The Pan-Pacific Entomologist

EDITORIAL BOARD

J. T. Sorensen, Editor

S. S. Shanks, Treasurer Susan M. Sawyer, Editorial Assistant R. M. Bohart

J. T. Doyen J. E. Hafemik, Jr. J. A. Powell

Published quarterly in January, April, July, and October with Society Proceed¬

ings usually appearing in the October issue. All communications regarding non¬

receipt of numbers, requests for sample copies, and financial communications

should be addressed to: Sandra Shanks, Treasurer, Pacific Coast Entomological

Society, Dept, of Entomology, California Academy of Sciences, Golden Gate Park,

San Francisco, CA 94118-4599.

Application for membership in the Society and changes of address should be

addressed to: Curtis Y. Takahashi, Membership Committee chair, Pacific Coast

Entomological Society, Dept, of Entomology, California Academy of Sciences,

Golden Gate Park, San Francisco, CA 94118-4599.

Manuscripts, proofs, and all correspondence concerning editorial matters (but

not aspects of publication charges or costs) should be sent to: Dr. John T. Sorensen,

Editor, Pan-Pacific Entomologist, Insect Taxonomy Laboratory, California Dept,

of Food & Agriculture, 1220 N Street, Sacramento, CA 95814. See the back cover

for Information-to-Contributors, and volume 66 (1): 1-8, January 1990, for more

detailed information. Refer inquiries for publication charges and costs to the

Treasurer.

The annual dues, paid in advance, are $20.00 for regular members of the Society,

$10.00 for student members, or $30.00 for institutional subscriptions. Members

of the Society receive The Pan-Pacific Entomologist. Single copies of recent num¬

bers or entire volumes are available; see 67(1):80 for current prices. Make checks

payable to the Pacific Coast Entomological Society.

Pacific Coast Entomological Society

OFFICERS FOR 1990

Leslie S. Saul, President

Norman D. Penny, President-Elect Wesley A. Maffei, Managing Secretary

Sandra S. Shanks, Treasurer Larry G. Bezark, Recording Secretary

THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly by the Pacific

Coast Entomological Society, c/o California Academy of Sciences, Golden Gate Park, San Francisco,

CA 94118-4599. Second-class postage is paid at San Francisco, CA and additional mailing offices.

Postmaster: Send address changes to the Pacific Coast Entomological Society, c/o California Academy

of Sciences, Golden Gate Park, San Francisco, CA 94118-4599.

This issue mailed 4 April 1991

The Pan-Pacific Entomologist (ISSN 0031-0603)

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

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

PAN-PACIFIC ENTOMOLOGIST

67(1): 1-6, (1991)

PACIFIC COAST ENTOMOLOGICAL SOCIETY

Organized 15 August 1901

ARTICLES OF INCORPORATION

Know all men by these presents:

That we, the undersigned, do hereby voluntarily associate ourselves together for the purpose of

forming a nonprofit corporation under the laws of the State of California, and we hereby certify that:

First: The name of this corporation shall be the Pacific Coast Entomological Society.

Second: The purposes for which this corporation is formed are to promote, sponsor, foster, and

stimulate research in the field of entomology; to provide facilities and opportunities for research and

analysis of any and all phases of entomology by lectures, forums, discussions, symposia, and analyses;

to publish books, pamphlets, monographs, and in general to disseminate information about entomology

and the results of entomological research; to accumulate and record historical facts and materials

relating to entomology in general; to give, grant, and sponsor the granting of fellowships and awards

for study or research in the field of entomology; and research in entomology between the members of

this corporation and scientists and entomologists throughout the world. To do any and all things

necessary, suitable, convenient, or proper for or in connection with or incidental to the accomplishment

of any of the purposes, or the attainment of any one or more of the objects herein enumerated or

designed, directly or indirectly to promote the interests of this corporation, and to do any and all

things and to exercise any and all powers which may now or hereafter be lawful for this corporation

to do or to exercise under the laws of the State of California; provided, however that this corporation

is and shall be a corporation which does not contemplate pecuniary gain or profits to the members

thereof. No part of the net earnings of this corporation shall inure to the benefit of any member

whatsoever, and no part of the activities of this corporation shall be or constitute carrying on prop¬

aganda or otherwise attempting to influence legislation.

Third: The county in the State of California where the principal office for the transaction of the

business of this corporation is to be located is the City and County of San Francisco.

Fourth: The number of this corporation’s directors who shall be known as the members of the

Executive Board of this corporation is eleven (11), and the names and addresses of the persons who

are to act in the capacity of members of the Executive Board until election of their successors are:

E. Gorton Linsley, University of California, Berkeley

H. M. Armitage, State Dept, of Agriculture, San Francisco

Peter C. Ting, State Dept, of Agriculture, San Francisco

E. R. Leach, 217 Hillside Avenue, Piedmont, California

R. L. Usinger, University of California, Berkeley

E. O. Essig, University of California, Berkeley

Gordon F. Ferris, Stanford University, California

Edward S. Ross, California Acad, of Sciences, San Francisco

Carl S. Duncan, San Jose State College, California

The number of members of the Executive Board may be changed from time to time by the bylaws

duly adopted; provided, however, that the number of the members of the Executive Board shall not

be less than three (3).

Fifth: The authorized number and qualifications of the members of this corporation, the different

classes of membership, if any, the property, voting, and other rights and privileges of each class of

membership, and the liability of each and all classes of members to dues and assessments, and the

method of collection thereof, shall be as set forth in the bylaws of this corporation.

Sixth: The name of the existing unincorporated association which is hereby being incorporated is

Pacific Coast Entomological Society.

Seventh: The property of this corporation is irrevocably dedicated to educational and scientific

purposes and no part of net income or assets of this organization shall ever enure to the benefit of

any director, officer, or member thereof or to the benefit of any private persons. Upon dissolution or

winding up of the corporation, its assets, remaining after payment of, or provisions for payment of,

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

all debts and liabilities of this corporation, shall be distributed to the California Academy of Sciences,

located in San Francisco, California, a nonprofit corporation which has established its tax-exempt

status under Section 501(c)(3) of the Internal Revenue Code of 1954. In the event the Academy or

any successor organization is not in existence or is not tax-exempt under Section 501 (c)(3), distribution

shall be made to a nonprofit fund, foundation, or corporation which has established its tax-exempt

status under Section 501(c)(3) of the Internal Revenue Code of 1954.

If this corporation holds any assets in trust, or the corporation is formed for charitable purposes,

such assets shall be disposed of in such a manner as may be directed by decree of the superior court

of the city in which the corporation has its principal office, upon petition therefore by the Attorney

General or by a person concerned in the liquidation, in a proceeding to which the Attorney General

is a party.

In Witness Whereof, the president and secretary of the Pacific Coast Entomological Society, an

existing unincorporated association, have hereunto set their hands this 28th day of August, 1940.

E. Gorton Linsley

President of Pacific Coast Entomological

Society, an Existing Unincorporated

Association

Peter C. Ting

Secretary of Pacific Coast Entomological

Society, an Existing Unincorporated

Association

Endorsed and filed in the

office of the Secretary of

State of the State of California,

11 September 1940.

Paul Peek, Secretary of State

By Chas. J. Hagerty, Deputy

BYLAWS

Incorporating Amendments dated 16 February 1951; 28 November 1953; 29 October 1955; 15 Feb¬

ruary 1963; 12 November 1965; 15 October 1971; 19 December 1975; 19 January 1979; 16 November

1979; 14 December 1990.

Article I

Offices

Section 1. The principal office for the transaction of the purposes of the corporation is hereby fixed

at the California Academy of Sciences, Golden Gate Park, in the City and County of San Francisco,

State of California. The Executive Board is hereby granted full power and authority to change said

principal office from one location to another in said county.

Article II

Executive Board

Section 1. Subject to the limitations of the Articles of Incorporation, the Bylaws, and the laws of

the State of California, all corporate powers shall be exercised under the authority of the board of

directors who shall be known as and who are herein designated as the members of the Executive

Board of the corporation. Without prejudice to such general powers, but subject to the same limitations,

it is hereby expressly declared that the Executive Board shall have the following powers:

First: to select or remove all other officers, agents, and employees of the corporation, prescribe such

powers and duties for them as may not be inconsistent with law, with the Articles of Incorporation,

or Bylaws, fix their compensation and require from them security for faithful service.

1991

ARTICLES OF INCORPORATION

Second: To make such rules and regulations in respect to the management of the affairs of the

corporation not inconsistent with law, with the Articles of Incorporation, or Bylaws, as they deem

best.

The action of the members of the Executive Board shall be subject to the holders of the membership

of the Society and none of its acts shall conflict with action authorized by such membership.

Sec. 2. The authorized number of members of the Executive Board shall be eleven (11) until changed

by an amendment of the Articles of Incorporation or by an amendment of this section as provided

by law.

Sec. 3. The Executive Board members shall consist of the chairpersons of standing committees, the

officers of the Society, and the editors of The Pan-Pacific Entomologist. In the event that one person

holds two or more of the above positions, additional members-at-large shall be elected in the same

manner as the regular elective officers, to fill the Executive Board of eleven (11) members. The members

of the Executive Board who shall be officers of the Society shall serve until the next annual meeting

of the members of the Society, and until their successors are duly elected and qualified. The members

of the Executive Board who serve as such because of their positions as chairpersons of the standing

committees shall serve until their successors have been appointed by the president and confirmed by

the Executive Board, and have qualified.

Sec. 4. Meetings of the members of the Executive Board for any purpose or purposes shall be called

at any time by the president, or if he is absent, unable or refuses to act, by any three (3) members of

the Executive Board. Notice of the time and place of any such meetings shall be given so that it will

be delivered to the Board members or to their addresses at least twenty-four (24) hours prior to the

time fixed for such meetings but preferably seven (7) days or more in advance.

Sec. 5. The Executive Board shall fill, by appointments, any vacancies occurring in any elective

office between annual elections.

Sec. 6. At least six (6) members of the Executive Board shall be necessary to constitute a quorum

for the transaction of business. Every act or decision done or made by a majority of the members of

the Executive Board shall be the act of the Executive Board unless a greater number be required by

law or by the Articles of Incorporation.

Article III

Officers

Section 1. The officers of the Society shall be a president, president-elect, managing secretary,

recording secretary, and treasurer.

Sec. 2. At the last regular meeting prior to the annual meeting, a nominating committee of at least

three members shall be appointed by the president from among those members who are not holding

any elective office at the time and who did not serve on the nominating committee of the previous

year. It shall be the duty of this committee to nominate candidates for the offices to be filled at the

annual meeting. Nominations may likewise be made from the floor.

Sec. 3. The president, president-elect, managing secretary, recording secretary, and treasurer shall

be elected by a majority of the members present at the annual meeting to serve for one (1) year, their

term in office beginning with the close of the annual meeting at which they were elected, and until

their successors are elected and qualified.

Sec. 4. The president and president-elect shall hold office for no longer than two consecutive years

but are eligible for re-election after the lapse of one year from the time of leaving office. The managing

secretary and treasurer shall be selected with a view to continuity of tenure of their offices, and hence,

other things being equal, may be re-elected each year, subject to their own desires in the matter.

Sec. 5. No member shall hold more than one elective office at one time.

Sec. 6. Vacancies occurring in any elective office between annual elections shall be filled by action

of the Executive Board.

Article IV

Duties of Officers

Section 1. The president shall preside at all meetings of the Executive Board and regular meetings

of the membership. He/she shall appoint temporary committees including a nominating committee,

appoint members to fill vacancies in standing committees, act as ex officio member of all standing

committees, appoint a new Program Committee and its chairperson soon after taking office and during

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

his/her term of office may replace any members as necessary, arrange details of meetings in cooperation

with the Program Committee, and call any special meetings. The president shall, at the regular meeting

prior to the annual meeting, appoint an Auditing Committee, no member of which is an officer of the

Society, to review the financial records of the Society and to report to the members of the Society at

the annual meeting. At the annual meeting he/she shall appoint two (2) members to the Publication

Committee to replace retiring members, and name the Chairperson, report on the state of the Society,

and deliver an annual address.

Sec. 2. The president-elect shall assume all responsibilities of the president in the event of the

temporary absence of the latter. The president and president-elect being absent, a chairperson may

be selected pro tempore from among the members present. The temporary chairperson shall conduct

such business as is absolutely necessary, postponing consideration of other matters until such time as

the regular officers may be in attendance.

Sec. 3. The managing secretary shall send out notices of all meetings at the order of the president,

handle all correspondence of the Society, and keep a record of past and current members. He/she shall

maintain a supply of copies of the Articles of Incorporation and the Bylaws, including new amendments,

and shall furnish a copy to each new member of the Executive Board, and to other members of the

Executive Board at their request. The recording secretary shall keep a fist of members and visitors

present at each meeting, and of the proceedings of said meetings, and submit an edited manuscript

of the proceedings for each year to be published preferably in the October number of the following

year’s Pan-Pacific Entomologist.

Sec. 4. The treasurer shall handle all financial matters of the Society and shall maintain bank accounts

in the Society’s name. Current receipts, unless otherwise specified, become a part of the general funds

of the Society, administered by the Executive Board through the treasurer who is hereby granted

authority to expend the money necessary to send out notices of meetings and to defray expenses, send

out bills, etc., in connection with publications of the Society and such other necessary expenses of the

Society as are approved by the Executive Board.

Article V

Committees

Section 1. Standing committees shall include the following: Publication Committee, Program Com¬

mittee, Historical Committee, and Membership Committee. The president shall be invited to each

meeting of a standing committee as an ex officio member. The chairperson shall call the meetings and

preside. The membership of these committees may be reviewed at any meeting of the Executive Board,

which may make such changes as may be deemed necessary. Committee vacancies occurring between

Executive Board meetings may be filled by presidential appointment, subject to confirmation by the

Executive Board at its next meeting.

Sec. 2. The Publication Committee shall be responsible for the publications of the Society. It shall

consist of six members appointed by the president for three-year terms, groups of two expiring in

successive years. The Publication Committee shall meet at least once a year before the annual meting

to formulate an annual report for the Society. Four members shall constitute a quorum.

Sec. 3. The Historical Committee of five members shall cooperate with the California Academy of

Sciences in accumulating historical entomological material to be deposited with the Academy.

Sec. 4. The Membership Committee of three members shall consult with applicants or prospective

members, acquaint them with the purposes of the Society and with the obligations incurred with

membership, and consider such persons and propose their names, if satisfactory, to the Society for

election to membership. It shall further be the duty of the committee to consider members’ applications

for student status and to review the membership from time to time for persons to be nominated as

Honored Members. The Committee shall meet at least once, early in the year, to establish a new list

of nonmembers, including organizations, to whom Society meeting announcements shall be sent, in

an effort to increase membership and meeting attendance. The chairperson shall provide the managing

secretary with a copy of the list and retain a copy for the committee records.

Sec. 5. The Program Committee shall plan the various meetings, including the field days, arrange

for speakers, encourage members to bring exhibits, and contribute short notes and observations, and

improve the meetings of the Society in other ways. The chairperson shall call all meetings of the

committee after consultation with the president'. The committee shall inform the managing secretary

of the Society as to the programs of the meetings in sufficient time to permit the issuing of the specific

notices of the meetings.

1991

ARTICLES OF INCORPORATION

Sec. 6. Prior to the annual meeting of the Society, the chairperson of each standing committee shall

contact the members of his/her committee and determine whether or not each wishes to continue his/

her committee service for another year. Each chairperson shall then immediately notify the president,

in writing, whether or not members have resigned for the following year. Committee vacancies known

at this time shall be filled by appointment by the president not later than the next regular meeting of

the membership.

Article VI

Members

Section 1. Membership shall be open to persons interested in insects or other Arthropoda.

Sec. 2. An applicant for membership will communicate with a member of the Membership Com¬

mittee and pay his or her dues in advance. The Membership Committee shall then nominate the

applicant for membership at the next regular meeting of the Society. An affirmative vote of two-thirds

of the members present at any meeting is required, whereupon the applicant becomes a member. The

chairperson shall furnish the managing secretary and treasurer with the names and mailing addresses

of the newly-elected members.

Sec. 3. The Executive Board shall fix the annual dues for regular members and student members,

the annual subscription rate for The Pan-Pacific Entomologist, and the prices for back issues of The

Pan-Pacific Entomologist and other Society publications. Changes in dues, rates, and prices shall be

announced at the next regular meeting of the Society and published in the proceedings. The dues are

payable in advance. Each additional member (called a family member) in the immediate family of a

regular member may pay dues of only $ 1.00 per year, but this shall entitle said family to only one

subscription to The Pan-Pacific Entomologist.

Sec. 4. Members at retirement with at least ten (10) years of active membership may, on request,

be continued as active members without payment of dues. Moreover, such members, if they desire,

may receive The Pan-Pacific Entomologist upon payment of one-half regular membership dues per

year.

Sec. 5. Students registered at any school or recognized institution of higher learning are eligible for

student membership. Student members shall have all the privileges of regular members.

Sec. 6. Any member who, through long and distinguished service to the Society, is deemed worthy

of special recognition may be elected an Honored Member. Candidates for honored membership shall

be nominated by the Membership Committee at the Society meeting prior to their election by the

membership, or they may be elected directly by the Executive Board, with announcement of the

election made at the next Society meeting and recorded in the Society’s proceedings. Such members

shall be free from the payment of dues or other financial obligations but shall retain all privileges of

regular members.

Sec. 7. All members, except retired members or as herein otherwise provided, shall receive The

Pan-Pacific Entomologist with no additional charge.

Article VII

Meetings

Section 1. At least four regular meetings shall be held each year, preferably with two in the fall and

two in the spring, with an annual Field Day or equivalent. Regular notice of such meetings shall be

given to each member at his or her request either personally or by mail, or other means of commu¬

nication, charges prepaid, addressed to such member at his or her address, if any, appearing upon the

books of the Society, or given by him or her to the Society for the purpose of notice. All such notices

shall be sent to the members not less than ten (10) days before such meeting.

Sec. 2. Special meetings may be called at any time at the discretion of the president, or shall be

called at the request of five members, provided that written notice is sent to all members at least two

weeks in advance of the meeting.

Sec. 3. The December meeting of each year shall be known as the Annual Meeting. At this time,

in addition to the election of officers and the transacting of other regular and special business, annual

reports of all committees shall be submitted as well as an annual report of the treasurer.

Sec. 4. The presence in person of ten (10) members of the Society at any regular or special meeting

shall constitute a quorum for the transaction of business, but less than a quorum may adjourn any

meeting from time to time and the meeting may be held as adjournment without further notice.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(1)

Article VIII

Delegates

The Society may accredit delegates, appointed by the president, to national and international meet¬

ings such as the American Association for the Advancement of Science and the International Ento¬

mological Congresses. Members may be accredited by a two-thirds vote of the members present at

any meting.

Article IX

Amendments

These Bylaws may be amended or repealed by the Executive Board, subject always to the power of

the members to change or repeal such Bylaws; or by the vote or written assent of a majority of the

members; or by a majority of a quorum at a meeting of the members duly called for the purpose of

amending the Bylaws.

PAN-PACIFIC ENTOMOLOGIST

67(1): 7-19, (1991)

OMANONABIS : A NEW GENUS OF NABIDAE

(HETEROPTERA) FROM WESTERN NORTH AMERICA,

WITH A REVIEW OF NABIS ED AX BLATCHLEY

Adam Asquith and John D. Lattin

Systematic Entomology Laboratory, Department of Entomology,

Oregon State University, Corvallis, Oregon 97331

Abstract. —Omanonabis NEW GENUS is proposed for the species Nabis lovetti Harris. Distin¬

guishing features including illustrations of genitalia and scanning electron micrographs are pro¬

vided. Nabis edax Blatchley is reviewed and distinguishing features, including illustrations of

genitalia, are provided. Its morphology is compared with that of other species of Nabis and its

taxonomic position is reevaluated.

Key Words. — Insecta, Heteroptera Nabidae, Omanonabis, Nabis lovetti, Nabis edax

Two species of Nabidae found along the Pacific coast of North America have

puzzled heteropterists for some time, Nabis lovetti Harris and Nabis edax Blatch¬

ley. Harris recognized the distinctiveness of lovetti when he described it from

California and Oregon in 1925, noting especially the dorsal pubescence, the spine¬

like tubercles on the genital capsule and the lance-like parameres. Even though

Harris recognized subgenera of Nabis in his monograph of North American Nabi¬

dae (1928), he left lovetti in the nominal subgenus Nabis despite its unique char¬

acters. Although Mitri (1960) suggested that lovetti might deserve separate sub¬

generic placement based on its distinctive seminal depository, Henry & Lattin

(1988) left it in the subgenus Nabis pending clarification of its proper position.

Nabis edax was described by Blatchley from Los Angeles, California in 1929,

just after the appearance of Harris’ monograph (1928). No other references to the

species were published until Henry & Lattin (1988) retained it in the subgenus

Nabis based on information received from Dr. Kerzhner of Leningrad. However,

Mitri (1960) illustrated the female genitalia of a specimen from Alameda County,

California, that could not be placed in any recognized species of North American

Nabis; we now believe that this specimen represents N. edax. In this paper, we

compare the morphology of these two species with other members of the genus

Nabis and discuss their probable relationships, creating a new genus for N. lovetti.

Omanonabis Asquith & Lattin, NEW GENUS

(Figs. 1-7)

Type-Species.—Nabis lovetti Harris 1925:205 (California).

Description. —Macropterous male: Large, broadly linear, widest across middle of clavus; length (apex

of tylus to apices of hemelytra) 7.9-9.1 mm. Head: Short, eyes large; ocelli raised above surface of

vertex, but not prominent; head testaceous with broad median longitudinal black stripe. Ventral surface

of head yellow, with two broad, longitudinal, pale fuscous stripes either side of midline. Antennal

segment I slightly curved laterally, distal half slightly enlarged, yellow with ventral surface infuscated,

with long, inclined, yellow setae. Antennal segment II linear, distinctly thinner than segment one,

yellow with apex fuscous to black, with semierect, yellow setae. Segment III linear, testaceous; segment

IV slightly but evenly enlarged distally, testaceous to black; both segments III and IV with semierect

yellow setae. Pronotum: Campanulate, broader than long, widest across posterior margin; lateral

Figure 1. Omanonabis lovetti (Harris). Dorsal habitus.

1991

ASQUITH & LATTIN: NEW GENUS OF NABIDAE

Figure 2. Hemelytral setae, a-b. Omanonabis lovetti. c-d. Nabis roseipennis Reuter.

margins slightly sinuous; posterior margin straight or slightly convex; posterior angles bluntly rounded;

lateral and posterior margins weakly carinate. Collar distinct, length equal to width of eye, weakly

punctate. Dorsal surface of anterior lobe with paired transverse rectangular areas either side of midline,

surface slightly depressed, granulose. Posterior lobe with minute shallow punctures, dull yellow or

testaceous, with fuscous median stripe; surface with short, reclined, golden setae and elongate, erect

hairs. Scutellum broad, wider than long, two distinct fovea present either side of midline; fuscous,

with anterior comers yellow; surface with short reclined golden setae and elongate erect hairs. Hemely-

tra: Elongate, tapering to apices; sides slightly arcuate, extending beyond end of abdomen; all veins

distinctly elevated; anterior surface minutely, shallowly punctate; membrane with three closed cells,

middle cell very narrow, gray-yellow, veins pale, bordered with fuscous; dorsal surface densely covered

with short, reclined, golden setae (Fig. 2). Venter: Connexivum very broad, dorsolateral and midventral

fuscous stripes running length of ventral surface. Lateral and ventral surface of abdomen with long,

silver, silky pubescence. Channel of ostiolar peritreme very broad, surface smooth, dorsal half of

posterior margin elevated and convex (Fig. 3). Tibiae relatively short. Apical pads on pro- and

mesotibiae large, reaching to base of second tarsal segment. Profemora swollen, tapering distally.

Mesofemora slightly enlarged, tapering distally; ventral surface with numerous long black teeth. Meta¬

femora thick but not swollen, short, just reaching end of abdomen, slightly curved posteriorly. All

appendages dotted with fuscous. Base of claws slightly swollen, occasionally with blunt tooth at base.

Genitalia: Genital capsule extremely large, length equal to all anterior segments combined; anterolateral

surface with small tooth on either side, oriented anteriorly (Fig. 4a). Posterior surface of genital capsule

concave, with posterior comers rounded and slightly enlarged. Parameres very narrow and elongate

with apex strongly recurved dorsally (Fig. 4b); lateral margin with strong keel; base of lateral surface

with broad band of erect pale setae. Aedeagus long and narrow, twisted at middle, with four large

sclerites (Fig. 5).

Brachypterous Male.—As macropterous male, except: more oval in outline, total length 7.2-8.3

mm. Pronotum more evenly triangular, less campanulate; posterior lobe narrower. Hemelytra narrow,

connexivum visible in dorsal view, reaching to anterior margin of genital capsule; apex narrowed to

blunt point; membrane with only two closed cells. Hind wings reaching to 4th or 5th abdominal

tergite.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Figure 3. Left metathoracic scent gland of Omanonabis lovetti.

Female.— Similar to male, slightly smaller (Table 1), total length 7.2-8.6 mm. All antennal segments

shorter than male. Hemelytra of macropterous form reaching to last abdominal tergite. Genitalia:

Seminal depository oblong oval, basally constricted (Fig. 6); oviducts arise separately from dorsal

surface, separated by a raised sclerotized structure; spermatheca prominent, arising baso-medially and

directed basally to cover base of ovipositor. Two paired, elongate-oval sclerotized rings present on

ventro-lateral surface.

Diagnosis. — This genus is readily distinguished by the short, dense, golden

pubescence on the hemelytra (Fig. 2a); the broad, flat, anterior surface of the

ostiolar peritreme (Fig. 3); the extremely large male genital capsule with spines

present on the anterolateral margins (Fig. 4a), and the elongate, lanceolate par-

ameres (Fig. 4b).

Etymology. — This genus is named in honor of our esteemed colleage, Dr. Paul

Oman, Professor Emeritus of Entomology at Oregon State University. He has

shared with us his vast knowledge of entomology and his friendship.

Omanonabis lovettii (Harris) 1925:205, NEW COMBINATION

Type. — Holotype male, Nabis lovetti Harris. CALIFORNIA. SAN FRANCIS¬

CO Co.: Ingleside, 20 Dec 1920, H. Dietrich; deposited U.S. National Museum

of Natural History, Washington, D.C.

Redescription. —Male. — Most characters as in generic description. Genitalia: Aedeagus with four

large sclerites (Fig. 5); basalmost sclerite smallest, linear, gradually tapering distally with apex oriented

towards apex of aedeagus; second sclerite just basad of twist in aedeagus, linear, slightly curved, widest

at base, tapering to a sharp apex, oriented towards apex of aedeagus; third sclerite largest, linear, blade¬

shaped, distal half incompletely bifid into two blades, oriented towards apex of aedeagus; distalmost

sclerite just basad of sclerotized “cap,” comb-shaped, with three large teeth of unequal shape and size.

Aedeagus with two patches of minute denticles, one area just basad of middle twist, the other basad

and underneath sclerotized “cap.”

Distribution. — This species occurs along the Pacific coast from southwestern

1991

ASQUITH & LATTIN: NEW GENUS OF NABIDAE

Figure 4. Genitalia of Omanonabis lovetti. a. Male genital capsule, dorsal view. b. Paramere,

lateral view.

British Columbia (Scudder 1985) south to Monterey, California (Harris 1925,

1928) (Fig. 7). It is most common in the coastal salt marshes, and in the Willam-

ette-Puget Lowlands, but is also found in a few marshy habitats east of the Cascade

Mountains in southern Oregon. This species has also been reported from Lake

Provo, Utah (Torre-Bueno 1934). This locality is greatly disjunct from the known

range of lovetti and needs to be confirmed.

Figure 5. Genitalia of Omanonabis lovetti, Aedeagus.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Figure 6. Genitalia of Omanonabis lovetti, Seminal depository, a. Ventral view. b. Dorsal view.

Material Examined.— USA. CALIFORNIA. DEL NORTE Co.: Lake Earl, 24 Nov 1959, J. Schuh

(AMNH). MARIN Co.: 1.7 km (1 mi) S of Inverness, 11 Dec 1953, P. D. Ashlock, salt marsh; Point

Reyes, 21 Aug 1952, P. D. Ashlock; (5 mi) SW of Point Reyes Sta., 14 Nov 1952, J. D. Lattin.

MONTEREY Co.: Carmel, 27 May 1922. SAN FRANCISCO Ca.Ingleside, 20 Dec 1920, H. Dietrich.

SAN MATEO Co.: San Bruno Mts, 16 Aug 1969, R. P. Rapp. SONOMA Co.: Sonoma. OREGON.

BENTON Co.: Corvallis, 1918, A. L. Lovett (Harris 1925); Corvallis, lOOct 1957; MacDonald Forest,

1 Oct 1975; McFadden Pond, 16.7 km (10 mi) S of Corvallis, 17 Oct 1967, P. Oman; Peavy Arboretum,

21 Oct 1980, G. Cassi\ sweeping; 5.8 km (3.5 mi) NE of Summit, 5 Oct 1961, J. D. Lattin; Willamette

Riv, 12 Oct 1977, J. D. Lattin, ex willow; Winkle Lake, 16.7 km (10 mi) S of Corvallis, 8 Oct 1959,

J. D. Lattin. COLUMBIA Co.: Scappoose, 16 Apr 1958, K. Gray; Vemonia, 21 Apr 1936, K. Gray.

CURRY Co.: South Slough Sanctuary, S of Charleston, 24 Sep 1988, A. Asquith. DESCHUTES Co.:

Indian Ford, 6 Jun 1957, B. Malkin. JACKSON Co.: Buckhom Mineral Springs, 18.3 km (11 mi) ESE

of Ashland, 853 mm (2800 ft), 19 May 1960, J. D. Lattin. KLAMATH Co.: Klamath Falls, Algoma,

10 May 1955, J. Schuh; Lost River, 13.3 km (8 mi) SE ofDairy, 4 Aug 1966, W. Gagne & J. Haddock.

LANE Co.: 6.7 km (4 mi) N of Noti, 19 Mar 1963, J. D. Lattin; Siltcoos Park, 2 Nov 1968, M. Stock;

Winchester Bay, 13 Jun 1947, B. Malkin & I. M. Newell. LINCOLN Co.: 5.8 km (3.5 mi) NE of

Harlan, 7 Apr 1960, J. D. Lattin; South Beach St. Park, 1.6 km (2 mi) S of Newport, 7 Nov 1988,

A. Asquith, in deflation plain; Yachats, flood plain of Yachats Riv, 14 Apr 1970, P. Oman. POLK

Co.: 4H Education Center, 7 Jul 1973, L. Russell. TILLAMOOK Co.: Cape Lookout St. Park, 11 Sep

1988, A. Asquith; 1.6 km (2 mi) SE of Pacific City, 1 May 1973, J. D. Lattin; Sandlake, peat bog, 20

Sep 1962, K. Goeden; 1.6 km (2 mi) S of Sand Lake, 13 Jun 1972, W.N. Mathis; hwy 101, N of Siletz

Riv, 11 Sep 1988, A. Asquith. WASHINGTON Co.: Hillsboro, Apr 1919, L.P. Rockwood, ex skunk

cabbage. WASHINGTON. KING Co.: Bothell, 28 Apr 1964; Lake Sammamish St Park, 6 May 1969,

R. Levenson.

The Relationships of Omanonabis

We have seen neither the dense pubescence nor the teeth on the genital capsule

in any other genus of nabids, and these characters may be unique to Omanonabis.

The setae are not unusual in their structure and are only slightly larger in diameter

than those of Nabis roseipennis (Fig. 2). Also distinctive is the broad, flat anterior

surface of the peritreme channel. In species of Nabis and Halonabis the channel

is clearly much narrower (Fig. 11).

Omanonabis does not appear to be related to any other North American genus

of Nabini. Although it shares several characters in common with Palaearctic

genera, we suspect that these characters are plesiomorphic. For example, several

to many small sclerites in the aedeagus are found in the Palaearctic genera Hi-

macerus, Aptus, Stalia, Halonabis, Aspilaspis and in Omanonabis. In almost all

1991

ASQUITH & LATTIN: NEW GENUS OF NABIDAE

species of Nabicula and Nabis, the small distal sclerites have been lost, and one

or two large basal sclerites are retained (Fig. 8). In some species (N. ( Reduviolus )

alternatus ) even the two basal sclerites have been lost (Fig. 9). We believe that

the coalescence and loss of the distal sclerites in Nabicula and Nabis is more

parsimonious than several independent origins of mutliple, small sclerites. There-

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Table 1. Comparison of macropterous and brachypterous male and female Omanonabis lovetti.

Measurements are given in millimeters as mean plus range.

Measurement

Jc (range)

Male

Female

Macropterous

Brachypterous

Macropterous

Brachypterous

Total length

8.7

(7.9-9.1)

7.8

(7.2-8.3)

8.2

(8.0-8.6)

7.4

e i.i-i

■6)

Pronotum length

1.78

(1.60-1.88)

1.68

(1.55-

-1.79)

1.74

(1.69-1.81)

1.61

(1.55-

-1.67)

Pronotum width

2.26

(2.04-2.36)

2.02

(1.91-

-2.14)

2.22

(2.13-2.28)

1.93

(1.85-

-1.99)

Head width

1.03

(0.95-1.01)

1.04

(0.99-

-1.09)

0.97

(0.92-1.01)

0.99

(0.95-

-1.02)

Antennal segment I

0.90

(0.83-0.95)

0.92

(0.87-0.96)

0.78

(0.53-0.87)

0.86

(0.77-0.89)

Antennal segment II

1.46

(1.37-1.51)

1.56

(1.06-

-1.41)

1.32

(1.17-1.27)

1.22

(1.15-

-1.29)

Antennal segment III

1.30

(1.26-1.40)

1.31

(1.06-

-1.41)

1.21

(1.17-1.27)

1.22

(1.15-

-1.29)

Antennal segment IV

1.04

(0.92-1.10)

1.07

(1.03-

-1.22)

1.03

(1.01-1.03)

1.06

(0.96-

-1.18)

fore, although Omanonabis retains several distal sclerites, it cannot be grouped

with the other genera that display this morphology, but it can be removed from

the genus Nabis, which has lost this condition.

Superficially, the parameres of Omanonabis closely resemble those of Halonab-

is. In the latter genus, however, a wide, longitudinal groove is present which begins

basally on the ventral margin of the paramere, but turns dorsal to run along the

lateral margin distally. In Omanonabis, this groove is present, but it is much

narrower and is restricted to the ventral margin of the paramere. Besides lacking

the other diagnostic features of Omanonabis, Halonabis also differs by lacking

spinules on the ventral surface of the fore femora, having proportionately much

larger eyes and having the apical third of the head weakly curved ventrally. Species

of Aspilaspis also have a rather elongate, linear paramere. If the lanceolate par¬

amere is homologous, then Omanonabis may be related to Halonabis and Aspi-

lapsis. The two Palaearctic genera are Mediterranean in distribution and are also

ground inhabiting, preferring moist habitats (Pericart 1987).

The Relationships of Nabis edax

Nabis edax Blatchley was described from a single male specimen from Los

Angeles, California (Blatchley 1929). This species has not been reported since,

and was apparently known only from the specimen used in the original description.

Based on an examination of this specimen, Kerzhner (reported in Henry & Lattin

1988) placed this species in the subgenus Nabis, with N. roseipennis Reuter and

rufusculus Reuter. Here, we provide a description of the female of N. edax, detail

the male and female genitalia, and reassess its taxonomic position.

In an unpublished Master’s thesis dealing with the female genitalia of the Nabi-

dae, Mitri (1960) figured the seminal depository of a specimen from Milpitas,

Alameda Co., California. He noted that the structure of the seminal depository

was unlike that of any other North American species for which females were

known. Recently, additional single females were discovered in the collections of

the California Academy of Sciences and the University of California, Berkeley

that also possess this unique seminal depository structure. Mitri (1960) stated that

he could not discern sclerotized rings in this species, but presumed they were

present and merely lightly sclerotized. We have reexamined the specimen he

dissected and one additional specimen. We also could not identify sclerotized

1991

ASQUITH & LATTIN: NEW GENUS OF NABIDAE

Figures 8-9. Aedeagus. 8. Nabis ( Nabis ) roseipennis. 9. Aedeagus of Nab is ( Reduviolus ) alternatus.

rings, although a thin structure at the ventral surface of the base may represent

the vestiges of the rings. In addition, male specimens of N. edax were also dis¬

covered. All specimens were collected from central and southern California, from

San Francisco to Los Angeles.

There are two possible explanations for this scenario. First, that there are two

species in southern California, N. edax, known only from male specimens, and

an undescribed species, known only from female specimens. We consider this

very unlikely, because most North American Nabidae have relatively large ranges,

and two closely related species confined to the same restricted area would be most

unusual. Second, that the female specimens are N. edax. We believe this to be

the case, because both male and female specimens share combinations of char¬

acters not possessed by any other North American species.

Taxonomic Position. — Kerzhner assigned edax to the subgenus Nabis (Henry

and Lattin 1988) with the other two North American members, N. roseipennis

Reuter and N. rufusculus Reuter. However, N. roseipennis has two sclerites in the

aedeagus both with large hooks at the base (Fig. 8), and N. rufusculus has two

large sclerites at the base and one small sclerite at the distal end of the aedeagus.

The seminal depositories of roseipennis and rufusculus are broadly spindle shaped,

and both have a single, large, bilobed sclerotized ring covering the anterior and

antero-ventral surface of the depositories. N. edax shares no characters in common

with these species and therefore clearly is not a member of the subgenus Nabis.

Remane (1964) designated several species groups in the genus Nabis, two of

which contain species with a single sclerite in the aedeagus similar to N. edax. In

the rugosus group, N. brevis Schultz has only a single sclerite, but it bears a tooth

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Figure 10. Dorsal view of the pronotum of Nabis edax Blatchley. Female.

near its base and it is situated near the base of the aedeagus. In addition, this

group contains predominately short-winged forms, and N. edax is known only

from macropterous specimens. In the ferus group, both N. ferns (L.) and N. palifer

Seid. have but a single sclerite. In ferus, however, the sclerite has a large, well

developed hook at the base, and in palifer, the sclerite is very small, only half the

length of that in edax. In addition, the seminal depository of the ferus group is

large, membranous, with a large sclerotized ring that occupies most of the antero-

ventral surface of the seminal depository.

The subgenus Reduviolus Kirby which includes N. alternatus Parshley and N.

americoferus Carayon, is predominately a North American group, with the ex¬

ception of N. inscriptus (Kirby) which is Holarctic in distribution. The female

depository in this group is short and symmetric with a small thin sclerotized ring

on the postero-ventral surface, very similar to N. edax. However, all members of

the inscriptus group lack any sclerites in the aedeagus.

It appears that N. edax Blatchley cannot be placed in any of the taxonomic

subdivisions presently recognized in the genus Nabis. It is most similar to members

of the subgenus Reduviolus, but is not closely related to any of these North

American species. Examinations of a few Mexican and South American species

of Nabis also failed to find any which appear to be related to N. edax. Rather

than create a new subgeneric taxon for N. edax, we believe it more prudent simply

to recognized its uniqueness and label this species incertae sedis within the genus

Nabis pending a more detailed and broader geographic study of the genus.

Nabis edax Blatchley

(Figs. 10-14)

Type. —Holotype male. Data: “California]: Los Angeles, W.S.R., 12-6-27/ Pur¬

due Blatchley Collection/ TYPE/ LECTOTYPE Nabis edax Blatchley, Des. W.S.

Blatchley, 1930.” In the Purdue University Insect Collection.

1991

ASQUITH & LATTIN: NEW GENUS OF NABIDAE

Figures 11-14. Nabis edax 11. Left metathoracic scent gland. 12. Female seminal depository,

dorsal view. 13. Left paramere in lateral (a) and medial (b) views. 14. Aedeagus.

The Lectotype designation by Blatchley was never validated in a publication.

Although Blatchley did not designate a holotype in the original description, the

specimen should now be considered the holotype by monotypy (ICZN Recom¬

mendation 73F, ICZN 1985).

Redescription.—Female.— Medium sized species, total length (7.1-7.8 mm), broadly linear in out¬

line, widest just posterior of clavus. Head: Porrect, longer than wide; eyes large, dorsal edges reaching

just above vertex, ocelli prominent; pale yellow, area posterior of eye fuscous, ventral surface of head

immaculate; antennae linear, I (0.87-1.02 mm), II (1.51-1.69 mm), yellow, segment I with longitudinal

fuscous stripe on medial surface, fuscous stripe on dorsal surface of segment II. Pronotum: Broad,

length (1.43-1.64 mm), posterior width (1.77-2.04 mm), posterior width-collar width ratio >2.5,

strongly campanulate, abruptly expanded posteriorly with comers angular (Fig. 10), posterior lobe

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(1)

elevated, with numerous shallow punctures, posterior margin distinctly carinate, pale yellow. Collar

distinct, longer than distance from posterior margin of eyes to posterior margin of head, anterior

portion shallowly punctate. Scutellum very broad, wider than long, middle depressed and margins

slightly elevated, pale yellow. Hemelytra: Shiny, smooth or minutely granulose, anterior one third of

clavus and corium with small shallow punctures; clavus short and broad, straw yellow, small fuscous

spot at apex of claval suture; veins of membrane indistinct, membrane with indistinct, broad fuscous

stripe, more defined apically. Dorsum covered with long, inclined yellow setae. Venter: Ventral surface

pale yellow. Metathoracic scent gland distinct, elongate oval, apex evenly rounded (Fig. 11). Legs

immaculate or with light brown spots at base of setae. Fore and hind femora without black teeth or

denticles on ventral surface, middle femora occasionally with a single short row near apex. Hind legs

more robust than other Nabis spp.; hind tibia pilose, with erect hairs, the longest of which are almost

2 x the width of the tibia. Seminal depository short, thick and symmetric (Fig. 12), strongly wrinkled

along the sides and base; oviducts wide, arising from a meso-dorsal position on the seminal depository.

Male. — Similar in size and shape to females but much darker, with color patterns more distinct.

Length (7.18 mm). Head: Grey on dorsum, ventral surface testaceous. Pronotum: Less wide or cam-

panulate than female, length (1.54 mm) posterior width (1.86 mm), with a dark fuscous middorsal

stripe. Scutellum also with wide dark fuscous median stripe. Hemielytra: Midline of membrane with

an irregular, broad fuscous or smoky stripe. Propleuron fuscous; abdomen with testaceous to fuscous

lateral stripe. Meso- and metafemora and tibiae with fuscous spots. Genitalia: Paramere most similar

to that of N. rufusculus (Fig. 13). Apex produced, with a short, squared tip. Ventrolateral surface with

a wide row of erect setae, following curve of ventral margin of paramere. Aedeagus with a single large

sclerite without a tooth at its base (Fig. 14), situated in distal half of aedeagus.

Diagnosis. — To serve as a diagnosis, we have revised a portion of Harris’ key

to North American Nabidae (1928) to include N. edax :

19. Head beneath in greater part fuscous to black; posterior tibiae dotted

with fuscous; brachypterous form with closed cells in the membrane;

male paramere with long sinuate stem . roseipennis Reuter

Head beneath in greater part yellow to testaceous; posterior spotted or

immaculate; male paramere with short rectangular stem . 20

20. Maximum width of posterior femora > distance between eye and ocellus;

ventral surface of mesofemora lacking black denticles or with only a

short row near apex; posterior tibiae of male spotted with fuscous;

aedeagus with a single sclerite; posterior tibiae of female pilose, longest

setae nearly at right angles with axis of tibia and 2 x its width; seminal

depository without sclerotized ring; coastal California .... edax Blatchley

Maximum width of posterior femora < distance between eye and ocellus;

ventral surface of mesofemora with black denticles; posterior tibia of

male usually immaculate; aedeagus with two sclerites; posterior tibiae

of female not pilose, all setae short; seminal depository with sclerotized

ring present, wide spread . 20a

20a. First antennal segment thickened distally, somewhat sinuate above; bra¬

chypterous form common, usually with no closed cells in membrane;

diameter of blade of male paramere greater than that of an eye viewed

from above . rufusculus Reuter

First antennal segment scarcely thickened distally, from above almost

straight; brachypterous form unknown; diameter of blade of male

equal to that of an eye . kalmii Reuter

Material Examined. -USA. CALIFORNIA. CONTRA COSTA Co.: Pittsburg, 18 Nov 1923, J.D.

