xm
long and from 0.25 to 0.45 /xm in diameter on the thorax and para-
tergites (Fig. 4). They are considerably shorter and wider on the
abdominal sternites; on a sagittal cut through the fifth sternite,
where the entire length of a few hairs was visible, they were only
2.7 /xm long but 0.65 /xm in diameter.

Comparison of the thoracic and paratergal hairs with the plastral
hairs of Aphelocheirus (Fig. 5) shows similarities which strongly
suggest that those of Cryphocricos (Fig. 4) are also plastral devices.
The most striking similarities are ( 1 ) the presence of small nodules
on the basal portions of the hairs, (2) the densely-packed arrange-
ment of these hairs, and (3) the posterior bending of their apical
portions, which lack nodules. Using the light microscope and trans-
mission electron microscope, Thorpe and Crisp (1947a) observed the
bending of the hairs in Aphelocheirus but did not see the nodules.
The bending appears to be more pronounced in both specimens of
Aphelocheirus than in the three specimens of Cryphocricos. In both
genera the hairs tend to adhere to each other in clumps, making the
direction of the bending difficult to detect in surface view (Fig. 3) ;
the clumping is probably an artifact.

Fig. 3. Cryphocricos barozzii; surface view of apical parts of plastral
hairs on metathoracic episternum. More basal parts of hairs, with nodules,
visible at lower left. Clumping of tips is probably an artifact. Left = pos-
terior, right — anterior. Scale line = 2 /xm.

Fig. 4. C. barozzii; clump of plastral hairs on fifth ventral abdominal
paratergite. Leaf-like setae, which concealed these hairs, were scraped
off during preparation of specimen. Bases of hairs were concealed by tips
of surrounding hairs. Note posterior (to right) bending of apices and pres-
ence of nodules on proximal parts of hairs. Scale line = 1 /xm.

Fig. 5. A phelocheirus aestivalis ; cut edge of metathoracic episternum,
showing full length of plastral hairs. Note posterior (to right) bending
of apices and presence of basal nodules. Scale line = 1 /xm.

1974]

Parsons & Hewson — Cryphocricos

515

5 1 6

Psyche

[September-December

The thoracic plastral hairs of A phelocheirus are approximately
5-6 /x m in length, as Thorpe (1950) also noted. They are thus
somewhat shorter than those of Cryphocricos , but appear to be com-
parable to the latter in diameter. Their bases are somewhat thicker
(approximately 0.4 /xm) than their tips (approximately 0.3-0.35 /xm).
In both genera adjacent hairs seem to l;e very clcse to each other.
The nodules may help to hold them apart, allowing room for air to
circulate between them. Also, because of the thickness of the gold
coating (0.02-0.04 /xm), the hairs probably appear, under the scan-
ning electron microscope, somewhat thicker, and the intervening
spaces smaller, than they actually are. By means of the light and
transmission electron microscopes, Thorpe and Crisp (1947a) esti-
mated the space between adjacent hairs on the abdominal sternites of
A phelocheirus to be approximately twice as great (0.4 /xm) as the
diameter of the tip of a single hair (0.18-0.25 /xm). The discrepancy
between their observations and ours may be due, in part, to the dif-
ferent techniques employed. It is also possible that the hairs are less
densely packed on the abdominal sternites than on the thorax of
A phelocheirus.

The hairs of both Cryphocricos and A phelocheirus are much
shorter, narrower, and more densely packed than those which cover
the ventral surface of the abdomen of Pelocoris fernoratus , a “slow-
water” naucorid with air-bubble respiration (Parsons 1974). A brief
observation of one specimen of Pelocoris revealed that the hairs on
the second abdominal paratergite lack basal nodules and are 30-40
/xm long, 2-3 /xm wide at the base, and approximately 10 /xm apart.

(2) Leaf -like Setae

Leaf-like setae occur on the ventral paratergites of the abdomen
(Fig. 1, light stippling lateral to LS). They extend all the way to
the lateral edge of the body on the second abdominal paratergite, and
nearly to the lateral edge on the more posterior paratergites. On the
second paratergite (PT 2) a narrow anteromedial strip adjoining
the metathoracic episternum (ES III) and the metacoxal indentation
(MCI) lacks leaf-like setae (Fig. 6). This “nonfoliated” region is
larger in Cryphocricos hungerfordi than in C. barozzii. The second
abdominal spiracle (Fig. 1, SP 2) and sense organ (SO 2), like
those of the other abdominal segments, lie in the “foliated” region.

In C. barozzii the flattened setae are widest (8-20 /xm) near their
bases and are 20-40 /xm long. In C. hungerfordi a smaller insect,
they are approximately 75% this size. They appear to be socketed
(Fig. 8) and are arranged somewhat like shingles, in closely packed,

1974]

Parsons & Hewson — Cryphocricos

517

Fig. 6. Cryphocricos barozzii; portions of left raetathoracic episternum (upper left) and second ventral
abdominal paratergite, showing arrangement of leaf-like setae on latter. Note absence of leaf-like setae on por-
tion of paratergite immediately posterolateral to episternum. Top anterior, bottom =- posterior. Scale line
^ 50 fJ-m. FIG. 7. C. barozzii; several leaf-like setae on third ventral abdominal paratergite. Overlapping
arrangement largely conceals underlying plastral hairs, which are visible at right. Scale line = 5/*m.

5 1 8

Psyche

[September -December

somewhat irregular rows, each row overlapping the one posterior to
it (Figs. 6, 7). The tapered tip of each seta points posteriorly and
somewhat laterally on the body except in the spiracular regions, where
those immediately lateral to the spiracles point posteromedially. On
the eighth paratergite the setae are fewer and farther apart and do
not overlap.

Under the stereoscopic microscope the ventral paratergites, unlike
the “nonfoliated” regions of the body, have a faintly golden sheen.
The sheen is visible when specimens are immersed in alcohol, which
would remove any adherent air layer. It is probably the result of
light diffraction by the fine longitudinal ridges on the ventral surfaces
of the flattened setae (Figs. 7, 8).

In their shape and overlapping arrangement the setae resemble the
plastral devices of Phytobius (Coleoptera; Thorpe and Crisp 1949).
Unlike the latter, however, they do not bear plastral hairs on their
external surfaces. The plastral hairs arise from the exoskeletal sur-
face beneath the leaf-like setae and are largely concealed by them
(Fig. 7) but were clearly visible in regions where the setae had been
scraped off with watchmaker’s forceps (Fig. 4). Although the setae
do not completely cover the ventral abdominal spiracles they overlap
so much of their margins that we could not observe spiracular struc-
ture or determine whether or not they possess radiating “rosettes”
similar to those of the second through seventh abdominal spiracles of
Aphelocheirus (Thorpe and Crisp 1947a).

The second through seventh abdominal sense organs (Fig. 1, SO 2-
SO 7) are composed of leaf-like setae which are two to three times
longer than those covering the rest of each paratergite (Fig. 9).
Most of these elongated setae project more laterally than the shorter
ones, and have blunter tips and more pronounced longitudinal ridges

Fig. 8. Cryphocricos hungerfordi ; detail of one leaf-like seta on fifth
ventral abdominal paratergite. Note longitudinal ridges and tapering tip,
which points posterolaterally (lower left). Surrounding setae were rubbed
off, revealing tips of underlying plastral hairs and apparently socketed
base of seta. Scale line = 2 ^m.

Fig. 9. C. barozzii ; sense organ on third ventral abdominal paratergite
Sense organ is composed of elongated leaf-like setae. Tips of shorter
leaf-like setae, covering rest of paratergite, are visible at bottom and
upper left of figure. Right = lateral, bottom = posterior. Scale line =
30 /im.

Fig. 10. C. barozzii; detail of Fig. 9, showing longitudinal ridges on
elongated leaf-like setae. Scale line = 2 j^m.

1974]

Parsons £sf Hewson — Cryphocricos

519

520

Psyche

[September-December

(Fig. io). A considerable quantity of air is trapped among these
overlapping, elongated setae. On living specimens the sense organs
are the only parts of the body on which air can be seen with the
naked eye (Polhemus, personal communication), and the air on these
regions is easily seen with the stereoscopic microscope.

In A phelocheirus similar sense organs, which are probably pressure
receptors, occur only on the second abdominal segment (Thorpe and
Crisp 1947c). Like those of Cryphocricos they are composed of
flattened, elongated setae, approximately 75 /mi long, which point
posterolaterally and overlap each other. The setae appear to be longi-
tudinally ridged, although this could not be determined with certainty
because of the large amount of debris which covered them. Leaf-like
setae were not observed on any other region of the body of A phelo-
cheirus.

( 3 ) M icrotrichia

O11 the anteriormost portions of the fourth through sixth abdomi-
nal sternites (Fig. 1, ST 4-ST 6) plastral hairs occur only laterally ;
near the midline they are lacking. The only structures observed in
this hairless region are short, blunt microtrichia, approximately 1 /mi
wide at the bases and 2 /mi long, which project posteriorly. They are
arranged in widely-spaced groups of from two to six. Each group of
microtrichia arises from the slightly raised, curved, posterior edge of
a roughly semicircular region of the cuticle. The microtrichia lack
sockets and lie nearly parallel to the surface of the cuticle.

More posteriorly on the fourth through sixth sternites, but not as
far as the posteriormost edge, the microtrichia are more numerous
per group, longer (up to 3 /mi), and occur in more closely-spaced
groups (Fig. 11). Each group consists of 5-15 microtrichia and
these, along with the region of smooth cuticle to which they attach,
form a fringed, semicircular unit. The units are arranged like shin-
gles. On this central part of the sternite, unlike the more anterior
region, hairs appear to be present between adjacent rows of micro-
trichia. Since only the tips of these hairs are visible, we could not
determine whether they are similar to the short plastral hairs which
cover the lateral and posterior regions of the fourth through sixth
sternites; these regions lack microtrichia (Fig. 1, unstippled portions
of ST 4-ST 6) .

(4) Fringed Setae (Fig. 12)

Fringed setae may occur either singly or in large numbers. They
arise from sockets in the cuticle and project posterolaterally. A trails-

1974]

Parsons Plewson Cryphocricos

521

verse section through a fringed seta near its base would appear as a
nearly complete circle, with the open end facing away from the cuti-
cle. Towards the tips the setae are wider, more flattened, and would
appear as more widely open arcs in transverse section. Their outer
surfaces bear longitudinal grooves corresponding to pronounced
ridges on their inner surfaces. The apical portion of each of these
ridges subdivides into several short projections which give the seta a
fringed appearance. The sizes of the fringed setae vary considerably ;
on the lateral edges of the ventral paratergites their length ranges
from 12 to 38 fim, and their diameter from 6 to 15 ftm.

Single fringed setae occur on nearly all parts of the ventral sur-
face, but appear to be less common on the prothorax and the medial
portions of the abdominal paratergites than on other regions. Large
concentrations of them occur along the lateral edges of the third
through eighth ventral paratergites. On the second paratergite the
leaf-like setae extend all the way to the lateral edge, and there are
few fringed setae. On the most posterior segments, however, the ex-
treme lateral portions of the paratergites bear fringed rather than leaf-
like setae. They occur in largest numbers on the edges of the fifth,
sixth, and seventh abdominal segments and become increasingly less
numerous on each of the more anterior ones. The eighth paratergite
bears fewer of them than the seventh.

In Cryphocricos hungerfordi a narrow strip bearing only plastral
hairs separates the leaf-like and fringed setae, and the latter, which
occur in a single row, are fewer and farther apart than in C. harozzii.
The two types of setae are less clearly separated in C. harozzii, whose
fringed setae are more numerous, closer together, and in several
poorly-defined rows (Fig. 12). Their dense distribution on the lateral
edges of the fifth through seventh paratergites in this species sug-
gested, at first, that they might be hydrofuge devices. On the para-
tergites of C. hungerfordi , however, they appear to be too far apart
to function in this way. Their sporadic occurrence on nearly all the
exposed surfaces of the body suggests that they have some other func-
tion, probably a sensory one.

Dorsal Surface

The dorsal surface of the body was studied only briefly in one
specimen of each species of Cryphocricos. In both specimens this
surface is covered with considerably more silt and debris than the
ventral one. The detritus clings to all the parts of the dorsum which
are exposed to the water (Fig. 2, left side) ; these regions have a
roughened appearance under the stereoscopic microscope.

522

Psyche

[September-December

The surfaces which are concealed by the shortened forewings, how-
ever, are relatively free of debris. Except for the third abdominal
tergite (Fig. 2, T3) these regions appear smooth under the stereo-
scopic microscope (Fig. 2, light stippling on right side). The smooth
terga and the inner (ventral) surfaces of the forewings lack fringed
setae and are covered with densely-packed hydrofuge hairs. Those
on the metatergum (T III) and second abdominal tergite (T 2),
which were examined in more detail, possess nodules basally, are at
least 5 [im long, and have diameters similar to those of the plastral
hairs on the ventral surface of the body. The hairs on the inner
surface of the forewing are considerably shorter (approximately
1.5-2 fim in length) than those on the second abdominal tergite and
metatergum. Basal nodules were observed on these alar hairs only
in the region of the forewing which is externally visible in ventral
view (Fig. 1, FW) ; here the hairs are longer, but since their tips
appear to coalesce their height could not be estimated.