Martin. FRESNO Co.: Tranquillity, Fresno Slough, 8 Jul 1962, P. F. Torchio. LOS ANGELES Co.:

Los Angeles, Hancock Park, 6 Dec 1927, W. S. Blatchley; Milpitas, 23 Jun 1955, (Acc. No. 121), P.

F. Torchio. SANTA CLARA Co.: San Jose, 30 Sep 1954, D. R. Bale. COUNTY UNCERTAIN: South

Eastern California, 8 Jun 1962, light trap, USDA.

1991

ASQUITH & LATTIN: NEW GENUS OF NABIDAE

Acknowledgment

For assistance in examining specimens in their care and their suggestions we

thank P. H. Amaud (California Academy of Sciences, San Francisco); the late P.

D. Ashlock (University of Kansas, Lawrence); T. J. Henry (USDA, Systematic

Entomology Laboratory, Washington, D.C.); I. M. Kerzhner (Zoological Institute,

Leningrad, U.S.S.R.); J. A. Powell (University of California, Berkeley); R. T.

Schuh (American Museum of Natural History, New York); R. L. Westcott (Oregon

State Department of Agriculture) and D. W. Bloodgood (Purdue University, West

Lafayette, Indiana). We also thank B. B. Hall for the fine habitus drawing.

Literature Cited

Blatchley, W. S. 1929. Two new Heteroptera from southern California (Cydnidae, Nabidae). Entomol.

News, 40: 74-76.

Harris, H. M. 1925. A new species of Nabidae (Costa) from the Western United States (Hemiptera).

Entomol. News, 36: 205-206.

Harris, H. M. 1928. A monographic study of the Hemipterous family Nabidae as it occurs in North

America. Entomol. Amer., 9: 1-98.

Henry, T. J. & J. D. Lattin. 1988. Family Nabidae. pp. 508-520. In Henry, T. J. & R. C. Froeschner

(eds.). Catalog of the Heteroptera or true bugs of Canada and the United States. 1988. J. C.

Brill, Lieden, Netherlands.

International Code of Zoological Nomenclature. 1985. (3rd ed) International Trust for Zoological

Nomenclature (BM[NH]). University of California Press, Berkeley, California.

Mitri, T. K. 1960. The taxonomic value of the external female genitalia in the genus Nabis Latreille

(Hemiptera: Heteroptera: Nabidae). Master’s Thesis, Oregon State University.

Pericart, J. 1987. Hemipteres Nabidae: D’Europe Occidentale et du Maghreb. Faune de France.

France et regions limitrophes. 71. Federation frangaise des Societes de sciences naturelles.

Remane, R. 1964. Weitere Beitrage zur Kenntnis der Gattung Nabis Latr. (Hemiptera Heteroptera

Nabidae). Zool. Beitr. N.F., 10: 253-314.

Scudder, G. G. E. 1985. Heteroptera new to Canada. J. Entomol. Soc. Brit. Columbia, 82: 66-71.

Torre-Bueno, J. R. 1934. On some Heteroptera from the West and Southwest. Bull. Brooklyn

Entomol. Soc., 29: 155-157.

Received 8 June 1990; accepted 24 September 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 20-23, (1991)

LARVAL DEVELOPMENT OF DARGIDA PROCINCTA

(GROTE) (LEPIDOPTERA: NOCTUIDAE) IN THE

PACIFIC NORTHWEST

J. A. Kamm

National Forage Seed Production Research Center,

U.S. Department of Agriculture, Agricultural Research Service,

Corvallis, Oregon 97331-7102

Abstract.— Larval growth of D. procincta (Grote) was regulated by temperature and larvae re¬

quired 20.5 days at 21° C to complete larval development vs 51.2 days at 15° C. Duration of

the pupal stage was 11.7 days at 27° C vs 36.7 days at 15° C. Photoperiod had no observable

effect on larval development. The six instars were readily distinguished by the width of head

capsule. Third instars, observed in the field in early April, are believed to emerge as adults in

June and lay eggs for a second generation. Dense infestations of larvae were observed to develop

on the lush spring growth of grasses but not on the fall regrowth of established or seedling stands

of ryegrass.

Key Words.— Insecta, cutworm, grass pest, larval growth

Dargida procincta (Grote) is one of two representatives of the genus in the

continental United States (Holland 1903, Hodges 1983). In the Pacific Northwest,

the larvae are known to defoliate ryegrasses (Kamm 1985). Legumes are also host

plants (Tietz 1972). The seasonal flight of adults is unknown because they are

rarely captured in blacklight traps, even in fields with dense populations of larvae

(Kamm 1985). Dyar (1898) and Godfrey (1972) briefly described the larvae and

Spencer (1946) observed the oviposition behavior of females. Thompson (1943)

considered the species an important pest of grasses in the Pacific Northwest.

Dargida procincta was the most abundant cutworm species in commercial ryegrass

grown for seed, but nothing is known about its larval development or the number

of generations each year (Kamm 1985).

This paper reports the results of laboratory tests that document the number of

instars and the influence of photoperiod and temperature on the rate of larval

growth. Field observations were made and are discussed in relation to the seasonal

cycle of D. procincta in ryegrass grown for seed in the Willamette Valley of Oregon.

Materials and Methods

A laboratory culture of D. procincta was initiated from larvae collected in a

field of ryegrass near Corvallis, Oregon. All larvae were reared individually in

stender dishes (5.0 x 2.5 cm), the bottoms of which were lined with damp blotting

paper. Larvae were supplied with freshly cut leaves of ryegrass, Lolium multiflo-

rum Lamarck, every 1-3 days. Controlled environment chambers were used to

rear the stock cultures and to provide experimental treatments of light and tem¬

perature. A short day was a regime of 10 h light and 14 h dark (10:14 LD) and

a long day was a regime of 16 h light and 8 h dark (16:8 LD). Adults obtained

from larvae reared in the laboratory were confined in cages (20 x 20 x 50 cm)

on ryegrass grown in the greenhouse to mate and to obtain eggs. Eggs laid on the

1991

KAMM: LARVAL DEVELOPMENT OF DARGIDA

foliage were removed and kept in stender dishes until they hatched. Experimental

treatments were initiated the day of egg hatch.

Head capsule size was the criterion used to determine larval instars. Measure¬

ments were made across the widest part of the head capsule in dorsal view by

using an ocular micrometer in a stereoscopic microscope. Head capsules were

measured every other day during the test. Adults were dissected at different ages

to determine the stage of ovarial development by methods described elsewhere

(Kamm & Ritcher 1972). Specimens of reared adults have been deposited in the

insect museum at Oregon State University. A sweep net was used to sample fields

for larvae.

Results and Discussion

Description of Eggs and Larvae . — Females glue milky white eggs to the under¬

sides of grass leaves and within leaf sheaths. The eggs are pumpkin shaped. Several

days before hatch they develop a brown ring that encircles the egg, with a brown

dot in the center of the ring. Each ovary has four ovarioles about twice the length

of the abdomen. No mature eggs are visible in the ovaries when females emerge,

but after females were fed a solution of 10% honey and water for a week, eggs of

various sizes were present in the ovaries and females initiated oviposition. The

ovaries removed from females collected in the field during July and August con¬

tained mature eggs.

Dargida procincta is described as being green with pale stripes or black with

the pale stripes distinct (Crumb 1956, Godfrey 1972). When a large lot of larvae

was reared in the laboratory to fourth instars, the basic body color was numerous

shades of green (110), black (59), or brown (14). The color of the pale stripes was

usually tan or brown, which may or may not have interrupted black markings

that parallel the stripes. Because D. procincta is a leaf feeder, the different color

morphs probably conceal some larvae from predators during seasonal changes in

color of the grass.

Larval Growth. —Figure 1 shows the cumulative rates of growth of larvae ex¬

posed to a long-day regime at 21° C. Each instar was easily discerned by mea¬

surement of the head capsule. Progression through the instars was remarkably

uniform, both in time between molt and the size of head capsule, except for the

sixth instar. Unlike many Lepidoptera larvae (Beck 1980), photoperiod had no

significant effect on the rate of larval growth at either 15 or 21° C (Table 1). No

evidence was found that exposure of small larvae to short days and/or low tem¬

peratures induced diapause in either larvae or pupae. In temperature treatments

with the same photoperiod, the rate of larval and pupal development nearly

doubled (P < 0.05) when larvae were exposed to 15° C compared with those

larvae exposed to 21° C.

Another experiment confirmed that larval growth was regulated by temperature

when larvae were exposed to an expanded range of temperatures (Table 2). The

duration of development for larvae, from egg to pupal stage, ranged from 20.5

days to 51.2 days. The duration of the pupal stage was 11.7 days at 27° C vs 36.7

days at 15° C. The ability of D. procincta to develop in cool temperatures and

short daylengths without diapause or apparent pathological effect gives the species

the advantage of using the early spring and autumn growth of cool-season grasses

in the Pacific Northwest. During winter, larvae have been observed to feed during

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(1)

Table 1. Effect of photoperiod at 15° and 21° C on the development of D. procincta larvae from

time of egg hatch to adult.

Temperature

No. of

X Time to develop (days)

Photoperiod

insects

Larvae

Pupae

Total (± SD)

16:8 LD

29.5

16.9

46.4 ± 1.9 a

10:14 LD

28.8

18.5

47.3 ± 2.2

16:8 LD

54.4

34.4

88.8 ± 2.8

10:14 LD

52.0

39.7

91.7 ± 2.9

a Rate of development of larvae exposed to the same temperature and different photoperiods were

not significantly different, but that of larvae exposed to the same photoperiod and different temperatures

was significantly different (Mann-Whitney U- test, P < 0.05). Each treatment was initiated using 80

larvae.

warm periods in January. Second and third instars were easily collected on ryegrass

foliage where they feed by sweeping fields during early April and are believed to

be first-generation larvae that mature and pupate in ryegrass fields windrowed for

harvest in mid-June (Kamm 1985). The presence of gravid females from May

through August suggests a second generation of larvae in late summer. A few

mature larvae that developed during the typical arid summer have been collected

from September to December and are believed to be the wintering stage. In no

case have dense larval infestations been observed in established or seedling fields

of ryegrass during October and November suggesting that the species primarily

exploits the lush spring growth of grass.

c n

3.8

3.4

2.6

2.2

1.8

1.4

6TH

0 0 0

0000000 000

0000 0

000000000

0000000000 5TH

00000000000

0000000

4TH

000000

0000000000000

000000000

0000000000

2ND

3RD

0000000000

1ST

1 5 9 13 17

AGE OF LARVAE (DAYS)

Figure 1. Progressive increase in the size of head capsule of larvae of D. procincta exposed to long

daylengths at 21° C.

1991

KAMM: LARVAL DEVELOPMENT OF DARGIDA

Table 2. Effect of temperature on rate of development when D. procincta larvae were exposed to

different temperatures and long days (16:8 LD) from time of egg hatch to adult.

Temperature (°C)

No. of insects 0

X Time to develop (days)

Larvae

Pupae

Total (± SD)

20.5

11.7

32.2 ± 2.1

24.6

14.9

39.5 ± 1.2

25.0

16.4

41.4 ± 0.8

35.9

24.4

60.3 ± 3.3

51.2

36.7

87.9 ± 2.7

a Each treatment was initiated using 60 larvae.

Acknowledgment

This article is a contribution of the USDA-ARS in cooperation with the Ag¬

ricultural Experiment Station, Oregon State University, Technical Paper 9069 of

the Agricultural Experiment Station, Oregon State University.

Literature Cited

Beck, S. D. 1980. Insect photoperiodism (2nd ed.). Academic Press, New York.

Crumb, S. E. 1956. The larvae of Phalaenidae. U.S. Dept. Agric., Tech. Bull., 1135.

Dyar, H. G. 1898. Descriptions of the larvae of fifty North American Noctuidae. Proc. Wash.

Entomol. Soc., 4: 315-332.

Godfrey, G. L. 1972. A review and reclassification of the subfamily Hadeninae (Lepidoptera: Noc¬

tuidae) of America North of Mexico. U.S. Dept. Agric., Tech. Bull., 1450.

Hodges, R. W. 1983. Checklist of the Lepidoptera of America, North of Mexico. E. W. Classey Ltd.

and Wedge Entomological Res. Foundation, London.

Holland, W. J. 1903. The moth book. Doubleday, Page & Co., New York.

Kamm, J. A. 1985. Cutworm defoliators of ryegrass. Pan-Pacifc. Entomol., 61: 68-71.

Kamm, J. A. & P. O. Ritcher. 1972. Rapid dissection of insects to determine ovarial development.

Ann. Entomol. Soc. Am., 65: 271-274.

Spencer, G. J. 1946. On the oviposition habits of Dargida procincta (Lepidoptera: Phalaenidae).

Proc. Brit. Col. Entomol. Soc., 43: 10.

Thompson, B. G. 1943. Cutworm control in Oregon. Oregon Agric. Exp. Sta., Circ. 147.

Tietz, H. M. 1972. An index of the described life histories, early stages and hosts of the macrolepidop-

tera of the continental United States and Canada. Allyn Museum of Entomology, Sarasota,

Florida.

Received 6 May 1990; accepted 20 October 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 24-27, (1991)

A NEW DROSOPHILA SPECIES IN THE MONTIUM

MEIJERE SUBGROUP OF THE MELANOGASTER MEIGEN

SPECIES-GROUP IN THE SUBGENUS SOPHOPHORA

STURTEVANT FROM TAIWAN

(DIPTERA: DROSOPHILIDAE)

Shun-Chern Tsaur and Fei-Jann Lin 1

Institute of Zoology, Academia Sinica, Nankang, Taipei, Taiwan 11529,

Republic of China

Abstract.— A new Drosophila from Taiwan, D. austroheptica NEW SPECIES is described and

illustrated. It belongs to the melanogaster species-group, and is closely allied to D. quadraria

Bock & Wheeler.

Key Words. — Insecta, Diptera, Sophophora, Drosophila austroheptica

The Drosophila montium Meijere subgroup was first established by Hsu (1949:

121) with the following characters: “A large tooth-bearing secondary clasper pres¬

ent, seemingly originated by separation from anal plate; marginal bristles of pri¬

mary clasper greatly enlarged.” This diagnosis was subsequently defined more

completely by Bock & Wheeler (1972).

The subgroup is by far the largest in the melanogaster species-group and pres¬

ently accomodates 79 taxa (Lemeunier et al. 1986), occurring from the Oriental

to Afrotropical regions. Seven species in the subgroup have hitherto been recorded

from Taiwan: D. ashahinai Okada, D. auraria Peng, D. bocki Baimai, D.jumbulina

Parshad & Paika, D. kikkawai Burla, D. lini Bock & Wheeler, and D. quadraria

Bock & Wheeler (Chen and Lin, unpublished data). This paper describes an

additional species, D. austroheptica.

Materials and Methods

All the isofemale laboratory culture stocks available for this study were obtained

by field collections in Nanjenshan (Renting National Park) by either sweeping on

grasses or using beer-banana traps beside streams. Stocks were reared on standard

com meal medium at 22° C and 75% RH. During the study, fresh specimens were

dissected directly in a weak solution of phenol, but pinned flies were first boiled

in IN potassium hydroxide for several minutes until clear and then washed in

tap water. All drawings were made with a Nikon Optiphot® drawing tube at the

same magnification (ca. 200 x). All dissected parts were slide-mounted.

The terminology employed in the descriptions is largely patterned after that

proposed by Sturtevant (1942), although more standardized terminology for the

Diptera is noted within parentheses.

Drosophila (Sophophora) austroheptica

Tsaur & Lin, NEW SPECIES

(Fig- 1)

Types. — Holotype: male; data: REPUBLIC OF CHINA. TAIWAN. Taipei:

Nankang, 25 May 1990, taken from isofemale line stock culture 0211.24 in the

1 To whom reprint request should be sent.

1991

TSAUR & LIN: NEW SPECIES OF DROSOPHILA

Figure 1. Genitalia and egg of Drosophila ( Sophophora) austroheptica NEW SPECIES. A. genital

arch (female), lateral view. B. caudal view (male). C. lateral view (male). Phallic organs: D. lateral

view of phallic organ (male). E. ventral (V) and dorsal (D*) views of phallic organs (male). F. egg. G.

egg guide (female). H. decastemum (male). (Scale lines = 1 mm.)

Institute of Zoology, Academia Sinica (originally provisioned with field material

from Nanjenshan, Pingtung Hsien, 13 Nov 1989, S. C. Tsaur and C. T. Ting).

Paratypes: 10 males, 10 females; same data as holotype. Holotype and paratypes

deposited in the Institute of Zoology, Academia Sinica, Taipei, Taiwan, Republic

of China.

Male. — Body length: 2.3 mm (live specimens); wing length: 1.9 mm. Head: Antennae, second

segment black-yellow, with two large setae, several medium to small setae, third segment darker.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Arista with four branches above, three below plus a terminal fork. Front pale brown, with a transverse,

yellow band between antennae orb-3 (proclinate orbital). Periorbitals moderately small, four to six.

Orbitals: orb-1 (posterior reclinate orbital) obviously nearer to orb-3 than to inner vertical bristles;

orb-2 (anterior reclinate orbital) situated between orb-1 and orb-3, closer to orb-1 than to orb-3 about

1:2.5, more than one-third length of orb-1, less than one-third length of orb-3. Three oral vibrissae,

second almost same length of first, t hi rd about two-thirds length of others. Carina yellow-white,

moderately elevated. Face dark brown. Palpi dark brown, with single apical and subapical bristles

pointed downward. Proboscis entirely yellow-brown. Ocellar triangle dark brown. Ocellars normal,

yellow. Ocelli orange. Eyes red, piled, slightly oblique to body axis. Maximum width of cheek about

one-seventh maximal diameter of eye. Mesonotum and scutellum: Shining brown, lacking markings.

Acrostichal hairs in six rows. Dorsocentral bristles with anterior four-fifths the length of the posteriors,

both sets parallel; cross distance between anterior dorsocentrals about 2 x distance from anteriors to

posteriors. Prescutellar bristles absent. Pleurae brown, lacking stripes. Humerals two, lower longer

than upper, about 1.5:1. Sternopleural bristles three. Stemo-index about 0.6. Anterior scutellar bristle

parallel to convergent, slightly shorter than posteriors. Posterior scutellar bristles crossed at mid¬

length. Halters tan. Legs yellow-brown, inner side of fore femur with three long bristles from mid¬

length to subapex, divided in equal length. Preapical bristles on all tibiae, apical bristles on first and

second tibiae. Sex-comb longitudinal along entire length of first and second tarsal segments; first tarsal

comb consisting of about 24 uniform teeth; comb on second tarsal segment with a similar row of

about 14 teeth, all nearly same length. Abdominal tergites: Brown-yellow, segments II-IV black,

segment V with or without inconspicuous yellow-brown band, all bands occupied varying from apical

one-fifth to one-half, except segment V that is only slightly pigmented posteriorly. Some stout bristles

on posterior margin of tergites in regular rows. Wings: Slightly grey diffused, veins brown-black. Second

vein (R 2+3 ) straight. Third (R 4+5 ) and 4th veins (M, +2 ) parallel. Two C-l bristles. C-3 fringe on its

basal half. Approximate indices: C index 2.1; 4c index 1.4; 4Y index 2.3; 5x index 2.2. Periphallic

organs: Genital arch (epandrium) broad laterally, ventral lobe with one long, four to five moderately

long bristles, without triangular process covering base of primary clasper. Primary (surstylus) and

secondary (ventral margin of cercus) claspers present. Primary clasper with a lateral row of five to six

small pointed teeth and medial cluster of about 11 larger teeth, one stout and elongated. Secondary

clasper with three large, curved black medial teeth and several fine setae ventrally and dorsally. Anal

plate dark brown, with long bristles. Median lobe of decastemum bicomuted. Phallic organs: Brown.

Aedeagus broadly rounded and finely hirsute at apex, slightly constricted subapically, lacking pointed

lateral expansions. Anterior parameres large, well separated, broadly subtriangular, without minute

apical sensilla. Posterior parameres spoon-shaped, long, surpassing tip of aedeagus. Ventral phragma

narrowed medially, shovel-shaped. Caudal margin of novastemum (hypandrium) with hexagonal

median truncated process, apically bearing two short spines.

Female. — Body length: 2.4 mm (live specimens); wing length 2.1 mm. As in male except for sexual

dimorphic characters, and abdominal tergites of segments II-V with dark brown bands. Egg guide

(oviscapt): Yellow-brown, with about 13 teeth and a subterminal hair. Egg filaments: Two long slender

filaments.

Pupae.— Anterior spiracles with 10 branches. Horn index: 0.08.

Diagnosis. —Drosophila austroheptica can be separated from the other Dro¬

sophila of the montium subgroup in Taiwan by its unique combination of the

following characters: secondary clasper with only two teeth and both the caudal

margin of the novastemum with a hexagonal medium truncated process plus the

posterior paramere extending slightly past the tip of the aedeagus.

Life Cycle. —One generation takes about 14 days at 22°C, 75% RH.

Distribution. — Only known from the type locality in Taiwan.

Etymology. — The specific name indicates that it has been collected from Southern

Taiwan.

Material Examined. — (Maintained Cultures ) Stock cultures in Academia Sinica: 0211.24 to 0211.25

(2 stocks, isofemale lines, collected from Nanjenshan, Pingtung Hsien, 13 Nov 1989 by S. C. Tsaur

and C. T. Ting).

1991

TSAUR & LIN: NEW SPECIES OF DROSOPHILA

Acknowledgment

We sincerely thank Miss Chau-Ti Ting (National Taiwan University) for as¬

sistance in collecting specimens and laboratory work. This work has been sup¬

ported by a grant from the National Science Council (NSC 79-0211-BOO 1-31).

Literature Cited

Bock, I. R. & M. R. Wheeler. 1972. The Drosophila melanogaster species group. Univ. Texas. Publ.,

7213: 1-102.

Hsu, T. C. 1949. The external genital apparatus of male Drosophilidae in relation to systematics.

Univ. Texas. Publ., 4920: 80-142.

Lemeunier, F., J. R. David, L. Tsacas & M. Ashbumer. 1986. The melanogaster species group, pp.

147-256. In Ashbumer, M., H. L. Carson & J. N. Thompson, Jr. (eds.). The genetics and

biology of Drosophila. Vol. 3e. Academia Press, London.

Wheeler, M. R. & H. Takada. 1964. Insects of Micronesia. Vol. 14, No. 6. Diptera: Drosophilidae.

Bernice P. Bishop Museum, Honolulu, Hawaii.

Wheeler, M. R. 1981. The Drosophilidae: a taxonomic overview, pp. 1-97. In Ashbumer, M., H.

L. Carson & J. N. Thompson, Jr. (eds.). The genetics and biology of Drosophila. Vol. 3a.

Academia Press, London.

Received 27 July 1990; accepted 1 November 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 28-54, (1991)

PHYLOGENETIC CHARACTER RESPONSES FOR SHAPE

COMPONENT VARIANCE DURING THE MULTIVARIATE

EVOLUTION OF EULACHNINE APHIDS:

REDESCRIPTION OF PSEUDESSIGELLA HILLE RIS

LAMBERS (HOMOPTERA: APHIDIDAE: LACHNINAE)

John T. Sorensen

Insect Taxonomy Laboratory, California Department of Food & Agriculture,

Sacramento, California 95814

Abstract. — Although suitable for conventional taxonomic purposes, previous descriptions of the

monotypic aphid genus Pseudessigella Hille Ris Lambers and its species Pseudessigella brachy-

chaeta Hille Ris Lambers are inadequate to characterize important phylogenetic trends in char¬

acter evolution in the Eulachnina; therefore, these taxa are redescribed. The evolutionary re¬

sponses of morphometric traits for Pseudessigella, Essigella (as a whole) and two primitive

Essigella species are deduced and interpreted with reference to Lande’s models of multivariate

phenotypic evolution. Twenty-six traits among these taxa were analyzed for their shape com¬

ponent variance using principal components analysis; ranks for the relative contribution of each

trait upon the dominant two shape vectors were assigned on the basis of character loading

coefficients. Between the genera, 42% of the traits were considered to be stable in their partitioned

shape variance over the implied evolutionary episode; 39% of the traits showed moderate sta¬

bility, while 19% were judged to be unstable. The average similarity of evolutionary response

for the traits on the dominant shape vectors between these genera was 0.56; the primitive Essigella

species showed higher similarities (0.71, 0.62) with Pseudessigella than did Essigella overall.

When discriminant function analysis was used as a measure of Landean minimum selective

mortality to determine the multivariately optimal traits that separate Essigella species, both

traits with predominantly general-size variance and traits with predominantly shape variance

appeared to be important in species separation. The most influential traits with the highest shape

variance on the discriminant vectors, however, appeared to largely separate clades of species

(i.e., species-groups or above) within the genus, rather than chiefly separating species within such

groups.

Key Words. — Insecta, multivariate evolution, phylogenetic characters, natural selection, principal

components analysis, discriminant function analysis

The monotypic aphid genus Pseudessigella Hille Ris Lambers and its type-

species, P. brachychaeta Hille Ris Lambers, were described from a single collection

in central Asia. This interesting species represents a phylogenetic intermediate

between Eulachnus del Guercio and Essigella del Guercio within the subtribe

Eulachnina (Sorensen 1990), and was used as an out-group to root the phylogenetic

network devised for Essigella (Sorensen 1987a). Because of Pseudessigella' s phy¬

logenetic position, it has assumed a new importance for the study of character

transformations within the Eulachnina, and for Essigella in particular. As a result,

Hille Ris Lambers’ (1966) descriptions for Pseudessigella and P. brachychaeta,

although sufficient for previous taxonomic and diagnostic purposes, now fail to

adequately characterize the variation, within that genus, for several traits that

have been found to be important in evolution at the species and species-group

levels in Essigella (Sorensen 1983, 1988).

This article, and Sorensen (1990), deal with the phylogenetic aspects of the

Eulachnina. The latter provided an analysis of discrete, coded phylogenetic traits

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

and determined the phylogenetic sequence among the genera of the subtribe. This

study provides analyses that indicate the multivariate evolutionary interaction of

traits in both Pseudessigella and Essigella, and it serves as a base for future studies

of character evolution and transformations within Essigella (unpublished data).

The paper also redescribes Pseudessigella and P. brachychaeta in the form pre¬

viously used for descriptions of Essigella species (Sorensen 1988), and that will

also be used in a pending generic revision of Essigella (unpublished data).

Taxonomy

Pseudessigella Hille Ris Lambers, 1966

Pseudessigella Hille Ris Lambers 1966, Tijdschrift voor Entomologie, 109: 219.

Type Species. —Pseudessigella brachychaeta Hille Ris Lambers 1966, Tijdschrift

voor Entomologie, 109: 219-220; by monotypy.

Redescription. — Viviparous Apterae. Body elongate, linear, with few hairs. Antennae five segmented,

processus terminalis short. Head very slightly wider than long, unfused with pronotum. Eyes without

distinct triommatidia. Rostrum retractile; last rostral segment not subdivided but short, blunt. Meso-

thoracic notum lightly but entirely sclerotic. Metathoracic notum mostly membranous laterally and

posteriorly but with a lightly sclerotized subhemispherical shaped patch anteromesally. Abdominal

dorsum with terga I-VII membranous with dorsal and marginal setae on defined sclerotic bases;

tergum VIII lightly but entirely sclerotic. Siphunculi as rimmed pores on short truncated sclerotic

cones, hairless, base well defined from membranous field of abdominal terga. Basitarsus with five

ventral setae, lacking dorsal setae in apterae; metabasitarsus ventrally 1.5 x as long as dorsally. Cauda

rounded, lacking median protuberance. Profemora dorsoproximad base moderately swollen. Tarsal

claws simple, not incised or bifid; tip sharp, relatively drawn out.

Other Morphs. —Unknown.

Diagnosis. —Pseudessigella is separated from Eulachnus by its five segmented

antennae, and from Essigella by its simple unincised tarsal claws.

Pseudessigella brachychaeta Hille Ris Lambers, 1966

(Figs. 1-3)

Pseudessigella brachychaeta Hille Ris Lambers 1966. Tijdschrift voor Entomo¬

logie, 109: 219-220.

Types.—ELolotype— viviparous aptera; data: (“WEST”) PAKISTAN. Murree,

4 Jul 1964, R. van den Bosch (P-VII-4d), Pinus griffithii McClelland. Paratypes—

viviparous apterae and nymphs. Holotype and paratypes slides deposited British

Museum (Natural History), London, ex Hille Ris Lambers aphid collection.

Redescription. — Viviparous Apterae. Morphology (n = 14, measurements and counts as: range [Jc ±

SD]): Body length 1.950-2.505 (2.225 ± 0.206) mm. HEAD—Primary rhinarium on terminal antennal

segment (V) not exceptionally distad, distance from tip of processus terminalis to distal face of rhinarial

rim slightly greater than 0.5 x diameter of rhinarium, rhinarial membrane not conspicuously pro¬

tuberant. Length of: antennal segment V 98-118 (106 ± 6) um, processus terminalis 25-40 (34 ± 5)

Mm, IV 75-95 (87 ± 5) III 180-220 (204 ± 11) um, II 73-85 (79 ± 4) pm. Longest frontal

setae 5-15 (12 ± 5) pm long, tips incrassate. Head width 226-390 (300 ± 46) pm. Length of: stylets

475-542 (509 ± 24) pm, ultimate rostral segment 70-93 (74 ± 6) pm\ rostral tip reaching slightly

past mesocoxae. Head and pronotum not fused, their combined length 390-447 (422 ±21) pm.

THORAX—Length of: mesothorax 304-371 (331 ± 17) ftm, metathorax 63-93 (76 ± 10) pm. Meso-,

metathorax and abdominal segment I not fused. ABDOMEN—Maximum distal width of flange on

1991 SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS 31

ABC

D E F G H

Figure 2. Expression of ventral abdominal sclerites on abdominal segments II-IV in Pseudessigella.

The sclerites, which serve as muscle attachment points, vary from a single, subelliptical to linear

sclerite (A-D), often with constrictions (B-D), to a bipartite (H) or tripartite (E-G) series of sclerites,

in which the anterior (E) or usually the central (F, G) sclerite is diminished. (Shading indicates

sclerotization; no transformation is implied.)

siphunculi 23-28 (24 ± 2) jum; siphunculi truncated sclerotic cones, protruding to 0.5 x maximal

distal width. Ventral abdominal sclerite shape on segments II-IV (Fig. 2) variable, often linear with

posterior enlarged but anterior narrow, sometimes broken into two to several minor plates: the posterior

then subquadrate, subcircular to subelliptical and the anterior plate(s) smaller, more irregular, size

0.2-0.5 x posterior plate and more variable in shape from subquadrate, subcircular or subelliptical

to asterisk-like, mere specks or absent; entire linear sclerite (or series of broken platelets) 35-95 (63

± 19) jum long, 1.0-2.0 x metatibial diameter. Dorsal setae on abdominal terga II-IV (Fig. 3) dividable

into major and minor series; six major dorsal setae across medial portion of tergum of each abdominal

segment, arranged in roughly transverse irregular, meandering or staggered row, positions (if numbered

left to right) of setae number 1, 3, 4 and 6 nearly in straight row, but setae number 2 and 5 slightly

to moderately anterior of that row, on relatively larger (12-15 u m diameter) scleroites (sclerotized

platelets at setal bases), with incrassate tips; minor dorsal setae on relatively smaller scleroites (8-10

m m diameter), like marginal setae but present mesally between lateral most dorsal abdominal muscle

DMAS (mes)

Figure 3. Setal positions on dorsum of abdominal terga II-IV in Pseudessigella. The abdominal

segment shown is slightly skewed to the right showing more of its left lateral face; items are indicated

on one side only. Dorsal muscle attachment sclerites occur as three sclerites on each side of the segment

and are represented by a single mesad sclerite, DMAS (mes), and a pair of laterad sclerites, DMAS

(lat); the dark bar oriented through the laterad sclerites on the left indicates the “territorial” line that

is used here to separate (define) the minor dorsal versus marginal setae. Dorsal setae occur in two

series, a transversely oriented and staggered set of six major setae, DS (maj), which cross the center

of the segment, and a cluster of no to four minor setae, DS (min), on each side of the segment just

mesad to the lateral dorsal muscle attachment sclerites. Marginal setae, MS, occur as a set of two to

six setae laterad of all dorsal muscle attachment sclerites. Spiracles, S, that occur ventrally may be

confused with other sclerites in slide preparations.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(1)

attachment plates and slightly mesad to them, all tips incrassate, arranged in loose cluster(s) antero-

laterally and also sometimes posterolateral of major dorsals, each side with 0-4 (1.7 ±0.8) component

setae. Marginal setae (lateral to lateral most dorsal abdominal muscle attachment plates) with incrassate

tips, on small scleroites (8-10 Atm diameter), 4-12 per segment, each side with 2-6 (4.6 ± 0.6),

apparently arranged in two to four subgroupings. Abdominal tergum VIII with 10-16 (14.3 ± 1.8)

hairs, 10-15(11 ±2) Atm long, tips incrassate, subregularly placed. Cauda rounded, caudal protuberance

absent, longest caudal setae 40-80 (67 ± 10) Atm long, tips sharp. LEGS—Length of: metafemora 561-

646 (624 ± 30) Atm, metatibiae 637-741 (700 ± 30) Atm; longest dorsal setae on central one-third of

metatibiae 8—18(12 ± 3) Atm long, 0.1-0.5 x metatibial diameter, tips incrassate, approximately equal

or very gradually increasing distally, no setal length dimorphism; longest ventral setae on metatibiae

23-30 (27 ± 3) Atm long, tips sharp. Length of: metabasitarsus 50-88 (69 ± 9) Atm, metadistitarsus:

225-255 (243 ± 8) Atm. Ratio of metabasitarsus to metadistitarsus averaging 1.0:3.6, ranging from

1.0:2.8 to nearly 1:5. Pigmentation. Color in life pale green (Hille Ris Lambers 1966). Prepared

specimens: background of sclerotic body dorsum pale (to approximately 40% pigment density [as solid

black in a 54 line/cm screen]), unicolorous. Frontal setal bases concolorous with surrounding terga,

undifferentiated. Thoracic muscle attachment plates dusky, inconspicuous to conspicuous. Dorsal and

marginal setal bases on abdomen dark, conspicuous, arising from distinct scleroites that contrast the

membranous abdominal field. Dorsal muscle attachment plates of abdomen, spiracular plates, si-

phuncular cones and ventral abdominal sclerites pale, usually dusky light brown (to 60% pigment

density), conspicuous. Cauda, anal and subgenital plates concolorous, slightly less dark than head and

mesothorax. Antennal segments V and IV slightly to moderately dusky over entire segment, to mod¬

erately brown distally, III pale to moderately dusky distally, II concolorous with proximad section of

III and I concolorous with frons. Pro-, meso- and metatibiae dusky, concolorous, slightly more dusky

than remaining sclerotic body dorsum.

Ultimate Stadium Nymphs of Viviparous Apterae. —Nonmorphometries and pigmentation of pre¬

pared specimens as described for viviparous apterae.

Viviparous Alatae, Males, Oviparae, Fundatrices.— Unknown.

Diagnosis.—Pseudessigella brachychaeta is the only species in the genus, and

the only known linear-bodied aphid feeding on the needles of Pinaceae which has

both a five segmented antennae in adults and simple, unincised tarsal claws.

Host.—Pinus griffithii McClelland, stated to be “ Pinus excels aP on the type

series slides and “ Pinus wallichiana [excelsa)” in the original description (Hille

Ris Lambers 1966). Pinus wallichiana A. B. Jacks and Pinus excelsa Wallich ex

Lambert are synonyms of P. griffithii (Critchfield & Little 1966, Mirov 1967, Little

& Critchfield 1969).

Range.— Known from one collection in (“West”) Pakistan at Murree. Potential

native range (see discussion), assumed to be the distribution of P. griffithii (Fig.

4), is throughout Himalaya Mountains to eastern Afghanistan, northeastern Bur¬

ma and Yunnan Province, China (Troup 1921, Vavilov 1959, Critchfield & Little

1966, Mirov 1967); in mid- and high elevation forests, especially in drier inner

valleys (Critchfield & Little 1966). Neither the collection sample nor description

quote an elevation, but Hille Ris Lambers (1966) described several other new

aphid species, also taken by van den Bosch in a sequence of field collections at

Murree between 30 Jun-5 Jul 1966, and cited elevations of 2132-2284 m (7000-

7500 ft) for those collections. These elevations agree with Mirov’s (1967) notation

that in the “Murree Hills, Rawalpindi,” P. griffithii grows from 1200 m (in cool,

moist conditions) to above 2000 m, where it occurs in pure stands or mixed with

broad-leaf trees.

Etymology.— The species was named for its short setae.

Discussion.—Pseudessigella is phylogenetically intermediate between Eulach-

nus and Essigella, the more ancestral and derived genera respectively, of the

subtribe Eulachnina (Sorensen 1990). Pseudessigella, like Eulachnus, appears to be

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

Figure 4. Potential range of Pseudessigella brachychaeta (= distribution of Pinus grijfithii [Critch-

field & Little 1966]).

paraphyletically defined; although monotypic, Pseudessigella lacks a unique de¬

fining autapomorphy. Pseudessigella shares with Essigella the following apomor-

phic character states: differentiation in size, shape, tip condition or projection

angle of dorsal versus ventral setae on antennal segments and tibiae (partially

shared with Eulachnus ); absence of secondary rhinaria on ultimate and penulti¬

mate antennal segments (a homoplasy with Schizolachnus)\ a reduction of antennal

segments to five in adults; and representation of abdominal tergum VIII as a

single, entire sclerotized field not apparently formed from fused lateral sclerites

associated with dorsal setal bases (see Sorensen 1990). Pseudessigella resembles

Eulachnus in having the following symplesiomorphies: unincised tarsal claws, a

membranous abdominal dorsum, and a head separated from the pronotum in

adult viviparous apterae.

Slide mounted adult viviparous apterae of Pseudessigella are somewhat super¬

ficially similar to late stadium nymphs of the apterae of Essigella, especially the

most plesiomorphic species (e.g., “group I” of Sorensen [1987a]: E. kathleenae

Sorensen and E. kirki Sorensen, although usually darker than both) or slightly

more derived species (e.g., “group II” of Sorensen [1987a]: E. fusca Gillette &

Palmer and its relatives). Adult Pseudessigella differ most noticeably from the

nymphs of any Essigella species (other than in adult genitalic traits) by their

elongated metabasitarsus to distitarsus ratio; averaging 1.0:3.6 for Pseudessigella

versus slightly over 1.0:2.05 for E. kathleenae, the longest among Essigella. This

“long distitarsus” ratio probably represents a relative plesiomorphy for Pseudes¬

sigella (see discussion of analyses below) within the Eulachnina, despite impli-

THE PAN-PACIFIC ENTOMOLOGIST

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cations of Mamontova (1972) to the contrary for the Cinarini, and is more similar

to conditions in Cinara and Schizolachnus.

The tarsal trait is homoplasious among the Cinarini, including the Eulachnina

(Sorensen 1990). In contrast to Pseudessigella, Eulachnus and Essigella appear to

have homoplasiously evolved an apomorphically shorted metatarsal ratio among

species-groups occupying differing niches in varying pine groups (unpublished

data). In at least the case of Essigella this may possibly represent a “resource¬

tracking” attribute (sensu Brooks 1981, see Moran 1986). A more detailed dis¬

cussion of the relationship of Pseudessigella within the Eulachnina is dealt with

in Sorensen (1990), which comments more fully upon character states thought to

be of phylogenetic significance for the genus and discusses Mamontova’s (1972)

contrasting phylogenetic concepts and character transformations, considered er¬

roneous in part, for the eulachnine aphids.

The phylogenetic relationships among (Sorensen 1990), and biological associ¬

ations within (Sorensen 1987a), the eulachnine genera make ecological predictions

possible for Pseudessigella allowing future corroboration. Among the Eulachnina,

Pseudessigella and Eulachnus are Palaearctic, although the latter has been intro¬

duced into the Nearctic where it apparently feeds chiefly on cultivated old world

pines; I have taken Eulachnus only on Palaearctic species of cultivated Pinus in

the western Nearctic (unpublished data), despite intensive collecting for Essigella

on natural and planted stands of native Nearctic pines (Sorensen 1983). Essigella

is Nearctic and shows phylogenetic associations among Pinus, but dissident de¬

rived species (in “group III,” Sorensen [1987a]) also feed upon Pseudotsuga and

Picea. Because nothing is known of the biology of P. brachychaeta, except that

the single collection of it in the Himalayas was on Pinus grifflthii, biological

predictions for Pseudessigella must be based on Eulachnus and especially Essi¬

gella, particularly “groups I and II” (Sorensen 1987a).

Based upon biological knowledge of “group I” Essigella (Sorensen 1983, 1987a,

1988, unpublished data), I suspect that P. brachychaeta shares the rather mo-

nophagous feeding habits of E. kathleenae and E. kirki. It, therefore, seems rea¬

sonable that the range of Pinus griffithii itself (Fig. 4) should be a probable indicator

of the potential native distribution of P. brachychaeta, as has been found in these

group I Essigella. The host of P. brachychaeta, Pinus griffithii, is a haploxylon

pine in the section Strobus, subsection Strobi of Pinus (Strobus ) (Little & Critch-

field 1969). Subsection Strobi also contains the hosts of these two most primitive

Essigella : Pinus lambertiana Douglas for E. kathleenae, and Pinus flexilis James

and P. strobiformis Engelmann (a slightly overlapping allopatric species previously

considered a variation [reflexa Engelmann] of flexilis [Critchfield & Little 1966:

7, map 8]) for E. kirki (Sorensen 1987a, 1988).

Species of Essigella and Eulachnus are fast moving when disturbed, and rela¬

tively solitary; these behavioral traits are assumed to be typical of Pseudessigella

also. I suspect the (now unknown) alatae of Pseudessigella are (will be found to

be) relatively uncommon, because alatae of E. kathleenae and E. kirki remain

unknown despite extensive sampling (Sorensen 1983), although alatae of more

derived Essigella groups, particularly some group III representatives (e.g., E.

californica (Essig) and E. hoerneri Gillette & Palmer) can be quite common.

In previous phylogenetic studies on Essigella, I have referred to P. brachychaeta

using the acronym “PLES” (Sorensen 1983) and “Sp. X” (Sorensen 1987a). In

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

the latter study, which used a maximum-likelihood algorithm (Felsenstein 1984)

to construct a phylogenetic network based upon centroid values on discriminant

functions, E. kirki (Sorensen 1987a: sp. K) was considered to be the most ple-

siomorphic Essigella species because it showed the shortest distance on the phy¬

logenetic pathway to Pseudessigella; E. kathleenae (Sorensen 1987a: sp. J) was

slightly more “advanced” on that network. This study, however, provides evi¬

dence that E. kathleenae, rather than E. kirki, displays several attributes that may

be more plesiomorphic in their similarity to those of Pseudessigella: (a) P. griffithii,

the host of Pseudessigella, is genetically close enough to be one of the few pines

that hybridizes with the Pinus lambertiana (Sorensen 1987a; W. B. Critchfield,

personal communication), the host of Essigella kathleenae : (b) E. kathleenae

displays the relatively highest (more plesiomorphic) ratio of the metadistitarsus

to metabasitarsus within Essigella (Sorensen 1988); and (c), perhaps most con¬

vincing, E. kathleenae, rather than E. kirki, shows a higher similarity value for

shape component variance with Pseudessigella (see multivariate evolution sec¬

tion).

Material Examined. —The type-series viviparous apterae and nymphs have been examined, and

represent the only known specimens.

Multivariate Evolutionary Responses for Characters

In any organism, continuous phenotypic variation is caused by genetically cor¬

related characters that are linked by many covarying genes (Lande 1980). The

evolutionary aspects of such correlated traits are assumed to be predominantly

under the genetic influence (Falconer 1981). Each gene has a relatively minor

influence in the evolutionary change of either the overall phenotype or any par¬

ticular phenotypic attribute, because these changes are due to an integrated, ad¬

ditive genetic effect, plus minor additive environmental influences (Lande 1985,

1988). The evolution of species and character interactions under such quantitative

genetic systems is based upon an (admittedly simplistic) phenotypic model for

the multivariate evolution of correlated morphological character suites developed

by Lande (1979). This model has been employed previously to determine the

phylogenetic relationships among aphids (Sorensen 1983, 1987a) and birds (Schlu-

ter 1984), as well as phenological shifts in phenotype between sibling species of

leafhoppers (Sorensen 1987b, Sorensen & Sawyer 1989).

Lande’s model allows reconstruction of the net gradients of genetic selection

(taken as phenotypic responses where the environmental component is effectively

null) within and between groups with approximately equal genetic covariance

matrices. The model permits an estimation of the degree to which net selective

pressures have acted upon characters within suites of correlated phenotypic traits

during evolution. Under this multivariate model of evolution the traits respond

to given historical net selection gradients, to drive the mean group phenotype for

a species or population through the dimensional multivariate evolutionary space

defined by all such gradients.

For a given individual’s phenotype, a single trait, z i5 contributes to the evolution

of that phenotype and thus itself evolves in relation to other traits, z j3 as (Lande

1979: equation 7b):

AZj = 2 G ii a ln W/dZj

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

where minor selective change on trait z £ is the sole cause of differences in Mal¬

thusian mean fitness (d In W/dz,); and z x shows no correlated effects, whereas other

traits, Zj, are constant with j A i (Lande 1979). Therefore, over each generation,

changes in the trait z ; include genetic gains from selection that act upon both it

and Zj, as other genetically correlated characters. The measurement of selection

on correlated traits is discussed in more detail by Lande & Arnold (1983).

Principal components analysis (PCA) is particularly suited as a practical, ana¬

lytical probe to determine the multivariate evolutionary responses of characters

in covarient suites as they react in unison to historically imposed selection regimes

(Sorensen 1987a, b; Sorensen & Sawyer 1989). In a usual (Q-type) PCA, each

eigenvector represents a historical net natural selection gradient (as: V In W, sensu

Lande 1979) in multivariate evolution; the mean group phenotype of a species

evolves along these gradients (eigenvectors) due to the concert action between its

genetic covariance matrix and the vector as a selection regime. The relative input

weight of each character on any given eigenvector shows the relative amount of

its partitioned and sequestered variance that can be attributed to the covariant

character suite that is defined by that vector (Sorensen 1987b, Sorensen & Sawyer

1989).

During multivariate evolution, the characters in any such covariant suite of

traits respond to the net evolutionary selective pressures of their eigenvector in

an integrated fashion, according to the relative degree and sign of their vector

loading coefficients. All traits belong in varying degrees to all covariant character

suites, with each suite responding to a particular vector’s selection gradient; this

is because the traits each have a contributing (weighted) input on each eigenvector

which varies among the vectors.

During evolutionary response to selection pressures, each covariant suite of

traits is driven chiefly by dominating traits (“drivers”) which are those characters

with the highest absolute loading coefficients (and correlations) on the given vector.

In contrast, traits with near zero loading coefficients on any vector are more

subordinate for that suite and accordingly more neutral in response to the net

selection pressures defined by that eigenvector as an evolutionary selection gra¬

dient. Therefore, when selection is imposed upon any suite, it acts largely upon

the dominating “driver” traits but causes modification of all traits in proportion

to their loading input and correlation for the given vector.

Although stabilizing selection maintains linkages, and thus integrated reaction

to selection, among the traits in each character suite, if enough disruptive selection

force is applied to a given trait in a suite it may cause the “shearing” of that trait

from the suite. The remaining traits in the suite would be initially modified also

by the disruptive selection, along with the trait to which it was applied, but they

would eventually “break-away” as an integrated unit and return to their former

expression values leaving the disrupted strait to evolve separately from the suite.

Zeng (1988) discusses the complicating effects of stabilizing and disruptive selec¬

tion upon characters in multivariate evolution.

Character interaction during multivariate evolution can be examined using an

R-type analysis in PCA. In an R-type analysis, each of the individuals, rather

than the attributes (as in Q-type PCA), provides the input coefficient along the

orthogonal eigenvectors, and the positions of the original characters are plotted

along these vectors in the I-space defined by the values derived from individuals

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

(Williams & Dale 1965, Sneath & Sokal 1973). An R-type analysis thus depicts

spatially the relative degree of association and interaction between each trait along

orthogonal eigenvectors as character, rather than populational, responsive models

of Landean gradients of net natural selection.

R-type PCA also allows a perspective dependant portrayal of the relative in¬

tegrated response of functional (virtual) groups of characters to several indepen¬

dent selection gradients (as orthogonal eigenvectors). Such functional groups of

characters may show an orchestrated response to simultaneously imposed multiple

selection regimes during multivariate evolution. These virtual groups of characters

are recognizable by their relative spatial affiliations when more than one R-type

eigenvector is plotted; the apparent groups of characters may react relatively

similarly to the equally induced selection pressures on the vectors. Such virtual

character groups defined by multiple vector plots may exist over any partial subsets

of the summarized n- 1 eigenvectors.

Because of its usefulness as an analytical probe, R-type PCA was used here to

determine the historical interaction of characters among the taxa during the mul¬

tivariate evolution of the eulachnine aphids.

Methods

Determination of “Shape” as a Character Component.— To determine the rel¬

ative influence and stability of shape component variance among traits during

the multivariate evolutionary episode between Pseudessigella and Essigella, and

within group I (Sorensen 1987a) of Essigella, data were analyzed for all taxa and

compared. Although many controversial criteria and experimental methods have

been advanced to completely partition multivariate “shape” from “size” in data

matrices (e.g., Burnaby 1966; Bookstein 1986, 1989; Bookstein et al. 1985; Hum¬

phries et al. 1981; Rohlf & Bookstein 1987; Somers 1986, 1989; Sundberg 1989)

none have been completely successful, and all adulterate the generated evolu¬

tionary space by various methods, such as shearing. To avoid the philosophical

problems inherent in such methods, especially with reference to existing models

of multivariate evolution, it was decided to represent “shape” here as simply

principal component vectors 2-n, with vector 1 taken as (chiefly) a size factor if

there appeared to be high loadings and correlations for traits such as segmental

body lengths, which normally are overtly general-size dependant.

Material and Data Examined.— Analysis of Pseudessigella brachychaeta em¬

ployed the paratype adult viviparous apterae (n = 14), which represent the only

known collection (see taxonomy: material examined). Analyses of Essigella used

exemplars of adult viviparous apterae that represented the nonclonal, intraspe¬

cific variance shown within E. kathleenae (n= 13, morphometric data in Sorensen

[1983: appendix El, “KATH”]), E. kirki (n = 12, morphometric data in Sorenson

[1983: appendix El, “HOTT”]), and all Essigella species ( n = 255, morphometric

data in Sorensen [1983: appendix El, all acronyms excluding “PLES”]); hereafter,

the use of “Essigella” refers to the collective concept of its pan-generic variance,

including that shown by E. kathleenae and E. kirki. Locality data for examined

material of E. kathleenae and E. kirki is listed in Sorensen (1988); it is available

for all examined material of Essigella in Sorensen (1983) by cross-checking the

collection numbers in appendix El with locations in appendix Al.

All specimens were measured from the slide mounted material, employing the

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Yol. 67(1)

Table 1. Morphometric characters.

Number

Acronym”

Character descriptions

LBODY

length of body, vertex to posterior of abdominal segment VIII ex¬

cluding cauda.

LANT5

length of total antennal segment V.

LANT5PT

length of processus terminalis, proximal rim of accessory rhinaria

to tip.

LANT4

length of antennal segment IV.

LANT3

length of antennal segment III.

LANT2

length of antennal segment II.

LHFRON

length of longest seta on the frons.

WO ANT

(body) width taken at lateral most (outer) portion of the rim of

the antennal sockets.

LSTY

length of the stylets.

LUROST

length of the ultimate rostral segment.

LHEAD

combined length of the unfused head plus pronotum, along the

median dorsal body axis.

L2THOR

length of the mesothoracic notum, along the median dorsal body

axis.

L3THOR

length of the metathoracic notum, represented as a sclerotic plate

in the otherwise membranous metathoracic field, along the me¬

dian dorsal body axis.

WSIPH

maximal distal width of the siphuncular flange.

LVABSC

maximum length of the longest ventral abdominal muscle attach¬

ment sclerite(s) on abdominal segments II-IV (combined length

in the case of bi- or multi-partite subsclerites in quasilinear ar¬

ray).

NHAB2DT

maximum total number of combined major and minor dorsal se¬

tae (between dorsal muscle attachment sites) on abdominal ter-

ga II-IV.

NHAB2M

maximum number of marginal setae (lateral to dorsal muscle at¬

tachment sites) on abdominal terga II-IV.

NHAB8

number of setae on abdominal tergum VIII.

LHAB8

length of longest seta on abdominal tergum VIII.

LHCAUD

length of longest seta on cauda.

L3FEM

length of metafemur.

L3TIB

length of metatibia.

LH3TIBD

length of longest seta along the central one-third of the dorsal sur¬

face of the metatibia.

LH3TIBV

length of longest seta along the central one-third of the ventral

surface of the metatibia.

L3TAR1

length of the metabasitarsus.

L3TAR2

length of the metadistitarsus.

a Acronym for trait used in Sorensen (1983).

26 traits (Table 1) used previously for Essigella (Sorensen 1983, 1987a, 1988).

Measurements were done with a Zeiss compound microscope using an ocular

scale at up to 400 x, and are assumed accurate to 2 gm for the smallest structures

observed. The data for Pseudessigella, E. kathleenae and E. kirki were log-trans¬

formed to minimize the influence of scaling. The data for Essigella, however,

were not log-transformed; analyses of log-transformed versus nonlog-trans¬

formed data for sequestered parts of that matrix suggested that the obtained vector

loading coefficients did not appear to differ significantly.

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

Algorithms and Analyses.—Ml data were subjected to an R-type principal com¬

ponent analysis. Analyses for Pseudessigella, E. kathleenae and E. kirki were

conducted on a Macintosh® computer and used Statview512 + ® (Brainpower,

Inc., Calabasas, California). Analyses for Essigella were conducted on a CDC6400

computer using the algorithm “PNCOMP” (Duncan & Phillips 1980); analytical

checks of smaller subsections of the Essigella data indicated that the output for

PNCOMP was equivalent to that of Statview512+, as used for the other data.

All PCA solutions yielded unrotated, orthogonal vectors, and the derived loading

coefficients for the characters were used to portray each trait’s influence on the

vectors. To determine the multivariate diagnostic value of characters for sepa¬

rating all species within Essigella, that matrix underwent discriminant function

analysis (SPSS version 7 subprogram “DISCRIMINANT,” direct selection, Wilks-X

criterion; Klecka [1975]).

Results of the analysis on Pseudessigella are presented using histogram illus¬

trations of the loading coefficients were created with Excel® (Microsoft Corpo¬

ration, Redmond, Washington). For Pseudessigella, the virtual groupings of char¬

acters that occur in the evolutionary space defined by multiple vectors, were

assessed by transferring the vector loading coefficients derived from the unrotated

factor matrix to MacSpin® (D 2 Software, Austin, Texas), a three-dimensional

X-Y-Z coordinate rotation program on the Macintosh; scattergrams composed

of orthogonal combinations of multiple vectors were then rotated to assess the

potential groupings of characters in the differing rotational perspectives.

To determine the relative importance of each trait upon any generated eigen¬

vector, the vector loading coefficients of the traits were grouped and ranked into

three classes. This was done by dividing the scale between the highest and lowest

absolute (sign-less) vector loading coefficients for a given vector into equal sized

class partitions, and then assigning each trait to its appropriate class, based upon

its loading coefficient for the vector. The values of shape influence for traits were

determined for all taxa by assigning to each trait its greatest ranking on either

vectors 2 or 3 of the PCA (as the dominant shape vectors) for the taxa. The ranking

of traits as multivariate separators within Essigella was similar, except that dis¬

criminant vectors 1-3 were ranked.

Shape Stability between Taxa. — The evolutionary stability of shape components

between pairs of taxa was calculated as intertaxon similarities. These were deter¬

mined by assigning a distance value of 0.5 to any one-rank-step change for a

character between two taxa; a value of 1.0 was assigned for any two-riink-step

change. The summed total distance between any given taxon-pair was then divided

by 26, which represented the total potential distance for changes among all char¬

acters. The derived distance fraction was converted to a similarity fraction by

subtracting it from 1.0, which represented absolute similarity.

Results and Discussion

Histograms in Figs. 5 and 6 depict the relative contributions of the traits for

Pseudessigella, among the character suites defined by vectors 1-6. In those figures,

the trait contributions for each suite are represented as the absolute values for

loading coefficients of the traits on vectors 1-3 (Figs. 5A-C) and 4-6 (Figs. 6A-

C). Despite the log-transformation conducted on the data, vector 1 (Fig. 5 A) seems

to retain a quite high affiliation as a general-size component. The relative influence

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

0.9

0.8

0.7

0.6

1 3 5 7 9 1113151719212325

Figure 5. Histograms indicating each trait’s relative influence in covariant character suites. Each

suite is represented as an orthogonal PCA vector upon which the trait responds in proportion to its

vector loading coefficient value; a black bar indicates positively loading traits, a white bar indicates

negatively loading traits. (A) vector 1, (B) vector 2, (C) vector 3.

of allometric size versus shape, as separable components of total variance, is

illustrated in Fig. 7 A, which compares the percentage of summed absolute vector

loading coefficients for the traits on vector 1 (Fig. 7 A: white), as a size component,

versus vectors 2-6 (Fig. 7A: black), as shape components. The relative contri¬

bution of each trait to size or shape variance is also shown in Fig. 7B, where all

influences have been scaled to 100% rather than represented as the summation

of vector loading coefficients, as in Fig. 7A. In Fig. 7B the contribution of general

size to each trait is shown in white, while its shape contributions are in nonwhite;

shape contributions on vectors 2 and 3 (the two most dominant shape vectors)

are in gray, and those of vectors 4-6 are in black. An approximation of “unique¬

ness,” or differentiation, among the traits is depicted in Fig. 7C, as variance not

due to common factors in the sense of pleiotropy. A spatial portrayal of character

groupings based upon the dominant three shape factors, vectors 2-4, is shown

for Pseudessigella in Figs. 8A-B. Figure 9 depicts, in tabular form, the rankings

of shape variance across all taxa, and the diagnostic value of traits for group

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

0.7

0.6

0.5

0.4

V 4

0.3

0.2

0.1

1 3 5 7 91113151719212325

1 3 5 7 9 1113151719212325

Figure 6. Histograms indicating each trait’s relative influence in covariant character suites. Each

suite is represented as an orthogonal PCA vector upon which the trait responds in proportion to its

vector loading coefficient value; a black bar indicates positively loading traits, a white bar indicates

negatively loading traits. (A) vector 4, (B) vector 5, (C) vector 6.

separation within Essigella. Figure 10 shows the relative similarity in shape com¬

ponent stability of the taxa analyzed.

Multivariate Character Suites within Pseudessigella.—The dominant six vectors

derived from the Pseudessigella matrix accounted for 84.0% of the total variance

(34.1, 15.2, 11.8, 8.9, 7.7 and 6.3%, respectively for vectors 1-6), indicating that

character variation within Pseudessigella is reasonably complex, as found for

various species-groups within Essigella (Sorensen 1983).

Vector 1 (Fig. 5 A) is interpretable as a size factor because most body appendage

lengths (characters 1-6, 11, 15, 21, 22 and 26) load and correlate highly and

positively upon it. Only those traits poorly related or unrelated to size were not

influential on vector 1. Notable exceptions are those traits (13 and 25 as lengths

for the metathoracic nota and the metabasitarsus) which might have been expected

to load more strongly with general-size; trait 7 (frons setal length) loads highly

but negative; traits 8, 9, 13, 14, 16-20, and 23-25 show relatively minor and

often negative relations with general-size.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Vector 2 (Fig. 5B) represents a covariant suite driven largely and positively by

traits 13, 16-18, 24 and 25, and to a lessor extent by traits 15 and 23; the suite

is driven negatively by trait 7 and less so by trait 20. This suite involves the

metathoracic nota, the ventral abdominal sclerites, the number of setae on ab¬

dominal segments, the metatibial setae and the metabasitarsus, all of which in¬

teract negatively with the length of the frons setae. The character suite shown by

vector 3 (Fig. 5C) is driven positively by traits 8 and 9, and less so by trait 16;

it is driven most negatively by traits 19 and 23. Vector 3 involves body width,

stylet length and the number of dorsal setae on the abdomen, all of which interact

negatively with the length of setae on abdominal tergum VIII and the length of

the dorsal setae on the metatibiae. The suite for vector 4 (Fig. 6 A) is more complex,

with several traits giving a moderate interactive influence; the suite is driven most

strongly, however, by trait 14, a siphuncular width function. The suite for vector

5 (Fig. 6B) is dominated positively by traits 10 and 20, and negatively by trait

18. It involves the lengths of the ultimate rostral segment and the caudal setae,

which interact antagonistically with the number of setae on abdominal tergum

VIII. The suite for vector 6 (Fig. 6C) is driven positively by traits 7, 14, and 17,

with traits 13 and 19 reacting negatively and less strongly. The setae on the frons,

the width of the siphunculi and the number of marginal setae on the abdominal

segments react positively in this suite, while the metathoracic notum and the

length of setae on abdominal tergum VIII react negatively.

For this Pseudessigella matrix, the summation of the absolute values for the

factor loading coefficients of traits on vectors 2-6 functionally represents the

proportion of variance attributable to multivariate “shape” for each trait (Fig.

7A). “Shape” variance, in this sense, is defined as (usually) variance that is sub¬

ordinate and orthogonal to “size,” or the residual variance after the general-size

allometric function is divorced from the matrix. Thus the summed coefficients

across vectors other than 1 can be taken to represent shape, in as much as vector

1 actually represents a true approximation of theoretical general-size factor for

the matrix. This is quite important from the view point of multivariate evolution,

because such models generally assume that genetic covariance matrices are static

and involve little or ideally no changes in “shape” over short periods of evolu¬

tionary time. Functionally, it is relatively easy for groups that differ only in “size”

to evolve “apart” in response to selection; all that is required is a differential,

allometric evolutionary response to selection where the differing group means

slide away from one another, up or down a multivariate regression line that

represents the general-size factor. This kind of evolutionary response does not

change “shape” because it does not modify the slope or relative elevation of the

allometric multivariate regression line in ^-dimensional evolutionary space.

In contrast, evolving a difference in “shape” requires expending considerably

more “evolutionary energy.” This is because the elevation and/or slope of the

multivariate regression in space must be changed by disruptive and stabilizing

selections, which realign the genetic correlations among characters on the shape

vectors that are orthogonal to the general-size factor. The amount of difficulty

involved in shape evolution depends on the strength of disruptive selection, the

amount of pleiotropic linkage for those traits being selected, and the degree of

stabilizing selection exerted upon the character suite involved under selective

pressure (see Zeng 1988).

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

2.5

30-f

1 3 5 7 9 1113151719212325

Figure 7. Histograms for Pseudessigella indicating: (A) the relative contributions of multivariate

“shape” (black), as the summed absolute values of the vector loading coefficients [a] for vectors 2-6,

and “size” (white) as the absolute values of vector loading coefficients for the traits on vector 1; (B)

the scaled contribution of each trait, as a percentage of size/shape [S], to size vector 1 (white), shape

vectors 2 and 3 (gray), and shape vectors 4-6 (black); (C) “uniqueness” among the traits, as the

percentage of their differentiation [D], derived as the reciprocal of communality (an indication of the

degree of genetic linkage) for the traits.

The cost of shape evolution is indicated by Lemen & Freeman (1984), who

note that differing bat genera within the Phyllostomidae are quite conservative

for intrageneric shape variance. In those genera intrageneric shape averaged only

1.5% of total variance, while size averaged 35%. The conservation of shape change

is further demonstrated intraspecifically by the conclusions of Holman & Kin-

dlmann (1987), whose data indicate that shape changes are ontogenetically fixed

between instars of the pea aphid, Acyrthosiphon pisum (Harris), in the allometry

of the developing antennal and leg segments.

In Pseudessigella, the summarized “shape” variance among traits varies con¬

siderably, as depicted by the “skyline” atop the black bars in Fig. 7A. This figure

shows the stacked summation of each trait’s absolute loading coefficients for all

vectors. The white bar component for each trait shows the character’s size con-

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

tribution, and the overall “skyline” atop the white bars shows the degree of total

contribution by each trait to the overall variance represented by vectors 1-6 (80%

of the total variance). Traits that load high on vector 1 (Fig. 5 A) generally have

larger proportions of white to black in Fig. 7A. These traits, which tend to be

segmental lengths, form a character group which isolate on vector 1 (Fig. 5A);

they are strongly linked pleiotropically and basically respond to selection for

allometric size variation. Other traits that load heavily on vectors 2-6 (Figs. 5B-

C, 6A-C) show a much higher ratio of shape to size in Fig. 7A; for example, note

that trait 25, metabasitarsus length, and trait 20, the length of caudal setae, among

others, show predominantly shape variance.

Because Fig. 7A shows that the traits differ in their absolute contributions to

the variance sequestered among the vectors, Fig. 7B was created to more easily

portray how the relative contributions of each trait are partitioned. The contri¬

butions in Fig. 7B are scaled to 100%. Again, the vector 1 general-size contribution

is white, but the shape components (Fig. 7B: nonwhite) are partitioned into the

contributions of vectors 2 + 3 (Fig. 7B: gray), and those of vectors 4-6 (Fig. 7B:

black). Because each subsequent eigenvector decreases in its proportion of total

variance in the data matrix, vectors 2 and 3 together represent 27% of the overall

variance, after the 34.1% from size vector 1 is accounted for; the three less

dominant shape vectors, 4-6, together account for an additional 22.9% of the

variance. All other shape vectors that are potentially extractable from the matrix

used (19 more beyond vector 6, [as character n — 1]) together account for the

remaining 20% of variance in the matrix.

Figure 7C shows the proportion of “uniqueness” (differentiation) as a percentage

for each character. In this sense, the uniqueness variance is defined as the reciprocal

of communality, which is the variance component for a character that is attrib¬

utable to common factors (Sneath & Sokal 1973). Genetically, communality en¬

tails the variance attributable to pleiotropy or other linkage mechanisms between

traits; this operationally assumes environmental effects (including measurement

errors) are null, as do most quantitative genetic models. Thus, uniqueness is a

measure of the independence of the variance of a character, after its linkage

correlations are removed. This independence can result from genetic discord,

sampling (measurement) error or, of course, environmental variances which are

operationally assumed to be zero here. Note that because the vectors here are

derived from PCA rather than discriminant function analysis (DFA), the com¬

munality referred does not represent plesiomorphy in the sense of Sorensen (1987a,

b) and Sorensen & Sawyer (1989), because PCA does not divorce the information

common among groups to provide apomorphy, in the sense of group-divergence

information, as does DFA when calculating intergroup Mahalanobis’ distance

using the Wilks-A criterion. Because of this important difference, the present figures

differ from similar histogram portrayals of variance used elsewhere (Sorensen

1987b: fig. 3, Sorensen & Sawyer 1989: fig. 5) to show the character contributions

of apomorphic anagenic distance between groups.

Predictive statements can be made for trait interaction during evolution in

eulachnine aphids by comparing the summarized information contained in Figs.

7A-C. Such predictions, however, are based entirely upon the intraspecific vari¬

ance within Pseudessigella, and are dependant upon the stability of the traits that

is shown across the genera in the subtribe. This stability is a function, at least in

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

part, of the genetic independence of the traits (e.g., Fig. 1C), in response to both

the disruptive and stabilizing selection pressures that act upon them. Because the

information in Figs. 7A-C is multivariately based, however, it should be a better

estimator of character interactions during evolution than simple bivariate com¬

parisons of correlations between traits would be; the latter involve a nonmulti¬

variate approach that has been shown to be inferior to multivariately based in¬

formation in assessing the evolutionary interplay among traits (Willig et al. 1986).

Comparing Figs. 7A-C, several observations can be made. For example, of char¬

acters 1-7, which show strong general-size affiliation, characters 4, 5 and 7 show

relatively elevated levels of uniqueness and are thus assumed to be under less

strong genetic linkage; they may, therefore, be more prone to successful disruptive

selection. Of characters 2 versus 3 (length of antennal segment V versus the pro¬

cessus terminalis, respectively), 3 shows slightly more total variance of which

more is attributable to shape parameters (Figs. 7A, B), although it also shows less

genetic independence (Fig. 7C) than trait 2.

Consider the implications for interaction of traits 25 and 26 (metabasitarsal

and metadistitarsal length, respectively), during evolution. In comparison to trait

26, trait 25 shows slightly less total contributed variance on vectors 1-6 (Fig. 7A),

but its shape influence is larger and thus is more general-size independent; it also

shows the greatest level of uniqueness, due to less linkage, of any of the 26 analyzed

traits. The metabasitarsal to metadistitarsal ratio shows significant variance

throughout the Cinarina and Eulachnina (Mamontova 1972) and within Essigella

(Sorensen 1983, 1988, unpublished data); previous interpretations of polarity for

the interaction of these traits has been questioned (Sorensen 1990). The expression

of the variance relationships of the tarsi in Pseudessigella is important to potential

interpretations for these characters within Essigella. By inferring from Pseudes¬

sigella, if the metatarsal ratio were to evolve differentially among species-groups

within Essigella, as it has (unpublished data), the metabasitarsus would have

necessarily had to undergo shape related evolutionary change via disruptive se¬

lection, while the change in the metadistitarsus should be more size related.

Although evolution of the metabasitarsus would be more difficult because of its

shape variance, it might on the other hand be more easily accomplished because

of its lower apparent linkage.

If selective pressure were applied to the metabasitarsus, vector 2 (Fig. 5B), as

the suite upon which it loads most heavily, shows that of the linkages that do

exist for this trait, it would necessarily have to respond in the most similar fashion

to changes in traits 13, 16-18, and 24 (the length of the metanotum [13], the

number of dorsal [16] and marginal [17] setae on abdominal terga II-IV and

abdominal tergum VIII [18], and the length of ventral setae on the metatibiae).

Alternatively, however, if selective pressure were applied to this suite of traits,

the metabasitarsus would be the most likely of any of the traits to respond in

discord, becoming unstable because of its lower linkage. Disruptive selection

would, therefore, most easily act upon this trait.

Character Groups Defined by Multiple Vectors of Shape Select ion . — Figures 8 A

and 8B show the 26 traits in the three-dimensional evolutionary I-space that is

generated by the dominant three shape vectors, 2-4. These figures show the same

scattergram, but it is rotated 90° in yaw to demonstrate character separation. The

original, unrotated, orthogonal vectors were used to define the evolutionary space

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

V 3

Figure 8. Character groupings for Pseudessigella apparent in the evolutionary space defined by a

combination of vectors 2—4, as the dominant shape components. Perspectives (A) and (B) represent

the same scattergram rotated 90° in yaw. The relative size of each dot and its number shows the

relative depth of the point in the third dimension (depth perpendicular to the page). The apparent

“groupings” of characters (I and II) are divided with dashed line for emphasis. Traits are assigned to

1991 SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS 47

shown, but the space itself was then rotated to the perspectives depicted. The

I-space shown portrays the dominant selective pressure gradients that act upon

shape as a component of character variance; it represents 35.9% of the total

variance within the Pseudessigella matrix, after the elimination of the 34.1% of

general-size related variance.

The vectors in Fig. 8 are indicated as intersecting lines, where the distal (labeled)

end represents the most positive coefficients on the vectors, and the vector in¬

tersection point represents a zero loading coefficient. Note that for this three-

dimensional space, as a multivector selection regime for shape, trait 1 occurs the

closest to the vector line intersection in both the A and B perspectives. In contrast,

traits 12 and 4 also appear close in perspectives A and B, respectively, but are

clearly not adjacent to the vector line intersection in the alternative perspectives.

Because the location of that intersection represents a zero loading coefficient on

all three shape vectors shown, trait 1, overall body size, as the most influential

trait in the general-size suite (Fig. 5 A) shows almost no residual shape component

variance. In contrast, those traits with the highest rank for shape on vectors 2 or

3 occur mostly along the periphery of the distribution field shown.

In Fig. 8, the traits have been ordered into three ranks, showing their relative

importance, as drivers, derived from the absolute value of their loading coefficients

on the vectors. Although the I-space defined in Fig. 8 represents vectors 2-4, the

rank assigned to each trait is the best rank (lowest rank number) that the trait

achieved on either vector 2 or 3, as the two most dominant shape vectors. Under

this criterion, traits 6, 8, 9, 13, 16-18, and 23-25 have the highest shape influence;

traits 2, 3, 15, 19, 20 and 26 are intermediate; and traits 1, 4, 5, 7, 11, 12, 14,

21 and 22, are lowest in shape influence.

The perspectives shown in Fig. 8 depict two character groups are apparent as

rough, flattened, somewhat oblong “disks,” each with a horizontal orientation;

these groups are labeled I and II in the figures. The disks demonstrate that the

26 traits are not randomly distributed in this three-dimensional I-space, rather

the characters apparently separate into two “affiliations” as seen from these per¬

spectives. These trait distributions reflect their responses to selection pressure

imposed upon them by the vectors as multiple gradients of net natural selection

for shape change in multivariate evolution. The relative rankings of the traits are

also nonrandomly distributed in this space. Note that character group I has a

preponderance of the Rank I characters (6 of 10); group II contains all (9) rank

III traits and nearly all (6 of 7) rank II traits as well. If the traits or their rankings

were distributed between these groups randomly, group I would be expected to

have 27% of the traits in each rank, or 2.7, 1.9 and 2.4 traits from ranks I—III,

respectively. The evolutionary meaning of these perspective-dependant character

affiliations, if any, is unclear.

Character Stability During the Evolution of Essigella.—Because Pseudessigella

is the nearest approximation to a progenitor of Essigella, and because (except for

mutation) variance must be present for selection to result in evolution (Mayr

one of three ranks determined by the relative importance of their contributions to the shape variance

sequestered by vectors 2 or 3: rank I (black)—traits with the highest shape variance; rank II (gray)—

traits with intermediate shape variance; rank III (white)—traits with the lowest shape variance.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

1963), it was postulated that the pattern of variance for traits shown between

groups within Essigella should also theoretically occur as intraspecific variance

among those traits in Pseudessigella. This would imply somewhat stable genetic

covariance matrices between these genera, as operational mother and daughter

groups (sensu Sorensen 1987a). To explore this hypothesis the dominant shape

traits occurring on vectors 2 and 3 were analyzed and ranked for Essigella, E.

kathleenae and E. kirki, as was done for Pseudessigella.

Figure 9 shows these character shape rankings, as color codings, for these taxa

was well as Pseudessigella. The relative degree of shape instability among the taxa

can be deduced from Fig. 9 by comparing the color codings of the traits between

the taxa. In that figure, stable traits, regardless of their ranking, maintain their

color between taxa; moderately stable traits, which change their ranking by only

one rank, are shown by white/gray or gray/black color changes between taxa;

unstable traits, which change their ranking by two ranks, are shown by white/

black color changes.

In comparison to Pseudessigella, less than half of the traits on vectors 2 or 3

remained stable during the evolution to Essigella. Traits 16-18, 23 and 24 re¬

mained unchanged in rank I (Fig. 9: black) as attributes high in shape variance

on vectors 2 or 3. Traits 1, 11, 12, 14, 21, and 22 were unchanged in rank III

(Fig. 3: white), but no traits remained unchanged in rank II (Fig. 9: gray). Several

traits, however, changed dramatically between the genera: trait 7 changed from

rank III (Fig. 9: white) in Pseudessigella, from poor in shape variance, to rank I

in Essigella ; traits 6, 9, 13 and 25 behaved the opposite. Many traits (2-5, 8, 10,

15, 19, 20, and 26) changed only one rank, in various directions, between the

genera. Thus, between Pseudessigella and Essigella nearly 42% of traits (11 of 26)

are stable with regard to shape variance; 39% (10 of 26) show moderate stability,

and 19% (5 of 26) are unstable.

Note that in Essigella the metabasitarsus, trait 25, has apparently changed shape

from its variance in Pseudessigella, becoming unstable and moving from rank I

to III. The metabasitarsus, therefore, has decoupled in part from the other dom¬

inant traits, 16-18 and 24, in the suite represented by vector 2 (Fig. 6B) in

Pseudessigella. The same is true for trait 13, although the latter showed consid¬

erably less independence in Pseudessigella (Fig. 1C). Unfortunately, stability in

evolutionary traits remains more easily observable than predictable.

Figure 9 also shows the shape component rankings for E. kathleenae and E.

kirki. For E. kathleenae, traits 17, 18 and 23 are rank I; traits 2, 3, 6, 7, 9-11,

15, 19, 20, 22 and 24 are rank II; and traits 1, 4, 5, 8, 12-14, 16, 21, 25 and 26

are rank III. For E. kirki, traits 3, 4, 6, 10, 14, 16, and 18 are rank I; traits 2, 5,

8, 20 and 23-25 are rank II; and traits 1, 7, 9, 11-13, 15, 17, 19, 21, 22 and 26

are rank III.

The similarity of shape component variance among all the taxa was calculated

for taxon-pairs and represents a measure of shape stability during evolution (Fig.

10). Pseudessigella and Essigella have a shape similarity of 0.62. Essigella kath¬

leenae shows the greatest degree of shape stability in common with Pseudessigella

(0.71), more so in fact than with Essigella (0.67); E. kirki has an equal degree of

shape stability (0.62) with either genus. Interestingly, however, shape stability

between E. kathleenae and E. kirki (0.60), both group I species, is lower than that

between either species versus either genus. These values suggest that E. kirki has

Analysis/Taxon

PCA: Pseudessigella

•

PCA: Essigella kathleenae

•

PCA: Essigella kirki

•

PCA: Essigella (all)

•

DFA (overall): Essigella (all)

•

DFA (shape): Essigella (all)

•

Character

•

Figure 9. Relative ranking and stability of the shape component variance for each trait on PCA vectors 2 or 3 among taxa, and the DFA diagnostic ranking

of traits for Essigella. For PCAs (either vector 2 or 3): rank I (black)—traits with the highest shape variance; rank II (gray)—traits with intermediate shape variance;

rank III (white)—traits with the lowest shape variance. For DFA (overall) (vectors 1-3): rank I (black)—best multivariate separators; rank II (gray)—intermediate

multivariate separators; rank III (white)—poorest multivariate separators. For DFA (shape) (vectors 1-3), rankings for “shape-based-diagnostics” are: rank I

(black)—trait is a rank I multivariate separator with either rank I or II shape variance; rank II (gray)—trait is a rank II multivariate separator with either rank I

or II shape variance; rank III (white)—trait is a multivariate separator of any rank but has rank III shape variance, or trait is a rank III multivariate separator of

any rank shape variance.

'•O

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Essigella

kathleenae

Essigella

kirki

Figure 10. Similarity among the taxa, for shape components on PCA vectors 2/3. The similarity

values (= 1 - distance) can be interpreted as an index of the stability of the relative rankings for the

shape component of the traits on the two most dominant shape vectors.

had more shape selection imposed upon it than has E. kathleenae. It might be

considered to be more “wayward” in shape, along a possible evolutionary “shape”

continuum between the genera. This is despite previous findings that E. kirki was

the closest in total anagenic distance to Pseudessigella on the phylogenetic network

for these taxa and, therefore, represented the most primitive Essigella species

(Sorensen 1987a).

The variance shown in shape stability between the taxa also indicates shape

components may not, therefore, be as stable during multivariate evolution as

might be thought, at least within the Eulachnina. Stabilizing and disruptive se¬

lection among the covariant character suites may be rather important during

multivariate evolution, both within Essigella and between it and Pseudessigella.

Because orthogonal shape is more theoretically conservative as a phylogenetic

component during the evolution of a genus than is size (Leman & Freeman 1984),

one would expect that shape variance should also be more important between

differing groups of species, as clades, within a genus also (e.g., species-groups and

monophyletic subgenera). Variance in shape components between species within

a genus should occur mostly at the species-group level or above, because of the

more significant evolutionary effort required to modify differing orthogonal char¬

acter suites. In contrast, the differences between species within species-groups or

complexes should most efficiently evolve by simple allometric change along their

common general-size component; this would suggest that monophyletic species-

groups should show mostly size differences between their closely related species,

subspecies, etc.

This is not shown, however, between the group I species here. Using this logic,

if E. kathleenae and E. kirki were parts of a tight monophyletic group, they should

have shown a higher similarity in shape variance, with their major differences

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

occurring in general-size variance. Because group I of Essigella is paraphyletically

defined and plesiomorphically sequestered within genus (Sorensen 1987a), how¬

ever, the relatively low stability in shape between these species is not that sur¬

prising. They do not form a clade and because of their early “divorce” from each

other and the remaining Essigella lineage(s), they have the least phyletic com¬

monality of any pair of species in the genus. Although they are phenotypically

quite similar, their similarity is largely based in plesiomorphy with reference to

other Essigella.

Because both the size and shape components of traits should be important

during radiant evolution of species within a genus, a DFA of all species of Essigella

was conducted to explore which traits that are highest in shape variance are also

the most multivariately important in differentiating monophyletic groups (clades)

within the genus. Because the functions of DFA represent Landean minimum

selective mortality measures (as: [V In W] r z, sensu Lande 1979), they are parsi¬

monious indices of the apomorphic anagenic distance that is necessary to account

for the observed contemporaneous positions of the group centroids for the species

in the ^-dimensional space defined by the DFA vectors (Sorensen 1987a, b;

Sorensen & Sawyer 1989). In this context, the vector loading coefficients of the

characters on the functions determine the relative importance of each trait as a

parsimonious contributor to the observed evolution (separation) of species for

the matrix employed.

The results of the DFA are also shown in Fig. 9. Traits 1, 5, 8, 14-16, 22 and

25 in rank I have the highest multivariate contributions in optimally distinguishing

(and evolving, sensu Lande 1979) Essigella species. Traits 2, 4, 7, 9, 10, 12, 13,

17, 24 and 26 show intermediate multivariate importance in rank II, and traits

3, 6, 11, 18-21 and 23 have the least importance in rank III. From these results,

it is apparent that a mixture of traits that show their strongest variance in both

size and shape seem important in the overall historical evolution of Essigella

species.

To determine if traits with high shape variance (present criteria) best multi¬

variately distinguish groups of species, as clades, within Essigella rather than

individual species within such groups, a new set of rankings was created (shape-

based-diagnostics) to reflect both the multivariate contribution of each trait as a

diagnostic in DFA, and the relative importance of its shape contribution in Es¬

sigella. Three ranks were assigned (Fig. 9): rank I traits were the best (rank I)

separators in DFA that also had either the highest or intermediate shape variance

(ranks I or II) in PCA; rank II traits were intermediate separators (rank II) in

DFA which also had either the highest or intermediate shape variance (ranks I

or II) in PCA. Rank III traits were poor DFA separators (diagnositc rank III) of

any shape rank, or were either the best or intermediate rank separators (diagnostic

ranks I or II) that showed poor shape variance (shape rank III).

As rank I shape-based-diagnostics, traits 5, 8, 15 and 16 appear to best separate

groupings of species within Essigella rather than simply individual species (Soren¬

sen 1983). Trait 16, the number of dorsal setae on the terga of the anterior

abdomen segments, separates the three major groupings of Essigella (Sorensen

1987a: groups I, II, III); groups II and III care clades, as is the combination of

II + III. Trait 16 also differentiates E. pini Wilson homoplasiously from the

monophyletic species-group involving E. californica and E. hoerneri (Sorensen

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(1)

1987a: spp. H, A, B, respectively), within group III. Trait 8, a body width com¬

ponent, separates the monophyletic species complex involving E. knowltoni Hottes

(Sorensen 1987a: spp. E, F, G) and to a lesser extent its monophyletic cohort E.

alyeska Sorensen (Sorensen 1987a: sp. D), as a clade within group III; this as¬

semblage involves a major portion of the orthogonal shape covariance shown

among the species of group III. Trait 15, the maximal length of a set of sclerites

that serve as muscle anchorages on the venter of the anterior abdominal segments,

serves to separate many clades within Essigella, notably: group II, the E. knowltoni

complex of group III, and the E. californica complex of group III. Trait 5, the

length of the third antennal segment, shows both size and shape variance; it

separates the E. californica complex as a clade, and also separates individual

species within several other species-groups.

As rank II shape-based-diagnostics, traits 4, 7, 17 and 24 are less apparent in

distinguishing groupings above the species level (Sorensen 1983). Trait 4, the

length of the fourth antennal segment, behaves much as trait 3 in that some

groupings of species show differential expression for the segment (e.g., group II,

and some complexes within group III) to a degree, but the trait varies between

species within some species-groups (e.g., the E. knowltoni complex of group III).

Traits 7 and 24, the length of setae on the frons and ventral setae on the metatibia,

are more difficult to interpret because of homoplasy and intraspecific variance.

Both sets of setae have characteristic patterns of variance within species; they are

always (plesiomorphically ?) short in some Essigella species (e.g., E. kathleenae

and E. kirki of group I), but can vary widely in their lengths among individuals

of other Essigella species (e.g., the E. knowltoni and particularly the E. californica

complexes of group III) showing an apomorphically longer mean setal length for

such species. Trait 17, the number of marginal setae on the anterior abdominal

segments, parallels trait 16 and shows the same group defining properties, except

that the variance in trait 17 is slightly more within several species.

Rank I shape-based-diagnostics, as a class, therefore, appear to best distinguish

clades at various levels within Essigella, more than individual species within those

groupings. Rank II shape-based-diagnostics, however, appear less evident in their

separation of clades rather than individual species, and these traits tend to show

variance that is either more homoplasious between species(-groups) or is subject

to intraspecific variance in Essigella.

Traits within Essigella that are rank I diagnostics, exclusive of those that are

also rank I or II shape-based-diagnostics, should be expected to portray higher

levels of (homoplasious) importance in separating species within species-groups.

Because the analysis of such size-based-diagnostics requires a local partitioning

of the overall variance within Essigella at the level of its species, the examination

of size-based-diagnostics will be treated elsewhere (unpublished data).

Acknowledgment

I thank the following people for their help: R. L. Blackman and V. F. Eastop

(British Museum [Natural History], London) for loans of Pseudessigella\ R. G.

Foottit (Biosystematics Research Institute, Agriculture Canada, Ottawa) for dis¬

cussions concerning abdominal tergum VIII in Eulachnus; the late D. Hille Ris

Lambers for his comments and discussions on character transformations and

phylogenetic placements during the earlier phases of the research; S. M. Sawyer

1991

SORENSEN: MULTIVARIATE EVOLUTION IN APHIDS

(Insect Taxonomy Laboratory, CDFA) for illustrating the outline of Figure 1; and

K. H. Sorensen for editorial assistance.

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Received 15 October 1990; accepted 29 November 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 55-64, (1991)

ENTOMOPATHOGENIC FUNGI

(ZYGOMYCOTINA: ENTOMOPHTHORALES) INFECTING

CEREAL APHIDS (HOMOPTERA: APHIDIDAE)

IN MONTANA

Ming-Guang Feng , 1 Robert M. Nowierski,

Albert L. Scharen , 2 and David C. Sands 3

Entomology Research Laboratory, Montana State University, Bozeman, Montana

59717

Abstract. — A field survey of dryland and irrigated crops of small grains was conducted in Montana

during the 1990 growing season to search for entomopathogenic fungi for use in the biological

control of the Russian wheat aphid, Diuraphis noxia (Mordvilko). Dryland crops were heavily

infested by D. noxia, but no evidence of fungal infection was found among the populations of

this aphid species. Diuraphis noxia populations were undetectable on spring grains grown under

irrigation. However, other cereal aphids, mainly Metopolophium dirhodum (Walker) and Rho-

palosiphum maidis (Fitch), occurred abundantly. Mycoses caused by Entomophthorales were

observed killing both aphid species from late July through August. Among 292 aphid cadavers

collected during the occurrence of mycoses, 45.9% were killed by Pandora neoaphidis (Remau-

diere & Hennebert) Humber, 31.8% by Conidiobolus obscurus (Hall & Dunn) Remaudiere &

Keller, 21.6% by Entomophthora planchoniana Cornu, and 0.7% by Zoophthora radicans (Bre-

feld) Batko. Among the aphids killed by C. obscurus, the proportion of cadavers producing resting

spores inside the body, rather than primary conidia on the body surface, increased quickly as

the incidence of infection decreased on both aphid hosts. Distinguishing features of the two true

Entomophthora species, E. planchoniana and E. chromaphidis Burger & Swain, infecting aphids

in North America are discussed in comparison with information reported from different geo¬

graphic areas. The four fungal species, plus Conidiobolus thromboides Drechsler, which was not

found in 1990, but was successfully isolated from Diuraphis tritici (Gillette) in 1989, are new

records as pathogens of cereal aphids for Montana.

Key Words.— Insecta, cereal aphids, Aphididae, Entomophthorales, aphid-pathogenic fungi,

biological control agents

Aphids are often subject to attack by entomopathogenic fungi, including nu¬

merous species of Entomophthorales (Latge & Papierok 1988, Waterhouse &

Brady 1982) and at least two species of Hyphomycetes (Feng & Johnson 1990;

Feng et al. 1990a, b; Hall 1981). These fungi are considered to have great potential

as candidates for use in microbial control of various aphids (Latge & Papierok

1988).