The surfaces which are exposed to the water are covered with
small, wart-like, cuticular protuberances which give them a roughened
texture. The raised surfaces of these protuberances, which are ap-
proximately 20-35 fim in diameter and one to two times this distance
apart, lack hairs. Each “wart” bears a small central indentation con-
taining a curious fringed papilla, possibly a sensory device. The large
spaces between the protuberances were, unfortunately, covered with
debris on all of the exposed abdominal tergites. The anterior third
of the third abdominal tergite, however, is overlapped by the posterior
edge of the forewing, which largely protects it from debris. In this
region, densely-packed hairs, at least 5 fim long and 0.3 to 0.4 /xm
wide at the tips, cover the areas between the cuticular “warts”.
Similar hairs occur between the protuberances on the exposed portion
of the mesothoracic tergum (Fig. 2, T II). Quite probably hydro-
fuge hairs are present between the protuberances on the prothoracic
and fourth through seventh abdominal terga, but are hidden beneath
the debris. Occasional fringed setae occur on all of the exposed dorsal
surfaces of the body, and are especially numerous along the lateral
edges of the abdominal terga and in the abdominal intersegmental
sulci.

Discussion

Although in most areas the fringed setae appear to be too widely
spaced to retain an air layer, at least two of the other three types of
fine structures on the cuticular surface of Cryphocricos appear to be

1974]

Parsons & Hewson — Cryphocricos

523

plastral devices. The densely-packed hairs on the ventral surfaces of
the thorax and abdomen strongly resemble those of Aphelocheirus ,
which has plastral respiration (Thorpe and Crisp 1947a) and differ
from those of Pelocoris, which has “air-bubble” respiration (Parsons
1974). The microtrichia in the central portions of the fourth
through sixth abdominal sternites have a shingle-like arrangement
which strongly resembles that of the plastral devices on the sternites
of T orridincola (Coleoptera, Hinton 1969b) ; these microtrichia,
along with the hairs between them, probably also retain a thin layer
of air.

The function of the leaf-like setae on the paratergites is more
problemmatic. They may serve merely to protect the underlying
plastral hairs from becoming too clogged with debris to allow the
adherent air layer to carry on gas exchange with the water. Although
they may, indeed, do this, it is unlikely that it is their sole function,
since they are absent in other regions with plastral hairs. In Aphelo-
cheirus, which is also covered with debris, leaf-like setae occur only
on the second abdominal sense organs, but the experiments of Thorpe
and Crisp (1947a, 1947b) have shown that the insect is able to
maintain efficient plastral respiration.

The presence of large amounts of air among the elongated leaf-
like setae on the sense organs of Cryphocricos suggests that the
shorter ones on the rest of the paratergites are similarly capable of
retaining air, although in smaller and less visible amounts. They
may well act in conjunction with the underlying plastral hairs, in
much the same way that long and short hydrofuge hairs hold a
“macroplastron” external to a “microplastron” in some aquatic Cole-
optera such as Hydrophilus piceus (Thorpe and Crisp 1949). Like
the long hairs which retain the “macroplastron” in the latter, the
leaf-like setae may, under increased hydrostatic pressure, help to
protect the thin layer of air, which is trapped in the underlying short
hairs, against water entry. When the insect dives to regions of in-
creased pressure the posterolateral slope of the setae may increase,
so that the setae become pressed together and form a barrier against
water penetration. Their presence only on the paratergites, and their
absence in other regions, may be connected with the need to protect
the second through eighth abdominal spiracles, which lie on the sur-
faces of the paratergites, against flooding. The three more anterior
pairs of spiracles (mesothoracic, metathoracic, and first abdominal)
are concealed within invaginations of the exoskeleton and are thus
more protected than the posterior ones against the entry of water.

524

Psyche

[September -December

Thorpe and Crisp (1947c) have described how the elongated
setae on the abdominal sense organs of A pKelocheirus become com-
pressed under increased pressure, stimulating shorter sensory hairs
which are interspersed among them, and thus warning the insect
against diving to dangerous depths. The elongated leaf-like setae on
the six pairs of abdominal sense organs of Cryphocricos probably
have a similar function, and the shorter leaf-like setae on the rest
of each paratergite may have evolved from these longer ones.

The efficiency of the plastron in Cryphocr'cos cannot be deter-
mined without experiments similar to those which Thorpe and Crisp
(1947a, 1947b) performed on A phelocheirus. Nearly all of the
ventral surface of Cryphocricos , and at least a part of the exposed
dorsal surface, bear apparent hydrofuge devices. If these devices do,
indeed, retain a thin, permanent, plastral air layer, a considerable
surface area of air is exposed to the water and is capable of gas ex-
change with the latter. Dissolved oxygen could pass directly to the
mesothoracic, metathoracic, and second through eighth abdominal
spiracles, all of which face portions of the ventral plastron. Although
the dorsal subalar space is not exposed to the water, it communicates
with the ventral air layer by means of the narrow gap between the
ventrally exposed portion of the forewing (Fig. 1, FW) and the
ventrolateral edge of the body. The hydrofuge hairs l ning this gap
would permit a small amount of oxygen to pass from the ventral
plastron to the first abdominal spiracle, which faces the subalar space
(Parsons 1974). The extent to which the spiracles, especially the
first abdominal ones, are capable of inhalation is not known, however,
and will require experimentation and further fine-structural studies.

There is a . strong possibility that well-developed plastral respira-
tion has evolved in other Naucoridae, as Hinton (1969a) claims.
Further investigation will probably reveal it not only in other mem-
bers of this family but quite possibly in other families of aquatic
Heteroptera as well.

Fig. 11. C. barozzii; shin,gle-like arrangement of several groups of
microtrichia on median portion of fourth abdominal sternite, approximately
midway between its anterior (towards top) and posterior (towards bottom)
boundaries. Arrow indicates where tips of hairs, mostly concealed by debris,
appear to be interspersed among groups of microtrichia. Scale line = 5 ^m.

Fig. 12. C. barozzii; large numbers of fringed setae on lateral edge of
sixth ventral abdominal paratergite, left side of body. Arrow indicates
edge of body. Scale line = 10 |Um.