Fungal pathogens infecting cereal aphids have been reported in Europe (Dean

& Wilding 1971, 1973; Dedryver 1983; Papierok & Havukkala 1986) and South

America (Lassari 1985). Some regional lists of aphid fungi have also been pub¬

lished in Australia (Milner et al. 1980), Finland (Papierok 1989) and Israel (Ben-

Ze’ev et al. 1984). In North America, five entomophthoralean species have been

recorded from 34 aphid host species in the eastern United States and Canada

1 Permanent address: China National Rice Research Institute, Hangzhou, Zhejiang Province, Peo¬

ple’s Republic of China.

2 USDA-ARS, Department of Plant Pathology, Montana State University, Bozeman, Montana

59717.

3 Department of Plant Pathology, Montana State University, Bozeman, Montana 59717.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

(Remaudiere et al. 1978). Recently, Feng et al. (1990b) recovered 10 species of

fungal pathogens from several species of cereal aphids infesting grain crops grown

under irrigation in southwestern Idaho. One of these aphid hosts is the Russian

wheat aphid, Diuraphis noxia (Mordvilko), a recently-introduced, devastating pest

of small grains in North America (Burton 1988). Microbial control, involving the

use of entomopathogenic fungi, is one promising strategy that will contribute to

the management of this pest in the future (Burton 1988, 1989).

This study surveyed for fungal pathogens of cereal aphids in dryland and ir¬

rigated small grains in Montana and generate regional strains for use in both

laboratory and field studies.

Materials and Methods

Field Survey. — A field survey conducted in Montana during the 1990 growing

season included dryland and irrigated crops. The dryland crops were fall-sown

winter wheat in two fields and spring-sown barley in one field located 25 or 35

km west of Billings. The irrigated crops were several small experimental plots of

late-sown spring wheat and barley on the Montana State University campus in

Bozeman. The survey was initiated in mid-May and continued through harvest.

The fields were visited each week.

On each field date, numerous aphid colonies throughout each of the fields were

examined carefully to determine if any aphids were infected with fungal pathogens.

On the dryland crops, this examination was made while samples, ranging from

100 to 960 tillers (decreasing with increasing aphid density), were randomly taken

in each field for purposes of estimating aphid population levels. Aphid cadavers,

when found in the colonies, were placed in plastic vials in a cooler and transported

to the laboratory for identification and isolation of fungal agents responsible for

aphid mortality.

Identification of Aphid-Pathogenic Fungi. — External symptoms and fungal re¬

productive structures from each of the aphid cadavers were examined under a

dissecting microscope as soon as possible following the collection (within the same

day). Desiccated specimens were maintained in a moist chamber for several hours

to allow for development of reproductive structures. Cadavers were then indi¬

vidually mounted on glass slides using cotton blue or aceto-orcein stains. Iden¬

tification was based on external symptoms and the morphology of spores and

sporulating structures (Waterhouse & Brady 1982). A recent revision of the clas¬

sification for the Entomophthorales (Humber 1989) was used. The fungi associated

with the cadavers were considered to have caused the death of their hosts if they

had been previously documented as pathogens of aphids in the literature.

Isolation of Aphid-Pathogenic Fungi. — Saubouraud dextrose agar (DIFCO Lab¬

oratories, Detroit, Mich.) supplemented with 1% yeast extract (DIFCO, Labo¬

ratories, Detroit, Mich.) was used as a basic medium (SDAY) to isolate fungal

pathogens from infected aphids. Some fresh aphid cadavers were individually

attached to a small piece of double-faced sticky film in the center of a Petri dish

cover. Cadavers were suspended over SDAY medium in the bottom of a Petri

dish for a period of 10-20 h and then removed. Discharged spores from the

cadaver would thus fall down to the medium for initiation of an in vitro culture.

The Petri dishes containing inoculum were incubated at approximately 20° C with

a photoperiod of 16:8 (L:D).

1991

FENG ET AL.: ENTOMOPHTHORALES INFECTING APHIDS

Results and Discussion

The dryland crops were heavily infested by D. noxia during the 1990 growing

season. Mean population densities (± SE) on average reached 14.4 (± 46.5) and

47.9 (± 87.4) aphids per tiller in the two winter wheat fields, respectively and

136.2 (± 102.9) on the spring barley. The percentage of tillers infested was 50%,

86%, and 96%, respectively. Other cereal aphids were occasionally found, but no

consistent populations were detected. Despite the high levels of D. noxia popu¬

lations on the dryland crops, no aphids were found to be infected by any fungal

pathogens during the growing season.

Diuraphis noxia populations were undetectable on regularly sown grain crops

in the Bozeman area during the growing season. However, other cereal aphids,

including the com leaf aphid, Rhopalosiphum maidis (Fitch), and the rose-grass

aphid, Metopolophium dirhodum (Walker), occurred abundantly in the small plots

of late-sown spring grains grown under irrigation after early July. Mycoses induced

by entomophthoralean fungi were observed killing both aphid species from late

July through late August.

Four species of entomophthoralean fungi were identified from among the 292

cadavers of M. dirhodum and R. maidis collected during the four-week period of

mycosis development. Nearly half of those cadavers (45.9%) were killed by Pan¬

dora neoaphidis (Remaudiere & Hennebert) Humber, 31.8% by Conidiobolus

obscurus (Hall & Dunn) Remaudiere & Keller, 21.6% by Entomophthora plan-

choniana Comu, and 0.7% by Zoophthora radicans (Brefeld) Batko. The identity

and descriptions of these fungi are given separately below.

Conidiobolus spp. —Although only a single species, C. obscurus, was found

among the Conidiobolus- killed aphids in the 1990 growing season, another fungus,

Conidiobolus thromboides Drechsler, was successfully isolated from an infected

individual of the western wheat aphid, Diuraphis tritici (Gillette), on winter wheat

in Bozeman during the fall of 1989. Hence, these two species are listed here

together as Montana records. Conidiobolus obscurus was recovered from 30 M.

dirhodum and 63 R. maidis cadavers. Several isolates of C. obscurus have been

obtained from both aphid hosts on SDAY and sent to ARSEF (USDA-ARS

Collection of Entomopathogenic Fungal Cultures, USDA-ARS Plant Protection

Research Unit, U.S. Plant, Soil and Nutrition Laboratory, Ithaca, New York) for

permanent storage.

Aphids recently killed by C. obscurus were grey or grey-brown, and were attached

to the plant by their probosces. Neither rhizoids nor cystidia were present. Co-

nidiophores were simple, unbranched (Fig. la). Primary conidia containing many

globules were nearly spherical (Fig. lb), and measured as 33.8 (28.8-37.5) x 28.0

(22.5-33.8) im l {n = 100). Secondary conidia formed from primary conidia by

germination (Fig. lc). Resting spores (Fig. Id) were spherical with a diameter of

36.5 (27.5-45.0) x 35.4 (27.5-43.8) pm {n = 50). No differences were significant

in the dimensions of the primary conidia (t x = 1.41, P > 0.10; t 2 = 0.53, P >

0.50; df = 98) and resting spores (t x = 1.10, P > 0.20; t 2 = 0.50, P > 0.50; df =

98) between the two aphid hosts. However, the primary conidia [39.6 (28.8-33.7)

x 33.7 (25.0-44.3) u m, n = 50] and resting spores [38.9 (36.3-43.8) x 37.4 (35.0-

40.0) ^m, n= 15] from in vitro cultures were significantly larger than the primary

conidia (t x = 10.03, P < 0.01; t 2 = 11.09, P < 0.01; df = 148) and resting spores

(C = 4.55, P < 0.01; t 2 = 3.74, P < 0.01; df = 113) collected in vivo, respectively.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(1)

Figure 1. (a-d) Conidiobolus obscurus, (e-f) C. thromboides, and (g-k) Entomophthora planchoni-

ana. (a) Developing conidiophores (CP), (b) A primary conidium. (c) A secondary conidium (SC) being

produced from a primary conidium (PC), (d) Resting spores of C. obscurus. (e) A primary conidium.

(1) A secondary conidium (SC) being produced from a primary conidium (PC) of C. thromboides. (g)

A conidiophore. (h-i) Primary conidia imbedded in cytoplasms, (j) Secondary conidia (SC) being

produced from primary conidia (PC), (k) Hyphal bodies germinated from conidia of E. planchoniana.

Scale bars: 10 /im. The bar on (a) applies to (c) and (g); the bar on (b) applies to (e), (h-j); and the bar

on (d) applies to (k).

The primary conidia of C. thromboides were subspherical, but smaller (Fig. le)

than those of C. obscurus. The measurements from the in vitro culture were 26.4

(22.0-31.3) x 20.9 (17.5-23.8) /im (n = 50). The resting spores were also similar

in shape but smaller than those of C. obscurus.

1991

FENG ET AL.: ENTOMOPHTHORALES INFECTING APHIDS

Entomophthora planchoniana.—This fungus was recovered from 37 M. dirho-

dum and 26 R. maidis cadavers. No attempt was made to obtain a pure culture

because true E. planchoniana has rarely been successfully cultured using an ar¬

tificial medium.

Aphids killed by E. planchoniana were yellow-brown or brick brown and stuck

to the plant by rhizoids in a thick bundle or several bundles forming a platform.

Cystidia were absent. Conidiophores were unbranched (Fig. lg). Primary conidia

were typically discharged with a mass of cytoplasm creating a ‘halo’ (Fig. lh, li).

Primary conidia were bell-shaped and measured as 18.9 (15.0-22.5) x 15.5 (12.0-

17.8) n m (n = 75). Secondary conidia (Fig. lj) that germinated from the primary

conidia were more rounded and smaller than the primary conidia: 15.1 (13.3-

17.5) x 12.6 (10.8-14.8) mm (n = 50). No resting spores were observed from the

materials examined.

There are only two true Entomophthora species that have been documented as

aphid pathogens in the world: E. planchoniana and E. chromaphidis Burger &

Swain. Entomophthora planchoniana is well known in Europe (Waterhouse &

Brady 1982; Papierok 1989) and also has been reported in Israel (Ben-Ze’ev et

al. 1984) and Australia (Milner et al. 1980). In North America, however, this

species has been recorded only from two aphid cadavers of unknown identity that

were collected from the eastern coast (Remaudiere et al. 1978). On the other hand,

E. chromaphidis was originally described from the walnut aphid, Chromaphidis

juglandicola (Kaltenbach), in southern California (Burger & Swain 1918) and

recently recovered from cereal aphids in southwestern Idaho (Feng 1990). The

two species can be distinguished from each other by the size of primary conidia

and the number of nuclei in each conidium (R. A. Humber, MGF, unpublished

data). Primary conidia of E. chromaphidis contain 2-8 nuclei each and are gen¬

erally smaller than those of E. planchoniana, which contain 2-6 nuclei (Water-

house & Brady 1982). There was no significant difference in the size of the primary

conidia from M. dirhodum and R. maidis cadavers, killed by Entomophthora in

Montana, compared to those seen in other reports (Table 1) from Europe and

Australia. However, the primary conidia of the Entomophthora from Montana

are significantly larger than those of the real Entomophthora species reported from

California and Idaho. The comparison did not include the materials from Israel

(Ben-Ze’ev et al. 1984) and the east coast of North America (Remaudiere et al.

1978) because the authors did not provide measurements. Unfortunately, our

specimens were not stained well enough to allow the counting of the nuclei inside

the conidia because almost all the primary conidia were imbedded in the cyto¬

plasm. Even so, the Montana fungus could be E. planchoniana rather than E.

chromaphidis because of the similarity to the European fungus in conidial size.

Pandora neoaphidis.—A total of 134 cadavers, including 115 M. dirhodum and

19 R. maidis, were attributed to infection by P. neoaphidis. No isolates were

obtained using SDAY.

Aphids freshly killed by P. neoaphidis turned pale brown or yellow-brown and

adhered to the plant by numerous rhizoids, each of which consisted of a thin stalk

ending in a disk-like terminal expansion. Cystidia, present under moist conditions,

were distally tapered, thicker than conidiophores, and extended far beyond them

(Fig. 2a). Conidiophores were digitately branched at their apices (Figs. 2a, 2b).

Primary conidia contained a single large nucleus, were ovoid to cylindrical in

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Table 1. Comparison of the sizes of primary conidia of true Entomophthora species from aphids

in Montana and other geographic areas.

Region

Fungus 3

Primary conidia, /wn

LAV 6 Mean' Range

Reference

Australia

Epl

(17.0)

14-20

1.61

>0.10

Milner et al. 1980

(15.0)

13-18

0.47

>0.50

England

Epl

19.0

14-23

-0.09

>0.50

Waterhouse & Brady 1982

14.0

12-20

1.32

>0.10

Finland

Epl

18.1

13-22

0.68

>0.40

Papierok 1989

14.9

10-18

0.55

>0.50

California

Ech

(12.5)

11-14

5.42

<0.01

Burger & Swain 1918

(10.5)

10-11

4.33

<0.01

Idaho

Ech

14.4

13-16

3.79

<0.01

Feng 1990

12.3

10-13

2.82

<0.01

a Epl = E. planchoniana, Ech = E. chromaphidis.

b L = length (including papillae), W = width.

c Table entries in brackets estimated from the range of original measurements but not given directly

by the authors.

d i-test under null hypothesis H 0 : x MT = y COM , where x MT is the measurement from Montana material

(L: 18.89 ± 1.17; W: 15.54 ± 1.16; n = 75), Ycqm the measurement from the materials compared,

and t = (x MT — } ; C om)/SE x [(/7 + 1 )/n] vl with df = n - 1.

shape (Fig. 2c) and measured 23.6 (18.8-30.0) x 12.4 (10.0-15.0) pm (n = 80).

Secondary conidia developed from primary conidia (Fig. 2d) and were similar in

shape but smaller than the primary conidia. No resting spores were present in the

specimens examined.

Zoophthora radicans.—This fungus was recovered from two M. dirhodum ca¬

davers only. Because of the paucity of specimens, no attempt was made to isolate

Z. radicans, although it usually grows quite well on SDAY.

Aphids killed by Z. radicans were pale brown or orange-brown. Rhizoids were

present. Cystidia were also present, but were relatively few in number compared

with those of P. neoaphidis. Conidiophores were branched (Fig. 2e). Primary

conidia were uninucleate, long and ellipsoid in shape (Fig. 2f): 18.3 (15.3-21.3)

x 7.4 (6.5-8.8) pm (n = 50). Secondary conidia germinated apically or laterally

from primary conidia and were similar to primary conidia. No capilliconidia and

resting spores were observed in the limited number of specimens.

Development of Aphid Mycosis.— The relative proportion of infections by P.

neoaphidis tended to increase during the 4 week period of mycosis development,

whereas that of C. obscurus appeared to decrease (Fig. 3). The fungus, Z. radicans,

was observed only at the beginning of the period. Thereafter, it remained unde¬

tected or possibly its influence on the aphids was masked by other fungi. The

infection by E. planchoniana remained about the same among the aphids after

early August (Fig. 3). More M. dirhdodum cadavers were attributed to infection

by P. neoaphidis than were R. maidis cadavers. The reverse was true for C.

obscurus.

Many of the aphids killed by C. obscurus produced resting spores rather than

primary conidia, which are the primary form infecting live aphids. As shown in

Fig. 4, the proportion of cadavers containing resting spores in the field appeared

to increase during the 4 week period, irrespective of the species of aphid host.

1991

FENG ET AL.: ENTOMOPHTHORALES INFECTING APHIDS

Figure 2. (a-d) Pandora neoaphidis and (e-f) Zoophthora radicans. (a) Cystidia (CYST), (b) Co-

nidiophores. (c) Primary conidia. (d) A secondary conidium (SC) being produced from a primary

conidium (PC), (e) Conidiophores. (f) Primary conidia and secondary conidium (SC). Scale bars: 10

ium. The bar on (c) applies to (d-f).

This phenomenon has not been reported by previous researchers (e.g., Feng et al.

1990b, Milner et al. 1980, Papierok 1989) although resting spores of Conidiobolus

spp. from aphids are frequently observed from in vitro cultures. The production

of resting spores is often thought to require a short photoperiod or low temperature

or a combination of both for initiation (Wilding & Latteur 1987). However, during

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

1.0

0.8

P. neoaphidis

E. planchoniana

Late Jul

C. obscurus

Z. radicans

::::

in! iiiii!

::::: ::::: :

::::::::::: : ^

■

Early Aug Mid Aug

TIME

Late Aug

Figure 3. Proportion of each fungal pathogen among the cadavers of M. dirhodum and R. maidis

in irrigated spring grains in Bozeman, Montana during the 1990 growing season.

the period from late July to late August in 1990, the natural photoperiod and

temperature in Bozeman remained in a summer pattern: 14-15 h day length and

mild temperature conditions; the monthly mean temperature for August was 19.4°

C (11.0-27.9° C). It is unclear whether any other environmental factors may have

been involved in this phenomenon.

The production of resting spores in the field implies an interruption of the fungal

infection cycle, because generally following resting spore formation a minimum

period of vernalization is required before germination is initiated (Wilding &

Latteur 1987). This may explain why the level of infection by C. obscurus decreased

among the aphids during the survey (Fig. 3). This also suggests that for Conidiobo-

lus species to be successfully used against cereal aphids in Montana, additional

understanding of the mechanisms for the production and germination of resting

spores will be required.

The mortality of cereal aphids resulting from fungal infection in Bozeman was

at low levels (< 10%) compared with those reported from cereal aphids in irrigated

spring wheat in southwestern Idaho (up to 90%; Feng 1990). The frequent thun¬

dershowers from late July through early August as well as the irrigation water

provided may have enhanced the development of fungal infection in the aphid

populations in Bozeman. Arid weather (low moisture and high temperature) may

have prevented fungal diseases from developing in the aphid populations on

dryland crops.

The four species of Entomophthorales, found during the 1990 season, plus the

C. thromboides isolate, obtained in the fall of 1989, are new records as pathogens

of cereal aphids for Montana. Entomophthora planchoniana represents the first

record from cereal aphids in North America.

1991

FENG ET AL.: ENTOMOPHTHORALES INFECTING APHIDS

1.0

0.8

I—

Q_ 0.4

0.2

0.0

Late Jul Early Aug Mid Aug Late Aug

TIME

Figure 4. Proportion of aphid cadavers producing resting spores in irrigated spring grains in

Bozeman, Montana during the development of mycosis.

Acknowledgment

The authors wish to thank K. M. O’Neill, J. L. Littlefield (Entomology Research

Laboratory, Montana State University, Bozeman), and S. A. Woods (USDA-ARS,

Rangeland Insect Laboratory, Montana State University, Bozeman), for their

review and comments on this manuscript. This study was supported by APHIS

Cooperative Agreement Number 0348-0039 and a grant from the Wheat and

Barley Commision, Montana. This paper is a joint contribution from the Agri¬

cultural Experiment Station, Montana State University, Bozeman and the Agri¬

cultural Research Service, U.S. Department of Agriculture, and is published as

Journal Series No. J-2577.

R. maidis

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

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Feng, M. G., J. B. Johnson & L. P. Kish. 1990a. Virulence of Verticillium lecanii and an aphid-

derived isolate of Beauveria bassiana (Fungi: Hyphomycetes) for six species of cereal aphids

(Homoptera: Aphididae). Environ. Entomol., 19: 815-820.

Feng, M. G., J. B. Johnson & L. P. Kish. 1990b. Survey of entomopathogenic fungi naturally infecting

cereal aphids (Homoptera: Aphididae) of irrigated grain crops in southwestern Idaho. Environ.

Entomol., 19: 1534-1542.

Hall, R. A. 1981. The fungus Verticillium lecanii as a microbial insecticide against aphids and scales,

pp. 483-498. In Burges, H. D. (ed.). Microbial control of pests and plant diseases 1970-1980.

Academic Press, New York.

Humber, R. A. 1989. Synopsis of a revised classification for the Entomophthorales (Zygomycotina).

Mycotaxon, 34: 441-460.

Lassari, S. N. 1985. Natural enemies of aphids (Homoptera, Aphididae) on barley ( Horderum sp.)

in Parana. Ana. Soc. Ent. Braz., 14: 5-15.

Latge, J. P. & B. Papierok. 1988. Aphid pathogens, pp. 323-336. In Minks, A. K. & P. Harrewijn

(eds.). Aphids, their biology, natural enemies and control, vol. 2B. Elsevier, Amsterdam.

Milner, R. J., R. E. Teakle, G. G. Lutton & F. M. Dare. 1980. Pathogens (Phycomycetes: Ento¬

mophthorales) of the blue-green aphid Acyrthosiphon kondoi Shinji and other aphids in Aus¬

tralia. Aust. J. Bot., 28: 601-619.

Papierok, B. 1989. On the occurrence of Entomophthorales (Zygomycetes) in Finland. I. Species

attacking aphids (Homoptera, Aphididae). Ann. Entomol. Fennici, 55: 63-69.

Papierok, B. & I. Havukkala. 1986. Entomophthoraceous fungi parasitizing cereal aphids in Finland.

Ann. Entomol. Fennici, 52: 36-38.

Remaudiere, G., J. P. Latge & W. A. Smirnoff. 1978. Considerations ecologiques sur quelques

Entomophthorales pathogens d’aphids communes dans Test des U.S.A. et du Canada. Phyto¬

protection, 59: 150-156.

Waterhouse, M. & B. L. Brady. 1982. Key to the species of Entomophthora sensu lato. Bull. Br.

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and their utilisation. Med. Fac. Landbouww. Rijksuniv. Gent., 52: 159-164.

Received 13 November 1990; accepted 3 December 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 65-71, (1991)

NEW SPECIES OF ASPIRHINA KIRBY FROM THE

NEOTROPICAL REGION

(HYMENOPTERA: CHALCIDIDAE)

Jeffrey A. Halstead

296 Burgan, Clovis, California 93612 1

Abstract.— Four new Aspirhina Kirby are described from the Neotropical region: A. alvarengai

NEW SPECIES, A. bifurca NEW SPECIES, A. deceptor NEW SPECIES, and A. spinosa NEW

SPECIES. Aspirhina dubitator (Walker), the only previously described species, is redescribed and

its female is described for the first time. Characters are presented to distinguish Aspirhina from

other Neotropical Chalcididae. No biological information is known for members of the genus

Aspirhina.

Key Words.— Insecta, Aspirhina, Neotropical, Chalcididae, Chalcidoidea, Hymenoptera

The genus Aspirhina was proposed by Kirby (1883) for a Brazilian species that

Walker (1862) originally described as Halticella [correctly Haltichella ] dubitator.

Since then, no other species were added or described in Aspirhina. Four new

species from the Neotropical region are described in this paper. A key to the

species of Aspirhina and diagnostic characters for the genus are presented. No

biological or host information is known for Aspirhina, and their potential as

biological control agents is unassessed. Overall, a total of 25 specimens were

examined, indicating that Aspirhina are rare in collections.

Aspirhina is distinguished from other genera of New World Chalcididae by

the following character combination: hind tibia truncate apically, two apical spurs

present (Haltichellinae); marginal vein reaching anterior margin of forewing, post¬

marginal and stigmal veins present (Haltichellini); vertex of head not produced

into horns; preorbital carina strong, arch-like above anterior ocellus; pronotum

with a carina along anterior margin except at median; forewing clouded, a darker

spot under marginal vein; scutellum with a posteriorly projecting triangular to

finger-like process, never with a small median tooth; tergum I dorsally with at

least three longitudinal carinae at base, originating from a transverse carina.

Museum acronyms for specimen depositories are: American Museum of Natural

History, New York (AMNH); British Museum of Natural History, London

(BMNH); Canadian National Collection, Ottawa (CNC); Florida State Collection

of Arthropods, Florida Department of Agriculture and Consumer Affairs, Gaines¬

ville (FSCA); J. A. Halstead, personal collection (JAHC); and U.S. National Mu¬

seum of Natural History, Washington, D.C. (USNM). Abbreviations include “T”

for tergum or terga (e.g., T1 for tergum I). Distributional information for paratypes

and material examined is listed alphabetically by country and location, respec¬

tively.

1 Correspondence: % Kings River Conservation District, 4886 E. Jensen Avenue, Fresno, California

93725.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Key to Species of Aspirhina

1 Tergum I dorsally punctate, with four longitudinal carinae (rarely five

carinae) in which outermost carinae are about 0.5 x length of innermost

carinae (Fig. la); scutellar projection apically emarginate to forked

(Figs. 12a, 12b) and dorsally punctate (Fig. lb) .

. alvarengai NEW SPECIES

- Tergum I dorsally smooth, with three or four longitudinal carinae of

about equal length (Figs. 2a, 3a, 4a, 5a); scutellar projection usually

not as above . 2

2(1) Tergum I dorsally with three longitudinal carinae (Fig. 3a); scutellar

projection apically rounded (Fig. 11) and dorsally striate (Fig. 3b) ..

. spinosa NEW SPECIES

Tergite I dorsally with four longitudinal carinae; scutellar projection not

as above . 3

3(2) Scutellar projection apically rounded (Fig. 14); hind femur with teeth on

ventral margin extending 0.66 x hind femur length (Fig. 9).

. dubitator (Walker)

- Scutellar projection apicially emarginate to forked (Figs. 13, 15); hind

femur with teeth on ventral margin extending 0.5 x or less hind femur

length. 4

4(3) Scutellar projection apically emarginate (Fig. 13) and dorsally punctate

(Fig. 4b). deceptor NEW SPECIES

- Scutellar projection apically forked (Fig. 15) and dorsally striate (Fig. 5b)

. bifurca NEW SPECIES.

Aspirhina dubitator (Walker)

(Figs. 2a, 2b, 9, 14)

Halticella dubitator Walker, 1862: 366.

Aspirhina dubitator (Walker); Kirby 1883: 60.

Types. — Holotype, male; data: “Brazil, Aspirhina dubitator Walker, B.M. TYPE

5.555.” Examined by author. Holotype resides in the British Museum of Natural

History, London.

Description.—Male (holotype). Body length: 5.2 mm. Color: black with tegula, fore leg, mid leg

(except coxa), hind trochanter, and apical half of hind tibia orange. Pronotum and Mesoscutum: dorsally

without a median carina. Scutellum: in lateral view with dorsal margin straight and apex of projection

finger-like, blunt (Fig. 14); in dorsal view apical projection 0.25 x maximum scutellar length (Fig.

2b); sculptured with oval setigerous punctures separated by 0.5 to 1.0 x puncture diameter; 16

punctures along midline between base and apex; integument smooth and polished except weakly

coriaceous basally, apical projection punctate except at apex. Hindfemur: oval; teeth on ventral margin

extending 0.66 x hind femur length (Fig. 9). Gaster: pubescence silver on Tl-2, remainder of terga

with gold pubescence. Tl: dorsally with four longitudinal carinae, extending 0.66 x T1 length (Fig.

2a), integument polished.

Female. Like male, but pubescence of gaster entirely silver and hind tibia and hind femur (basally

and apically) orange.

Diagnosis. —Aspirhina dubitator may be distinguished from other Aspirhina by

its unique combination of the characters: tergum I dorsally smooth and with four

longitudinal carinae which are about equal in length (Fig. 2a), scutellar projection

1991 HALSTEAD: NEW NEOTROPICAL ASPIRHINA 67

Figures 1-5. Aspirhina spp., males. Tergum 1 (a) and scutellum (b), dorsal views. Sculpture of

apical projection of scutellum and punctation of tergum 1 (circle insert) illustrated. 1. A. alvarengai

NEW SPECIES. 2. A. dubitator. 3. A. spinosa NEW SPECIES. 4. A. deceptor NEW SPECIES. 5. A.

bifurca NEW SPECIES. Scale line 1.0 mm.

apically rounded (Fig. 14) and dorsally punctate (Fig. 2b), and teeth on ventral

margin of hind femur extending 0.66 x hind femur length (Fig. 9).

Variation.—Male. Longitudinal carinae of T1 vary in length from 0.6 to 0.8 x

T1 length. Two specimens with only fore and mid leg brown and tarsi orange.

One specimen with hind leg (except coxa and central area of outerside of femur)

orange. Female. Known only from specimen described, which is deposited in the

AMNH.

Distribution. — Brazil.

Material Examined.— BRAZIL. AMAZONAS ESTIRAR DE EQUADOR: Rio Javari, Sep 1979,

M. Alvarenga, 4 males (CNC). MATO GROSSO: Sinop, Oct 1974, Malaise trap, M. Alvargena, 1

male (JAHC). RODONIA: Rio Guapore opposite mouth of Rio Baures, 1-5 Oct 1964, Bouseman and

Lussenhop, 1 female.

Aspirhina bifurca Halstead, NEW SPECIES

(Figs. 5a, 5b, 10, 15)

Types. — Holotype, male; allotype, female; data: “BOLIVIA, Beni: Rio Itenez

opposite Costa Marques (Brazil), Sept. 4-6, 1964, Bouseman & Lussenhop.”

Holotype and allotype reside in the American Museum of Natural History, New

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Figures 6-15. Aspirhina spp., males. Figs. 6-10. Hind femur, lateral view. Figs. 11-15. Scutellum,

lateral view. 6, 11. A spinosa NEW SPECIES. 7, 12a, 12b. A. alvarengaiNEW SPECIES, 12b-variation

in apex of scutellum. 8, 13. A. deceptor NEW SPECIES. 9, 14. A. dubitator. 10, 15. A. bifurca NEW

SPECIES. Scale line 1.0 mm.

York. Paratypes: 1 male with same data and depository as holotype; BRAZIL.

GOIAS: Jatai, Nov 1972, F. M. Oliveira, 1 male (USNM). COSTA RICA. San

Isidro de General, 700-800 m, Aug 1980, N.L.H. Krauss, 1 male (JAHC, from

AMNH). ECUADOR. PICHINCHA: 47 km S of Santo Domingo Rio Palenque

Station, 21-25 Feb 1979, S. and J. Peck, 1 male (CNC).

Description. —Male (holotype). Body length: 3.8 mm. Color: black with tegula and legs (except central

area of outerside of hind femur) orange; antenna, apical projection of scutellum, hind femur centrally,

and hind coxa brown. Pronotum and Mesoscutum: dorsally without a median carina. Scutellum: in

lateral view with dorsal margin convex and apex of projection forked (Fig. 15); in dorsal view apical

projection 0.35 x maximum scutellar length (Fig. 5b); sculptured with oval setigerous punctures

separated by 0.12 to 0.33 x puncture diameter; 12 punctures along midline between base and apex;

integument sculpture coriaceous, anterior projection faintly striated to apex. Hind femur: oval; teeth

on ventral margin extending 0.42 x hind femur length (Fig. 10). Gaster: pubescence silver on Tl-2,

remainder of terga with gold pubescence. 77: dorsally with four longitudinal carinae, extending 0.4

x T1 length (Fig. 5a), integument polished.

Female (allotype). Like male, but body length 4.2 mm, legs completely orange, and antenna slenderly

filiform.

Diagnosis. —Aspirhina bifurca may be distinguished from other Aspirhina by

its unique combination of the characters: tergum I dorsally smooth and with four

longitudinal carinae which are about equal in length (Fig. 5a), scutellar projection

apically forked (Figs. 15) and dorsally striate (Fig. 5b), and teeth on ventral margin

of hind femur extending 0.42 x length of hind femur (Fig. 10).

1991

HALSTEAD: NEW NEOTROPICAL ASPIRHINA

Variation. —Male. One paratype with hind femur completely orange; another

paratype with hind femur completely brown. Apical projection of scutellum varies

slightly in length and width. Some specimens with striation on projection confined

to apical half. Female. Known only from allotype.

Distribution.— Bolivia, Brazil, Costa Rica, and Equador.

Etymology. — The specific name (Latin) refers to the forked apical projection of

the scutellum.

Material Examined. —This species is known only from the type specimens.

Aspirhina alvarengai Halstead, NEW SPECIES

(Figs, la, lb, 7, 12a, 12b)

Types. — Holotype, male; data: “BOLIVIA: Dept. Beni, Romansos, 1 km. N.

Junction Rio Itenez & Rio Paragua, VII-30-1964, J. K. Bouseman and L. Lus-

senhop Collectors.” Holotype resides in the American Museum of Natural His¬

tory, New York. Paratypes: BOLIVIA. SANTA CRUZ: Gral. Saavedra Est. Ex¬

perimental, Aug 1973, Malaise trap, C. Porter and L. Stange, 1 male (FSCA).

BRAZIL. Pedra Azul, M. Gerais, Nov 1972, Seabra and Oliveira, 1 male (CNC).

CEARA: Brasalha, May 1969, M. Alvarenga, 1 male (CNC). RONDONIA: Rio

guapore opposite mouth of Rio Baures, 1-5 May 1964, Bouseman and Lussenhop,

1 male (AMNH). BAHIA: Encruzilhada, 960 m, Nov 1974, M. Alvarenga, 1 male

(CNC). PANAMA. Barro Colorado Island Canal Zone, 14 Mar 1956, C. W. and

M. E. Rettenmeyer, 1 male (USNM). SURINAME. Saramaca, Prov. Raleigh

Vallen N. P. Foengoe Is., 22-31 Jan 1985, T. Thormin, 1 male (JAHC, from

CNC).

Description.—Male (holotype). Body length: 6.0 mm. Color: black with apices of tibia and tarsus of

fore and mid legs orange. Pronotum and Mesoscutum: dorsally without a median carina. Scutellum:

in lateral view with dorsal margin nearly straight but steeply angled and apex of projection longer

dorsally than ventrally (Fig. 12a); in dorsal view apical projection 0.29 x maximum scutellar length

(Fig. lb); sculptured with oval setigerous punctures separated by 0.25 to 0.5 x puncture diameter; 14

punctures along midline between base and apex; integument sculpture coriaceous, apical projection

punctate except at apex. Hind femur: oval; teeth on ventral margin extending 0.52 x hind femur

length (Fig. 7). Gaster: pubescence silver on Tl-2, remainder of terga with gold pubescence. 77:

dorsally with four longitudinal carinae, extending 0.33 x T1 length (Fig. la), sculpture punctate.

Female. Unknown.

Diagnosis. —Aspirhina alvarengai may be distinguished from other Aspirhina

by its unique combination of the characters: tergum I dorsally smooth and with

four longitudinal carinae in which the outermost carinae are about 0.5 x length

of innermost carinae (Fig. la), scutellar projection apically emarginate to forked

(Figs. 12a, 12b) and dorsally punctate (Fig. lb), and teeth on ventral margin of

hind femur extending 0.52 x length of hind femur (Fig. 7).

Variation. —Male. Apical projection of scutellum varies from weakly to strongly

emarginate (Figs. 12a, 12b). Outermost longitudinal carina of T1 rarely shorter

than the innermost carinae. Tergum I commonly smooth and polished at base.

Two paratypes with punctures of T1 vague (Panama) to absent (Bolivia); integ¬

ument smooth and polished though indications of punctures evident. One para¬

type (Bolivia) with five longitudinal carinae on Tl.

Distribution. — Bolivia, Brazil, Panama, and Suriname.

Etymology.— Named for M. Alvarenga who collected much of the Aspirhina

material.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

Material Examined. — This species is known only from the type specimens.

Aspirhina spinosa Halstead, NEW SPECIES

(Figs. 3a, 3b, 6, 11)

Types. -Holotype, male; data: “BRAZIL, Serrade Bocaina S. Paulo, XI-1968,

1600 m, M. Alvarenga.” Holotype resides in the Canadian National Collection,

Ottawa. Paratype: 1 male with same data and depository as holotype.

Description. —Male (holotype). Body length: 4.5 mm. Color: black with apices of tibia, tarsus of fore

and mid legs, and club of antenna orange. Pronotum and Mesoscutum: dorsally with a median carina.

Scutellum: in lateral view with dorsal margin convex and apex of projection rounded (Fig. 11); in

dorsal view apical projection 0.4 x maximum scutellar length (Fig. 3a); sculptured with oval setigerous

punctures separated by 0.12 to 0.17 x puncture diameter; 10 punctures along midline between base

and apex; integument sculpture coriaceous; apical projection with a median, striated carina, punctate

laterally below carina. Hind femur: oval; teeth on ventral margin extending 0.37 x hind femur length

(Fig. 6). Gaster: pubescence silver on Tl-2, remainder of terga with gold pubescence. Tl: dorsally

with three longitudinal carinae, extending 0.33 x Tl length (Fig. 3b), integument polished; on each

side of lateral carina a long furrow extending length of longitudinal carinae; Tl somewhat fused with

T2 but separation between them evident.

Female. Unknown.

Diagnosis. —Aspirhina spinosa may be distinguished from other Aspirhina by

its unique combination of the characters: tergum I dorsally smooth and with three

longitudinal carinae which are about equal in length (Fig. 3a), scutellar projection

apically rounded (Fig. 11) and dorsally striate (Fig. 3b), and teeth on ventral

margin of hind femur extending 0.37 x length of hind femur (Fig. 6).

Variation. — Paratype male with antenna brown, hind tarsus orange, and median

carina on mesoscutum vague.

Distribution. —Brazil and Bolivia.

Etymology. — The specific name (Latin) refers to the long, slender apical pro¬

jection of the scutellum.

Material Examined. —This species is known only from the type specimens.

Aspirhina deceptor Halstead, NEW SPECIES

(Figs. 4a, 4b, 8, 13)

Types. —Holotype, male; data: “BOLIVIA, Beni: Rio Itenez opposite Costa

Marques (Brazil), Sept. 4-6, 1964, Bouseman & Lussenhop.” Holotype resides

in the American Museum of Natural History, New York.

Description.—Male (holotype). Body length: 4.1 mm. Color: black with antenna, tegula, and legs

brown. Pronotum and Mesoscutum: dorsally without a median carina. Scutellum: in lateral view with

dorsal margin convex and apex of projection slightly incised (Fig. 13); in dorsal view apical projection

0.29 x maximum scutellar length (Fig. 4b); sculptured with oval setigerous punctures separated by

0.12 to 0.5 x puncture diameter; nine punctures along midline between base and apex; integument

sculpture coriaceous, apical projection punctate to apex. Hind femur: oval; teeth on ventral margin

extending 0.5 x hind femur length (Fig. 8). Gaster: pubescence silver on Tl-2, remainder of terga

with gold pubescence. Tl: dorsally with four longitudinal carinae, extending 0.33 x Tl length (Fig.

4a), integument polished.

Female. Unknown.

1991

HALSTEAD: NEW NEOTROPICAL ASPIRHINA

Diagnosis. —Aspirhina deceptor may be distinguished from other Aspirhina by

its unique combination of the characters; tergum I dorsally smooth and with four

longitudinal carinae which are about equal in length (Fig. 4a), scutellar projection

apically emarginate (Fig. 13) and dorsally punctate (Fig. 4b), and teeth on ventral

margin of hind femur extending 0.5 x length of hind femur (Fig. 8).

Variation. —Unknown. Species known only from type specimen.

Distribution. — Bolivia.

Etymology. — The specific name (Latin) refers to this species’ ability to elude

collectors.

Material Examined.— This species is known only from the type specimen.

Acknowledgment

I thank the staff of the museums listed for the opportunity to examine their

specimens; Gary Gibson and John Huber (CNC) for gathering chalcidid material;

Z. Boucek for assistance identifying the genus Aspirhina, and J. S. Noyes (British

Museum of Natural History, London) for loaning the type of A. dubitator, K. J.

Woodwich for the use of his drawing tube to prepare the figures, and D. J. Burdick

(California State University, Fresno) for the use of laboratory facilities and for

reviewing this paper. I thank also two anonymous reviewers for their editorial

comments which improved this manuscript.

Literature Cited

Kirby, W. F. 1883. Remarks on the genera of the subfamily Chalcidinae, with synonymic notes and

descriptions of new species of Leucospidinae and Chalcidinae. J. Linn. Soc. London, Zoology,

17: 53-76.

Walker, F. 1862. Notes on chalcidites, and characters of undescribed species. Trans. Entomol. Soc.

London (3), I: 345-397.

Received 1 February 1990; accepted 6 December 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 72, (1991)

Scientific Note

FIRST RECORD OF THE AUSTRALIAN PSYLLID

BLASTOPSYLLA OCCIDENTALIS TAYLOR

(HOMOPTERA; PSYLLOIDEA) ON EUCALYPTUS

(MYRTACEAE) IN MEXICO

The occurrence of Australian Myrtaceae-feeding psyllids, including Blastopsylla

occidentalis Taylor, has recently been reported from California on introduced but

long-established host plants such as Eucalyptus spathulata Hooker and Lophos-

temon confertus R. Brown (Taylor, K. 1985. J.’Aust. Entomol. Soc., 24: 17-30;

Taylor, K. 1987. J. Aust. Entomol. Soc., 26: 229-233).

It has generally been assumed that these records represent recent direct intro¬

ductions from Australia. However, Blastopsylla occidentalism also well established

in Mexico. Large populations were discovered on mature Eucalyptus trees growing

1 km south of San Juan Carapan, Michoacan on 12 Feb 1989, raising the possibility

that Californian infestations may have originated from within the New World.

A number of pest species of psyllid have recently extended their ranges over

considerable distances (Hodkinson, I. D. 1988. New Scientist, 118: 47-51). For

example, the Leucaena psyllid, Heteropsylla cubana Crawford, has spread, over

the last few years, from an origin in Central America, throughout the Pacific Basin

as far as India. Psyllids can form a significant element of the aerial plankton

(Spalding, J. B. 1979. Arctic Alpine Res., 11: 83-94) and wind dispersal or passive

transport in or on aircraft have been suggested as mechanisms of spread, in

addition to the dissemination of infested plant materials. The potato psyllid,

Paratrioza cockerelli Sulc is already known to disperse seasonally on prevailing

winds from breeding areas in northern Mexico, New Mexico, Arizona and Texas

into the central states of the U.S.A. (Wallis, R. L. 1955. U.S. Dept. Agric., Tech.

Bull., 1107). The normal direction of the prevailing winds, however, poses a

greater obstacle to direct wind-assisted dispersal flights by B. occidentalis from

Mexico to California, although more roundabout routes remain a distinct pos¬

sibility. The importance of Central America as a potential source of psyllid in¬

festation cannot be overstated. The recent introduction into Honduras (Burck-

hardt, D. & M. Martinez. 1989. Bull. Soc. Entomol. Fr., 94: 65-66) of a major

disease vector of citrus, the Asian citrus psyllid, Diaphorina citri Kuwayama,

raises the spectre of a rapid northwards expansion into the major citrus growing

areas of the U.S.

I. D. Hodkinson, School of Natural Sciences, Liverpool Polytechnic, Liverpool,

L3 3AF, United Kingdom.

Received 17 May 1990; accepted 1 November 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 73, (1991)

Scientific Note

LOCATION OF HOLOTYPE OF ARAEOSCHIZUS

ANTENNATUS BLAISDELLI PAPP

(COLEOPTERA: TENEBRIONIDAE)

Charles S. Papp recently described two new Araeoschizus from our Baja Cali¬

fornia, Mexico collections (Papp, C. S. 1989. Notes on the Stenosini genus Araeo¬

schizus LeConte from Baja, [sic] California, Mexico. (Coleoptera: Tenebrionidae.)

Entomography, 6: 335-340). Papp (p. 338) incorrectly listed deposition of the

holotype of Araeoschizus antennatus blaisdelli Papp as “In the Natural History

Museum of the College of Idaho, Caldwell, Idaho” [CIDA]. Since we do not retain

primary types, the specimen has been forwarded to the California Academy of

Sciences as Type #16623. The holotype of Araeoschizus antennatus clarki Papp

is also at the collection of the California Academy of Sciences (Papp, p. 335) as

Type #16624. Additional specimens of both taxa are also deposited in CIDA.

William H. Clark, Orma J. Smith Museum of Natural History, College of Idaho,

Caldwell, Idaho 83605.

Received 31 October 1990; accepted 15 November 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 73-74, (1991)

Scientific Note

NEW HOST FOR ACANTHOSCELIDES A UREOLUS HORN

(COLEOPTERA: BRUCHIDAE)

Acanthoscelides aureolus Horn forms a species-complex (Johnson, C. D. 1970.

Biosystematics of the Arizona, California, and Oregon species of the seed beetle

genus Acanthoscelides Schilsky (Coleoptera: Bruchidae). Univ. Calif. Publ. En-

tomol., 59; Johnson, C. D. 1983. Misc. Publ. Entomol. Soc. Amer., 56) of small

(< 3 mm) beetles that range from the Alberta and Manitoba to Chiapas, Mexico,

and from California to Minnesota and Nebraska. It commonly feeds on several

species of Astragalus (Leguminosae).