1974]

Parsons & Hewson — Cryphocricos

525

526

Psyche

[September-December

Acknowledgments

Acknowledgment is made for the use of the scanning electron
microscope in the Royal Ontario Museum, established through a
grant from the National Research Council of Canada to the Depart-
ment of Zoology, University of Toronto. We wish to thank John T.
Polhemus, Randall T. Schuh, Fritz Plaumann, and D. T. Crisp,
who provided specimens and/or data on habitats. We are also grate-
ful to Eric Lin and Kian Chua for their technical help, to W. H.
Thorpe and Thomas S. Parsons for their advice and encouragement,
and to Donald A. Chant and the Department of Zoology, University
of Toronto, who made available the laboratory facilities for this
study. The investigation was made possible by a grant in aid of
research to the senior author from the National Research Council of
Canada.

Summary

The scanning electron microscope shows that in Cryphocricos
barozzii and C. hungerfordi most of the ventral surface of the body
and at least a part of the dorsal surface are covered with fine struc-
tures which appear to be hydrofuge devices. Both surfaces bear short,
fine hairs which are very similar to those of Aphelocheirus aestivalis ,
the only aquatic heteropteran in which plastral respiration has thus
far been demonstrated. These hairs are much smaller and more
densely-packed than those of Pelocoris fernoratus , which carries a
large air bubble and relies upon atmospheric oxygen. On the ventral
abdominal paratergites of Cryphocricos the plastral hairs are covered
by curious, leaf-like, ridged setae which overlap each other and per-
haps retain a “macroplastron”. Very elongated leaf-like setae occur
on the six pairs of abdominal pressure receptors of Cryphocricos.
A phelocheirus possesses leaf-like setae only on its single pairs of ab-
dominal sense organs. The fourth through sixth abdominal sternites
of Cryphocricos bear groups of microtrichia with a shingle-like ar-
rangement; they resemble the plastral devices of some aquatic Cole-
optera. These fine structural observations strongly suggest that Cry-
phocricos has plastral respiration and utilizes dissolved rather than
atmospheric oxygen.

1974]

Parsons & Hewson — Cryphocricos

527

Literature Cited

Hinton, H. E.

1969a. Algunas pequerias estructuras de insectos observadas con micro-
scopio electronico explorador. Acta Politecnica Mexicana 10:
181-201.

1969b. Plastron respiration in adult beetles of the suborder Myxo-
phaga. J. Zool., Lond. 195: 131-137.

Larsen, O.

1955. Spezifische Mechanorezeptoren bei Aphelocheirus aestivalis Fabr.
nebst Bemerkungen uber die Respiration dieser Wanze. Lunds
Univ. Arsskr. 51(11): 1-58.

Parsons, M. C.

1969. The food pump of Aphelocheirus aestivalis F. as compared with
that of typical Naucoridae (Heteroptera). J. Morphol. 129: 17-
30.

1974. Anterior displacement of the metathoracic spiracle and lateral
intersegmental boundary in the pterothorax of Hydrocorisae
(aquatic Heteroptera). Z. Morphol. Tiere 79: 165-198.

Thorpe, W. H.

1950. Plastron respiration in aquatic insects. Biol. Rev. 25: 344-390.

Thorpe, W. H. and D. J. Crisp

1947a. Studies on plastron respiration. I. The biology of Aphelocheirus
aestivalis [Hc-miptera, Aphelocheiridae (Naucoridae)] and the
mechanism of plastron retention. J. Exp. Biol. 24: 227-269.

1947b. Studies on plastron respiration. II. The respiratory efficiency of
the plastron in Aphelocheirus. J. Exp. Biol. 24: 270-303.

1947c. Studies on plastron respiration. III. The orientation responses
of Aphelocheirus [Hemiptera, Aphelocheiridae (Naucoridae)] in
relation to plastron respiration ; together with an account of
specialized pressure receptors in aquatic insects. J. Exp. Biol.
24: 310-328.

1949. Studies on plastron respiration. IV. Plastron respiration in the
Ccleoptera. J. Exp. Biol. 26: 219-260.

Usinger, R. L.

1956. Aquatic Hemiptera. In Aquatic insects of California. Edited by
R. L. Usinger. University of California Press, Berkeley and
Los Angeles.

Young, B.

1944. Respiratory adaptation in an aquatic Hemipteron Cheirochela.
Sinensia 1 5 : 141-144.

HOME RANGES OF

MALE CERCERIS SIMPLEX MACROSTICTA
(HYMENOPTERA, SPHECIDAE)*

By John Alcock and George Gamboa

Department of Zoology, Arizona State University, Tempe 85281 :
and Department of Zoology, University of Iowa, Iowa City, 52242.

Although it is well known that many sphecid males visit flowers
and shrubs and attempt to mate with females found there (e.g.
Peckham, Kurczewski, & Peckham, 1973), many unanswered ques-
tions about this general pattern of male behavior remain. For
example, do flower- visiting males move randomly from area to area
or do they remain in one location for substantial periods of time?
If they show an attachment to a particular site, do they defend the
area or do they share it with other males?

We addressed these questions for one species by studying males of
Cerceris simplex macrosticta Viereck and Cockerell at two separate
sites along State Line Rd., p2 km west of Rodeo, New Mexico.
Study area 1 consisted of a strip of roadside ditch, 75 m X 2 m,
covered with a dense stand of flowering milkweeds ( Asclepias sub-
verticillata) . Two mesquites and a sprawling multiflora rosebush
were the only large plants at this location. Study site 2, located
several hundred meters from study site 1, was an area roughly
25 nr X 10 m in an uncultivated pear orchard overrun with clumps
of Russian thistle ( Salsola kali) and dotted with scattered flowering
weeds. From 23-27 July 1974 sixteen males were captured in these
two areas and marked on the dorsum of the thorax with acrylic paints,
each individual receiving a distinctive color or color combination.
One or more censuses were made at the two sites on all days from
23 July to 8 August with the exception of 2, 4, and 7 August.

Results

Home ranges of C. simplex

Nine of the marked males were seen again at or near the location
where they had been originally captured and seven of the sixteen
males were followed for a week or more (Table 1). The failure to
relocate six marked individuals could be attributed to a variety of
factors including mortality and the passage of transient males from
the study areas.

* Manuscript received, by the editor January 20, 1975.

528

1974]

Alcock & Gamboa — Cerceris

529

Table 1. Site attachment of male C. simplex. A total of sixteen individuals
marked 23-27 July.

1

2

3

4

Days At Site
5 6 7 8

9

10

11

6

,

1

Number of Wasps

1 2

,

2

2

The resident males were repeatedly observed pursuing regular
circuits between 0839-1230 hrs. and rarely as late as 1400 M.S.T.
Four carefully studied individuals possessed routes ranging from
7.0-18.5 m in length and from 2-3 m in width (Fig. 1). However,
marked wasps also made occasional forays up to six m from their
customary route. Males in the roadside ditch cruised in a fairly
rapid and erratic flight going down one side of the ditch skimming
over flowering Asclepias , and then back up the other. Orchard males
flew from one clump of Salsola to another sailing around and over
them, sometimes visiting flowering weeds along the route (Fig. 1).

Wasp “yellow” had the smallest circuit of the males we studied
and required a mean of 32 sec to complete one flight about its home
range (N — 9; range, 22-68 sec; 28 July, 0820-1115). Most of a
male’s time was spent in flight, although the wasps regularly alighted
on perches in their cruising area (Fig. 2). For example, “white”
landed on one branch of a rose bush 16 times in 35 minutes (23
July) remained perched an average of 6:5 sec (range, 1-19 sec).
Several males had a primary perch to which they commonly returned
as well as several secondary perches that were used less frequently.

Males pursued any medium-sized flying insect including a wide
variety of wasps, beeflies, and brilliant black and red hemipterans as
well as male and female conspecifics. Males made physical contact
only with conspecific females. The indiscriminate nature of the pur-
suit flight and the absence of grappling or contact between males
leads us to believe that male C. simplex lack behavior designed to
defend a territory. The great overlap in routes travelled by different
males (at least four males regularly entered “blue’s” circuit and
easily that many were present in “white’s” area) further suggests
that males of this species occupy home ranges and not territories.

Attempted matings

Despite many hours of observation, attempted copulations were
seen only three times (at 0915, 0925, 1030), all in the orchard.
In one case a female cruised slowly through a Salsola clump occa-

530

Psyche

[September-December

Fig. 1. The overlapping home ranges of three males at the orchard
study area. “Blue’s” home range is outlined with a dashed line; “red’s”
home range is outlined with a dot-dashed line ; “white-dot’s” home range
is outlined with a solid line. The circular stippled areas represent prominent
Salsola bushes. One large flowering weed is shown schematically in the
lower left hand corner of “blue’s” home range.

sionally alighting and walking up stems apparently searching for
prey, the tenebrionid beetle Metapoloba pruinosa (Alcock, 1974),
which were sometimes seen in this plant. As she flew slowly past a
perched male he flew up and cruised behind her. The female alighted
and the male quickly dropped onto her back facing in the same direc-
tion as his potential mate. While stroking her antennae with his the
male held the female’s wings out at right angles to her body by sand-
wiching each wing between his fore- and midlegs, presumably making
escape more difficult for the female. His hindlegs grasped the fe-
male’s abdomen. After several minutes of unsuccessful attempts to
copulate, the male released his partner and both flew away. The
other two attempted matings also followed this general pattern. One
took place in a Salsola plant, the other on a flower head of a weed.
The related wasp C. frontata also mates at flowers, initially adopting
the position used by C. simplex (pers. obs.) with the male dismount-
ing after coupling is achieved (Scullen, 1965).

1974]

Alcock Cff Gamboa — Cerceris

53

Sleeping sites

In the late afternoon, especially before impending storms, males
were seen flying slowly along the large rose bush. The wasps eventu-
ally alighted and walked under groups of tightly packed leaves and,
in one case, into a tube formed by a rolled leaf, evidently in search
of a sheltered sleeping site. One male (“red”) that had been marked
on the morning of 27 July at a spot 50 m to the south of the rose
bush was found by this plant at 1610 seeking a resting place before a
thunderstorm. The next morning it had returned to the area where
it had been marked indicating that some males may fly modest dis-
tances from their home range in order to exploit a sleeping site.

Fig. 2. A male Cerceris simplex perched on a rose leaf and resting
between cruising flights about its home range.

532

Psyche

[September-December

Discussion

To the best of our knowledge C. simplex provides the only docu-
mented example of a wasp species whose males occupy fairly exten-
sive home ranges in areas away from an aggregation of nesting
females. Doubtless this only reflects the lack of research of male
wasps and it is probable that home ranges embracing patches of
flowers are a common phenomenon in male sphecids.

The spacing system of male C. simplex does, however, differ from
that of various close relatives, among them several species of Eucer-
ceris as well as Philanthus and Clypeadon (Alcock, 1975 and in
prep.). Males of these related species form relatively dense aggrega-
tions with each individual applying an attractant pheromone from
abdominal glands to a perch and vigorously repelling all intruders
from an area roughly J4-I m in radius around the perch. Whether
all the members of one genus share the same basic mating strategy is
unknown and will require much further study, although the very
limited current information hints that this may be the case (Alcock,
in prep.) .

Despite the differences between C. simplex and other philanthinine
wasps there are some similarities as well. In all cases males occupy a
perch to which they return after bouts of flying around. All pursue
a remarkably wide range of flying insects evidently in an effort to
determine the species and sex of the passerby or intruder. The key
variable between different species is the area inspected by an indi-
vidual. Those males that use a perch as a calling post defend it and
a small area around it while C. simplex , which does not appear to
employ an attractant pheromone, covers a much larger area (up to
at least 40 m2), so large as to be essentially indefensible. Attempts
to drive all other males from a strip of roadside ditch 15 m X 2 m
would require an extraordinary investment in aggressive territorial
patrolling. Instead, C.' simplex males make no special effort to locate
and repel conspecific males. Time and energy are devoted to re-
peated investigation of a large area in a location likely to attract
females, one which is rich in nectar-producing flowers or beetle-
producing bushes. Having found a potentially good area, males re-
main there becoming familiar with its prime spots. Presumably this
is a superior strategy than one that would take a male on a random
journey through the countryside in search of a mate. Because natu-
rally occurring attractants (e.g. flowers) that draw in females tend
to be patchily distributed, one would predict that males of other
species that use such attractants to focus their search for females will

1974]

Alcock & Gamboa — Cerceris

533

also possess home ranges. Additional comparative data based on
studies of marked populations of males are required to test this pre-
diction.

Acknowledgments

We thank Dr. Howard E. Evans for his review of a draft of this
paper. This study was supported by National Science Foundation
Grant GB-42865.

References

Alcock, J.