I discovered a new host for Acanthoscelides aureolus at a record high elevation

of 3620 m (11,800 ft) in White-Inyo Range of Mono Co., California. The new

host is Astragalus kentrophyta Gray var. implexus (Canby) Bameby, an unusual

species of Astragalus with fruits that rarely contain more than one seed (though

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

there are an average of six ovules per pistil). Its seeds are small, averaging 0.0018

g (SD = 0.0005 g), and typically mature in late August and early September.

Adults of the beetle have been reared from laboratory stored seeds of the plant

one or two months after collection. Adults have been observed emerging yearly

from seeds collected between 1987 and 1990.

Voucher specimens of the beetle are deposited at Northern Arizona University,

Flagstaff, Arizona, and the California Department of Food and Agriculture, Sac¬

ramento. Specimens of A. kentrophyta implexus are deposited at the Department

of Botany Herbarium, University of California, Davis.

Material Examined.— CALIFORNIA. MONO Co.: White-Inyo Range, Sheep Mountain, alpine

dolomite barrens above Patriarch Grove, 3620 m (11,880 ft) W. R. Owen, Astragalus kentrophyta

Gray var. implexus (Canby) Bameby.

Acknowledgment.— I thank C. D. Johnson (Northern Arizona University, Flag¬

staff) for confirming the identification of A. aureolus and T. N. Seeno (California

Department of Food and Agriculture, Sacramento).

Wayne R. Owen, White Mountain Research Station, University of California,

Los Angeles and Graduate Group in Ecology, University of California, Davis 95616.

Received 21 November 1990; accepted 7 December 1990.

PAN-PACIFIC ENTOMOLOGIST

67(1): 75-76, (1991)

Book Review

Oman, P. W., W. J. Knight & M. W. Nielson. 1990. Leafhoppers (Cicadellidae):

a Bibliography, Generic Check-List and Index to the World Literature 1956-

1985. CAB International, Wallingford, Oxon, United Kingdom, 368 p. (ISBN

0-85198-690-0).

One problem in systematics, as an academic field, is its rigorous dependence

upon both information and priority. A systematist without a set of references on

the taxonomic nomenclature and bibliography of his group is as handicapped as

a lawyer without access to a legal library. Fortunately for leafhopper workers, a

monumental catalogue for the cicadellids was published in several parts, largely

by Z. P. Metcalf, and later by V. W. Burnside (e.g., [in part], Metcalf, Z. P. 1962—

68. General catalogue of the Homoptera. Fascicle VI. Cicadelloidea. Parts 1-17.

U.S. Dept. Agric., Agric. Res. Service, Washington D.C.). Metcalfs historic work

provided an information base for leafhopper systematics that covered 200 years,

ending in 1955. During the last 35 years, cicadellid researchers have severely felt

the absence of Metcalfs coverage.

The current, very credible and welcomed volume by Oman, Knight and Nielson

correct the deficient of easily accessible information since Metcalf. The work was

largely compiled by Paul Oman over the years, and was completed when W. J.

Knight and M. W. Nielson came to his aid to finish the formidable volume. These

workers form the planet’s most impressive and illustrious team in leafhopper

taxonomy to date. Oman has achieved the position of world patriarch in cicadellid

systematics; his catalogue of the genera of North America (Oman, P. W. 1949.

The Nearctic leafhoppers [Homoptera: Cicadellidae], a generic classification and

checklist. Mem. Entomol. Soc. Wash., 3) remains the taxonomic “bible.” Knight

and Neilson both have extensive experience and major systematic works to their

credit; Neilson’s compendium on leafhopper vectors (Neilson, M. W. 1968. The

leafhopper vectors of phytopathogenic viruses [Homoptera, Cicadellidae]. Tax¬

onomy, biology, and virus transmission. U.S. Dept. Agric. Tech. Bull., 1382) is

a staple for the held of plant disease vectors.

The present work continues Metcalfs broad based bibliography through 31 Dec

1985, and covers all fields that are relevant to leafhopper biosystematics in its

broadest sense, from behavior and biogeography to natural enemies and vector/

pathogen interaction. The 7000 citations (all 164 pages of them in double bed,

quarto format) contained in it were gathered both manually, and by computer

searches of the Abstracts of Entomology, the Bibliography of Agriculture, Bio¬

logical Abstracts, Entomology Abstracts, the Review of Applied Entomology and

Zoological Record. The format employed assigns each citation an author-year-

letter designation (i.e., “Bairyamova, V. 1970b”), as did Metcalf; this allows

authors to make their literature citations in reference to this volume, rather than

having to provide complete bibliographic information in their articles.

The taxonomic portion of the work covers 8000 new species and 1100 new

genera. It provides information on: the family-group names proposed between

1956-1985, and the use of such names in the Cicadellidae during that period; a

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

provisional classification for the family; misassigned generic names; misidentified

type-species; names placed on the official lists for generic names and family-group

names in zoology; and names placed on the official index of rejected and invalid

family-group names in zoology. It also provides an annotated, alphabetical check¬

list of the names at and above the generic level within the family. This information

is indispensible, and will prove invaluable to leafhopper taxonomists, and anyone

doing work on biodiversity.

Because of the wealth of important information that this tome contains, my

criticisms of the volume are few; I’m sure, however, that some taxonomists will

have individual points of contention with particular taxonomic treatments that

are listed (all taxonomists seldom think completely alike). Diacritical marks have

been omitted throughout the book; an unfortunate, but not major, problem for

such a reference. Although the format is attractive and book is easily used, it

might have been physically produced with a more archival quality in mind;

especially because it will serve as a reference for generations. For example, I find

no mention that the paper used is acid-free. Because of marketability, however,

books today are seldom made to better standards.

If you work on leafhoppers, particularly their systematics, you cannot afford to

be without this volume, which retails for $86.50 US (£49.50 UK).

John T. Sorensen, Insect Taxonomy Laboratory, California Dept, of Food &

Agriculture, 1220 N Street, Sacramento, California 94815.

PAN-PACIFIC ENTOMOLOGIST

67(1): 77, (1991)

1989 SPONSORING MEMBERS OF THE

PACIFIC COAST ENTOMOLOGICAL SOCIETY

Phillip A. Adams

Robert P. Allen

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Wayne C. Fields Jr.

John G. Franclemont

E. Eric Grissell

John E. Hafemik Jr.

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Robert L. Mangan

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Robert W. L. Potts

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Norman E. Gershenz & Leslie S. Saul

Evert I. & Marion E. Schlinger

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(1)

PACIFIC COAST ENTOMOLOGICAL SOCIETY

STATEMENT OF INCOME, EXPENDITURES AND

CHANGES IN FUND BALANCES

YEARS ENDED 30 SEPTEMBER 1989 AND 1988

1989 1988

Income

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Increase (Decrease) in fund balances . ($ 9,848) $ 5,912

Fund balances October 1, 1989 and 1988 . 110,709 104,797

Fund balances September 30, 1989 and 1988 . $100,861 $110,709

STATEMENT OF ASSETS

30 SEPTEMBER 1989 AND 1988

1989 1988

Cash in bank

Commercial account . $ 16 $ 11,161

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General Fund. 12,876 20,727

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Total cash in bank . . $ 85,358 $100,511

Investment in 80 shares of American Telephone

& Telegraph Co. common stock and 264 shares

of Pacific Telesis Group at market value. $ 15,503 $ 10,198

$100,861 $110,709

See accompanying notes to the financial statements.

1991

1988 & 1989 FINANCIAL STATEMENT

PACIFIC COAST ENTOMOLOGICAL SOCIETY

NOTES TO THE FINANCIAL STATEMENTS

YEAR ENDED SEPTEMBER 30, 1989

SUMMARY OF SIGNIFICANT ACCOUNTING POLICIES.

Accounting Method: Income and expenses are recorded by using the cash basis of accounting.

Capital Expenditures: Annual capital expenditures of $5,000 or less are charged to expense.

Marketable Securities: American Telephone & Telegraph Co. and Pacific Telesis Group common stock

are carried at market value. Increases and decreases in value are reflected in income.

Income Tax: The Society is exempt from Federal income and California franchise tax.

As Chairman of the Auditing Committee, and in accordance with its bylaws, I have reviewed the

financial records of the Society.

During the course of this review nothing was noted which indicated any material inaccuracy in the

foregoing statements.

H. Vannoy Davis

Chairman of the Auditing Committee

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Information for Contributors

See volume 66 (1): 1-8, January 1990, for detailed format information and the issues thereafter for examples. Manuscripts must be

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Literature Cited. — Format examples are:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

Blackman, R. L., P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometries

provide some answers? pp. 233-238. In Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics

and taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985.

SPB Academic Publishing, The Hague, The Netherlands.

Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.

Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.

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Volume 67

THE PAN-PACIFIC ENTOMOLOGIST

January 1991

Number 1

Contents

Pacific Coast Entomological Society, articles of incorporation and bylaws. 1

ASQUITH, A. & J. D. LATTIN— Omanonabis: a new genus of Nabidae (Heteroptera) from

western North America, with a review of Nabis edax Blatchley. 7

KAMM, J. A. —Larval development of Dargida procincta (Grote) (Lepidoptera: Noctuidae) in

the pacific northwest. 20

TSAUR, S. C. & F. J. LIN—A new Drosophila species in the montium Meijere subgroup of

the melanogaster Meigen species-group in the subgenus Sophophora Sturtevant from

Taiwan (Diptera: Drosophilidae).. 24

SORENSEN, J. T.—Phylogenetic character responses for shape component variance during the

multivariate evolution of eulachnine aphids: redescription of Pseudessigella Hille Ris

Lambers (Homoptera: Aphididae: Lachninae). 28

FENG, M. G., R. M. NOWIERSKI, A. L. SCHAREN & D. C. SANDS-Entomopathogenic

fungi (Zygomycotina: Entomophthorales) infecting cereal aphids (Homoptera: Aphidi¬

dae) in Montana. 55

HALSTEAD, J. A.—New species of Aspirhina Kirby from the Neotropical region (Hymenop-

tera: Chalcididae). 65

SCIENTIFIC NOTES

HODKINSON, I. D.—First record of the Australian psyllid Blastopsylla occidentalis Taylor

(Homoptera: Psylloidea) on Eucalyptus (Myrtaceae) in Mexico. 72

CLARK, W. H. —Location of holotype of Araeoschizus antennatus blaisdelli Papp (Coleoptera:

Tenebrionidae). 73

OWEN, W. R.—New host for Acanthoscelides aureolus Horn (Coleoptera: Bruchidae). 74

BOOK REVIEW

SORENSEN, J. T.-Oman, P. W., W. J. Knight & M. W. Nielson. 1990. Leafhoppers (Cica-

dellidae): a bibliography, generic-check-list and index to the world literature 1956-1985.

CAB International, Wallingford, Oxon, United Kingdom, 368 p. 75

Sponsoring members of the Pacific Coast Entomological Society: 1989 . 77

Pacific Coast Entomological Society, financial statement for 1988, 1989 . 78

Announcement—publications of the Pacific Coast Entomological Society. 80

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 67

April 1991

Number 2

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

(ISSN 0031-0603)

The Pan-Pacific Entomologist

EDITORIAL BOARD

J. T. Sorensen, Editor

S. S. Shanks, Treasurer Susan M. Sawyer, Editorial Assistant R. M. Bohart

J. T. Doyen J. E. Hafernik, Jr. J. A. Powell

Published quarterly in January, April, July, and October with Society Proceed¬

ings usually appearing in the October issue. All communications regarding non¬

receipt of numbers, requests for sample copies, and financial communications

should be addressed to: Sandra Shanks, Treasurer, Pacific Coast Entomological

Society, Dept, of Entomology, California Academy of Sciences, Golden Gate Park,

San Francisco, CA 94118-4599.

Application for membership in the Society and changes of address should be

addressed to: Curtis Y. Takahashi, Membership Committee chair, Pacific Coast

Entomological Society, Dept, of Entomology, California Academy of Sciences,

Golden Gate Park, San Francisco, CA 94118-4599.

Manuscripts, proofs, and all correspondence concerning editorial matters (but

not aspects of publication charges or costs) should be sent to: Dr. John T. Sorensen,

Editor, Pan-Pacific Entomologist, Insect Taxonomy Laboratory, California Dept,

of Food & Agriculture, 1220 N Street, Sacramento, CA 95814. See the back cover

for Information-to-Contributors, and volume 66 (1): 1-8, January 1990, for more

detailed information. Refer inquiries for publication charges and costs to the

Treasurer.

The annual dues, paid in advance, are $20.00 for regular members of the Society,

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Pacific Coast Entomological Society

OFFICERS FOR 1990

Leslie S. Saul, President

Norman D. Penny, President-Elect Wesley A. Maffei, Managing Secretary

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THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly by the Pacific

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PAN-PACIFIC ENTOMOLOGIST

67(2): 81-85, (1991)

MATERNAL EFFECTS AND EGG HATCHABILITY IN

MELANOPLUS (ORTHOPTERA: ACRIDIDAE)

W. Chapco and P. W. Riegert

Department of Biology, University of Regina,

Regina, Saskatchewan, S4S 0A2, Canada

Abstract.— Continuous culturing of nondiapause strains of Melanoplus is possible because eggs

are able to hatch within a short period of time and without the requirement of cold exposure.

Analysis of intercross data involving field and nondiapause strains reveals that the proportion

of eggs with this property is under maternal influence and, in some cases, a function of the egg’s

genotype. The problem of loss of genetic variation that is often associated with laboratory strain

maintenance is discussed and a solution that capitalizes on maternal effects is offered.

Key Words.— Insecta, Orthoptera, Acrididae, Melanoplus, hatchability, maternal effects

The central and desirable feature of the much used nondiapause strain of Me¬

lanoplus sanguinipes Fabr., developed by Pickford & Randell (1969), and of the

recently established nondiapause strain of M. differentialis (Thomas) (Oma et al.

1990), is the ability of eggs to hatch in a relatively short period of time (about 21

days at 30° C for M. sanguinipes ) without the requirement of cold treatment. In

this paper the term “egg hatchability” is restricted to this usage, which in reality

is equivalent to the proportion of nondiapausing eggs; indeed, it is this trait that

was selected for by the developers of these nondiapause cultures. There is no

doubt that egg hatchability has a heritable basis, and although the mode of in¬

heritance is presently unknown, polygenic inheritance is likely involved judging

from the slow responses to selection for the trait (Slifer & King 1961). Environ¬

mental factors, such as temperature and photoperiod (Oma et al. 1990), can also

influence hatchability; these agents probably interact with genotype to account

for latitudinal variations in voltinism that are observed in the United States.

Maternal effects, genic or nongenic (mediated, for example, through maternal

nutrition), result in offspring with phenotypes more similar to those of the female

parent than to those of the male parent (Mather & Jinks 1982). Such effects were

recently suspected during hybridization experiments (WC, unpublished data) when

the species status of distant populations of M. sanguinipes and the nondiapause

strain noted were examined. To support the claim for maternal effects, an analysis

of these data and results collected 27 years ago (PWR, unpublished data) on M.

sanguinipes and M. packardii Scudder is presented.

Materials and Methods

Data were gathered separately by the authors. In studies by WC, the following

field populations were sampled. Fourth and fifth instars of M. sanguinipes were

collected near Bethune, Saskatchewan (Bet); Rigaud, Quebec (Que); and Green

Mountain Road, British Columbia (BC) during the spring and summer of 1981.

Insects were sexed, separated and allowed to mature in laboratory cages until

commencement of matings. Adult M. sanguinipes were sampled from West La¬

fayette, Louisiana (Lo); Trinity Texas (Txl); Oakhurst, Texas (Tx2); and Prescott,

Arizona (Az) during the spring and summer of 1980. Insects were allowed to

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Table 1. Mean (±SE) hatchability of field strains and of crosses with nondiapause strains of

Melanoplus sanguinipes (NDwc and NDpr) and M. packardii (NDpr-pac).

Strain

Strain values

Crossed with

ND 99

Crossed with

ND 33

NDwc

Bet

78.8 ± 2.4 (11 l) a

0.3 ± 0.3 (22)

87.2 ± 3.4 (26)

22.4 ± 5.9

(31)

Que

2.6 ± 1.3 (36)

54.0 ± 5.1 (43)

0.0 ± 0.0

(11)

25.2 ± 4.5 (49)

79.4 ± 6.3 (24)

5.1 ± 2.8

(7)

30.4 ±6.4 (38)

55.3 ± 5.9 (42)

42.7 ± 8.6

(24)

Txl

38.9 ± 6.9 (21)

50.7 ± 8.2 (14)

21.7 ± 13.6 (5)

Tx2

30.6 ± 13.6(7)

70.5 ± 7.6 (21)

9.7 ± 7.6

(8)

38.7 ± 6.8 (34)

91.8 ± 2.6 (22)

43.1 ± 7.4

(28)

NDpr

Wyn

81.7 ± 2.7 (100)

83.5 ± 2.7(61)

23.1 ± 2.3

(168)

NDpr-pac

Dav

76.4 ± 1.5 (305)

26.2 ± b (49)

68.9 ± 4.9 (53)

25.8 ± 3.4

(67)

a n = number of pods.

b —no standard error because pods hatched in groups.

NA—not available.

deposit eggs in cages available in the laboratories of resident colleagues (see Ack¬

nowledgment). Harvested eggs (F, of field animals) were sent to the University

of Regina and the emergent offspring used for these studies. All strains were crossed

reciprocally with the nondiapause strain (NDwc) developed by Pickford & Randell

(1969) beginning with material collected near Delisle, Saskatchewan (Pickford

1958). In addition, two of the above field populations, Que and BC, were mated

reciprocally. Experimental conditions were essentially the same as those in Chapco

(1984). About 10 pairs of virgin adults were introduced into standard Hunter-

Jones cages. Dead insects were removed and replaced with virgins, if available.

Egg pods were collected daily until most or all egg layers were dead and then

individually transferred to shell vials with moistened vermiculite and incubated

at 30° C for a maximum of 50 days, a length well beyond the period required for

the nondiapause strain. Hatchlings were counted daily and removed. Hatchability

for each pod was obtained by dividing hatchling number by the total number of

eggs, determined by summing the former figure and the number of unhatched

eggs.

In studies by PWR, fifth instars of M. sanguinipes and M. packardii were

sampled from Wynyard (Wyn) and Davidson (Dav), Saskatchewan, respectively

and allowed to mature in the laboratory. The former was crossed reciprocally (14

pairs) with Pickford and Randell’s nondiapause strain (in its 45th generation of

selection at the time of the study) and the latter (10 pairs) with a nondiapause

strain (in its seventh generation of selection for nondiapause) initiated by PWR.

To avoid confusion with NDwc used by WC, the nondiapause strains of PWR

are labelled NDpr and NDpr-pac, respectively. Experimental conditions were the

same as those of WC except that egg pods were transferred to petri dishes con¬

taining moistened filter paper.

Statistical significance of reciprocal differences and of other linear contrasts (see

below) was assessed by a modified Mest that takes into account unequal sample

variances (Zar 1984), which is the case here. Nonparametric tests of significance

1991

CHAPCO & RIEGERT: MELANOPLUS BIOLOGY

T able 2. Analysis (i-tests) of reciprocal differences (maternal effects) and homogenic vs heterogamic

mating differences (egg effects).

Strain

Reciprocal differences

ND strain vs ND

22 x field 6S

Field strain vs field

22 x ND S6

Bet

***

Que

***

Txl

Tx2

***

Wyn

***

—

Dav

***

—

ns—not significant.

*-P < 0.05.

**-P < o.Ol.

***_p < 0 . 001 .

(Campbell 1967) essentially yielded the same conclusions as the (-tests, attesting

to the robustness of the latter.

Results and Discussion

Mean percent hatchabilities along with standard errors are presented in Table

1 for the field strains and reciprocal crosses with nondiapause strains. Hatchabil¬

ities for the WC field material ranged from almost 0 (Bet) to 39% (Txl), far below

the value for the nondiapause strain, 79%. A comparison of values for reciprocal

crosses suggests a maternal influence on the trait. In general, hatchabilities of eggs

laid by nondiapause females mated with field males are consistently greater (sig¬

nificantly so in 5 cases; Table 2) than values for the reciprocal crosses. For example,

the hatchability of eggs produced by the cross, NDwc 29 x Bet 86, exceeds the

value for Bet 22 x NDwc 88 by about 65%. The phenomenon is apparently not

restricted to the nondiapause strain. Que and BC are two field strains that differ

in hatchability and exhibit significant reciprocal differences (P < 0.01): values for

Que 22 x Bet and Bet 22 x Que 66 are 0.7 ± 0.7 (10) and 39.2 ± 9.9 (9),

respectively. Analysis of the PWR data on M. sanguinipes and M. packardii also

reveals significant differences between reciprocal crosses (Table 2) adding further

support to the claim for maternal effects. It is difficult to assess what these effects

really represent. Had all strains been reared under the same conditions, the claim

could be made that maternal effects are genotypically mediated on the female

parent’s side. This conclusion may be valid for the southern populations since

insects were uniformly treated (at the University of Regina) and reared from eggs

that had been sent although transgenerational effects are not unknown (Sander et

al. 1985). In any case, the experimental design precludes a proper separation of

genetic and environmental factors that underly the maternal effects.

Despite the ambivalence regarding inheritance and hatchability at the maternal

level, a genetic analysis is possible, but at the level of the egg. The availability,

on the whole, of two sets of homogamic and heterogamic crosses permits an

examination of the possible role of the egg’s genotype on hatchability. If maternal

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

effects, genic or nongenic, were entirely responsible for the trait, then it would be

expected that values would be the same irrespective of the source of the male.

To test this possibility, differences between means for crosses with the same

maternal type were examined (Table 2, last two columns). Out of a total of nine

contrasts involving nondiapause females, there were five significant differences

and out of a total of seven contrasts involving field females, there were three

significant differences. The magnitude of these differences varies. To illustrate, in

the Quebec study, the hatchability of eggs laid by NDwc females mated to Que

males is in the same direction as that for the NDwc strain (i.e., indicating a

maternal effect), but the value (54%) is significantly less than the value for NDwc

(79%). In a few situations the hatchability of F t eggs is greater than the corre¬

sponding parental value, a result that suggests heterotic effects. For instance, the

hatchability associated with the cross, NDwc 22 x Az 66, is 92%, a value signif¬

icantly greater than the figure for NDwc. It would appear, therefore, that the

genotype of the egg can, in some cases, influence its hatchability.

The findings with respect to maternal effects are germane to those wishing to

maintain cultures of nondiapause grasshoppers that serve as model systems for

purposes of investigating natural populations. It is not uncommon for laboratory

strains to experience a reduction in numbers and a subsequent loss of genetic

variation, or experience inadvertent selection as a result of adaptation to labo¬

ratory conditions. After a number of generations, a laboratory strain may no

longer be representative of its species. The problem can be ameliorated to some

extent by mating virgin females from the nondiapause strain with field-caught

males thereby increasing the storehouse of genetic variation. By virtue of the

maternal influence revealed in this paper, the inconvenience that diapause would

otherwise present can be circumvented. Indeed, two such “hybrid strains” had

been successfully propagated with the Louisiana and Arizona materials for seven

generations, when they were terminated.

Acknowledgment

We thank M. Dakin of the University of Southwestern Louisiana, J. R. Hilliard

of Sam Houston State University, and R. N. Foster and B. Thomley of USD A,

Phoenix, Arizona for their unselfish help in the collecting of grasshoppers. Fi¬

nancial support from the Natural Sciences and Engineering Research Council

(Strategic Grant G-0399 to WC) is gratefully acknowledged.

Literature Cited

Campbell, R. C. 1967. Statistics for biologists. Cambridge University Press, Cambridge.

Chapco, W. 1984. Genetics of grasshoppers. I. Linkage studies of albino, reduced eye, and a new

trait, bloody, in Melanoplus sanguinipes. J. Hered., 75: 362-364.

Mather, K. & J. L. Jinks. 1982. Biometrical genetics. Chapman and Hall, London.

Oma, E. A., D. A. Streett & F. E. Henry. 1990. Establishment of a nondiapause Melanoplus differ-

entialis (Thomas) (Orthoptera: Acrididae). Can. Entomol., 122: 583-584.

Pickford, R. 1958. Observations on the reproductive potential of Melanoplus bilituratus (Wlk.)

(Orthoptera: Acrididae) reared on different food plants in the laboratory. Can. Entomol., 90:

483-485.

Pickford, R. & R. L. Randell. 1969. A non-diapause strain of the migratory grasshopper Melanoplus

sanguinipes (Orthoptera: Acrididae). Can. Entomol., 101: 894-896.

Sander, K., H. O. Gutzeit & H. Jackie. Insect embryogenesis: morphology, physiology, genetical and

1991

CHAPCO & RIEGERT: MELANOPLUS BIOLOGY

molecular aspects, pp. 319-385. In Kerkut, G. A. & L. I. Gilbert (eds.). Comprehensive insect

physiology, biochemistry and pharmacology. Vol. 1. Pergamon, New York.

Slifer, E. H. & R. L. King. 1961. The inheritance of diapause in grasshopper eggs. J. Heredity, 52:

39-44.

Zar, J. H. 1984. Biostatistical analysis. Prentice-Hall, Englewood Cliffs, New Jersey.

Received 31 August 1990; accepted 3 December 1990.

PAN-PACIFIC ENTOMOLOGIST

67(2): 86-98, (1991)

LIFE HISTORY AND DESCRIPTIONS OF IMMATURE

STAGES OF TEPHRITIS BACCHARIS (COQUILLETT)

ON BACCHARIS SALICIFOLIA (RUIZ & PAYON)

PERSOON IN SOUTHERN CALIFORNIA

(DIPTERA: TEPHRITIDAE)

Richard D. Goeden and David H. Headrick

Department of Entomology, University of California,

Riverside, California 92521

Abstract. — Tephritis baccharis (Coquillett) is bivoltine and monophagous on Baccharis salicifolia

(Ruiz & Pavon) Persoon in southern California. The egg, second and third instar larvae, and

puparium are described and illustrated. Eggs are inserted singly into terminal buds of main and

axillary branches in late winter or early spring. First instar larvae hatch and tunnel into the pith

just basad to the apical bud where they initiate gall formation. Gall and larval growth continue

slowly into the fall, when pupation occurs. Most Fj flies emerge, mate, and after about one week

begin oviposition. Some F, flies that emerge in the fall may overwinter as adults; however, a

few F, individuals may not emerge and instead overwinter as pupae in puparia in mature galls.

Second and third instar, F 2 larvae also overwinter in developing galls. Flies were long-lived

under laboratory conditions; males and females lived an average of 140 days and 83 days,

respectively. This longevity and the long fecundity period of females allow T. baccharis to attack

the new buds and branch growth produced by B. salicifolia, because this phraetophyte is capable

of protracted, nearly year-round vegetative growth and flowering in southern California.

Principal natural enemies of T. baccharis include three, solitary, primary, parasitoids, Pnigalo

sp. (Hymenoptera: Eulophidae), Halticoptera sp. and Pteromalus sp. (Hymenoptera: Pteromal-

idae), and unidentified birds as important predators on overwintering larvae in galls.

Key Words.— Insecta, biology, gall, parasitoids, bird predation, mating behavior, chemosensilla

Among 18 described species of Tephritis indigenous to North America (Foote

1960, Stolzfus 1977, Foote & Blanc 1979, Jenkins & Turner 1989), only the

biology of T. stigmatica (Coquillett) on Senecio spp. heretofore was well known

(Tauber & Toschi 1965, Goeden 1988a). This paper details the biology of a second

Nearctic species, T. baccharis (Coquillett), on Baccharis salicifolia (Ruiz & Pavon)

Persoon [= B. glutinosa Persoon and B. viminea deCandolle (McVaugh 1984)],

Asteraceae in southern California.

Interest has been expressed in the biological control of B. salicifolia (Boldt &

Robbins 1990), other Baccharis spp. (Boldt & Robbins 1987, Boldt et al. 1988),

and other indigenous weeds (Pemberton 1985) in the U.S. with natural enemies

obtained from South America. Also, stenophagous Baccharis, feeding insects from

North America have been imported to Australia for the biological control of B.

halimifolia L. (Palmer 1987, Julien 1987, Palmer & Bennett 1989).

Materials and Methods

Field observations, laboratory examination, and the rearing of field collected

galls from several locations (see below) in southern California during 1983-1990

provided most of the information reported herein. These field data were supple¬

mented by laboratory cagings, in 1989, of single males and females or paired

reared adults of opposite sex. The adults were reared in 850 ml, clear plastic cages

1991

GOEDEN & HEADRICK: TEPHRITID LIFE HISTORY

fitted with screened lids for ventilation and basal water reservoirs in which ab-

sorbant cotton wicks and bouquets of excised vegetative branches of B. salicifolia

were emersed through a hole in each cage bottom. Honey striped with a bulb and

syringe on the underside of the lids provided food for the flies. Flies or parasitoids

were reared from larvae and puparia dissected from galls held separately in cotton-

stoppered, glass, shell vials within humidity chambers at 22-24° C and 76% RH.

Plant names follow Munz (1974) and McVaugh (1984); tephritid names follow

Jenkins & Turner (1989). Voucher specimens of T. baccharis from each study site

are located in the research collection of RDG (Department of Entomology, Uni¬

versity of California, Riverside). RDG also has established a separate collection

of hymenopterous parasitoids of California Tephritidae; DHH has established a

separate collection of immature Tephritidae.

Immature stages were described from two eggs dissected from buds (laboratory

cagings), a second and third instar larva, and a puparium dissected from galls.

Larvae and eggs were treated for scanning electron microscopy (SEM) as described

in Headrick & Goeden (1990a). Specimens were examined and micrographs pre¬

pared at 15 kV accelerating voltage, using Polaroid SS P/N film on a JEOL JSM-

C35 SEM, located in the Department of Nematology, University of California,

Riverside. The third instar larva is described in detail using the nomenclature

and format adopted by Headrick & Goeden (1990a); the second instar larva

description is limited to observed differences. Means ± SE are provided through¬

out the paper.

Taxonomy

Coquillett (1894) described T. baccharis as a Trypeta. Jenkins & Turner (1989)

revised the Baccharis- infesting tephritids of North America, reviewed the tax¬

onomy, designated the lectotype, illustrated and measured ova, and described and

illustrated important characters of adult T. baccharis.

Egg. — Five eggs dissected from oviposition punctures (Fig. 1A) were white,

fusiform, 0.68 ± 0.02 mm long and 0.26 ± 0.01 mm wide. The chorion is

reticulated and the apex bears a 0.04 mm button-like, honey-combed pedicel (Fig.

2A). Polygonal reticulation of the chorion (Fig. 2B) is common to all eggs of

Tephritidae examined to date (by DHH), but is highly developed in Tephritis

species, especially T. baccharis (Jenkins & Turner 1989). An unusual feature of

the reticulation is its differential complexity. The end embedded in plant tissue

has more shallow fenestrations (Fig. 2C), which increase in height and become

topped with papillae toward the heavily ridged pedicel (Fig. 2D). The plant itself

may provide some measure of protection for the smooth end of the egg, which

probably has little gas exchange function for the developing embryo, because it

is inside plant tissue. The polygonal reticulation strengthens the exposed chorion,

protecting not only the embryo, but also the pedicel through which most gas

exchange probably takes place (as seen in a more exaggerated form in the eggs of

Paracantha gentilis Hering [Headrick & Goeden 1990a]).

Third Instar Larva. — A single third instar larva measured 5.3 mm long and 2.5

mm wide (Fig. 3A). It is creamy white, superficially smooth, elongate, cylindrical,

tapered and bluntly rounded anteriorly, and gradually increasing in width to its

truncate posterior end.

The gnathocephalon is less conical and more reduced, and the rugose pads are

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(2)

Figure 1. Life stages and galls of Tephritis baccharis on Baccharis salicifolia : (Bar = 1 mm) (A)

egg protruding from ovipositional puncture in bud; (B) lateral view of small, immature gall; (C) sagittal

section through small immature gall exposing feeding chamber; (D) third instar larva in feeding

chamber in full-size gall; (E) gall with apical meristem killed by larval feeding; (F) puparium in feeding

chamber below exit tunnel for adult; (G) lateral view of mature gall with round window through which

adult emerges; (H) mating adults, dorsal view; (I) mating adults, ventral view.

1991

GOEDEN & HEADRICK: TEPHRITID LIFE HISTORY

Figure 2. Egg of T. baccharis : (A) habitus, dissected from B. salicifolia; (B) detail of polygonal

reticulation; (C) detail of point of insertion into plant tissue; (D) pedicel at exposed end of egg.

more dorsoventrally elongate than in most species of tephritid larvae examined

by us to date (Fig. 3B: 1). The gnathocephalon bears flattened anterior sensory

lobes separated by a medial depression (Fig. 3C). The paired dorsal sensory organs

lie just dorsad of the anterior sensory lobes and are composed of a single papilla

(Fig. 3C: 1). The sensory lobes bear the lateral sensory organ, the pit sensory organ

and the terminal sensory organ (Fig. 3C: 2, 3, 4), which share the same structure

and placement as with other tephritid larvae examined to date (DHH, unpublished

data). The integumental petals which surround the mouth lumen (Fig. 3B: 2) are

much reduced in comparison with other tephritid larvae (e.g., Stenopa affinis

Quisenberry [Goeden & Headrick 1990]). The mouth hooks are heavily sclerotized

and tridentate. The teeth are stout and bluntly conical (Fig. 3B: 3). A median oral

lobe, which was not visible with SEM, was observed with a dissecting microscope;

this brings the total number of nonfrugivorous tephritid larvae with a median

oral lobe to 15 species (Headrick & Goeden 1990a; Goeden & Headrick 1990;

DHH, unpublished data). Lateral lobes bearing several sensilla were located dor-

solaterally on the edge of the mouth lumen (Fig. 3B: 4), are similar in placement,

and share similar types of sensilla to those observed in S. affinis (Goeden &

Headrick 1990).

The prothorax is smooth and bears several flattened sensilla. Larvae of Te-

phritidae typically have flattened sensilla on the prothorax (Foote 1967, Headrick

& Goeden 1990a); however, T. baccharis has at least four sensilla, each with a

finger-like projection surrounded by a collar. These sensilla may be mechanically

1991

GOEDEN & HEADRICK: TEPHRITID LIFE HISTORY

stimulated by deflection (Fig. 3D). The anterior thoracic spiracle is located dor-

solaterad on the posterior margin and bears four papillae, each in turn topped

with a distinct, smooth, rounded smaller papilla (Fig. 3E).

The succeeding segments are superficially smooth and demarcated by a de¬

pression that circumscribes the body. The integument adjacent to the segmental

line is reticulated with shallow depressions and has intersegmental bands of minute

acanthae (Fig. 3F: 1). Each segment bears a group of three sensilla arranged in a

vertical row, posterior to the segmental line on the lateral aspect of the body (Fig.

3F: 2). The sensilla are smooth with a central pore, and are similar in shape and

placement to the lateral sensilla described for S. affinis (Goeden & Headrick 1990).

The caudal segment bears the posterior spiracular plates composed of three

elongate oval rimae about 0.07 mm long (Fig. 3G: 1), and four interspiracular

processes with two to five branches; the longest process measured 0.03 mm in

length (Fig. 3G: 2). The caudal segment also has the typical arrangement of sensory

papillae that includes dorsal, lateral and ventral pairs of finger-like projections in

a basal collar located on the posterior margin of the body (Fig. 3H: 1). A tuberculate

chemosensillum is associated with each of the two dorsal sensilla in a shallow

depression (Fig. 3H: 2). On the apex of the chemosensillum is a raised crown that

bears several open pores. This sensillum has not been observed or described for

any other tephritid larva to date (DHH, unpubl. data).

Second Instar Larva. — A second instar larva dissected and examined from an

overwintering gall measured approximately 3.5 mm long and 1.5 mm wide (Fig.

4A). It is a translucent white, cylindrical, with a cone-shaped gnathocephalon,

and rounded posteriorly. Most structures were similar in shape and placement to

those of the third instar larva. The gnathocephalon is slightly rugose, with elongate

pads as seen in the third instar. The anterior lobe is flattened and bears all three

sensory organs as well as the closely associated dorsal sensory organ. The mouth

hooks are bidentate and not bluntly rounded at the apex as in the third instar

larva. The caudal segment has dorsal, lateral and ventral finger-like sensilla around

the posterior margin. The posterior spiracular plates bear three rimae and inter¬

spiracular processes with one or two blades about 0.01 mm in length.

Puparium. -Fifty-two of 61 puparia of T. baccharis dissected from galls av¬

eraged 1.99 ± 0.03 (range, 1.50-2.31) mm in widest width, and 40 of these that

were intact averaged 4.37 ± 0.07 (range, 3.34-5.59) mm in length. The puparia

are superficially smooth, distinctly segmented, elongate ellipsoidal, anteriorly

rounded, and slightly flattened posteriorly and dorsoventrally. The anterior end

bears the invagination scar of the mouth and the anterior thoracic spiracles (Fig.

4B: 1, 2). The posterior end is finely wrinkled without protruding spiracular plates

(Fig. 4C). One of 61 puparia dissected from galls was mostly unpigmented and

black only at both ends; all other puparia were uniformly black. Goeden (1988a)

Figure 3. Third instar larva of T. baccharis : (A) habitus, anterior end at left; (B) detail of gnath¬

ocephalon, 1—rugose pads, 2—integumental petals, 3—mouth hooks, 4—lateral sensory lobes; (C)

anterior sensory lobes (dorsal at left), 1 — dorsal sensory organ, 2—lateral sensory organ, 3—pit sensory

organ, 4—terminal sensory organ; (D) sensillum on anterior of prothorax; (E) anterior thoracic spiracles;

(F) lateral aspect of the body, 1—acanthae, 2—lateral sensilla; (G) posterior spiracular plate (dorsal

at right), 1—rimae, 2—interspiracular processes; (H) posterior sensilla, 1—finger-like process, 2—

dome sensillum.

Figure 4. Second instar larva and puparium of T. baccharis: (A) habitus, second instar (anterior

at right); (B) anterior end of puparium, 1—mouth invagination, 2—anterior thoracic spiracles; (C)

posterior end of puparium.

1991

GOEDEN & HEADRICK: TEPHRITID LIFE HISTORY

reported incomplete pigmentation of some T. stigmatica puparia. This color vari¬

ation also has been observed among some flower head infesting Trupanea (Goeden

1988b), gallicolous Procecidochares spp. (RDG, unpublished data), and was re¬

ported by Headrick & Goeden (1990a) among puparia of Paracantha gentilis

Hering.

Distribution, Host, and Study Sites

Jenkins & Turner (1989) described the range of T. baccharis as “eastcentral

Mexico northwest to westcentral California.” The distribution of its only known

host plant, B. salicifolia (Wasbauer 1972, Jenkins & Turner 1989) was described

by Boldt & Robbins (1990) as “common along waterways in the southwestern

United States and northern Mexico as well as western and southern South Amer¬

ica.”

Tephritis baccharis has been reared only from bud galls on B. salicifolia, never

from capitula of this or any other species of Baccharis or Asteraceae widely

sampled throughout California since 1980 (RDG, unpublished data). Nearctic

species of Tephritis are either obligate or facultative gall formers on branches or

stems, or ovule feeders in capitula (Foote 1960; Tauber & Toschi 1965; Jenkins

& Turner 1989; Goeden 1988a; RDG, unpublished data). Tephritis baccharis is

a monophagous, obligate gall former.

Our field study locations for flies and galls augment the California and south¬

western U.S. distributions plotted by Foote & Blanc (1963) and Jenkins & Turner

(1989), respectively: W end of Central Valley, Laguna Canyon, and Cebada Can¬

yons (Goeden 1983), Santa Cruz Island, Santa Barbara Co., 8-13 Oct 1983; Big

Morongo Canyon, Riverside Co., 7 Mar 1985, 3 Apr 1986, 23 Jan 1989, 30 Jan

1990; Oriflamme Canyon, NW San Diego Co., 8 Mar 1989. Additional locations

where the characteristic galls on B. salicifolia were observed or collected included:

Canebrake Creek, N of Spring Canyon, NW Kern Co., 3 Mar 1987; Mission

Canyon, Riverside Co., 2 Apr 1987; Scissors Crossing, NW San Diego Co., 19

Dec 1989 and 9 Jan 1990. This fly, like B. salicifolia, is widely distributed in

riparian habitats in southern California; however, its galls were not observed at

many other locations where flower heads of this dioecious shrub sampled yielded

other Tephritidae (Goeden 1983; RDG, unpublished data).

Biology

Egg. — In laboratory cagings, eggs were inserted for all or most of their lengths,

mainly into apical buds, and a few into axillary buds. Only the highly reticulated,

pedicellar ends slightly projected beyond or below the edges of the longitudinal

egg punctures (Fig. 1A). Three egg punctures measured 0.55 mm long by 0.15

mm wide. The long axes of the egg bodies, buds, and branches were parallel. Bud

tissues turned necrotic in areas immediately surrounding the eggs, as reported for

Eutreta diana (Osten Sacken), another gallicolous tephritid (Goeden 1990). One

unhatched egg contained an embryo with its partly pigmented cephalopharyngeal

skeleton located at the pedicellar end; however, the first instar larva always ecloses

through a slit in the chorion near the end opposite the pedicel, and immediately

begins tunneling basipetally. Apparently the embryo of T. baccharis turns 180°

just before eclosion, like several other species of nonfrugivorous Tephritidae (Var-

ley 1937, Cavender & Goeden 1982, Goeden 1987, Goeden et al. 1987). This

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

behavior apparently evolved to allow instant entry by the delicate, newly eclosed

first instar larvae into moist, host plant tissues, and also to accommodate embryo

respiration via the exposed pedicel (Headrick & Goeden 1990a).

All galls examined contained only one larva or puparium, indicative of only

one egg having been oviposited in each bud in nature; however, in cagings, where

ovipositional sites were scarce, as many as 10 eggs were inserted in a single apical

bud. Similarly, in the field, most current generation galls were solitary at ends of

separate branches; only one branch from Oriflamme Canyon bore two similar

aged galls connected by a narrow neck of ungalled intemode. Both sexes of B.

salicifolia bore galls of T. baccharis.

Larva. — Two newly hatched larvae averaged 0.49 mm in length. The first instar

tunneled until it reached the pith of the branch just basad to the apical meristem.

The necrotic tunnel of one first instar was traced for 9 mm from this incipient

gall site distally to its empty egg chorion. The young larva settles, molts, and

initially excavates a small ellipsoidal cell about 3 mm long and <2 mm wide in

the juvenile gall, evidenced externally as only a slight swelling of a branch terminal

(Figs. IB, 1C). Here, it feeds and develops (perhaps sporadically) through the

summer or the following fall and winter, depending upon which generation it

represents, F l or F 2 , respectively (see below). The molt to the second instar could

be ascertained by the presence of a small, intact, cephalopharyngeal skeleton

discarded at one end of the cell. Once plant growth begins in mid-winter to early

spring, or is triggered once again later in the year by late summer rainfall as

sometimes occurs in southern California, gall and larval growth are accelerated

and proceed concurrently.

Twenty (87%) of 23 fully formed F 2 galls of the previous year’s overwintering

generation sampled at Oriflamme Canyon were fusiform; the remaining galls,

subovoidal. The former shape resulted from continued growth of the bud at each

gall apex; whereas, the latter shape resulted from death of the terminal bud from

larval feeding by T. baccharis. Such intragenerational differences in shape also

were noted among galls of T. stigmata by Goeden (1988a), and the causes for this

difference were discussed relative to galls of E. diana by Goeden (1990). The galls

of T. baccharis, like those of T. stigmatica, are shortened, thickened, succulent

terminal parts of main or axillary branches (Figs. ID, IE). The 23 mature galls

incorporated an average of 3.8 ± 0.3 (range, 1-6) nodes, and were smooth surfaced,

light green and longitudinally striped or unilaterally colored red-purple when

occupied. When empty, the galls turned tan, shriveled, and dehisced, or became

woody and persisted as branch swellings. Externally, the 23 galls averaged 13.1

± 0.5 (range, 8-19) mm in length, and 5.6 ± 0.2 (range, 4-7) mm in width. The

larva feeds on the expanded pith parenchyma which comprises the bulk of the

gall (Figs. 1C, ID). The feeding larva eventually hollowed out an ellipsoidal,

central, longitudinal, basally rounded cavity. This open feeding cavity was irreg¬

ularly surfaced with yellow-white callose tissue, and averaged 5.5 ± 0.2 (range,

4.4-7.8) mm in length and 2.7 ± 0.1 (n = 23; range, 2.1-3.5) mm in width (Fig.