1974. The nesting behaviour of Cerceris simplex macrosticta (Hymen-
optera: Sphecidae). J. Nat. Hist., 8: 645-652.

1975. Territorial behaviour by males of Philanthus multimaculatus

(Hymenoptera : Sphecidae) with a review of territoriality in

male sphecids. Anim. Behav., in press.

Peckham, D. J.. F. E. Kurczewski & D. B. Peckham

1973. Nesting behavior of Nearctic species of Oxybelus (Hymenoptera:
Sphecidae). Ann. Ent. Soc. Amer., 66: 647-661.

Scullen, H. A.

1965. Review of the genus Cerceris in America north of Mexico
(Hymenoptera: Sphecidae). Proc. U. S. Nat. Mus., 1 16: 333-548.

ADDITIONAL NOTES ON
STEPHANITIS TAKEYAI IN NEW ENGLAND
(HETEROPTERA: TINGIDAE)

By Norman S. Baiiley
Kingston, New Hampshire 03848*

In 1950 I reported this Asiatic lace bug as a serious pest of Pieris
japonica (Thunberg) Don in Greenwich, Connecticut, and the
surrounding area. Since 1956 I have been teaching at Bradford
College, in northeastern Massachusetts. Although there are several
specimens of Pieris on the rather nicely landscaped campus and in
the planting around a college house I lived in for many years, I saw
no evidence of lace bugs until the fall of 1973. On that occasion I
searched the plants showing considerable injury but failed to find
any tingids.

On September 9, 1974, however, I noted that the same plants
showed typical signs of severe feeding injury and found numerous
nymphs and adults of Stephanitis takeyai on the undersides of the
leaves of two Japanese Andromedas.1 These plants grow in a mixed
shrub planting along the north foundation of our main lecture hall.
In the same foundation planting are such other evergreen ericaceous
shrubs as Pieris floribunda Bentham & Hooker, Rhododendron caro-
linianum Rehder, Kalmia latifolia L., Leucothoe catesbaei Gray, and
a few deciduous azaleas. With the exception of an azalea, whose
branches intermingle with one of the infested Pieris japonica , there
was no evidence of lace bug feeding nor did I find any tingids on
the other ericaceous plants. This particular azalea, which may be
R. calendulaceum Torrey, does show lace bug injury and specimens
were collected from it. Many of its leaves had also been almost
skeletonized in a way that suggested Japanese beetle damage.

A few days later I found another heavily infested Pieris japonica
in a planting near a small building towards the south end of the
campus. This suggested checking further. On October 1st, two more
plants were found on the front campus with tingids present in some

*Address after July 1, 1975: Mohler Laboratory, Swans Island Marine
Station, Swans Island, Maine 04685.

The original name of this species, globulifera Matsumura, turns out to
be a junior homonym, and it was renamed takeyai by Drake and Maa,
1955. See also Drake and Ruhoff, 1965.

Manuscript received by the editor November 2, 1974.

534

1974]

Bailey — Stephanitis

535

numbers. One grows on the north corner of our auditorium and the
other on the west side of the same building. The latter grows beside
a hybrid Rhododendron that also showed considerable leaf discolora-
tion, especially on leaves of the previous year. This year’s growth
was only thinly specked with fine whitish dots on the upper surface
and no lace bugs were located on either the older or the newer
foliage of this plant.

On October 3rd I extended the search and inspected Pieris japonica
specimens in the front foundation plantings of five college owned
houses situated on the opposite sides of the two streets that parallel
the east and the west sides of the main campus. A sixth house is on
a road beyond the tennis courts at the southern edge of the college
property and about a half a mile from the front of the campus where
specimens were first found. All of the plants observed showed some
evidence of injury and several were heavily infested. In spite of
light frosts on at least two nights recently, temperatures below 40°F
for another two nights, and only in the low fifties during the two
consecutive days, adults (some teneral) and nymphs of Stephanitis
were numerous and active on some of these Pieris on October 3rd
and 4th.

On the 4th a collection was taken from a Pieris japonica by the
northeast entrance to the campus. This plant had numerous nymphs
and some adults on the leaves. At least four nymphal instars are
represented in this one late season collection. The smallest nymph,
without apparent spines, was 0.7 mm. long. Others measured 1.2
mm., 1.8 mm. and 2.4 mm. in length and these all had prominent
spines on the dorsal side of head, thorax, and abdomen Three adults
found with them measured from 3.9 to 4.2 mm.

Although many leaves were examined (several with the aid of a
stereoscopic microscope), no evidence of eggs was discovered. There-
fore, the late occurrence of early nymphal instars and of teneral
adults strongly indicates that S. takeyai may over-winter in these
stages, contrary to my earlier suggestion (1951). Obviously this
point needs clarification.2

On October 26th, specimens of Pieris japonica showing character-
istic discoloration were seen in Haverhill at least two miles from the

2While this paper was in press, I received a copy of Dr. Dennis Dunbar’s
recent (1974) article on this tingid, in which mention is made of the gen-
eral occurrence of takeyai in southern New England, without giving specific
localities in Massachusetts. The same paper provides a full account of the
life history in Connecticut and indicates the species lays eggs that over-
winter in the Pieris leaves there.

536

Psyche

[September-December

campus and north of the Merrimac River. The campus is in a resi-
dential area south of the river. Obviously this lace bug is now well
established locally. The leaves of Pieris japonica, much discolored
and damaged by the heavy feeding of Stephanitis , tend to drop pre-
maturely. Consequently, gardeners and nurserymen should be con-
cerned with the spread of this lace bug from southwestern Connecti-
cut into northeastern Massachusetts.

References

Bailey, L. H.

1949. Manual of Cultivated Plants. Macmillan Co. New York.
Bailey, Norman S.

1950. An Asiatic Tingid New to North America (Heteroptera) . Psyche
57: 143-145.

1951. The Tingoidea of New England and Their Biology. Ent.
Americana 3 1 (n.s.) :5 3-62.

Drake, C. J. and T. Maa

1955. Chinese and Other Oriental Tingoidea (Hemiptera). Part III.
Quart. Journ. Museum Taiwan 8:1-11.

Drake, C. J. and F. A. Ruhoff

1965. Lace Bugs of the World: A Catalogue. U.S. Nat. Mus. Bull.
213 :364.

Dunbar, Dennis M.

1974. Bionomics of the Andromeda Lace Bug, Stephanitis takeyai.
Mem. Conn. Ent. Soc. 1974:277-289.

PSYCHE

INDEX TO VOLUME 81, 1974

INDEX TO AUTHORS

Alcock , John and George Gamboa. Home Ranges of Male Cerccris simplex
macrostricta (Hymenoptera ; Sphecidae). 528
Bailey, Norman S. Additional Notes on Stephanitis takeyai in New Eng-
land (Heteroptera : Tingidae). 534

Brothers, Denis J. The First Recent Species of Protomutilla (Hymenoptera:
Mutillidae: Myrmosinae). 268

Brown, JVilliam L., Jr. A Remarkable New Island Isolate in the Ant Genus
Proceratium (Hymenoptera: Formicidae). 70
Burns, John. The Polytypic Genus Celotes (Lepidoptera : Hesperiidae:
Pyrginae) from the Southwestern United States and Northern Mexico.
51

Cooper, Kenneth JV . Sexual Biology, Chromosomes, Development, Life
Histories and Parasites of Boreus, Especially of B. notoperaies . A
Southern California Boreus. II. (Mecoptera: Boreidae). 84
Coyle, Frederick, A. Systematics of the Trapdoor Spider Genus Aliatypus
(Araneae: Antrodiaetidae) . 431

Eisner, Thomas, Daniel Aneshansley , Maria Eisner, Ronald Rutowski,
Berni Chong, and Jerrold Meinwald. Chemical Defense and Sound
Production in Australian Tenebrionid Beetles ( Adelium spp.). 189
Evans, H. E., R. tV . Matthews , and E. C. McC. Callan. Observations on
the Nesting Behavior of Rubrica surinamensis (DeGeer) {Hymen-
optera: Sphecidae). 334

Foelix, Rainer F. Application of the Transmission Electron Microscope
to the Examination of Spider Exuviae and Silk. 507
Harper, P. P. New Protonemura (s.l.) from Nepal (Plecoptera; Nemouri-
dae). 367

Kukalova-Peck, Jarmila. Wing-folding in the Paleozoic Insect Order Dia-
phanopterodea (Paleoptera) , with a Description of New Representa-
tives of the Family Elmoidae. 315

Kukalova-Peck, Jarmila. Pteralia of the Paleozoic Insect Orders Palaeo-
dictyoptera, Megasecoptera and Diaphanopterodea (Paleoptera). 416
Haskins, Caryl P. and Edna F. Haskins. Notes on Necrophoric Behavior in
the Archaic Ant Myrmecia vindex (Formicidae: Myreciinae). 258
Hlavac, Thomas F. Metope tuber: A Wing-body Interlocking Mechanism.
303

Hunt, James IP. Temporal Activity Patterns in Two Competing Ant Species
(Hymenoptera: Formicidae). 237

Jeanne, Robert L. and Robert Fagen. Polymorphism in Stelopolybia areata
(Hymenoptera: Vespidae). 155

Matthews , Janice R. History of the Cambridge Entomological Club. 3
Moglick, Michael and Bert Hoildobler. Social Carrying Behavior and Divi-
sion of Labor During Nest Moving of Ants. 219

537

Moglich, Michael and Bert Holldobler. Erratum. 415

Moore , Ian. A New Species of Opithes from Mexico, with a Key to the
Species. 353

Nutting, IF. L., M. S. Blunt, and H. M. Fales. Behavior of the North
American Termite., T enuirostriterm.es tenuirostris , with Special Refer-
ence to the Frontal Gland Secretion, Its Chemical Composition, and
Use in Defense. 167

Parsons, Margaret C. Modification of the Intersegmental Region in the
Pterothorax of Cryphocricos (Heteroptera : Naucoridae). 42

Parsons, Margaret C. and Rosemary J. Hewson. Plastral Respiratory
Devices in Adult Cryphocricos (Naucoridae: Heteroptera). 510

Peck, Stewart B. The Eyeless Catopocerus Beetles (Leiodidae) of Eastern
North America. 377

Peck , Stewart B. A Review of the Agyrtes (Silphidae) of North America.

501

Quintero, Diomedes , Jr. The Unusual Web of Spilasma tubulofaciens, with
Taxonomic Notes of the Species (Araneae: Araneidae). 307
Reiskind, Jonathan. The Castianeirinae of Mexico. I. Castianeira dugesi
(Becker) (Araneae: Clubionidae) . 178

Roth, Louis M. A New Cockroach Genus ( Gurneya ) Previously Confused
with Pinaconata (Blaberidae: Epilamprinae) . 288

Ro<vner, Jerome S. and Susan J. Knost. Post-immobilization Wrapping of
Prey by Lycosid Spiders of the Herbaceous Stratum. 398
Shapiro, Arthur M. Host-Plant Utilization by Pieris napi Populations in
California (Lepidoptera : Pieridae). 361
Shear, IVilliam A. The Milliped Genus Bollmanella (Diplopoda, Chore-
umida, Conotylidae) . 134

Slansky, Frank. Relationships of Larval Food-plants and Voltinism Pat-
terns in Temperate Butterflies. 243

Smiih, Robert L. Life History of Abedus herberti in Central Arizona
(Hemiptera; Belostomatidae) . 272

Valerio, Carlos E. Prey Capture by Drymusa dinora (Araneae, Scolytidae).
284

Waldorf, Elizabeth S. Variations in Cleaning Between the Sexes of Sinella
coeca (Collembola: Entomobryidae) . 254

Werner, Floyd G. A New Genus of Primitive Meloidae from West Texas
(Coleoptera) . 147

Wheeler, George C. and Jeanette Wheeler. Supplementary Studies of Ant
Larvae: T eratomyrmex. 38

Willey, Ruth L. Emergence Pattern of the Subalpine Dragonfly, Somato-
chlora semicircularis (Odonata: Conduliidae) . 121

Wilson, Edward O. The Soldier of the Ant, Camponotus ( Colobobsis )
fraxinicola, as a Trophic Caste. 182

W ussow, Paul A., Robert B. Willey, and June B. Steinberg. Synchronous
Visualization of Video-taped Sounds and Motions of Insects. 209

538

INDEX TO SUBJECTS

All new genera, new species and new names are printed in capital type.