ID).

Twenty (66%) of 30, F t galls collected on Santa Cruz Island were fusiform; the

remainder, subovoidal. These 30 galls incorporated 3.8 ±0.1 (range, 3-5) axils

and externally measured 20.4 ± 1.0 (range, 12-33) mm long by 8.3 ± 0.3 (range,

6-13) mm wide.

1991

GOEDEN & HEADRICK: TEPHRITID LIFE HISTORY

When fully grown, the larva extends the feeding cavity into an exit tunnel 1.7

± 0.1 (n = 23; range, 0.8-3.2) mm long, which may end apically (and thus kill

the terminal bud, Fig. IE), or more commonly ends subapically (Fig. IF) in an

oval to round, thin, cuticular window 1.6 ± 0.1 (range, 1.1-2.1) mm in diameter

(Fig. 1G).

Puparium.— Pupation occurs in the feeding chamber with the posterior end of

the puparium resting on, or up to, about 1 mm above the bottom of the feeding

chamber. The anterior end of the puparium normally faces the window to allow

egress of the emerging adult (Fig. IF).

Adult.— Five males and five females (F x ) emerged between 13 Nov and 6 Dec

1985 from galls collected on Santa Cruz Island during the previous October. Six

males and three females emerged from 22 Mar to 5 Apr (overwintered F 2 ) from

galls collected in Oriflamme Canyon in Mar 1989. Both sexes emerged throughout

the emergence periods. Also, two females emerged from F! galls collected at Big

Morongo Canyon on 23 Jan 1985. One male and one female emerged on 15 Jan

1990 from F t galls apparently containing these individuals as fully developed

pupae within puparia collected at Scissors Crossing on 9 Jan 1990. These latter

two rearing records demonstrate that some F, individuals also overwinter as

puparia in galls.

Apparently females are polygamous, males are polyandrous. Females began to

mate in laboratory cagings (« = 3) 1 to 2 days after their emergence. Pairs mated

repeatedly; one pair mated on at least 17 different days, and for the last time

when the female was 121 days old. Another pair caged together at emergence

mated on each of the first 7 consecutive days after they emerged. Mating by

virginal males and females in laboratory cagings was protracted, involving little

courtship behavior and no apparent postmating behavior. The male mounted the

female after a short period of mutual recognition evidenced when the flies stood

face-to-face and alternately, slowly waved their wings three to seven times. The

anal edge of each wing was turned downward as each wing was brought forward

until nearly perpendicular to the substrate in supination (Headrick & Goeden

1990b). The male then jumped over and atop the female or mounted her from

one side, forcibly spread her wings, and rapidly initiated coitus. The mating

posture (Figs. 1H, II) was much like that illustrated and described by Tauber &

Toschi (1965) for T. stigmatica. The hind tarsi of the male rest on the substrate

or move back and forth along the oviscape to help position the female’s terminalia,

his midtarsi grasp the middle of her abdomen laterally, and his foretarsi clasp her

abdomen at its juncture with the thorax. The mouthparts of the male are posi¬

tioned above her scutellum; the mouthparts of both flies pump rapidly during

copulation (four to five pumps/sec). The wings are held parallel to the substrate.

The wings of the female are spread at about 80° and centered over her abdomen;

those of the male are overlapped atop his dorsum or are spread up to about 30°

(Figs. 1H, II). Except for their mouthparts, both flies generally remain motionless

during copulation, although the female may groom her head and mouthparts with

her foretarsi, occasionally bring one wing forward, or even walk about while

carrying the male with his hind tarsi on or off the substrate. Matings were observed

throughout the daylight hours, and in one instance was extended over 2 days and

nights under artificial lighting. To avoid mating, a nonreceptive female turned in

tight circles, flew at the male, or turned and walked away. Males also mounted,

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

rode, but did not couple with nonreceptive females. For example, one male daily

mounted and remained in a mating posture atop a nonreceptive female for up to

6 h on each of 4 consecutive days after they last mated, but without coupling with

her.

Females began oviposition in bouquets in cages 1-2 weeks after emergence.

Probing did not always result in oviposition. An average of about one egg per

day was laid by each of seven females (range, 0.4-2 eggs daily). The oldest female

was 65 to 72 days old when she last oviposited. Five males lived an average of

140 (range, 58-196) days; three females, 83 (range, 38-142) days. Tauber & Toschi

(1965) reported the laboratory longevity of T. stigmatica adults when fed water,

sugar, and enzymatic protein hydrolysate of yeast as 6 to 187+ days. Long-lived

adults occur more widely among nonfrugivorous Tephritidae than is generally

recognized (Christenson & Foote 1960, Freidberg 1984). Whether longevity among

nonfrugivorous species correlates with the presence or absence of symbionts in

the adult as suggested by Harris (1989) remains to be addressed experimentally.

Seasonal History. — Tephritis baccharis usually is bivoltine on B. salicifolia in

southern California, but may be univoltine at some desert locations during dry

years, as reported with Procecidochares sp. by Silverman & Goeden (1980). Te¬

phritis baccharis does not additionally reproduce in capitula of the same host

species or of an alternate host, as do T. stigmatica (Coquillett) (Tauber & Toschi

1965, Goeden 1988a), T. arizonaensis Quisenberry (Goeden 1983; Jenkins &

Turner 1989; RDG, unpublished data), T. palmeri Jenkins, and T. subpura (John¬

son) (Jenkins & Turner 1989). However, the combination of long-lived adults

and long-fecund females in T. baccharis and a host phraetophyte capable of nearly

year-round growth and flowering under southern California conditions (Munz

1974), can lead to localized overlapping and obscuring of fly generations. This

reproductive flexibility on the part of T. baccharis facilitates utilization of buds

and new branch growth on its host plant, which itself shows reproductive flexi¬

bility, traits useful for survival in drainages with high water tables and riparian

habitats in otherwise harsh, arid environments.

Natural Enemies. — Three species of chalcidoid Hymenoptera were reared from

T. baccharis during this study. Three females of Pteromalus sp. (Pteromalidae)

were recovered from puparia as solitary endoparasitoids. Two males of Halti-

coptera sp. (Pteromalidae) were reared as solitary, larval ectoparasitoids. One

female of Pnigalo sp. (Eulophidae) was recovered from a small gall as a solitary,

ectoparasitoid of an early stage larva of T. baccharis.

Large holes in the sides of empty galls matched symptoms of predation by birds,

probably bush tits, Psaltriparus sp., recently documented for galls of Eutreta diana

(Goeden 1990). Five (14%) of 35 galls sampled from Oriflamme Canyon yielded

parasitoids; five more galls suffered bird predation. Fifty-nine (69%) of 86, mainly

F 2 galls of all sizes collected in Big Morongo Canyon on 30 Jan 1990 had been

opened by birds and the larvae within removed.

Acknowledgment

We sincerely thank D. W. Ricker, recently retired, and R. E. Orth for technical

assistance. We also thank F. L. Blanc, J. A. McMurtry, J. A. Teerink, and W. J.

Turner for their helpful comments on early drafts of this manuscript; and J. LaSalle

1991

GOEDEN & HEADRICK: TEPHRITID LIFE HISTORY

and I. Luhman, who identified the parasitoids when they were with the Depart¬

ment of Entomology, University of California, Riverside.

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(Compositae) in Texas. Environ. Entomol., 16: 887-895.

Boldt, P. E. & T. O. Robbins. 1990. Phytophagous and flower-visiting insect fauna of seepwillow,

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Boldt, P. E., W. Woods & T. O. Robbins. 1988. Phytophagous insect fauna of Baccharis sarothroides

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Cavender, G. L. & R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae)

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Christenson, L. D. & R. H. Foote. 1960. Biology of fruit flies. Annu. Rev. Entomol., 5: 171-192.

Coquillett, D. W. 1894. New North American Trypetidae. Can. Entomol., 26: 71-75.

Foote, B. A. 1967. Biology and immature stages of fruit flies: the genus Icterica (Diptera: Tephritidae).

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Foote, R. H. 1960. The genus Tephritis Latreille in the Nearctic region north of Mexico: descriptions

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71-85.

Foote, R. H. & F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect

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Foote, R. H. & F. L. Blanc. 1979. New species of Tephritidae (Diptera) from the western United

States, Mexico, and Guatemala, with revisionary notes. Pan-Pacif. Entomol., 55: 161-179.

Freidberg, A. 1984. Gall Tephritidae (Diptera). pp. 129-167. In Ananthakrishnan, T. N. (ed.). Biology

of gall insects. Oxford & IBH Publ. Co., New Delhi, India.

Goeden, R. D. 1983. Initial host-plant records for five species of fruit flies from southern California

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Goeden, R. D. 1987. Life history of Trupanea conjuncta (Adams) on Trixus californica Kellogg in

southern California. Pan-Pacif. Entomol., 63: 284-291.

Goeden, R. D. 1988a. Gall formation by the capitulum-infesting fruit fly, Tephritis stigmatica

(Diptera: Tephritidae). Proc. Entomol. Soc. Wash., 90: 37-43.

Goeden, R. D. 1988b. Life history of Trupanea imperfecta (Coquillett) on Bebbia juncea (Bentham)

Greene in the Colorado Desert of southern California (Diptera: Tephritidae). Pan-Pacif. En¬

tomol., 64: 345-351.

Goeden, R. D. 1990. Life history of Eutreta diana (Osten Sacken) on Artemisia tridentata Nuttall

in southern California (Diptera: Tephritidae). Pan-Pacif. Entomol., 66: 24-32.

Goeden, R. D., & D. Headrick. 1990. Notes on the biology and immature stages of Stenopa ajfinis

Quisenberry (Diptera: Tephritidae). Proc. Entomol. Soc. Wash., 92: 641-684.

Goeden, R. D., T. D. Cadatal & G. A. Cavender. 1987. Life history of Neotephritis finalis on native

Asteraceae in southern California. Proc. Entomol. Soc. Wash., 89: 552-558.

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10: 9-16.

Headrick, D. & R. D. Goeden. 1990a. Description of the immature stages of Paracantha gentilis

(Diptera: Tephritidae). Ann. Entomol. Soc. Am., 83: 220-229.

Headrick, D. & R. D. Goeden. 1990b. Life history of Paracantha gentilis Hering (Diptera: Tephriti¬

dae). Ann. Entomol. Soc. Am., 83: 776-785.

Jenkins, J. & W. J. Turner. 1989. Revision of the Baccharis- infesting (Asteraceae) fruit flies of the

genus Tephritis (Diptera: Tephritidae) in North America. Ann. Entomol. Soc. Am., 82: 674-

685.

Julien, M. C. (ed.). 1987. Biological control of weeds (2nd ed.). CAB International Institue of

Biological Control, Wallingford, U.K.

McVaugh, R. 1984. Flora Novo-Galiciana, a descriptive account of the vascular plants of western

Mexico. Vol. 12. Compositae. University of Michigan Press, Ann Arbor.

Munz, P. A. 1974. A flora of southern California. University of California Press, Berkeley.

Palmer, W. A. 1987. The phytophagous insect fauna associated with Baccharis halimifolia L. and

THE PAN-PACIFIC ENTOMOLOGIST

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B. neglecta Britton in Texas, Louisiana, and northern Mexico. Proc. Entomol. Soc. Wash., 89:

185-199.

Palmer, W. A. & F. D. Bennett. 1989. The phytophagous insect fauna associated with Baccharis

halimifolia L. in the eastern United States. Proc. Entomol. Soc. Wash., 90: 216-228.

Pemberton, R. W. 1985. Native weeds as candidates for biological control research, pp. 869-877.

In Delfosse, E. S. (ed.). Proceedings VI international symposium on biological control of weeds,

Vancouver, Canada, Aug. 1984. Agriculture Canada, Ottawa, Ontario.

Silverman, J. & R. D. Goeden. 1980. Life history of a fruit fly, Procecidochares sp., on the ragweed,

Ambrosia dumosa (Gray) Payne, in southern California (Diptera: Tephritidae). Pan-Pac. En¬

tomol., 56: 283-288.

Stolzfus, W. B. 1977. The taxonomy and biology of Eutreta (Diptera: Tephritidae). Iowa State J.

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(Coquillett). Pan-Pacif. Entomol., 41: 73-79.

Varley, G. C. 1937. The life-history of some trypetid flies, with descriptions of the early stages

(Diptera). Proc. R. Entomol. Soc. Lond. (A), 12: 109-122.

Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico

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Received 16 August 1990; accepted 13 December 1990.

PAN-PACIFIC ENTOMOLOGIST

67(2): 99-106, (1991)

PARASITOIDS OF BLACK SCALE IN CALIFORNIA:

ESTABLISHMENT OF PROCOCCOPHAGUS PROBUS

ANNECKE & MYNHARDT AND COCCOPHAGUS RUSTI

COMPERE (HYMENOPTERA: APHELINIDAE)

IN OLIVE ORCHARDS

K. M. Daane, M. S. Barzman, C. E. Kennett,

and L. E. Caltagirone

Division of Biological Control, Department of Entomology,

University of California, Berkeley, California 94720

Abstract.— A survey of the parasitoid species of black scale, Saissetia oleae (Olivier), in California

central valley olive orchards revealed that two recently imported parasitoids, Prococcophagus

probus Annecke & Mynhardt and Coccophagus rusti Compere, have become established. These

South African parasitoids were released from 1979 to 1983. During the release period, neither

parasitoid was recovered. However, subsequent to the release, surveys of distribution of these

species revealed that both had become established. In the first survey (1984-1987), 23 commercial

olive orchards located in Tehama, Glenn, Madera, and Tulare Counties, were sampled. A second

survey (1988-1990) included seven olive orchards located in Tehama and Madera Counties,

and black scale infestations on selected ornamental olive and oleander plants in Tehama, Colusa,

Sutter, Glenn, and Butte Counties. Prococcophagus probus and C. rusti were reared from black

scale infested olive and oleander plants in Tehama County. Recovery of these species, over the

three years after the last release, provides evidence of their establishment in the Sacramento

Valley.

Key Words.— Insecta, Aphelinidae, Prococcophagus, Coccophagus, Homoptera, Saissetia oleae,

olive, biological control

Black scale, Saissetia oleae (Olivier), believed native to South Africa (De Lotto

1976), is found throughout the tropical and subtropical regions of the world. It

is a very polyphagous species with a host range that includes such common plants

to California as oleander (Nerium oleander L.), pepper tree ( Schinus molle L.),

coyote brush (Baccharis pilularis Wolf), and toy on {Heteromeles arbutifolia Lin¬

dahl) (Argyriou 1963). In California, S. oleae has been a pest of olive (Olea

europaea L.) and Citrus spp. since the 1880s. As a result of its wide host range

and high reproductive rate, the scale has spread throughout California and is

ubiquitous in the state’s central valley olive orchards, resulting in frequent infes¬

tations.

Black scale is exotic to most olive growing regions and has been the target of

numerous classical biological control programs (Bartlett 1978). Studies of popu¬

lation dynamics in the Mediterranean region have attributed significant scale

mortality to natural enemies (Argyriou & Katsoyannos 1976, Monaco 1976, Para-

skakis et al. 1980, Roberti 1981). In California, biological control programs against

Saissetia oleae began in 1891 and natural enemies have been imported since from

Asia, Africa, Australia, Central and South America, Europe, and the Middle East

(Bartlett 1978, Kennett 1986). Parasitoid importation and establishment in Cal¬

ifornia resulted in relatively successful control of black scale on ornamentals and

citrus in coastal regions. However, parasitoid establishment in the central valley

100

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

has not been as successful. Over 50 natural enemies of black scale have been

introduced in California and less than 15 have become permanently established

in the central valley (Table 1).

Establishment of black scale parasitoids is best understood in light of the pop¬

ulation dynamics of their host. Seven black scale stages can be differentiated from

egg to adult (Argyriou 1963), although most black scale parasitoids are specific

to only two or three consecutive host stages (Smith & Compere 1928, Compere

1940). Thus, it is difficult for some parasitoids to become established where scale

development is univoltine and synchronous because of the long periods in which

suitable host stages are not commonly found. Due to harsh climate in the central

valley, black scale development there is more synchronous than in coastal Cali¬

fornia (Daane 1988). For example, Metaphycus lounsburyi (Howard) is abundant

at coastal locations but is rarely found in the interior valleys (Kennett 1986).

Further, many of the initial attempts to establish imported species of black

scale parasitoids failed because of inadequate biological information (Bartlett

1978). For example, until the 1930s many primary parasitoids with obligatory

male hyperparasitic habits were discarded as undesirable secondary parasitoids.

Differences between parasitoid biotypes may also affect establishment. Panis &

Marro (1978) showed that there were sufficient behavioral differences between

two biotypes of Metaphycus lounsburyi (Howard) for them to be used in the same

orchard for biological control.

We report here the establishment of two black scale parasitoid species, Pro-

coccophagus probus Annecke & Mynhardt and Coccophagus rusti Compere, in

olive orchards in the Sacramento Valley, where they were previously thought to

have failed to become established. Prococcophagus probus was first released in

California in 1979. Coccophagus rusti was first released in 1937 and was reported

as established (Flanders 1952) in coastal locations, but not within the central

valley. It was imported from South Africa again in 1978 and released in central

valley olive orchards in 1980.

Materials and Methods

Parasitoid Release. —From 1979 to 1982, five parasitoid species were imported

to California from South Africa. Prococcophagus saissetiae Annecke & Mynhardt,

P. probus, C. rusti, Aloencyrtus saissetiae (Compere), and Metaphycus inviscus

Compere were collected by S. Neser (South African Ministry of Agriculture) and

sent to the quarantine station at the Division of Biological Control, University

of California, in Albany. After release from quarantine, colonies were established

in the insectary on black scale reared on oleander. From 1979 to 1983, 1180 P.

probus (67% female), 3265 P. saissetiae (59% female), 535 C. rusti (88% female),

265 A. saissetiae (98% female), and 1352 M. inviscus were released at five olive

orchards in the central valley (three orchards near Coming, Tehama County, one

near Parlier, Fresno County, and one near Porterville, Tulare County) and on

selected ornamental host plants of the black scale in the east bay region of the

San Francisco Bay area (Alameda and Contra Costa Counties). Releases were

concentrated in the fall of each year when the proper black scale host stages were

most common.

Recovery Attempts.— During the parasitoid release period (1979-1983), olive

twigs infested with black scale were collected at each release site in the fall and

1991

DAANE ET AL.: PARASITOIDS OF BLACK SCALE

101

Table 1. A partial list of introduced primary black scale parasitoids (source: Smith & Compere

1928, Bartlett 1978, Kennett 1986).

Species

Aphelinidae

Aneristus ceroplastae Howard

Coccophagus anthracinus Compere

Coccophagus baldassarii Compere

Coccophagus basalis Compere

Coccophagus capensis Compere

Coccophagus caridei (Brethes)

Coccophagus cowperi Girault

Coccophagus eleaphilus Silvestri

Coccophagus hawaiiensis Timberlake

Coccophagus japonicus Compere

Coccophagus lycimnia (Walker)

Coccophagus lycimnia (Walker) 3

Coccophagus mexicensis Girault

Coccophagus nigritus Compere

Coccophagus ochraceous Howard

Coccophagus eritreaensis Compere

Coccophagus rusti Compere

Coccophagus scutellaris (Dalman)

Lounsburyi trifasciatus (Compere)

Prococcophagus saissetia Annecke & Mynhardt

Prococcophagus probus Annecke & Mynhardt

Encyrtidae

Aloencyrtus saissetia (Compere)

Anicetus annulatus Timberlake

Coccidoxenus niloticus Compere

Diversinervus elegans Silvestri

Diversinervus smithi Compere

Encyrtus fuliginosus Compere

Encyrtus infelix Embelton

Metaphycus angustifrons Compere

Metaphycus bartletti Annecke & Mynhardt

Metaphycus citrinus Compere

Metaphycus flavus (Howard)

Metaphycus gilvus Compere

Metaphycus helvolus (Compere)

Metaphycus lichtensiae Compere

Metaphycus lounsburyi (Howard)

Metaphycus luteolus (Timberlake)

Metaphycus stanleyi Compere

Metaphycus zebratus Mercet b

Microterys flavus (Howard)

Microterys okitsuensis Compere

Microterys saissetiae Compere

Microterys tricoloricornis (DeSoto)

Eupelmidae

Lecanobius utilis Compere

Estab-

Origin

Date introduced

lished

Taiwan

1933, 1952

South Africa

1923

Eritrea

1953

Brazil

1958

South Africa

1924

yes

Brazil, Argentina

1934, 1935

East Africa

1937

yes

Eritrea, Morocco

1953,1953

Taiwan

1951

Japan

1951

Africa

1900s (accidental)

yes

Brazil, Argentina

1935,1935

Australia

1972

Africa, Mexico

Mexico

1956

Eritrea

1954

Africa

1990s (accidental)

yes

South Africa

1925, 1937

Kenya, Uganda

1937

yes

South Africa

1932,1981

yes

Africa

1900s (accidental)

yes

South Africa

1924

yes

South Africa

1978

South Africa

1978

yes

South Africa

1979

Australia, Taiwan

1931, 1951

Kenya

1937

Eritrea, Lebanon

1953,1964

yes

South Africa

1937

South Africa

1937

Hawaii

1921

yes

Taiwan

1952, 1957

South Africa

1956

yes

Eritrea

1953

Spain

1954,1955

North Africa

1953

South Africa

1937

yes

South Africa

1958

Australia

1916-1918

yes

Mexico

1954,1955

yes

South Africa

1937

yes

Spain

1986

Pakistan

1957

China, Japan

1951, 1952

Uganda

1937

Mexico

1956

Brazil, Argentina

yes

102

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Table 1. Continued.

Species

Origin

Date introduced

Estab¬

lished

Pteromalidae

Any sis saissetiae (Ashmead)

China, Pakistan

1924, 1957

Lecaniobus cockerelli Ashmead

West Indies

1913, 1915, 1940

Mesopelitita atrocyanea (Masi)

East Africa

1937

South Africa

1957

Mexico

1958

Scutellista caerulea (Fonscolombe)

South Africa

1901-1902

yes

Scutellista cyanea Motschulsky (red larval race)

Taiwan

1952

a Coccophagus lycimnia has been introduced from a number of countries, however, establishment

of this species may be from accidental introductions prior to importation efforts.

b Metaphycus zebratus has been recovered continually since its release; documentation of permanent

establishment is in progress.

spring of each year. Samples were brought into the laboratory and held in glass-

topped sleeve cages from four to six weeks. Emerging parasitoids were collected

three times a week and stored in 70% ethanol for later identification. These samples

were taken as part of a central and northern California survey of black scale

parasitoids that included a majority of the counties within the Sacramento and

San Joaquin valleys and several coastal counties extending from Napa County to

San Luis Obispo County (Kennett 1986). Nearly 70% of the samples were taken

from urban and rural landscapes (olive; Citrus cultivars of grapefruit, orange;

Modesto ash, Fraxinus velutina var. glabra Rehder; English holly, Ilex aquifolium

L.), 20% from natural stands of native shrubs (coyote brush and toyon), and 10%

from commercial olive orchards. Detailed results from these collections were

reported by Kennett (1986).

From 1984 to 1987, a separate parasitoid survey was conducted in conjunction

with a black scale population dynamics study on olive. All samples were taken

from commercial olive orchards located in the central valley: seven orchards near

Coming, Tehama County; one near Orland, Glenn County; one near Madera,

Madera County; one near Parlier, Fresno County; and 11 near Lindsay, one near

Strathmore, and one near Exeter, Tulare County. Included in this 23 orchard

study were four of the five orchards that received the five imported black scale

parasitoid species from 1979 to 1983. The release orchard in Porterville was

removed in 1984 and is, therefore, not included in the survey. Likewise, the

orchard in Parlier was removed in 1986 and data presented for parasitoids col¬

lection in 1987 does not include this orchard.

In each orchard sampled, except in Glenn County, collections were made every

six to eight weeks. In the Glenn County orchard, samples were taken each fall

and spring, when black scale parasitoids were most common. Olive twigs infested

with black scale were collected and the scales were counted and recorded by

developmental stage (first, second, third, pre-ovipositional, and adult) and con¬

dition (alive or parasitized). Infested twigs were placed in parasitoid emergence

containers and held between 22 and 24° C for three to four weeks. Emerging par¬

asitoids were collected and later identified.

From 1987 to 1989, seven orchards (five near Coming and two near Madera)

were sampled, as described above, in each season of each year. An additional

1991

DAANE ET AL.: PARASITOIDS OF BLACK SCALE

103

survey for parasitoids on black scale infested olive and oleander plants was carried

out in the spring of 1990 at seven sites in the Sacramento Valley: Red Bluff,

Tehama County; Willows, Glenn County; Yuba City, Sutter County; Zamora,

Yolo County; and Williams and Maxwell, Colusa County. Infested twigs were

taken to the laboratory and parasitoids reared from the material as described

above.

Results and Discussion

New Parasitoids. — Of the five parasitoid species released, only P. probus and

C. rusti were found to have become established in central valley olive orchards.

Establishment is defined here as parasitoid recovery three years after the last

insectary release.

Prococcophagus probus was collected in 19 8 5 and 198 6 in two of the four original

release orchards. Both orchards were located near Coming, Tehama County.

Prococcophagus probus comprised 0.7% of the total parasitoids collected between

1984 and 1987 (Table 2). It was most commonly found in winter samples. In one

orchard, P. probus comprised 8.0% (n = 274) and 1.1% {n = 1235), respectively,

of parasitoids in November and December collections combined for 1985 and

1986. It was not recovered from any orchards in Glenn, Madera, or Tulare Coun¬

ties, nor was it recovered from any orchards other than the original release sites.

This suggests that P. probus did not disperse from the release areas.

Coccophagus rusti was collected from orchards near Coming, Tehama County

from 1985 to 1987. The parasite was most commonly reared from third instar

and pre-ovipositional stage scale, collected in spring and early summer months.

Overall, this parasitoid comprised only 0.4% of the total parasitoids collected

between 1984 and 1987 (Table 2). When considering release orchards only, the

percent species composition of C. rusti was often higher. In one release orchard,

C. rusti comprised 8.2% ( n = 205) of the parasitoids reared from two collections

in June and July, 1987. Coccophagus rusti was also recovered from two nonrelease

sites: 1986, in an olive orchard over 1 km from the nearest release site, and 1990,

in Red Bluff on oleander over 64 km from the nearest known release site. At the

latter site, C. rusti accounted for 58%, the greatest number, of 19 parasitoids

reared. This suggests that it either is increasing its geographic distribution or that

a population has been present, without detection, in some central valley locations

since the releases in 1937. Although C. rusti has not previously been found in the

central valley, Flanders (1952) reported it to have been established in California

at coastal locations, on ornamental plants. Its low numbers in California, contrasts

reports that it was the most abundant and widely distributed of black scale par¬

asitoids collected in 1937 in South Africa, Uganda, and Kenya (Compere 1940).

Neither P. saissetiae nor A. saissetiae were recovered in the 1984 to 1987 or

1988 to 1990 surveys. Metaphycus inviscus was not recovered in either the 1984

to 1987 or the 1988 to 1990 surveys. These findings differ somewhat from those

of the 1979 to 1983 survey (during the release period) when M. inviscus was the

only newly released parasitoid to show continued and strong evidence of estab¬

lishment (Kennett 1986). Metaphycus inviscus is morphologically similar to Me¬

taphycus bartletti Howard, a commonly collected parasitoid, and can be easily

overlooked when large numbers of M. bartletti are present.

Abundance.— Relative to the other parasitoid species collected, both P. probus

104

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

T able 2. Rankings (in parenthesis) and percent total parasitiod abundance of primary and secondary

parasitoids reared from black scale samples taken in Tehama, Madera, and Tulare County olive

orchards between 1985 and 1987.

Percent Parasitoid Abundance

Sacramento

Valley San Joaquin Valley

Species Tehama Madera Tulare Total

Metaphycus bartletti a

Metaphycus helvolus 3

Scutellista cyanea a

Coccophagus lycimnia ab

Metaphycus zebratus a

Cheiloneurus inimicus 0

Marietta mexicanas

Coccophagus ochraceous a

Coccophagus scutellaris a ’ b

Tetrastichus minutum 0

Cheiloneurus noxius 0

Prococcophagus probus a

Coccophagus rusti*

Metaphycus luteolus a

Moranila californica*- 0

42.9 (1)

27.1 (2)

25.6 (2)

48.7 (1)

13.6 (3)

12.0 (3)

4.1 (4)

5.3 (4)

3.6 (5)

1-1 (7)

3.2 (6)

1.5 (6)

2.2 (7)

3.6 (5)

2.0 (8)

0.1 (10)

1.7 (9)

0.2 (9)

0.9 (10)

0.6 (8)

0.7 (11)

—

0.7 (12)

—

0.5 (13)

—

<0.1 (14)

—

<0.1 (15)

—

n = 8290 n = 936

51.9(1)

43.2 (1)

35.4 (2)

29.2 (2)

0.2 (10)

11.7 (3)

7.9 (3)

4.8 (4)

0.4 (8)

3.1 (5)

0.6 (7)

2.7 (6)

0.6 (6)

2.2 (7)

1.5 (5)

1.8(8)

2.7 (4)

1.7 (9)

0.2 (9)

0.8 (10)

—

0.5 (11)

—

0.5 (12)

—

0.4 (13)

—

<0.1 (14)

—

<0.1 (15)

n = 2469 n = 10,593

a Primary parasitoids

b Males developed as a secondary parasitoid on some similar species

c Secondary parasitoid

d Only one specimen found.

and C. rusti were not common, ranking 12th and 13th and comprising only 0.5

and 0.4%, respectively, of the total parasitoids collected (Table 2). The most

abundant parasitoids in the collections were Metaphycus bartletti Annecke &

Mynhardt and Metaphycus helvolus (Compere). The widespread distribution and

abundance of M. bartletti reveals its increasingly important economic role since

its introduction (Kennett 1980). Scutellista caerulea (Fonscolombe) (= Scutellista

cyanea Motschulsky) ranked third in abundance.

The remaining parasitoid species were found in low numbers, often sporadically

in time and space. Together, P. probus, C. rusti, Metaphycus luteolus (Timberlake),

and Moranila californica Howard comprised less than 1% of the total parasitoids

recovered. Metaphycus luteolus may have been underrepresented because of their

small number and close similarity to M. bartletti. Because only one specimen of

M. californica was found, while a colony was kept at the Division of Biological

Control insectary, contamination of the field emergence container may have oc¬

curred despite housings in separate buildings.

Metaphycus zebratus (Mercet), imported from Spain in 1985 and released from

1986 to 1988 (Daane & Caltagirone 1989), was recovered in Tehama, Madera,

and Tulare Counties. It ranked fourth in parasitoid species collected, despite being

released in large numbers in orchards being sampled. The parasitoid was recovered

in lower numbers each subsequent year after releases were discontinued.

Distribution. — Greater diversity and abundance of parasitoid species were found

in the Sacramento Valley than the San Joaquin Valley (Table 2), due, in part, to

1991

DAANE ET AL.: PARASITOIDS OF BLACK SCALE

105

differences in the synchronization of scale development found in the central valley.

Cultural practices typically used in the San Joaquin Valley exacerbate climatic

mortality and promote a more synchronous scale population than in the Sacra¬

mento Valley (Daane 1988). For S. caerulea, there is strong evidence that scale

phenology is limiting in the species abundance. Although the third most com¬

monly collected parasitoid in Tehama County, S. caerulea was rarely reared from

black scale collected on olive in Tulare County. Moreover, it has been found in

moderate numbers, in Tulare County, on oleander, where the scale has a less

synchronous development pattern (D. Bromberger, personal communication). It

is, therefore, not surprising that P. probus and C. rusti, as well as a number of

lesser abundant primary and secondary parasitoids, were found in the Sacramento

Valley rather than in the San Joaquin Valley.

Biology.— Documenting black scale parasitoid distribution and establishment

in often difficult because of the low number of specimens recovered and the

morphological similarity of other parasitoid species. Brief biological descriptions

and morphological diagnoses of P. probus and C. rusti are provided because of

the difficulty in identifying these two relatively unknown parasitoid species in

California.

Prococcophagus probus was described by Annecke & Mynhardt (1979) as a

solitary, primary parasitoid of black scale. Its reproductive biology is probably

similar to that of Prococcophagus varius Silvestri & P. saissetiae, two black scale

parasitoids that are morphologically very close to P. probus. In those species, both

male and female are primary parasitoids, but the diploid female larva develops

as an endoparasitoid, while the haploid male larva develops as an ectoparasitoid

(Mazzone & Viggiani 1983). The validity of Prococcophagus, as a genus separate

from Coccophagus, has been questioned by Mazzone & Viggiani (1984). Never¬

theless, Prococcophagus can be distinguished from Coccophagus by its ventrally

expanded antennal scape, laterally compressed funicle, and usually largely infus-

cated fore wings. The antennal scape of the female is more than twice as long as

wide, with two dark streaks on the outer surface; the first antennal funicle segment

is the longest, with rhinaria (Annecke & Mynhardt 1979). Superficially, P. probus

is similar to M. bartletti in color and size. In samples with many parasitoids,

misidentification can easily occur.

Coccophagus rusti is a solitary endoparasite of third instar and preovipositional

adult black scale (Flanders 1952). The male is hyperparasitic, developing as a

direct endoparasite of some primary parasitoids of black scale and other hosts.

This species can also develop in soft brown scale, Coccus hesperidum L. (Compere

1940). Coccophagus rusti is easily distinguished from other black scale parasitoids

by the following characters: the scutellum has three pairs of setae, the abdomen

is entirely black, the thorax is yellow-brown, the pedicel is shorter than first funicle

segment, and the forewing has an infuscation with a pale diagonal streak (Smith

& Compere 1928). The male differs from the female, with its whole body black

or brown-black (Annecke 1964).

Acknowledgment

We thank K. Hagen (University of California, Berkeley), G. Martin and L.

Ferguson (University of California, Davis), and S. Sibbett (Tulare County Co¬

operative Extension), K. Hagen, R. Gill, and L. Richards reviewed earlier manu-

106

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(2)

script versions and we appreciate their helpful suggestions. This research was

funded, in part, by the California Olive Committee, University of California

Statewide Integrated Pest Management Project, USDA-ES Smith-Lever IPM Proj¬

ect Development, W-84 Regional Project, and the Division of Biological Control.

Literature Cited

Annecke, D. P. 1964. The encyrtid and aphelinid (Hym: Chalcidoidea) parasites of soft brown scale,

Coccus hesperidum Linnaeus (Hemiptera: Coccidae) in South Africa. Rep. Africa Dept, of Agric.

Tech. Serv. Entomol. Mem., 7.

Annecke, D. P. & M. J. Mynhardt. 1979. On the type-species and three new species of Prococcophagus

Silvestri from South Africa (Hymenoptera: Aphelinidae). J. Entomol. Soc. South Afr., 42: 289-

297.

Argyriou, L. C. 1963. Studies on the morphology and biology of the black scale ( Saissetia oleae

(Bernard)) in Greece. Ann. Inst. Phytopath., Benaki (N.S.), 5: 353-377.

Argyriou, L. C. & P. Katsoyannos. 1976. Establishment and dispersion of Metaphycus helvolus

Compere in Kerkyra (Corfu) on Saissetia oleae (Olivier). Ann. Inst. Phytopath., Benaki (N.S.),

11: 200-208.

Bartlett, B. R. 1978. Coccidae. pp. 57-74. In Clausen, C. P. (ed.). Introduced parasites and predators

of arthropod pests and weeds: a world review. Agriculture Handbook, 480. U.S. Dept. Agric.,

Washington, D.C.

Compere, H. 1940. Parasites of the black scale, Saissetia oleae, in Africa. Hilgardia, 13: 399.

Daane, K. M. 1988. The effect of cultural practices on the population dynamics of black scale,

Saissetia oleae (Olivier) and its natural enemies in California olive orchards. Ph.D. Thesis,

University of California, Berkeley.

Daane, K. M. & L. E. Caltagirone. 1989. Biological control in California olive orchards: cultural

practices affect biological control of black scale. Calif. Agric., 43(Jan-Feb, 1): 9-11.

De Lotto, G. 1976. On the black scales of southern Europe (Homoptera: Coccoidea: Coccidae). J.

Entomol. Soc. S. Aff., 39: 147-149.

Flanders, S. E. 1952. Biological observations on parasites of black scale. Ann. Entomol. Soc. Am.,

45: 544.

Kennett, C. E. 1980. Occurrence of Metaphycus bartletti Annecke and Mynhardt, a South African

parasite of black scale, Saissetia oleae (Olivier) in central and northern California (Hymenoptera:

Encyrtidae; Homoptera: Coccidae). Pan-Pacif. Entomol., 56: 107-110.

Kennett, C. E. 1986. A survey of the parasitoid complex attacking black scale, Saissetia oleae (Olivier),

in central and northern California (Hymenoptera: Chalcidoidea; Homoptera: Coccidae). Pan-

Pacif. Entomol., 62: 363-369.

Mazzone, P. & G. Viggiani. 1983. Stato attuale dei parassitoidi di Saissetia oleae (Oliv.) in Italia,

con notizie biologiche preliminari sui Prococcophagus varius Silv. e P. saissetia Ann. e Myn.

(Hym.: Aphelinidae) di recente introduzione. Atti XIII Congr. Naz. It. Entomol., Sestriere.

Torino, 1983: 191-196.

Mazzone, P. & G. Viggiani. 1984. Osservazioni morfo-biologiche sulgi stadi preimmaginali di Pro¬

coccophagus varius Silv. e P. saissetia Ann. e Myn. (Hym.: Aphelinidae), parassiti di Saissetia

oleae (Oliv.) (Horn.: Coccoidea). Boll. Lab. Entomol. Agr. Filippo Silvestri, 41: 143-148.

Monaco, R. 1976. Nota su Metaphycus lounsburyi (How.) (Hym., Encyrtidae) parassita di Saissetia

oleae (Oliv.). Entomol. Bari, 12: 143-151.

Panis, A. & J. P. Marro. 1978. Variation du comportement chez Metaphycus lounsburyi (Hym.:

Encyrtidae). Entomophaga, 23: 9-18.

Paraskakis, M., P. Neuenschwander & S. Michelakis. 1980. Saissetia oleae (Oliv.) (Horn., Coccidae)

and its parasites on olive trees in Crete, Greece. Z. Ang. Entomol., 90: 450^164.

Roberti, D. 1981. Osservazioni sulla dinamica di popolazione e sulla parassitizzazione della Saissetia

oleae (Oliv.) su olivo in Puglia. Entomologica, 16, Bari, 25: 113-120.

Smith, H. S. & H. Compere. 1928. A preliminary report on the insect parasites of the black scale,

Saissetia oleae (Bern.). Univ. Calif. Pub. Entomol., 4: 231-334.

Received 6 November 1990; accepted 18 December 1990.

PAN-PACIFIC ENTOMOLOGIST

67(2): 107-114, (1991)

DESCRIPTION OF A NEW SPECIES OF EPIBLEMA

(LEPIDOPTERA: TORTRICIDAE: OLETHREUTINAE)

FROM COASTAL REDWOOD FORESTS IN

CALIFORNIA WITH AN ANALYSIS OF

THE FOREWING PATTERN

Richard L. Brown 1 and Jerry A Powell 2

department of Entomology, Drawer EM, Mississippi State, Mississippi 39762;

department of Entomological Sciences, University of California,

Berkeley, California 94720

Abstract. —Epiblema deverrae R. Brown, NEW SPECIES, is described from Monterey and Sono¬

ma Counties, California and is differentiated from the sympatric species, E. hirsutana (Wal-

singham) and E. radicana (Walsingham), and from similar allopatric species. A description of

its forewing pattern is based on an analysis of fasciae, color fields, and strigulae in the Olethreuti-

nae.

Key Words. — Insecta, Tortricidae, Eucosmini, Epiblema deverrae, forewing pattern, strigulae,

Asteraceae, Madia

An inventory of the Lepidoptera at the Landels-Hill Big Creek Reserve, Mon¬

terey County, California, has been conducted during the past decade (JAP, un¬

published data). This is the first attempt to census the entire fauna of moths and

butterflies at any locality in western North America. The Big Creek Reserve, one

of 27 areas maintained by the Natural Reserve System (UCNRS) of the University

of California, is situated about 30 km southeast of Big Sur, in the Santa Lucia

Mountains, which end abruptly at the Pacific coast. The reserve encompasses

1550 ha, rising from sea level to 1200 m elevation (900 m in the areas surveyed).

Rugged coastal mountains rise abruptly from the shoreline and are dominated by

narrow ridges separated by deep, V-shaped canyons with walls that rise steeply

from narrow streamside terraces. Vegetation types include coastal scrub, Califor¬

nia sage scrub, Ceanothus and chamise chaparral, redwood forest, mixed hard¬

wood forest, ponderosa pine-manzanita woodland, and sparse chaparral on rocky

montane ridges (buckwheat [Eriogonum], yerba santa [Eriodictyon], and Yucca

whipplei Torrey).

The reserve has been surveyed on more than 80 dates, using three approaches:

diurnal net collecting, larval sampling and rearing, and blacklight attraction at

sheets and traps. To date about 650 species have been differentiated, of which

26% are represented by larval collections. Based on comparisons with the numbers

of flowering species (326) (Bickford & Rich 1984), as well as butterflies at Big

Creek (57), and macro moths known at other stations in coastal California

(McFarland 1965; Opler & Buckett 1971; JAP, unpublished data), a census of

800-850 species of Lepidoptera at Big Creek is anticipated.

The survey has yielded several species of moths that were previously unknown

in the central coast ranges and at least three undescribed species of microlepidotera

in better studied families (Momphidae, Tortricidae), including a distinctive new

108

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

species of Epiblema. The genus Epiblema includes more than 100 species in the

Holarctic region and southern Asia. Of these, 39 occur in America, north of Mexico

(Blanchard 1979, 1985; Powell 1983; Miller 1985). Epiblema appears to be related

to Eucosma, Pelochrista, Sonia, and other eucosmine genera that include species

with larvae boring in stems and roots of Asteraceae. The description of this new

Epiblema is facilitated with selected information from a recent analysis of forewing

patters in Tortricidae (RLB, unpublished data).

Methods and Materials

A stereomicroscope with an ocular micrometer was used to examine and mea¬

sure specimens. Specimens of all species mentioned in the diagnosis were ex¬

amined except E. simploniana (Duponchel), of which figures were examined. The

fore wing length was measured from base to apex, including the fringe. Comuti of

the aedeagus were counted by examination of their sockets with a compound

microscope. Valval length was measured as a straight fine from the ventral, prox¬

imal comer of the sacculus to the apex of the cucullus. Colors were described with

the standards of Komerup & Wanscher (1983). The forewing description is based

on a comparative study of pattern elements in more than 600 tortricid species

representing all tribes defined by Horak & Brown (in press). The following ab¬

breviations are used for depositories: Mississippi Entomological Museum, Mis¬

sissippi State University (MEM); University of California, Berkeley (UCB).

Forewing Pattern Elements in Epiblema

Nijhout (1978) provided the following terminology for the five major types of

pattern elements on Lepidoptera wings, as described by Suffert (1929): ripple

patterns, dependent patterns, crossbands (fasciae), eyespots (ocelli), and color

fields. These types, except ripple patterns, are present in Epiblema, including the

species described here.

Fasciae occur on ontogenetically determined areas of the wing (Nijhout 1978,

1990) and generally are more darkly pigmented than the surrounding ground

color. The areas between fasciae here are termed interfascial areas. The distinc¬

tiveness of fasciae is dependent on both the number of scales that are darkly

pigmented (degree of expression) and the degree to which dark pigment is lacking

in the interfascial areas. Based on an analysis of forewing patterns in the Tortrici¬

dae, the ancestral pattern is interpreted to include six fasciae: basal, subbasal,

medial, postmedial, subterminal, and terminal. These fasciae, except for the ter¬

minal, previously have been identified in the tortricine tribe Cochylini (Bradley

et al. 1973). In the Olethreutinae, the basal and subbasal fasciae are often confluent

to form a basal patch; the median fascia usually is well expressed, although often

broken near CuA 2 to form a separate pretomal spot; the postmedian fascia (termed

the preapical fascia by some authors) is expressed usually as one or more spots,

the largest and most anterior being the postmedian spot (termed the preapical

spot by some authors); and the subterminal and terminal fasciae are either not

expressed or are represented by narrow umbrae or small spots (Fig. 1).