Abedus herberti, 272
Additional Notes on Stephanitis ta-
keyai in New England, 534
Agyrtes longulus, 501
Agyrtes similis, 504
Aliatypus aquilonius, 458
Aliatypus calif ornicus , 446
Aliatypus erebus, 470
Aliatypus isolatus, 456
Aliatypus gnomus, 460
Aliatypus gulosus, 461
Aliatypus janus, 452
Aliatypus thompsoni, 463
Aliatypus plutonis, 473
Aliatypus torridus, 474
Aliatypus trophonius, 468
Application of the Transmission
Electron Microscope to the Exami-
nation of Spider Exuviae and
Silk, 507

Behavior of the North American
Termite, T enuirostritermes tenui-
rostris, with Special Reference to
the Frontal Gland Secretion, Its
Chemical Composition, and Use in
Defense, 167
Bollmanella, 134
Boreus, biology, 84
Boreus notoparates , 84

Cambridge Entomological Club, 3
Camponotus fraxinicola, 182
Castianeirinae of Mexico. I. Casti-
aneira dugesi (Becker), 178
Castianeira dugesi, 178
Catopocerus alabamae, 393
Catopoccrus appalachianus, 387
Catopocerus hamiltoni, 381
Catopocerus jonesi, 392
Catopocerus ulkei, 383
Celotes, 51
Celotes limpia, 51
Cerceris simplex macrostricta, 528
Chemical Defense and Sound Pro-
duction in Australian Tenebrionid
Beetles, 189
Cryphocricos , 42

DlAPHA CANDIDA, 323
Diaphanopterodea, 416
Drymusa dinora, 284

Elmodiapha ovata, 320
Emergence Pattern of the Subalpine
Dragonfly, Somatochlora semicir-
cular is, 121
Erratum, 415

Eyeless Catopocerus (Leiodidae) of
Eastern North America, 377

First Recent Species of Protomutilla,
268

Gurneya obliqua, 297

History of the Cambridge Ento-
mological Club, 3

Home Ranges of Male Cerceris sim-
plex macrostricta, 528
Host-Plant Utilization by Pieris napi
Populations in California, 361

Life History of Abedus herberti in
Central Arizona, 272
Lycosid spiders, 398

Megasecoptera, 416
Meloidae, 147

Merope tuber: A Wing-body Inter-
locking Mechanism, 303
Milliped Genus Bollmanella, 134
Modification of the Intersegmental
Region in the Pterothorax of
Cryphocricos, 42
Myrmecia vindex, 258

New Cockroach Genus ( Gurneya )
Previously Confused with Pina-
conata, 288

New Genus of Primitive Meloidae
from West Texas, 147
New Protonemura (s.l.) from Nepal,
367

New Species of Opithes from Mexico,
with a Key to the Species, 353

539

Notes on Necrophoric Behavior in
the Archaic Ant Myrmecia vin-
dex, 258

Observations on the Nesting Be-
havior of Rubrica surinamensis
(DeGeer), 334
Opithes CRANDALLI, 354

Palaeodictyoptera, 416
Paradiapha delicatula, 331
Permodiapha carpenteri, 325
Permodiapha lata, 325
Permodiapha specula, 327
Pinaconata, 288

Plastral Respiratory Devices in
Adult Cryphocricos , 510
Polymorphism in Stelopolybia areata,
155

Polytypic Genus Celotes from the
Southwestern United States and
Northern Mexico, 51
Post-immobilization Wrapping of
Prey by Lycosid Spiders of the
Herbaceous Stratum, 398
Prey Capture by Drymusa dinora,
284

Proceratium, 70
Protodiapha lineata, 321
Protodiapha maculifera, 321
Protonemura adunca, 372
Protonemura funicula, 370
Protonemura godavariensis, 369
Protonemura mastigophora, 372
Protonemura mira, 367
Protomutilla microsoma, 270
Pteralia of the Paleozoic Insect
Orders Palaeodictyoptera, Megase-
coptera and Diaphanopterodea
(Paleoptera), 416

Relationships of Larval Food-plants
and Voltinism Patterns in Tem-
perate Butterflies, 243
Remarkable New Island Isolate in
the Ant Genus Proceratium, 70
Review of the Agyrtes (Silphidae)
of North America, 501
Rubrica surinamensis, 334

Sexual Biology, Chromosomes, De-
velopment, Life Histories and
Parasites of Boreus, Especially of
B. notoparates. A Southern Cali-
fornia Boreus. II, 84
Social Carrying Behavior and Divi-
sion of Labor During Nest Moving
of Ants, 219
Sinella coeca, 254

Soldier Ant, Camponotus ( Colobop -
sis) fraxinicola, as a Trophic
Caste, 182

Somatochlora semicircularis, 121
Spider exuviae and silk, 501
Spilasma tubulofaciens, 307
Stelopolybia areata, 155
Stenodiapha angusta, 329
Stenodiapha moravica, 329
Stephanitis takeyai, 534
Supplementary Studies of Ant Lar-
vae: T eratomyrmex, 38
Synchronous Visualization of Video-
taped Sounds and Motions of In-
sects, 209

Systematics of the Trapdoor Spider
Genus Aliatypus, 431

Temporal Activity Patterns in Two
Competing Ant Species, 237
Tenebrionid beetles, 189
Te?iuirostritermes tenuirostris, 167
T eratomyrmex, 38
Thambospasta howdeni, 151

Unusual Web of Spilasma tubulo-
faciens with Taxonomic Notes of
the Species, 307

Variations in Cleaning Between the
Sexes of Sinella coeca, 254

Wing-folding in the Paleozoic Insect
Order Diaphanopterodea (Pale-
optera), with a Description of
New Representatives of the Family
Elmoidae, 315

540

CAMBRIDGE ENTOMOLOGICAL CLUB

A regular meeting of the Club is held on the second Tuesday of
each month October through May at 7:30 p.m. in Room B-455,
Biological Laboratories, Divinity Ave., Cambridge. Entomologists
visiting the vicinity are cordially invited to attend.

The illustration on the front cover of this issue is of Samuel Hubbard
Scudder of Cambridge, who, along with Professor Hagen, provided the
initiative for the founding of the Cambridge Entomological Club and a
Psyche in 1874. It is taken from a photograph in the archives of the
Museum of Comparative Zoology.

BACK VOLUMES OF PSYCHE

Requests for information about back volumes of Psyche should
be sent directly to the editor.

F. M. Carpenter

Editorial Office, Psyche
16 Divinity Avenue

Cambridge, Mass. 02138

FOR SALE

Classification of Insects, by C. T. Brues, A. L. Melander and
F. M. Carpenter, Published in March, 1954, as volume 108 of the
Bulletin of the Museum of Comparative Zoology, with 917 pages
and 1219 figures. It consists of keys to the living and extinct families
of insects, and to the living families of other terrestrial arthropods;
and includes 270 pages of bibliographic references and an index of
76 pages. Price $14.00 (cloth bound and postpaid). Send orders
to Museum of Comparative Zoology, Harvard College, Cambridge,
Mass. 02138.

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