Color fields have been defined as large areas of the wing surface that contrast

in coloration with other areas and are not a fascia or other pattern element (Nijhout

1978). Suffert (1929) and Nijhout (1978) noted that “color fields” represented a

lumping of more than one type of pattern element, and that little effort had been

1991

BROWN & POWELL: NEW SPECIES OF EPIBLEMA

109

Figure 1. Epiblema deverrae, forewing pattern indicating positions of expressed and nonexpressed

fasciae relative to coastal strigulae 1-10. ifs—interfascial spot, mf—median fascia, o—ocellus, pmf—

postmedian fascia, sbf—subbasal fascia, stf—subterminal fascia, tf—terminal fascia, ts—terminal stri¬

gulae.

made in the analysis of the various types of color fields. Two types of color fields

commonly occur in the Tortricidae, and here these are termed “patches” and

“spots.” The term patch is used to describe a color field that differs from the

ground color and that covers at least one fascia and one interfascial area. Although

patches include several subgroups, depending on their position and size, patches

often involve the expression of fasciae only on the costal or inner margins of the

wing and the intervening interfascial area. The term spot is used for contrasting

pigmentation that is confined either to a restricted portion of a fascia (i.e., pretomal

spot) or to a restricted portion of an interfascial area (i.e., interfascial spot).

Interfascial spots are present on the inner margin (dorsum) between the subbasal

and median fasciae in many species of the Olethreutinae, including the Epiblema

described here (Fig. 1). Patches are common on the inner margin is generalized

genera in the Eucosmini, especially in Ancylis and Epinotia (e.g., E. solandriana

(L.)) (Bradley et al. 1979: pi. 32, Figs. 5-8) and on the costa in various unrelated

genera (e.g. Ancylis, Pseudexentera and Acleris) (Bradley et al. 1973: pi. 40, Figs.

10-11). Although interfascial spots are present in many species of Epiblema,

Eucosma, and related genera, patches have not been detected in any representative

of this derived group of Eucosmini.

Strigulae, or short transverse marks, are present on the costa of most Olethreuti¬

nae, some Chlidanotinae, and some generalized Tortricinae (e.g., Phricanthini)

(Horak & Brown in press). These strigulae appear to be dependent patterns, as

they occur on the inner and outer margins of fasciae. In addition, strigulae apical

to R 2 occur between the veins at the wing margin. Danilevski & Kuznetsov (1968)

and Kuznetsov (1978) recognized seven pairs of costal strigulae, and these were

numbered one to seven from apex to base. Examinations of species in the tribes

Gatesclarkeanini, Phricanthini, and Chlidanotini by RLB indicate that the an¬

cestral condition is the presence of ten costal strigulae between the wing base and

R 5 ; these are numbered here 1-10 from wing base to apex. Strigulae are not limited

110

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

to the forewing costa; among various Tortricidae, a strigula may be present be¬

tween each pair of adjacent veins on the termen, and here these are termed terminal

strigulae.

Each of the costal strigulae in the Olethreutinae has a narrow stria extending

towards the inner or outer margin, and these striae, which often are a shade of

gray, appear to be silver when viewed through the microscope at an angle to the

light. Each of the ten costal strigulae are paired in representatives of primitive

taxa, but some are single in derived taxa. In derived taxa, some strigulae (e.g.,

numbers five and six in the species described here) are approximate or confluent

(Fig. 1). Confluence of two independent strigulae to form what appears to be a

single or paired strigula usually can be determined by examinations of related

species or the detection of two silver striae arising from one apparent strigula.

Suffert (1929), as translated by Nijhout (1978), recognized four classes of ocelli,

or eyespots, on Lepidoptera wings. Of these, band ocelli are present in the Ole¬

threutinae, especially in the Grapholitini and derived Eucosmini. These ocelli are

formed from the fragmentation of fasciae, usually the postmedian fascia, and the

resulting fascial spots are accentuated by the gray striae, often widened, that

originate from costal strigulae (Fig. 1).

Epiblema deverrae R. Brown, NEW SPECIES

Types. — Holotype, male, data: “CALIF: Monterey Co., Landels-Hill Big Creek

Res., 5 mi. N. Lucia, 4-6 June 1982; J. Powell, collector”; deposited at the

University of California, Berkeley. The holotype is in excellent condition except

for missing meso- and metathoracic legs on the right side. Paratypes, deposited

as noted in parentheses: USA. CALIFORNIA. MONTEREY Co.: same data as

holotype, 2 females, 4 males (genitalia slide JAP 5121) (UCB), 1 female (genitalia

slide R. L. Brown 1321) (MEM); Big Creek Reserve, UCNLWR, Brunette Creek,

60-180 m el, redwood-hardwood, 26-28 May 1987, J. Powell, 1 female, 3 males

(UCB); Big Creek Reserve, UCNLWR, headquarters, coastal to confluence area,

0-60 m, 26-28 May 1987, J. Powell, 1 female (UCB); Big Creek Reserve,

UCNLWRS, Brunette Creek, 60-180 m el, redwood-hardwood, 5-8 Jun 1989,

Y-F. Hsu and J. Powell, 1 female, 1 male (UCB), 1 male (MEM); Big Creek

Reserve, UCNRS, Devils Creek Flat, 120 m el, 7 Jun 1989, J. Powell, 1 male

(UCB). SONOMA Co.: Cazadero, 6 Jun 1979, J. Powell, 1 female (UCB).

Description. —Head: vertex and upper frons brown, brown-orange, or gray-brown, darker brown

between antennae in some specimens; labial palpus with basal one-half orange-white or orange-gray,

apical one-half brown or light brown. Mesonotum: brown to dark brown; tegulae brown to dark brown

basally, brown or brown intermixed with gray-orange apically. Forewing (Figs. 1, 2): Forewing length

5.3-7.1 mm in males, 5.5-7.0 in females; male costal fold extending from base to proximal margin

of median fascia, length of fold 0.33-0.40 forewing length. Upperside ground color brown to gray-

brown intermixed with orange, with orange increasing in apical one-half; pattern elements including

indistinct, broken, dark brown fasciae, white to orange-white interfascial spot on inner margin, white

to orange-white costal strigulae, and white terminal strigulae. Basal and subbasal fasciae not forming

distinct basal patch; basal fascia indistinct, basal area with small spots and broken, narrow umbrae;

subbasal fascia reduced to broken umbrae, umbrae broad and confluent near midwing and inner

margin, distal margin of fascia angled at midwing, extending to costa at about 0.30 x the forewing

length; median fascia broken by orange or brown near midwing into large spot on costa and large spot

on inner margin, some specimens with inner marginal portion of fascia broken by orange or brown

into small proximal spot and larger distal spot; postmedian fascia represented by large postmedian

spot (appearing as two or three spots in some rubbed specimens) near apex of discal cell and three

1991

BROWN & POWELL: NEW SPECIES OF EPIBLEMA

111

Figure 2. Epiblema deverrae, female, Sonoma Co., California.

small bars, three bars bordered by subequal, gray (silver when viewed at an angle to the light) spot

on inner margin near tomus and by broad, transverse, gray (silver) lines proximally and distally to

form ocellus; subterminal fascia indistinct; terminal fascia represented by small spot at wing apex in

some specimens. Interfascial spot present between subbasal fascia and median fascia, extending from

inner margin to Cu 2 , subquadrate with proximal margin longer than distal margin. Costa with four

to six conspicuous strigulae, strigulae one and two confluent with each other or indistinct; strigula

three single, marking distal margin of subbasal fascia; strigula four single, marking proximal margin

of median fascia, separated from strigula three by small dark brown spot, strigulae three and four

indistinct in some specimens, each strigula with a gray (silver) stria extending posteriorly, striae

becoming confluent at discal cell and extending to interfascial spot; strigulae five and six approximate

at distal margin of median fascia, appearing as a single paired strigula, strigula five with gray (silver)

stria extending to midwing, strigula six with gray (silver) stria extending to near tomus, becoming

broad at midwing; strigulae seven and eight apparently confluent, appearing as single strigula in some

specimens, with single gray (silver) stria extending to proximal margin of postmedian spot; strigula

nine single or paired, with gray (silver) stria extending to distal margin of postmedian spot; strigula

10 paired, with strigulae separated by dark brown spot, with gray (silver) stria extending to M, at

termen; strigulae 5-6, 7-8, 9, and 10 bordered by orange or dark brown, separated from each other

by dark brown spots. Termen with small strigulae between R 5 and M,, M, and M 2 , and CuA, and

CuA 2 , some specimens with orange-white, attenuate, fringe scales arising from strigulae. Proximal

fringe scales attenuate, short, brown with white to orange-white apices; distal fringe scales attenuate,

long, gray-brown. Forewing underside brown except white to orange-white costal strigulae and area

of hindwing overlap posterior to CuP. Hindwing: Upperside uniformly brown except area of forewing

overlap anterior to Sc+Rf, underside uniformly brown, concolorous with forewing underside. Male

genitalia (Fig. 3): Uncus reduced to rounded lobe, densely setose dorsally; socii slightly flattened,

moderately setose posteriorly; gnathos arising from triangular projections of tegumen; aedeagus with

patch of 13 or 14 comuti on vesica; anellus with pointed anterior ventral comers; valva with sacculus

moderately setose, clasper at base near dorsal margin with flat medial surface and beset with group

of short, moderately stout setae, a second clasper-like, rounded projection near sacculus with sparse,

slender, short to long setae, ventral margin of neck without setae, cucullus large relative to valva,

length of cucullus 0.60 x length of valva (one preparation examined). Female genitalia (Fig. 4): Sternum

VII with moderately dense scales throughout, posterior one-half with sparse, short setae, becoming

dense on posterolateral comers; tergum VIII with moderately dense, long setae on posterior fourth

and on triangular, lateral extensions, scales absent; papillae anales densely setose, without rugae or

papillose projections; lamella postvaginalis with dense microtrichia medially, with sparse, long setae

on lateral rims; lamella antevaginalis reduced to lightly sclerotized, smooth rim; ductus bursae weakly

spiraled, with moderately sclerotized colliculum posterior to inception of ductus seminalis; signa flat,

blade-like, bases with shallow invaginations (one preparation examined).

Diagnosis.—Epiblema deverrae is superficially similar to E. hirsutana (Wal-

singham), which was described from Sonoma Co., California. In E. hirsutana,

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Figure 3. Epiblema deverrae, male genitalia with aedeagus detached. Scale line = 0.5 mm.

the head and labial palpi appear bushy due to long, erect, narrow scales, and the

forewing lacks a large postmedian spot and a contrasting median fascia. The

interfascial white spot is less well defined in E. hirsutana, and is not suffused with

pale orange. The male genitalia of E. hirsutana has a valve with a broader cucullus

and a single clasper. The female genitalia of E. hirsutana has a longer ductus

bursae with a more lightly sclerotized colliculum. Epiblema radicana (Walsing-

ham), also occurring in western North America, is a larger species with a forewing

lacking an ocellus and having a paler interfascial area between the median fascia

and postmedian fascia and with genitalia differing in several characters (Obraztsov

1965: Figs. 1-10). Allopatric species that are superficially similar to E. deverrae

include E. walsinghami (Kearfott) and E. infelix (Heinrich), occurring in the

eastern United States, E. arctica Miller, occurring in Alaska, and E. graphana

Treitschke, E.farfarae Fletcher, E. simploniana, and some forms of E. scutulana

(Denis & Schiffermuller), occurring in the Palaearctic region. None of these latter

species has a forewing with a large postmedial spot combined with orange scales

in the apical area, and all differ in characters of the male genitalia, as figured by

Miller (1985, 1987) and Kuznetsov (1978).

Comments. — The new species was discovered in Big Creek Reserve at two sites,

about 2 km apart, in 1982: on the upper Brunette Creek trail at about 180 m

elevation in the Big Creek drainage, and along the road above Devil’s Creek Flat

at 120-130 m. Additional individuals were collected in later years, subsequent

to an extensive fire that burned both sites in July, 1985. Adults seem to be

univoltine and diurnal; all were flying during afternoons and none have appeared

at blacklights deployed in various areas of the reserve. They seem to be associated

with Madia madiodes (Nuttall) (Asteraceae) growing in loose rocky talus in open

areas in forests dominated by redwood ( Sequoia sempervirens (D. Don.)). No gall

formations were found on this herbaceous perennial, and we speculate that larvae

of E. deverrae feed on underground rootstocks.

1991

BROWN & POWELL: NEW SPECIES OF EPIBLEMA

113

Figure 4. Epiblema deverrae, female genitalia. Scale line = 0.5 mm.

Madia madioides ranges from Monterey County, northward, along the Pacific

coast, in mixed evergreen and coastal coniferous forests, to Vancouver Island

(Munz 1959).

Etymology. —The specific name deverrae is formed from Deverra, the Roman

goddess of brooms and sweeping.

Material Examined.—See types.

114

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(2)

Acknowledgment

We thank K. R. Tuck (British Museum [Natural History]) for comparing E.

deverrae with the type of E. hirsutana, J. A. MacGown for drawing the figures,

D. Reed for computer scanning, J. Tisdale for word processing, and S. Cho for

photographing the imago. Cooperation by John Smiley (Big Creek Reserve) has

facilitated the survey at Big Creek in many ways. The Mississippi Agricultural

and Forestry Experiment Station has approved this paper as contribution no.

7607.

Literature Cited

Bickford, C. & P. Rich. 1984. Vegetation and flora of the Landels-Hill Big Creek Reserve, Monterey

County, California (2nd ed). Envir. Field Program, University of California, Santa Cruz.

Blanchard, A. 1979. New status for Epiblema minutana (Kearfott) and new species of Epiblema

Hubner and Sonia Heinrich (Tortricidae). J. Lepid. Soc., 33: 179-188.

Blanchard, A. 1984 (1985). Epiblema luctuosana A. Blanchard, a homonym, is changed to Epiblema

luctuosissima, new name. J. Lepid. Soc., 38: 245.

Bradley, J. D., W. G. Tremewan & A. Smith. 1973. British tortricoid moths. Cochylidae and

Tortricidae: Tortricinae. Ray Society, London.

Bradley, J. D., W. G. Tremewan & A. Smith. 1979. British tortricoid moths. Tortricidae: Olethreu-

tinae. Ray Society, London.

Danilevski, A. S. & V. I. Kuznetsov. 1968. Fauna of the U.S.S.R., Lepidoptera, Yol. 5(1), Tortricidae,

Laspeyresiini. Akademiia Nauk SSSR, Leningrad.

Horak, M. & R. L. Brown, (in press). Taxonomy and phylogeny. In van der Geest, L. P. S. (ed.).

World crop pests, their biology, natural enemies and control. Elsevier Science, Amsterdam.

Komerup, A. & J. H. Wanscher. 1983. Methuen handbook of colour (3rd ed). Methuen and Co.,

London.

Kuznetsov, V. I. 1978. Family Tortricidae. In Medvedev, G. S. (ed.). Keys to the insects of the

European part of the USSR. Vol. 4, Lepidoptera, Part 1. Nauka Publ., Leningrad [1987. Trans¬

lation for U.S. Dept. Agric. and NSF. Amerind Publ. Co., New Delhi, pp. 279-956].

McFarland, N. 1965. The moths (macroheterocera) of a chaparral plant association in the Santa

Monica Mountains of southern California. J. Research Lepid., 4: 43-74.

Miller, W. E. 1985. Nearctic Epiblema : a new synonymy, a revised identity, and two new species

(Lepidoptera: Tortricidae). Great Lakes Entomol., 18: 33-38.

Miller, W. R. 1987. Guide to the olethreutine moths of Midland North America (Tortricidae). U.S.

Dept. Agric., Forest Service Agric. Handbook, 660.

Munz, P. A. 1959. A California flora. University of California Press, Berkeley.

Nijhout, H. F. 1978. Wing pattern formation in Lepidoptera: a model. J. Exp. Zool., 206: 119-136.

Nijhout, H. F. 1990. A comprehensive model for colour pattern formation in butterflies. Proc. R.

Soc. London B, 239: 81-113.

Obraztsov, N. S. 1965. On Paedisca radicana and the type species of the genus Griselda (Lepidoptera,

Tortricidae, Olethreutinae). Amer. Mus. Novit., 2213: 1-16.

Opler, P. A. & J. A. Buckett. 1970 (1971). Seasonal distribution of “macrolepidoptera” in Santa

Clara County, California. J. Research Lepid., 9: 75-88.

Powell, J. A. 1983. Tortricidae. pp. 31-41. In Hodges, R. W., et al. (eds.). Check list of the Lepidoptera

of America north of Mexico. E. W. Classey Ltd., London, and Wedge Entomological Research

Foundation, Washington, D.C.

Suffert, F. 1929. Morphologische Erscheinungsgruppen in der Fliigelzeichnung der Schmetterlinge,

insbesondere die Querbindenzeichnung. Wilh. Roux’ Arch., 120: 229-383.

Received 19 November 1990; accepted 7 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(2): 115-118, (1991)

A REVIEW OF THE GENUS MICROTHURGE

(HYMENOPTERA: MEGACHILIDAE)

Terry L. Griswold

USDA-ARS Bee Biology and Systematics Laboratory, Utah State University,

Logan, Utah 84322-5310

Abstract.— A new South American Microthurge, M. furcatus NEW SPECIES, is described. Mi-

crothurge boharti NEW SYNONYM is synonymized with M. corumbae. The male of M. phar-

cidontus is described. New records and keys to the species for males and females are presented.

Key Words.— Insecta, taxonomy, bees, floral records

In a recent revision of the megachilid subfamily Lithurginae (Michener 1983),

a new genus Microthurge Michener was described to accommodate a small group

of heriadiform bees from southern South America. Four species were included:

Microthurge pygmaeus (Friese), M. pharcidontus (Moure), M. boharti Michener

and M. corumbae (Cockerell), the last based only on the description, because

Michener was unable to locate the type. Here, I report on the identity of M.

corumbae, the discovery of a new species, describe the male of M. pharcidontus

and give new records for the genus. Depositories for specimens (cited by cities)

are: University of California (Davis), University of Kansas (Lawrence), American

Museum of Natural History (New York), Philadelphia Academy of Sciences (Phil¬

adelphia), Carnegie Museum of Natural History (Pittsburgh), USDA Bee Biology

and Systematics Laboratory (Logan).

Keys to Species

Males

1. Scutal surface roughened medially by transverse rugulae; scutellum rounded

in profile; mesopleural surface granular. pharcidontus (Moure)

Scutal surface not roughened by transverse rugulae; scutellum flat in

profile; mesopleural surface shiny . 2

2(1). Hind basitarsus with ventral hairs black in part; punctures on medial

area of mesopleuron nearly contiguous; basal depression of mandible

wide . pygmaeus (Friese)

Hind basitarsus with ventral hairs light; punctures on medial area of

mesopleuron well separated; basal depression of mandible linear ... 3

3(2). Punctures on upper part of clypeus foveolate, larger than those on frons;

frons with punctures contiguous; eyes slightly diverging above (ratio

of upper interorbital distance at level of lateral ocelli/minimal lower

interorbital distance < 1.35). furcatus NEW SPECIES

Punctures on upper part of clypeus fine, not foveolate, not larger than

those on frons; frons with punctures not contiguous; eyes strongly

diverging above (ratio of upper interorbital distance at level of lateral

ocelli/minimal lower interorbital distance >1.4) .

.. corumbae (Cockerell)

116

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Females

1. Upper margin of mandible with basal, dorsoventrally flattened projec¬

tion; abdominal scopa largely black . pygmaeus (Friese)

Upper margin of mandible without projection; abdominal scopa white

except sometimes on S5. 2

2(1). Scutum with broad median zone covered with coarse transverse rugulae;

mesopleural surface granular; scutellum rounded in profile; apical hair

of T6 dark brown . pharcidontus (Moure)

Scutum without transverse rugulae; mesopleural surface shiny; scutel¬

lum flat in profile; apical hair of T6 black. 3

3(2). Facial projection carinate, with deep V-shaped emargination in dorsal

view; labrum with conical basomedial projection; clypeus in part with

large shiny interspaces between punctures .... furcatus NEW SPECIES

Facial projection not carinate, with shallow rounded emargination in

dorsal view; labrum with low, notched basomedial projection; clypeus

contiguously punctate or essentially so. corumbae (Cockerell)

Microthurge pygmaeus (Friese)

Material Examined. — (New Records) ARGENTINA. SALTA: Rosario de Lerma, 10-14 Nov 1983,

malaise trap, M. Wasbauer, 5 males, 17 females (Logan); same except 4-8 Nov 1983, 6 females;

Rosario de Lerma, Oct 1984, M. Fritz, 1 male (Logan); same except Oct 1985, 1 male, 4 females

(Logan); same except Oct 1986, 2 males (Logan). CORDOBA: LaCumbre, 1140 m, 21 Nov 1975, R.

M. Bohart, 12 males, 7 females (Davis, Logan). JUJUY: Perico, S of Jujuy, 21 Oct 1968, L. E. Pena,

1 female (New York). CATAMARCA: Alijilan, 3 Nov 1972, Argemone sp., G. E. Bohart, 1 female

(Logan). TUCUMAN: Tucuman, 19 Oct 1972, Argemone subfusifor mis G. B. Ownbey, G. E. Bohart,

1 male, 2 females (Logan); Cadillal, 4 Dec 1975, R. M. Bohart, 1 male (Logan).

Microthurge pharcidontus (Moure)

Redescription. — Male. Length, 5-7 mm; forewing length, 4-5 mm. Basal depression of mandible

linear; pair of tubercles on supraclypeal area small and close or absent; punctures on upper part of

clypeus foveolate, larger than punctures on frons; scutum roughened medially by transverse rugulae;

mesopleural surface granular.

Diagnosis. — The transversely rugulose scutum, granular mesopleuron, and

rounded rather than flat scutellum are unique to M. pharcidontus. Males share

these unique characteristics, though the rugulae of the scutum are less well-de¬

veloped.

Discussion. — Additional material available for this study supports the tentative

association by Michener (1983) of males from Guayaramerin, El Beni, Bolivia

with M. pharcidontus.

Material Examined. —(New Records) ARGENTINA. SALTA: Dique Itiyuro, nr Pocitos, 14 Aug

1976, C. Porter, L. Stange (Lawrence). TUCUMAN: Tucuman, 19 Oct 1972, Argemone subfusiformis

G. B. Ownbey, G. E. Bohart, 1 male (Logan); San Jose, Rio Marija, 3 Nov 1972, Opuntia sulphurea

G. Don, G. E. Bohart, 1 male (Logan); Horco Molle, 4 Jan 1976, R. M. Bohart, 1 male, 2 females

(Davis, Logan).

Microthurge corumbae (Cockerell)

Lithurgus corumbae Cockerell, 1901. Proc. Acad. Nat. Sci. Phil., 1901: 216. Type

deposited in Pittsburgh.

1991

GRISWOLD: REVIEW OF MICROTHURGE

117

7 Vc.r

Figure 1. Head of Microthurge furcatus, female. Punctation shown only for clypeus.

Microthurge boharti Michener, 1983. Pan-Pacif. Entomol., 59: 186. NEW SYN¬

ONYMY. Type deposited in Buenos Aires.

Diagnosis. — Males have eyes more strongly diverging above than in M. furcatus

(ratio of upper interocular distance at level of lateral ocellus/minimal lower in¬

terocular distance 1.41-1.52; n = 8, X = 1.46).

Discussion. — The holotype female of Lithurgus corumbae Cockerell from Co¬

rumba, Mato Grosso, Brazil, was recently located in the entomological collection

of the Carnegie Museum of Natural History. It appears to be the same as Micro¬

thurge boharti from Cordoba Province, Argentina, despite the geographic sepa¬

ration and apparent ecological differences between the moist tropical region of

Corumba and the more xeric environment of Cordoba. The only discemable

difference is the slightly better developed frontal prominence in specimens from

Argentina. This may be size related, because there appears to be size-dependent

variation among the Corumba specimens.

Material Examined. —(New Records) BRAZIL. MATO GROSSO: Corumba, Apr, 4 males, 28

females (Philadelphia, Logan); same except Mar, 2 females (Philadelphia).

118

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Microthurge furcatus Griswold, NEW SPECIES

(Fig. 1)

Types. — Holotype, female, data: BOLIVIA. COCHABAMBA: Pena Colorada,

1800 m, 21 Feb 1976, L. E. Pena. Paratypes; 7 males, data same as holotype; 3

males, 1 female, data: BOLIVIA. COCHABAMBA: (no local data), Nov 1976, L.

E. Pena. Holotype deposited in American Museum of Natural History, New York;

paratypes deposited in the American Museum of Natural History and USDA Bee

Biology and Systematics Laboratory, Logan, Utah.

Female. —Length, 7.5-8 mm; forewing length 6 mm. Body black, apical tarsi red; wings heavily

infuscated. Pubescence white; scopa off-white except for few dark hairs laterally and apically on S5.

Mandible without projection basally on upper margin; labrum with basomedial cone-shaped projec¬

tion; clypeus with punctures on disk scattered; facial projection carinate, in dorsal view with wide

V-shaped notch; scutum without transverse rugulae; scutellum flat in profile; mesopleuron shiny, with

punctures separated.

Male. —Length, 6-7 mm; forewing length, 4.5-5 mm. Basal depression of mandible linear; punctures

on upper part of clypeus foveolate, larger than those on frons; punctures of frons contiguous; eyes

slightly diverging above (ratio of upper interocular distance at level of lateral ocelli/minimal lower

interocular distance 1.22-1.33; n — 10, x = 1.28); ventral hairs on hind basitarsus all light.

Diagnosis. —Microthurge furcatus seems most closely related to M. corumbae.

Males of the two species are very difficult to separate, differing as far as I can tell

only by slight differences in punctation and by the less divergent inner eye margins.

Females are more easily separated. In addition to the characters given in the key,

the mandible is covered with a few large punctures, has a very strong dorsal carina,

and is abruptly and strongly depressed basally. The mandible of M. corumbae

has numerous small punctures, a weak dorsal carina, and is only slightly depressed

basally.

Material Examined. — See types.

Acknowledgment

I thank John Rawlins and Robert Davidson (Carnegie Museum of Natural

History, Pittsburgh) for assistance in locating and studying the type of Lithurgus

corumbae and the following individuals and institutions for the loan of material:

Manfredo Fritz (Instituto de Investigaciones Entomologicas Salta, Rosario de

Lerma, Argentina), Jerome Rozen and Eric Quinter (American Museum of Natural

History, New York); Robert Schuster (University of California, Davis); Donald

Azuma (Academy of Natural Sciences, Philadelphia); Charles Michener (Uni¬

versity of Kansas, Lawrence). Marianne Cha Filbert produced the illustration.

This is a contribution from Utah Agricultural Experiment Station, Utah State

University, Logan, Utah 84322-4810, Journal Paper Number 4074, and USDA-

ARS Bee Biology and Systematics Laboratory, Utah State University, Logan,

Utah 84322-5310.

Literature Cited

Michener, C. D. 1983. The classification of the Lithurginae. Pan-Pacif. Entomol., 59: 176-187.

Received 30 October 1990; accepted 14 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(2): 119-123, (1991)

A NEW AND PRIMITIVE GENUS OF CRYPHOCRICINAE

(HETEROPTERA: NAUCORIDAE)

John T. Polhemus

University of Colorado Museum, Englewood, Colorado 80110

Abstract. —Procryphocricos perplexus NEW GENUS, NEW SPECIES is described and compared

to Cryphocricos Breddin. The predominantly plesiomorphic character states of this new genus

are discussed.

Key Words.— Insecta, Heteroptera, Naucoridae, Cryphocricinae

During an expedition to South America Dan A. Polhemus and I collected

numerous naucorids among which is a remarkable new cryphocricine genus from

Colombia that is superficially similar to Cryphocricos Breddin but possessing a

number of plesiomorphic and some apomorphic characters that distinguish it.

The subfamily Cryphocricinae was proposed by Montandon (1897) to hold

Cryphocricos Breddin and Cataractocoris Montandon. It was redefined by Usinger

(1941) to exclude Cataractocoris, which was placed in his new subfamily Am-

brysinae. Usinger proposed a family level classification of Naucoridae, based on

a number of characters, that was a radical departure from that of Montandon

(1897). Since then alternative arrangements the family-group classification of

Naucoridae have been proposed by a number of authors (e.g., Popov 1970, De

Carlo 1971, Lopez Ruf & Bachmann 1987, Stys & Jansson 1988; the latter over¬

looked the work by Lopez Ruf & Bachmann). In these proposals the rank and

relationships of many family group taxa have been shifted, and a new taxon,

Pelocoridae, was proposed by De Carlo (1971), supported by Lopez Ruf & Bach¬

mann (1987), but rejected by others (e.g., Polhemus 1979, Stys & Jansson 1988).

The various schemes have been discussed by the latter, who placed Cryphocricos

in the monotypic tribe Cryphicricini. Both D. A. Polhemus and I consider all of

these proposals to be poorly founded, based on too few characters, and none of

them have been supported by a convincing phylogenetic analysis; this matter will

be treated in detail later, and supported by a cladistic analysis. Although the

suprageneric classification of the Naucoridae remains in my view an open ques¬

tion, I have used the classification of Stys & Jansson (1988) in order to establish

a framework for this paper.

The addition of Procryphocricos NEW GENUS does not require redefinition

of the family group taxon that holds Cryphocricos, the Cryphocricini, but instead

supports Usinger’s decision to separate it. In Table 1 these two genera are com¬

pared and the polarity of characters shown. Their prostema, abdominal sense

organs, brachyptery and apparent plastron are so similar, but so different from

any other closely related naucorid genera (e.g., Cataractocoris Montandon) that

I consider it unnecessary to include other genera in the comparison. The mor¬

phology used here follows Parsons (1966) and Parsons & Hewson (1974).

The CL number following locality data refers to codes used by the author to

reference ecological data.

120

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(2)

Table 1. Comparison of Cryphocricos and Procryphocricos. Apomorphic character states are marked

with an asterisk.

Character Cryphocricos Procryphocricos

Setae on ventral surface

Mid and hind femora armed

(knobs, spines)

Distal tibial spines

Lateral pronotal margins

Male abdominal tergite 7

Aedeagus

Antennal segments, length

Prostemum, proepimeron

Male paratergite 8

Male abdominal stemite 7

Loral plates

Base of rostrum

Ocular setae

Abdominal tergites

very short; plastron only*

present*

2 close packed rows*

crenulate, without obvious

setae*

asymmetrical; spur on right

side*

strongly asymmetrical flagel¬

lum very long*

3 very short; 4 long*

separated by distinct suture

large, size equal to 7, ex¬

posed; projecting

fused with paratergites*

produced anterad of eyes*

ahead of eyes*

absent

distinct

scattered long silky setae plus

plastron

absent

1 short distal row, ragged sub-

distal row

smooth, set with short stiff se¬

tae

symmetrical; no spur

weakly asymmetrical; short fla¬

gellum

3 and 4 long, subequal

fused, junction faintly demar¬

cated*

small, mostly covered by 7;

angled medially*

free from paratergites

not produced anterad of eyes

between and below eyes

2 kinds present*

fused*

Procryphocricos J. T. Polhemus, NEW GENUS

Figs. 1-5

Type-species. —Procryphocricos perplexus J. T. Polhemus, NEW SPECIES.

Description.— Small, ovate, brachypterous. Length 5.1 mm, width 2.7 mm, widest across abdomen

(Fig. 1). Entire dorsum rugose, set with short appressed scale-like setae. Head narrow, produced

anteriorly 0.5 x eye length, anteclypeal margin almost straight; vertex slightly domed medially, pro¬

duced behind eyes 0.75 x eye length, with two (1 + 1) trichobothria adjacent to middle of eyes. Eyes

1.5 x as long as wide, with narrow posterior flange; raised, rounded, not forming smooth transition

to head, but set off by shallow sulcus on head; ocular setae of two kinds, many short mushroom¬

shaped setae scattered among the ommatidia, and longer slender setae, two of medium length on inner

eye margin, one long setae on lateral margin. Labrum small, arising at anterior margin of anteclypeus,

rounded; maxillary plates projecting anteriorly, tips pointed, extending to middle of rostral segment

three (second visible). Rostrum short, segment two (first visible) very short, recessed into head, segment

three much longer and slightly shorter than four. Antennae of moderate length, segments one and two

short, three and four subequal (Fig. 2). Pronotum convex, with broad shallow median depression,

margins depressed and demarcated posteriorly and posterolaterally (Fig. 1); lateral margins not cren¬

ulate, set with short stiff setae. Scutellum triangular, slightly raised above level of adjacent wing pads,

declivant anteriorly. Wing pads triangular, not touching medially, reaching posterior margin of ab¬

dominal tergite two; embolium poorly defined, set off mesad by a broad weak longitudinal carina not

reaching posterior margin. Abdomen moderately convex dorsally, transversely rugose, connexiva

evenly rounded, tergites three to five fused, intersegmental sutures weakly indicated only at lateral

margins; connexival margin depressed; posterior margin of tergite five broadly rounded posteriorly,

obscuring most of remaining tergites. Ventral surface sparsely set with long dark silky appressed setae

absent only on head and prostemum; anterior one-third of prostemum carinate medially, anteromedial

portions angled forward over base of carinate gula; prostemum completely exposed, very similar in

shape to Cryphocricos (see Usinger 1941: fig. 1), fused with proepistemum, demarcated by weak

1991

POLHEMUS: NEW GENUS OF NAUCORIDAE

121

Figures 1-5. Procryphocricos perplexus. 1. Habitus. 2. Antennae. 3. Male abdominal terminalia,

dorsal view.'4. Male abdominal terminalia, ventral view. 5. Aedeagus.

indication of suture. Mesostemal plate reflexed anteromedially forming tumescence, remainder me¬

dially carinate. Metastemal plate not carinate; metaxyphus triangular, posterior margins carinate.

Abdominal venter narrowly glabrous laterally; sense organs similar to and in same position as Cry-

phocricos (see Usinger 1941: Fig. 1). Fore legs with femur broad, tibia narrow, tarsi single-segmented;

single claw very short, triangular. Middle and hind femora slender, not modified, set with short stiff

appressed setae; tibia set with spines except basally, distally with ventral row of closely set spines,

subdistal ragged row of four spines, plus dorsal comb of many closely set stiff setae; tarsal segment

one short, two long, three longer. Mid and hind pretarsi with long setiform parempodia; claws long,

curved, each with large basal spur. Abdominal segments six to eight almost symmetrical (Figs. 3^1);

genital capsule similar to Cryphocricos, elongate; aedeagus slightly asymmetrical; parameres small

(Fig. 5).

122

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Diagnosis. —Procryphocricos differs from Potamocoris by its smaller antennae,

which are not visible in dorsal view, and by its brachyptery; only macropters of

the latter are known. It differs from Cryphocricos by its vestiture of silky hairs on

abdominal ventrites six through eight, and the characters given in Table 1.

Discussion.— As can be seen from Table 1, Cryphocricos possesses a number

of apomorphies in comparison to Procryphocricos. Because both occur in the

brachypterous form, both must possess a plastron, or there would be no way to

obtain and hold an air store [see Usinger (1941), Parsons & Hewson (1974) and

D. Polhemus (1986) for discussion]; this is an apomorphy for the Cryphocricini

within the Cryphocricinae but is not unique within the Naucoridae; see Polhemus

& Polhemus (1986) for discussion. In addition, all genera of the Cryphocricinae

have a dorsal vestiture of silky hairs on abdominal segments six, seven and eight,

but only in Cryphocricos these are absent ventrally.

To establish the polarity of characters in Table 1, the outgroup chosen for

comparison is the genus Potamocoris Hungerford, which in my view is close to

the ground plan for the Naucoridae [placed in a separate family by some authors;

see Stys & Jansson (1988) for discussion]. Procryphocricos shares some primitive

characters with Potamocoris, for instance similar antennal proportions, but in the

latter genus the antenna are much longer and visible in dorsal view. In Procry¬

phocricos one pair of facial trichobothria are present medially along the eyes, in

the same position as in Potamocoris, but the latter has three pairs; also the lateral

ocular setae are similar in the two genera, but the latter has no medial pair and

lacks the mushroom-shaped ocular setae which constitute an autapomorphy for

Procryphocricos.

Habitat.—The two known specimens of Procryphocricos were found among

tangles of fine roots along a small section of steep bank bathed by the moderate

current of the Rio Claro. A diligent search of other habitats of many kinds failed

to yield additional specimens. Midstream riffles of the Rio Claro and a nearby

tributary yielded Ambrysus, Cryphocricos and Limnocoris.

Material Examined .—See species types.

Procryphocricos perplexus J. T. Polhemus, NEW SPECIES

Figs. 1-5

Types. — Holotype, brachypterous male, data: COLOMBIA. ANTIOQIUA: Rio

Claro, 13 km W of Doradal, 250 m, water temp 25° C, CL 2405, 21 Jul 1989, J.

T. and D. A. Polhemus, in J. T. Polhemus Collection (JTPC). Paratype, 1 bra¬

chypterous male, same data as holotype (JTPC).

Description, Brachypterous Male.—(See generic description; only additional details given here.)

Ground color yellow-brown; pronotum anteromedially and laterally, wing pads, fore legs distally tinged

with brown; spines on mid and posterior legs orange-brown; eyes brown. Head deeply set into pro¬

notum; length 0.90 mm, width through eyes 1.10 mm. Eyes with many ommatidia; embraced pos¬

teriorly by pronotum; length 0.38 mm, width 0.25 mm, interocular space 0.60 mm. Pronotum broad,

maximum length 1.23 mm, length on midline 0.73 mm, width 2.13 mm. Scutellum triangular, roughly

2 x as wide as long, length 0.70 mm, width 1.48 mm. Wing pad length 1.03 mm. Abdominal tergites

three to seven each set laterally with two (1 + 1) long slender erect tufts of golden setae, on middle

of connexival segment three, on posterior one-quarter of four to seven; combined length of fused

abdominal tergites three to five, 2.13 mm. Antennal proportions as shown in Fig. 2; distal segment

set with a row of stiff evenly spaced setae. Proportions of legs in mm (femur: tibia: tarsus 1:2:3):

1991

POLHEMUS: NEW GENUS OF NAUCORIDAE

123

anterior, 1.43:1.10:0.30:0:0; middle, 1.18:1.10:0.08:0.18:0.30; posterior, 1.43:1.53:0.10:0.20:0.33. Ab¬

dominal terminalia and genitalia as in Figs. 3-5.

Female and Macropterous Forms. — Unknown.

Diagnosis.— Procryphocricosperplexus is the only species of the_genus.

Material Examined.—See types.

Acknowledgment

I am deeply grateful to D. A. Polhemus for assistance in the field, reviewing

the manuscript, and stimulating discussions on naucorid phylogeny. Gabriel Rol¬

dan Perez provided warm hospitality and invaluable assistance in many ways

during our stay in Colombia; without his help and that of Luisa Fernanda Alvarez

A. and Luis Fernando Roldan, we could not have succeeded in our endeavors

there. The field work was supported in part by the National Geographic Society,

to whom I am deeply grateful.

Literature Cited

De Carlo, J. A. 1971. Valor sistematico del estudio del aparato genital macho en los Hemipteros

acuaticos y semiacuaticos. Division de la Familia Naucoridae en dos familias. Caracteristica

externa de la capsula genital. Rev. Soc. Entomol. Arg., 38: 159-166.

Lopez Ruf, M. L. & A. O. Bachmann. 1987. Sobre la classification de los Naucoroidea Americanos

(Heteroptera). Rev. Soc. Entomol. Arg., 44: 341-344.

Montandon, A. L. 1897. Hemiptera Crytpocerata—Fam. Naucoridae. Sous-fam. Cryptocricinae.

Verh. Zool. Bot. Ges. Wien, 47: 6-23.

Parsons, M. C. 1966. Studies on the cephalic anatomy of Naucoridae (Heteroptera). Trans. R.

Entomol. Soc. Lond., 118: 119-151.

Parsons, M. C. & R. J. Hewson. 1974. Plastral respiratory devices in adult Cryphocricos (Naucoridae:

Heteroptera). Psyche, 81: 510-527.

Polhemus, D. A. 1986. A review of the genus Coptocatus Montandon (Hemiptera: Naucoridae).

Pan-Pacif. Entomol., 62: 248-256.

Polhemus, D. A. & J. T. Polhemus. 1986. Naucoridae (Hemiptera) of New Guinea. 2. A review of

the genus Idiocarus Montandon, with descriptions of three new species. J. New York Entomol.

Soc., 94: 39-50.

Polhemus, J. T. 1979. Family Naucoridae. pp. 131-138. In Menke, A. S. (ed.). The semiaquatic and

aquatic Hemiptera of California. Bull. Calif. Ins. Surv., 21.

Popov, Yu. A. 1970. Notes on the classification of the recent Naucoridae. Bull. Acad. Pol. Sci., Ser.

Sci. Biol., 18: 93-98.

Stys, P. & A. Jansson. 1988. Check-list of recent family-group and genus-group names of Nepomorpha

(Heteroptera) of the world. Acta Entomol. Fennica, 50: 1-44.

Usinger, R. L. 1941. Key to the subfamilies of Naucoridae with a generic synopsis of the new

subfamily Ambrysinae (Hemiptera). Ann. Entomol. Soc. Am., 34: 5-16.

Received 19 October 1990; accepted 16 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(2): 124-134, (1991)

THE MYDIDAE (DIPTERA) OF COSTA RICA

Judith L. Welch and Boris C. Kondratieff

Department of Entomology, Colorado State University,

Fort Collins, Colorado 80523

Abstract. — The species of Mydidae occurring in Costa Rica are reviewed. Four genera and seven

species are reported, including Nemomydas loreni, NEW SPECIES. Additionally, My das quad-

rilineatus Williston, NEW SYNONYM, is synonymized under M. rufiventris Macquart.

Key Words.— Insecta, Diptera, Mydidae, Costa Rica, insect fauna

Costa Rica occupies part of the isthmus of Central America and is a complex

ecological mosaic (Hall 1985). It has been estimated that upwards of 300,000

species of insects may occur in Costa Rica. This fauna is a complex mixture of

Mesoamerican and South American species. One group of flies that attract the

attention of even the most specialized collectors are mydas flies. Mydids are a

small, but worldwide, family composed of medium to very large, usually sparsely

pilose, flies that often resemble wasps (Fig. 1). Adults occupy a wide variety of

habitats and can be locally common, especially in arid areas. Many species are

restricted to hot, sandy habitats and can be found visiting blossoms or resting on

bare ground.

The preparation of a section (JLW, unpublished data) on the Mydidae of Costa

Rica for the “Guia para las familias de Insectos de Costa Rica,” stimulated this

review. There is no single publication available treating all genera and species.

Morphology and terminology follows Wilcox (1981); genitalic structures are

labeled in Figs. 4 and 12. Abbreviations for depositions that are used throughout

the paper are: British Museum of Natural History, London (BMNH); University

of California at Berkeley (UCB); Utah State University, Logan (USU); Museum

National d’Histoire Naturelle, Paris (MNHN); Illinois Natural History Survey,

Champaign (INHS); Museum of Comparative Zoology, Harvard University,

Cambridge, Massachusetts (MCZ); and Texas A&M University, Lubbock

(TXA&M); collection of J. L. Welch (JLW).

Key to the Genera of Mydidae of Costa Rica

(modified from Wilcox 1981)

1. Laterotergite completely bare; proboscis subequal in length to subcranial

cavity; labella broad; apex of female abdomen with setae; length vari¬

able . 2

Laterotergite pilose; proboscis 1.0-2.0 x length of subcranial cavity; apex

of female abdomen with circlet of spines; length 12-23 mm.

. Nemomydas Curran

2(1). Prementum of proboscis 0.5 x length of subcranial cavity; labella at¬

tached to prementum near its midpoint, and subequal with subcranial

cavity in length; anterior margin of subcranial cavity situated at about

0.4 x the distance from lower eye margin to base of antennae; length

15-29 mm. Messiasia d’Andretta

1991

WELCH & KONDRATIEFF: COSTA RICAN MYDIDAE

125

Prementum of proboscis and subcranial cavity subequal in length; labella

attached to mentum near its apical one-half, and extending at about

a 90° angle; anterior margin of subcranium level with lower margin

of the eye. 3

3(2). Carina on hind tibia partially developed (Fig. 2); apical spur of hind

tibia reduced or absent (Fig. 2); epandrium strongly arched in lateral

view (Fig. 4); process of gonocoxite digitiform (Figs. 4-5); gonostylus

saddle-shaped in caudal view (Fig. 5); length 29-45 mm.

. Protomydas Wilcox, Papavero & Pimentel

Carina on hind tibia completely developed (Fig. 3); apical spur of hind

tibia long and stout (Fig. 3); epandrium rounded in lateral view (Fig.

6); gonocoxite simple or falciform (Figs. 6-7); gonostylus bilobed in

caudal view (Fig. 7); length 25-26 mm. My das Fabr.

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Figures 2-5. 2. Protomydas rubidapex, hind leg. 3. Mydas clavatus, hind leg. 4. Protomydas rub-

idapex, male terminalia, lateral view. 5. Protomydas rubidapex, male terminalia, caudal view.

Messiasia d’Andretta

Messiasia d’Andretta 1951: 52.

Type Species. —Messiasia carrerai d’Andretta (by original designation).

Diagnosis. — See key to genera. Species included in this genus closely resemble,

in general appearance those of Mydas, but they can be readily distinguished from

Costa Rican Mydas species by their entirely black integument.

Discussion. —Messiasia is predominantly South American in distribution but

five species are known to range from southwestern United States south to Panama

(Wilcox & Papavero 1975).

Key to Costa Rican Messiasia

1. Wings pale amber; long white setae present above hind coxa; gonocoxite

of male elongate, tapering to hook-like process; gonostylus serrate (Figs.

8-9) . decor (Osten Sacken)

- Wings dull black; sparse black setae present above hind coxa; gonocoxite

1991

WELCH & KONDRATIEFF: COSTA RICAN MYDIDAE

127

of male truncate apically, upper process pointed; gonostylus broadly

truncate (Figs. 10-11) . perpolita (Johnson)

Messiasia decor (Osten Sacken)

Midas decor Osten Sacken 1886: 71.

My das decor. ; Williston 1898: 55.

Messiasia decor, d’Andretta 1951: 68.

Messiasia decor, Wilcox & Papavero 1975: 21 (includes specimens from La Suiza,

Costa Rica).

Types. — Holotype, male; from: PANAMA. Bugaba; deposited in the British

Museum (Natural History), London; examined.

Diagnosis. — The white setae above the hind coxae will usually separate both

sexes of M. decor from M. perpolita.

Discussion.— This species is known from Costa Rica and Panama. Wilcox &

Papavero (1975) indicate that this species occurs in forested areas. The illustration

of the male terminalia of M. decor provided by Wilcox & Papavero (1975) is

inaccurate.

Material Examined. — COSTA RICA. MONTEVERDE Prov.: 20-24 Jan 1986, W. Hanson and G.

Bohart, 1 female (USU).

Messiasia perpolita (Johnson)

My das perpolita Johnson 1933: 72.

Messiasia polita; d’Andretta 1951: 64.

Messiasia perpolita-, Wilcox & Papavero 1975: 31.

Types. — Holotype, male; from: MEXICO. YUCATAN: Chichen Itza; deposited

in the Museum of Comparative Zoology, Harvard University, Cambridge, Mass.;

examined.

Diagnosis. — The dull black wings and no white setae above the hind coxae will

distinguish both sexes of this species from M. decor.

Discussion. — This species is relatively common in Mexico. Janzen (1986) de¬

scribed the unique tropical dry forest habitat in Costa Rica where this species was

collected. Janzen’s specimens collected by malaise traps represent the southern¬

most range extension for this species.

Material Examined. —COSTA RICA. GUANACASTE Prov.: Santa Rosa National Park, 14-17 Jun

1977, D. H. Janzen, 1 female (AMNH); same data except, 300 m, May 1983, D. H. Janzen and W.

Hallwachs, 2 females (AMNH); same data except 1-15 Jun 1982, 300 m, D. H. Janzen and W.

Hallwachs, 1 male (AMNH).

Nemomydas Curran

Nemo my das Curran 1934: 165.

Type Species. —Leptomydas pantherinus Gerstaecker (by original designation).

Diagnosis. — See key to genera. Additionally, males are easily distinguished from

all other mydids by the digitate processes of the gonocoxite.

Discussion. —Nemomydas is restricted to North America and Central America

and presently includes 19 species (Steyskal 1956, Kondratieff & Welch 1990,

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Figures 6-11. Male terminalia. 6. My das rufiventris, lateral view. 7. My das rufiventris, caudal view.

8. Messiasia decor, lateral view. 9. Messiasia decor, caudal view. 10. Messiasia perpolita, lateral view.

11. Messiasia perpolita, caudal view. Abbrev.: ae, aedeagus; ep, epandrium; goncx, gonocoxite.

1991

WELCH & KONDRATIEFF: COSTA RICAN MYDIDAE

129

Figures 12-19. Male terminalia. 12. Nemomydas bequaerti, lateral view. 13. Nemomydas bequaerti,

ventral view. 14. Nemomydas sponsor, lateral view. 15. Nemomydas sponsor, ventral view. 16. Ne¬

momydas loreni, lateral view. 17. Nemomydas loreni, ventral view. 18. Nemomydas lamia, lateral

view. 19. Nemomydas lamis, ventral view. Abbrev.: ae, aedeagus; al, anal lamellae; ce, cercus; ddp,

dorsal digitate process; ep, epandrium; goncx, gonocoxite; vdp, ventral digitate process.

130

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Welch & Kondratieff 1990). Three species occur in Costa Rica including a new

species described here: N. lamia (Seguy), N. sponsor (Osten Sacken), and N. loreni

NEW SPECIES. These three species are very similar in appearance; females are

difficult to separate without associated males.

Key to Nemomydas males south of Mexico

1. Abdominal tergites black, posterior margins yellow . 2

- Abdominal tergites yellow-brown with middorsal black-brown spots or

dashes; terminalia as Figs. 12-13 (Honduras). bequaerti (Johnson)

2(1). Distal section of aedeagus in lateral view, abruptly expanded and re¬

curved medially (Figs. 14-15) (Guatemala, Costa Rica) .

. sponsor (Osten Sacken)

Distal section of aedeagus in lateral view, tube-like (Figs. 16-17) or

tongue-like (Figs. 18-19) . 3

3(2). Distal section of aedeagus in lateral view, tube-like (Figs. 16-17) ....

. loreni NEW SPECIES

Distal section of aedeagus in lateral view, tongue-like (Figs. 18-19) ..

. lamia (Seguy)

Nemomydas lamia (Seguy)

Nomoneura lamia Seguy 1928: 146.

Nemomydas lamia; Papavero & Wilcox 1968: 34.11.

Nemomydas lamia; Kondratieff & Welch 1990: 474.

Types. -Lectotype, male; from: COSTA RICA. La Caja; deposited in the Mu¬

seum National d’Histoire, Paris; examined.

Diagnosis. — See key to Nemomydas species.

Discussion.— Kondratieff & Welch (1990) provided descriptions of both sexes

of this uncommon species. The specimens examined are apparently the first ad¬

ditional material collected since its original description.

Material Examined. —COSTARICA. GUANACASTEProv.:S. Canas, 26-31 Jan 1989, F. D. Parker,

1 male (CSU), 2 males (USU). La Caja, Paul Serre, 1920, 1 male, 6 females (MNHN).

Nemomydas sponsor (Osten Sacken)

Leptomydas sponsor Osten Sacken 1886: 68.

Nemomydas sponsor, Kondratieff & Welch 1990: 475.

Types. — Holotype, female; from: GUATEMALA. San Geronimo; deposited in

the British Museum (Natural History), London; examined.

Diagnosis. — See key to Nemomydas species.

Discussion. — Kondratieff & Welch (1990) provided the first description of the

male of this species.

Material Examined.— COSTA RICA. SAN JOSE Prov.: San Jose, 27-29 Dec 1987, F. D. Parker,

4 males, 2 females (USU); Escazu, 14-16 Jan 1989, F. D. Parker, 1 male (USU); 1 male (MNHN).

Nemomydas loreni Welch & Kondratieff, NEW SPECIES

Types. —Holotype male, data: COSTA RICA. GUANACASTE Prov.: S. Canas,

1-3 Feb 1989, F. D. Parker; deposited: Utah State University, Logan. Paratypes

deposited in Utah State University, Logan, except as indicated; data: 16 males,

1991

WELCH & KONDRATIEFF: COSTA RICAN MYDIDAE

131

1 female, same data as holotype (1 male deposited CSU), (1 male deposited JLW);

same data except 1-5 Mar 1989, 1 male; same data except 21-25 Jan 1989, 4

males; same data except 26-31 Jan 1989, 3 females; same data except 9-14 Feb

1989, 1 male, 9 females (1 female deposited CSU); same data except 22-24 Feb

1989, 1 male (deposited CSU); Tamarindo, 6 Nov 1977, M. E. Irwin, ex. beach

sand, 1 male (deposited INHS). SAN JOSE Prov.: Escazu, 14-16 Jan 1989, F. D.

Parker, 2 females (1 female deposited JLW); Puntarenas, 19 Dec 1989, F. D. Parker,

2 males; Comelco Property, 30 Jan 1976, H. Reed, 1 male (deposited TXA&M).

Description.—Male. Length 12-16 mm. Head shiny black, with white, erect setae; orbital of eye

yellow pollinose; antenna 3.6-3.8 mm long, black, flagellomere two white pollinose dorsally, brown

pollinose ventrally; proboscis long, 2.6 times as long as subcranial cavity, black. Scutum black, with

a pair of submedian yellow pollinose stripes that converge posteriorly, and a pair of lateral yellow

pollinose stripes, setae yellow, long, erect, sparse; postpronotal lobe yellow pollinose; scutellum yellow

pollinose, black posteriorly; metanotum black, yellow pollinose anteriorly. Wings brown, longitudinal

veins brown, costa black with black setae; knob of halter black, stem brown. Foreleg and midleg brown

black, setae white dorsally, black ventrally; hind femur white basally, black apically; hind tibia white

dorsally, brown-black ventrally; with short, white setae, dorsally, black ventrally; tarsomeres brown-

black. Abdominal tergites shiny black, posterior margins yellow to brown; bulla black; tergite one with

setae white, long anteriorly, short, sparse posteriorly; tergite two with setae black, short, decumbent,

medially; setae white, long, divergent, anteriorly and laterally; tergites three to five with setae black,

short, decumbent; tergites six and seven with setae white, long, decumbent; stemites shiny black,

stemites two to five with posterior margins yellow to brown; stemites two to four with setae white,

long, decumbent; stemites five to seven with setae black, long, decumbent. Terminalia brown-black.

Gonocoxite with dorsal digitate process distinctly claw-like, ventral digitate process oblong, rounded,

slightly directed inwardly; a smaller rounded protrusion ventrally. Aedeagus tube-like distally, swollen

basally (Figs. 16-17).

Female. —Length 12-16 mm. Color and structure similar to male, except setae on abdominal tergites

short and sometimes tergites six and seven without yellow posterior margins.

Diagnosis. — The male of N. loreni can be distinguished from all other meso-

american species by the combination of black abdominal tergites with yellow

posterior margins and tube-like distal section of the aedeagus. The female cannot

be satisfactorily separated from N. lamia.

Etymology. — This species is named in memory of JLW’s father, who encouraged

her to take an interest in Diptera because of their intriguing life histories.

Material Examined.—Type, series.

Protomydas Wilcox, Papavero & Pimentel

Protomydas Wilcox, Papavero & Pimentel 1989: 13.

Type Species. —Mydas coerulescens Olivier (by original designation).

Diagnosis.—See key to the genera.

Discussion.— Wilcox et al. (1989) recognized three species in this primarily

Neotropical genus; only one is known from Costa Rica and listed here. D’Andretta

(1951) provided illustrations of the male terminalia and other characters for these

species.

Protomydas rubidapex (Wiedemann)

Mydas rubidapex Wiedemann 1830: 626.

Mydas dives Westwood 1841: 50. (Synonymized by Wilcox, Papavero & Pimentel

1989: 16.)

Protomydas rubidapex ; Wilcox, Papavero & Pimental 1989: 16.

132

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(2)

Types. — The type of P. rubidapex is from MEXICO, and is deposited in Hum¬

boldt Universitat, Berlin; it was not examined. The type of P. dives, from an

unknown locality, is deposited in Hope Museum, Oxford, England, and was not

examined.

Diagnosis.—Protomydas rubidapex is easily separated from all My das species

by the reduced or absent carina of the hind tibia, the arched epandrium, digitiform

process of the gonocoxite, and the saddle-shaped gonostylus.

Redescription.—Male. Length 29-45 mm long. Head shiny black, with long, erect, black setae;

mystax with long, decumbent, black and yellow setae; orbital margin of eye black; antenna 10-12

mm, orange or black with flagellomere 2 orange; proboscis black. Scutum dull black; setae sparse,

decumbent, black; wing opaque orange to brown, wing margin broadly brown to black; veins orange,

or sometimes brown apically, costa with black setae basally, and mixed black and orange setae apically;

calypter with long dense fringe of black squamose setae; halter black; legs black, setae and bristles

black; hind tibia with carina absent or reduced, apical spur weakly developed, shorter than width of

first hind tarsal segment. Abdominal tergite one black, with long, decumbent, sparse, golden setae;

tergite two black, with short, recumbent, golden setae; bulla black; tergites three to seven with short,

recumbent, black, setae; stemites with short, sparse, recumbent setae. Terminalia with epandrium, in

lateral view (Fig. 3) arched, concave above spur; gonocoxite with produced, “grasping” digitiform

process that is incurved in caudal view; gonostylus saddle shaped; aedeagus tube-like, ribbed dorsally,

with lateral rounded ridges (Fig. 4).

Female.— Length 35-40 mm. Antenna length 14.0-15.5 mm. Coloration and structure similar to

male.

Discussion. —Wilcox et al. (1989) did not provide a thorough description of

this species, and the redescription above is based on Costa Rican specimens. This

species ranges from northern Mexico to Venezuela and Brazil.

Material Examined. —COSTA RICA. GUANACASTE Prov.: Santa Rosa National Park, 21-24 Dec

1979, D. H. Janzen, ex malaise trap, 1 male (AMNH); same data as above except, 1-15 Jan 1982,

300 m, D. H. Janzen and W. Hallwachs, 1 male (AMNH); Sotobosque, Cerro El Hacha, 12 Sep-2

Jan 1988, Tacotal, ex malaise trap, 1 female (AMNH).

Mydas Fabr.

Mydas Fabr. 1794: 252.

Type Species. —Musca clavata Drury (by subsequent designation).

Diagnosis.—See key to the genera.

Discussion. — This genus and its allies were recently revised by Wilcox et al.

(1989); we do not agree with several of the taxonomic conclusions reached in that

study and will publish our interpretations elsewhere. Mydas ruftventris is the only

Mydas known from Costa Rica.

Mydas ruftventris Macquart

Mydas ruftventris Macquart 1850: 364.

Mydas vittatus Macquart 1850: 364.

Mydas militaris Gerstaecker 1868: 99. (New name for Mydas vittatus, preoccu¬

pied.)

Stratiomydas ruftventris ; Wilcox et al. 1989: 127.

Mydas quadrilineatus Williston 1898: 56. NEW SYNONYM.

Stratiomydas quadrilineatus; Wilcox et al. 1989: 125.

Types. — The holotype male of M. ruftventris is labeled “BRAZIL” (in error); it

is deposited in the Museum National d’Histoire Naturelle, Paris, and was ex-

1991

WELCH & KONDRATIEFF: COSTA RICAN MYDIDAE

133

amined. The holotvpe female of M. vittatus is labeled MEXICO; it is deposited

in the Museum National d’Histoire, and was examined. The syntypes of M.

quadrilineatus are labeled “MEXICO”; they are deposited in the British Museum

(Natural History), London, and were examined.

Diagnosis.— The white to yellow mystax and the four pollinose stripes of the

mesonotum will easily distinguish this species from all other possible Mydas

species.

Discussion. — In a recent review of the Mydini by Wilcox et al. (1989), M.

rufiventris was distinguished from M. quadrilineatus by the color of the abdominal

tergites which are primarily black in M. quadrilineatus and red in M. rufiventris.

Examination of specimens throughout the range of both species (Arizona, Mexico,

El Salvador, and Costa Rica) indicated the presence of only one variable species.

The male terminalia of both forms are alike. The key of Wilcox et al. (1989) to

this group is unusable, however, especially with females. The series of both sexes,

collected by Paul A. Opler near Canas, contains all color morphs, ranging from

specimens with all abdominal tergites completely black to some with tergites two

through five or two through seven red. This often extreme polymorphism in

abdominal color is common in other species groups of Mydas (e.g., Mydas ventralis

Gerstaecker of the southwestern United States).

Material Examined.—CO STA RICA. GUANACASTE Prov.: Canas, 6.4 km NW of La Pacifica, 14

May 1971, P. A. Opler, ex Casearia sp. (“ nitida ”), 1 male, 1 female (UCB); same data except 15 Nov

1971, 1 female; same data except 23-30 May 1972, ex Forsteronia sp. (“ spicata ”), 2 males, 2 females

(UCB), 1 male (CSU); same data except 6 Jun 1973, ex Asclepias sp. (“liana”), 2 males (UCB); same

data except 2-4 June 1973, Asclepias vine, 1 male, 2 females (UCB).

Acknowledgment

We thank the following people who made material available for study: J. E.

Chainey (British Museum [Natural History]); John T. Doyen (University of Cal¬

ifornia, Berkeley); Wilford J. Hanson (Utah State University); L. Matile (Museum

National d’Histoire Naturelle, Paris, France); Kathryn C. McGiffen (Illinois Nat¬

ural History Survey); Stephen Pratt (Museum of Comparative Zoology, Harvard

University); and Edward G. Riley (Texas A&M University). David Carlson, Con-

ery Calhoon, and Michael G. Kippenhan (Colorado State University) prepared

the illustrations. Howard E. Evans reviewed the manuscript before submission.

Two reviewers made many helpful suggestions that improved an earlier version

of this manuscript.

Literature Cited

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D’Andretta, M. A. V. 1951. Contribuicao para o conhecimento da familia Mydaidae. Generos:

Mydas F., 1794 e Messiasia n. gen. (Diptera). Papeis Avulsos Dep. Zool. (Sao Paulo), 10: 1-76.

Fabricius, J. C. 1794. Entomologia systematica emendata et aucta 4. Hafniae. Privately published,

Copenhagen.

Gerstaecker, A. 1868. Systematische iibersicht der bis jetzt bekannt gewordenen Mydaiden (Mydasii

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Hall, C. 1985. Costa Rica. A geographical interpretation in historical perspective. Dellplain Latin

American Studies, 17. Westview Press, Boulder, Colorado.

Janzen, D. H. 1986. Guanacaste National Park: tropical ecological and cultural restoration. Fun-

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Johnson, C. W. 1933. A new species of the genus Mydas. Psyche, 39: 72.

134

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Kondratieff, B. C. & J. L. Welch. 1990. The Nemomydas of Southwestern United States, Mexico,

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Macquart, J. 1850. Dipteres exotiques nouveaux ou peu connus. 4e. supplement (part). Mem. Soc.

des Sci., de l’Agr. et des Arts, Lille, 1849: 307^79.

Osten Sacken, C. R. 1886. Diptera, vol. 1 (part), pp. 1-128. In Godman F. D. & O. Salvin (eds.).

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Seguy, E. 1928. Etude sur quelques Mydaidae nouveaux ou peu connus. Encycl. Entomol., Ser. II,

Diptera, 4: 129-156.

Steyskal, G. C. 1956. The eastern species of Nemomydas Curran (Diptera: Mydaidae). Occ. Papers

Mus. Zool., Univ. Mich., 573.

Welch, J. L. & B. C. Kondratielf. 1990. A new species of Nemomydas (Diptera: Mydidae) from

Texas. J. Kans. Entomol. Soc., 63: 643-645.

Westwood, J. O. 1841. Arcana Entomologica, or Illustrations of new, rare, and interesting insects

1. Pis. XII and XIV. Synopsis of the dipterous family Mydasidae, with descriptions of numerous

species, pp. 49-56, pis. 13-14. Privately published, London.

Wiedemann, C. R. W. 1830. Aussereuropaische zweiflugelige Insekten. Vol 2. Privately published,

Hamm.

Wilcox, J. 1981. Mydidae. pp. 533-540. In McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J.

Teskey, J. R. Vockeroth & D. M. Wood. (eds.). Manual of Nearctic Diptera. Vol. 1. Agriculture

Canada Monograph, 27.

Wilcox, J. & N. Papavero. 1975. Studies of Mydidae (Diptera) systematics and evolution. III. The

genus Messiasia D’Andretta in the Americas (Mydinae). Arq. Zool. Sao Paulo, Brasil, 26: 1-48.

Wilcox, J., N. Papavero & T. Pimentel. 1989. Studies of Mydidae (Diptera). IVb. Mydas and allies

in the Americas (Mydinae, Mydini). Museu Paraense Emilio Goeldi, Belem, Brazil.

Williston, S. W. 1898. Notes and descriptions of Mydaidae. Trans. Kansas Acad. Sci., 15: 53-58.

Received 11 October 1990; accepted 16 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(2): 135-137, (1991)

POST-EGESTIYE SURVIVAL OF

SPHENOPHORUS PHOENICIENSIS CHITTENDEN

(COLEOPTERA: CURCULIONIDAE) EGESTED BY

WESTERN TOADS

Carl D. Barrentine

Department of Biology, California State University,

Bakersfield, California 93311

Abstract.— Billbugs (Coleoptera: Curculionidae) are eaten by toads and egested alive in fecal

pellets. This study examines post-egestive weight loss and survivorship of Sphenophorusphoenici-

ensis Chittenden after passing through the digestive tract of the western toad, Bufo boreas Baird

& Girard. Weight loss (mg/h) in egested billbugs exceeds that found for nonegested billbugs at

both low and high RH. Egested billbugs do not live as long as nonegested billbugs at the same

relative humidity.

Key Words.—Bufo boreas, Sphenophorus phoeniciensis, Curculionidae, weevils, billbugs, fecal

pellets, toads, survival

Toads (Bufonidae) are known to ingest billbugs (Coleoptera: Curculionidae).

Some billbugs ( Sphenophorus spp.) resist digestion by toads (Bufo boreas Baird

& Girard) and emerge alive from fecal pellets (Barrentine in press). Egested billbugs

have reduced survival at low humidities (Fair 1969). This is putatively due to

increased water loss resulting from epicuticular damage. This study compares

weight loss and survivorship between egested and nonegested billbugs (Sphenoph¬

orus phoeniciensis Chittenden) at both low and high relative humidities.

Materials and Methods

Billbugs, S. phoeniciensis, in treatment samples were collected from infested

turfgrass (2200-2300 h) and immediately fed to captive adult toads (B. boreas

halophilus Baird & Girard). After two to three days, billbugs were isolated from

freshly egested toad fecal pellets, individually weighed (0.1 mg) and then placed

in one of two Scheibler desiccators. Both desiccators were kept at room temper¬

ature at 23° C, but differed in < 5% versus > 95% RH. Humidity was controlled

using Drierite® (anhydrous calcium sulfate) or distilled water. Weight loss and

survivorship of individual weevils were recorded at 24 h intervals. Because cur-

culionids are capable of feigning death (DuPorte 1916, Weiss 1940), mortality

was confirmed by lack of antennal reflex (flexor response to mechanical stimu¬

lation). After death, billbugs were oven dried at 68° C to constant weight. Billbugs

in control samples were collected and handled as described above, for treatment

samples, except that these were not fed to toads. Billbugs in treatment and control

samples were paired by weight (± 0.1 mg, wet) before comparison using Student’s

7-test.

Results and Discussion

At low humidity, there was no difference in weights between paired treatment

and control samples at the time of death (Table 1). Although death weights were

136

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Table 1. Weights of Sphenophorus phoeniciensis (mg, x ± SE).

Wet weight (mg)

X ± SE

Death weight (mg)

X± SE

Dry weight (mg)

x ± SE

<5% RH

Control

18.62 ± 0.48

10.90 ± 0.36

7.93 ± 0.19

Treatment

18.62 ± 0.48

11.28 ± 0.39

7.89 ± 0.19

>95% RH

Control

18.57 ± 0.54

12.68 ± 0.351

6.97 ± 0.18

Treatment

18.57 ± 0.54

15.44 ± 0.53 J

6.90 ± 0.18

‘t = 4.29, P < 0.01.

similar for both samples (40%), billbugs in the treatment sample did not, on the

average, survive as long as those in the control sample (Table 2). Mean weight

loss for egested billbugs was 0.153 mg/h (7.34 mg/48.0 h) at < 5% RH. Mean

weight loss in nonegested (normal) billbugs was 0.134 mg/h (7.72 mg/57.6 h).

Assuming that weight loss is proportional to water loss, the desiccation rate for

egested billbugs is 15% higher than that for normal billbugs at low humidity.

At high humidity, death weights between paired treatment and control samples

differed (Table 1). Death occurred at 17% and 32% weight loss in treatment and

control groups, respectively. Billbugs in the treatment sample did not, on the

average, survive as long as those in the control sample (Table 2). Mean weight

loss for egested billbugs was 0.022 mg/h (3.13 mg/144.5 h) at > 95% RH. Mean

weight loss in nonegested (normal) billbugs was 0.020 mg/h (5.89 mg/295.7 h).

Again, assuming that weight loss is proportional to water loss, the desiccation

rate for egested billbugs is 10% higher than that for normal billbugs at high

humidity.

These results both corroborate and extend observations made by Fair (1969).

Fair observed that lowered humidity (39-43%) induced death in egested billbugs

and that reduced survival was probably the result of increased water loss caused

by damage to the epicuticle. This study thus documents reduced survivorship for

egested billbugs at both low and high humidities and quantifies weight (water)

loss for both egested and nonegested billbugs.

Table 2. Time of death for Sphenophorus phoeniciensis (hours, x ± SE and range).

Hours X ± SE

Hours (Min-Max)

<5% RH

Control

57.6 ± 1.81 1

(48-96)

Treatment

48.0 ± 1.94 J

(24-72)

>95% RH

Control

295.7 ± 17.07 1 b

(72-528)

Treatment

144.5 ± 20.29 J

(24-528)

a t = 3.58, P < 0.01.

b t= 5.65, P< 0.01.

1991

BARRENTINE: POST-EGESTIVE SURVIVAL OF BILLBUGS

137

Literature Cited

Barrentine, C. D. 1991. Survival of billbugs ( Sphenophorus spp.) egested by western toads (Bufo

bore as). Herpetol. Review 22: 5.

DuPorte, E. M. 1916. Death feigning reactions in Tychius picirostris. J. Animal Behavior, 6: 138—

149.

Fair, J. W. 1969. Survival of weevils through the digestive tract of an amphibian. Bull. So. Calif.

Acad. Sci., 68: 260-261.

Weiss, H. B. 1940. The death-feints of Sitophilus granarius Linn., and Sitophilus oryzae Linn. J.

New York Entomol. Soc., 48: 37^46.

Received 14 September 1990; accepted 28 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(2):, 138-144, (1991)

SEASONAL PATTERNS IN A SAN FRANCISCO BAY,

CALIFORNIA, SALT MARSH ARTHROPOD COMMUNITY

Steven S. Balling and Vincent H. Resh

Department of Entomological Sciences, University of California,

Berkeley, California 94720

Abstract.— Biomass, abundance, and species richness patterns of detritivore, herbivore, and

carnivore components of a San Francisco Bay, California, salt marsh arthropod community were

examined over an annual cycle. Abundance and species richness patterns for herbivores and

detritivores did not track the occurrence of their respective food resources, perhaps because of

the influence of salinity levels, and the frequency and duration of tidal inundation. In contrast,

abundance and species richness of carnivores corresponded well with occurrence of their her¬

bivore and detritivore prey, perhaps because carnivores are less susceptible to salt marsh en¬

vironmental extremes. Trophic biomass analysis showed a typical pyramid-shaped relationship

among producer, herbivore and detritivore, and carnivore components in spring; during autumn,

however, the biomass of carnivores exceeded that of herbivores and detritivores.

Key Words .—salt marsh, wetlands, seasonality, trophic, arthropods, insects, Salicornia

Over an annual cycle, salt marshes are subject to relatively large seasonal fluc¬

tuations in tidal inundation frequency and duration, and in soil, ground water,

and tidal water salinity. Does the composition of the arthropod fauna show similar

temporal fluctuations? If all species of salt marsh arthropods are well adapted to

the full range of tidal inundation and salinity that occurs over a year, the occurrence

of a trophic group and the availability of its food resources should correspond,

as has been reported for more seasonally constant environments (e.g., Hurd &

Wolf 1974, Brown & Southwood 1983). However, if physical conditions disrupt

the ability of a trophic group to track its food resources, poor correspondence

may result. In this paper, we describe the seasonal occurrence of arthropods (as

measured by biomass, abundance, and number of species), and relate these findings

to the above considerations.

Materials and Methods

The study was conducted in Petaluma Marsh, Sonoma Co., California (for map

of area, see Balhng & Resh 1983: Fig. 1), which is the largest of the San Francisco

Bay tidal marshes and one of the largest (1145 ha) estuarine marshes along the

Pacific Coast of North America. The marsh is dominated by pickleweed ( Sali¬

cornia virginica L.).

Arthropods were collected using a D-vac suction device with a 0.25 mm mesh

collection bag, and were separated from plant matter using a Berlese-Tullgren

funnel; a complete description of arthropod sampling methods is given in Balling

& Resh (1982). Samples for the present analysis were chosen from a series that

was taken approximately monthly from January to November 1978 (representing

12 sampling dates). Specimens were separated into three categories: detritivores,

herbivores, and carnivores. Primary feeding habits of the species were obtained

from taxonomic specialists in these groups and from existing literature (e.g., Cam¬

eron 1972). Although some species do not feed restrictively within a single trophic

1991

BALLING & RESH: SALT MARSH ARTHROPODS

139

group, we used the predominant feeding mode of a species to describe its trophic

status.

Arthropod abundance and species richness (number of species per 0.09 m 2

sample) were based on data from six samples collected each month. Arthropod

biomass was determined from two samples each month, which were individually

oven dried at 100° C to a constant weight.

Detailed measurements of physical variables (salinity, tidal inundation, and air

temperature) were not done in Petaluma Marsh until 1980; however, values during

that year matched periodic observations made during 1977-1979. In 1980, salinity

of tidal waters was measured weekly (during high tide) with a refractometer;

periodic measurements of interstitial water salinity (Balling & Resh 1983) indi¬

cated similar seasonal trends observed to those for tidal water salinity. Tidal in¬

undation frequency of the marsh surface was calculated using data from a tide

gauge located on a nearby slough. Average air temperature was determined from

constant temperature recordings.

Results and Discussion

Biomass. — Arthropod biomass during the year was characterized by three nar¬

row, well-defined peaks; each peak represented a different trophic level: detriti-

vores peaked in March, herbivores in May and June, and carnivores in September

and October (Fig. la). At least in part, these biomass patterns reflected the phe¬

nology of the large sized species in the arthropod community. For example, the

peak in detritivores closely followed peak occurrences of the amphipod Orchestia

traskiana Stimpson and the isopod Littorophiloscia richardsonae (Holmes & Gay).

Herbivores were not dominated by any single species; instead the peak largely

resulted from four species of brine flies [Ephydridae: Psilopa (Ceropsilopa) co-

quilletti (Cresson), Scatella ( Scatella ) stagnalis Fallen, Scatella (Neoscatella) setosa

Coquillett and Scatophila sp.] and two species of leafhoppers [Cicadellidae: Strep-

tanus confirms (Reuter) and Macrosteles sp. near fascifrons (St&l)]. The carnivore

peak reflected the maturation of the population of the large wolfspider (Lycosidae)

Pardosa ramulosa (McCook).

Abundance.—Of all trophic groups, more individuals of carnivores than her¬

bivores or detritivores were collected throughout the year; however, the abundance

peaks were less well defined than the biomass peaks (Fig. lb). Abundance of all

groups peaked in spring and then again in autumn. In contrast to the influence

of large individuals on biomass, changes in abundance are often biased toward

the smaller sized, more numerous species (e.g., Odum 1971). For example, the

spring detritivore peak was dominated by the sminthurid collembolan Smin-

thurides (Sminthurides) malmgreni Tullberg and the autumn peak was dominated

by the psocopteran Lachesilla pacifica Chapman. The spring peak in herbivores

was dominated by the same species of brine flies and leafhoppers as the herbivore

biomass peak. However, the earlier rise in abundance seen in March was caused

by early instar leafhoppers, whose small size made them proportionately more

important in abundance measures than in biomass measures. Carnivore abun¬

dance peaks in spring and autumn were dominated by the small predaceous

phytosieid mite Amblyseius scyphus Schuster & Pritchard, and also included a

broad array of other mites and spiders.

Species Richness. — Like abundance, the peaks in species richness were less well

140

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Figure 1. Mean (a) biomass, (b) abundance, and (c) species richness of terrestrial arthropods

collected in Petaluma Marsh during 1979; squares represent carnivores, triangles represent herbivores,

and circles represent detritivores.

defined than those for biomass. Detritivore species richness was highest in April,

with only a small increase in autumn. Herbivore species richness was highest in

June (although this was preceded by similarly high levels that began in March),

decreased in July and August, and increased again in September and October.

Carnivore species richness peaked in May, June and July, and October.

Trophic Pyramids.— Because biomass relationships among different trophic

groups changed seasonally (Fig. la), the shape of ecological pyramids that are

developed from biomass data vary depending on the time of year. For example,

1991

BALLING & RESH: SALT MARSH ARTHROPODS

141

CARNIVORES

HERBIVORES +

DETRITIVORES

PRODUCERS

CARNIVORES

HERBIVORES +

DETRITIVORES

PRODUCERS

a. Early May

b. Early October

Figure 2. Trophic pyramids of pickleweed and arthropod biomass in (a) early May and (b) early

October 1979. Plant biomass data are from Cameron (1972) and Balling & Resh (1983). Pyramid

widths are given as the log, 0 of the biomass (mg/m 2 ).

during early May the trophic relationships were pyramid shaped; biomass esti¬

mates of producers (pickleweed), herbivores/detritivores (combined to estimate

potential carnivore prey), and carnivores decreased at successively higher trophic

levels (Fig. 2a). In contrast, during early October the carnivore and herbivore/

detritivore relationship was inverted (Fig. 2b).

Interaction of Trophic Groups and Abiotic Factors. — Although biomass patterns

in this salt marsh arthropod community showed seasonal changes indicating tro¬

phic succession, the abundance and species richness patterns showed neither

trophic succession nor the typical numerical domination of herbivores. Perhaps

this is related to the seasonal patterns of physical features of the salt marsh

environment.

In Petaluma Marsh, water salinities were lowest in February, March, and April;

they steadily increased through the following summer, and then decreased with

the onset of rain at the beginning of the wet season in October (Fig. 3a). Tidal

inundation frequency (Fig. 3b) and air temperature (Fig. 3c) showed similar pat¬

terns.

Detritivores had their highest biomass, abundance, and species richness during

spring, even though their primary food resource (which in a pickleweed mono¬

culture is composed of litter from senescent stems) was most abundant in October

and November (Cameron 1972). The early spring flushing of the marsh surface

by rainfall and low salinity tides reduces the salt content of surface litter, which

may make it more palatable to insect detritivores (Foster & Treheme 1976). In

addition, microbial colonization may condition the litter by this time, which also

provides bacterial and fungal food sources.

Abundance, richness, and biomass of detritivores declined gradually until Au¬

gust, even though abundant litter still occurred in the marsh; perhaps this decline

resulted from high salinities of tidal water (over 20 ppt, Fig. 3a) and frequent tidal

inundation of the marsh (Fig. 3b). During July and August, the marsh surface is

inundated by tides approximately 10% of the time and for periods of up to 5.5

h. Such inundations are stressful to arthropods that remain under water, and also

to those that remain on the water surface or migrate to the tops of emergent

142

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

1980

Figure 3. (a) Mean salinity of tidal water, (b) frequency of tidal inundation (per month), and (c)

mean maximum air temperature in Petaluma Marsh during 1980.

vegetation because they will be more susceptible to predation or washout by

receding tides (Foster & Treheme 1976).

In autumn, the psocid L. pacifica was the only detritivore to increase in abun¬

dance. The apparent salt tolerance of this species allows it to use this abundant

food source at a time when few other detritivore species are present (e.g., Fig. lc

shows an average of only four detritivore species per sample during September

and October).

Abundance and biomass of herbivores is highest in late spring even though

their food, which consists mostly of surface algae and the annual succulent com¬

ponent of pickleweed, increases in quantity from March through October (Cam¬

eron 1972, Mahall & Park 1976, Zedler 1982, Josselyn 1983). The rise in her¬

bivores coincides with the most rapid increase in succulent plant biomass, and

with the seasonal increase in air temperature (Fig. 3c). The rather sudden decline

coincides with the combination of peak tidal inundation (Fig. 3b) and peak sa¬

linities (Fig. 3a).

Tidal salinities, which correlate with groundwater salinities, indirectly affect

sap-feeders such as leafhoppers through reductions in their food quality because

1991

BALLING & RESH: SALT MARSH ARTHROPODS

143

pickleweed sap salinity increases as groundwater salinity increases (Flowers et al.

1977). For example, Regge (1973) has shown that salt marsh aphids will seek

plants with lower sap salinity, and Vince et al. (1981) have shown that a positive

relationship between Spartina nutritional quality and herbivore abundance occurs

in an Atlantic coast salt marsh.

The decline of herbivores at the end of spring also coincided with the rise in

carnivore abundance; thus, predation may also be an important regulating factor.

Again, as with the detritivores, high tides of mid-summer may drive some insects

to the plant tops or water surface, and thus increase the chances of predation.

Vince et al. (1981) also indicated that spider predation may limit herbivore abun¬

dance.

The seasonal patterns of carnivore abundance and species richness corresponded

well with the occurrence of expected food sources (i.e., herbivores and detriti¬

vores). Carnivores, represented primarily by mites, reached their peak abundance

in spring, which coincided with the herbivore and detritivore abundance peaks.

Although carnivore abundance declined through the summer, biomass continued

to rise. Most of the autumn predators are spiders that are well adapted to traversing

water surfaces and capturing prey that are found there (Roth & Brown 1976).

Because most of their prey regulate the salinity of their hemolymph (at least to

some extent), predators may be less affected by the rise in salinities during summer

and autumn.

In conclusion, salt marsh herbivores and detritivores apparently do not closely

track changes in the quantity of their food. In fact, the inefficiency of the detri¬

tivores may help promote the rapid accumulation of peat that occurs in many

salt marshes. The results of this study suggest that the phenologies of salt marsh

herbivores and carnivores are determined less by the quantity of food than by

tidal inundation and the quality of food, which in turn is affected by tidal and

groundwater salinities (Collins et al. 1986, Collins & Resh 1989). Carnivores, in

contrast, appear to respond to food availability rather than to the physical extremes

of the salt marsh environment.

Acknowledgment

We thank M. A. Bamby and J. N. Collins for field and laboratory assistance;

C. Kennett, R. Gill, W. Mathis, R. Snider, and J. Chapman for confirming species

identifications; R. W. Merritt and R. F. Denno for their comments on this manu¬

script; and the California Mosquito and Vector Control Association’s Coastal

Region Mosquito Abatement Districts for their support and cooperation in this

project. Funding for this project was provided by University of California Mos¬

quito Research Funds.

Literature Cited

Balling, S. S. & V. H. Resh. 1982. Arthropod community response to mosquito control recirculation

ditches in San Francisco Bay salt marshes. Environ. Entomol., 11: 801-808.

Balling, S. S. & V. H. Resh. 1983. The influence of mosquito control recirculation ditches on plant

biomass, production and composition in two San Francisco Bay salt marshes. Estuarine Coastal

Shelf Sci., 16: 151-161.

Brown, V. K. & T. R. E. Southwood. 1983. Trophic diversity, niche breadth and generation times

of exopterygote insects in secondary succession. Oecologia, 56: 220-225.

144

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(2)

Cameron, G. N. 1972. Analysis of insect trophic diversity in two salt marsh communities. Ecology,

53: 58-73.

Collins, J. N., L. M. Collins, L. B. Leopold & V. H. Resh. 1986. The influence of mosquito control

ditches on the geomorphology of tidal marshes in the San Francisco Bay Area: evolution of

salt marsh mosquito habitats. Proc. Calif. Mosq. Vector Control Assoc., 54: 91-95.

Collins, J. N. & V. H. Resh. 1989. Guidelines for the ecological control of mosquitoes in non-tidal

wetlands of the San Francisco Bay Area. California Mosquito and Vector Control Association,

Sacramento, California.

Flowers, T. J., P. F. Troke & A. R. Yeo. 1977. The mechanism of salt tolerance in halophytes. An n.

Rev. Ecol. Syst., 28: 89-121.

Foster, W. A. & J. E. Treheme. 1976. Insects of marine salt-marshes: problems and adaptations,

pp. 5-42. In Cheng, L. (ed.). Marine insects. North-Holland, Amsterdam.

Hurd, L. E. & L. L. Wolf. 1974. Stability in relation to nutrient enrichment in arthropods of old-

field successional ecosystems. Ecol. Monogr., 44: 465^482.

Josselyn, M. 1983. The ecology of San Francisco Bay tidal marshes: a community profile. U.S. Fish

Wildl. Serv., Div. Biol. Serv., Washington, D.C. FWS/OBS-83/23.

Mahall, B. E. & R. B. Park. 1976. The ecotone between Spartina foliosa Trin. and Salicornia virginica

L. in salt marshes of northern San Francisco Bay. I. Biomass and production. J. Ecol., 64: 421-

433.

Odum, E. P. 1971. Fundamentals of ecology. W. B. Saunders, Philadelphia.

Regge, H. 1973. Die Blattlaus-Arten (Hexapoda, Aphidoidea) des Gezeitenbereichs de Nordseekueste

Schleswig-Holsteins. Faun. Oekol. Mitt., 4: 241-254.

Roth, V. D. & W. L. Brown. 1976. Other intertidal airbreathing arthropods, pp. 119-150. In Cheng,

L. (ed.). Marine insects. North-Holland, Amsterdam.

Vince, S. W., I. Valiela & J. M. Teal. 1981. An experimental study of the structure of herbivorous

insect communities in a salt marsh. Ecology, 62: 1662-1678.

Zedler, J. B. 1982. The ecology of southern California coastal salt marshes: a community profile.

U.S. Fish Wildl. Serv., Div. Biol. Serv., Washington, D.C. FWS/OBS-81/54.

Received 9 July 1990; accepted 31 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(2): 145-146, (1991)

Scientific Note

ATYPICAL SEX ROLE BEHAVIOR IN THE

BALL-ROLLING DUNG BEETLE, CANTHON PILULARIUS L.

(COLEOPTERA: SCARABAEIDAE)

During July, 1990, I observed unusual feeding and reproductive behavior in

the ball-rolling dung beetle, Canthon pilularius L. On three occasions, females

engaged in what have been described as exclusively male behaviors for this species.

These observations suggest that the behavioral roles of the sexes of this species

may not always be clear-cut.

The feeding and reproductive behavior of this species, and of the Scarabaeinae

in general, are described elsewhere (Matthews, E. G. 1963. Psyche, 70: 75-93;

Halffter, G. &E. G. Matthews. 1966. Folia Entomol. Mex., 12-14: 1—312; Hallfter,

G. & W. D. Edmonds. 1982. Publ. Inst. Ecol., Mex. D.F. 10). In all of these

accounts, the male is reported to be the active partner in reproducing pairs; only

males engaged in combat over possession of dung balls, and only males initiated

formation of brood balls. I report females engaging in both of these activities at

the Central Plains Experimental Range near Nunn, Colorado.

Groups of four and 12 beetles were maintained in 0.84 m 2 outdoor screen cages.

The cages were open to the ground to allow normal ball-rolling and digging

behavior, and the beetles were provided with a continuous supply of fresh cow

dung. The sides of the cages restricted the distances that beetles could roll their

dung balls, but this did not appear to alfect their behavior in any other way. All

individuals were marked with dots of paint to facilitate field identification without

disturbance. I observed the beetles on 31 days at 1-4 day intervals throughout

the summer of 1990.

On 11 Jul 1990,1 observed a female cut and roll a dung ball that later became

a brood ball. The cutting process lasted about 20 min, after which the female

beetle continued to shape the ball, with occasional attempts at rolling it. Some

difficulty was encountered in rolling due to a small stick and a maggot that

protruded from the ball. More than 3 h after initiating ball formation, the female

began rolling the ball, and 17 min later she began burying it. Six min later, when

the ball was already partially buried, she was joined by a male beetle; the two

immediately began rolling the ball as a pair, with the male rolling and the female

riding atop the ball. After 32 min of rolling, they began burying the ball together.

It is not certain that the female initially intended this ball to be a brood ball rather

than a food ball, although the extraordinarily long time taken to create the ball

is suggestive.

On two other occasions I observed female beetles engage in combat; in at least

one of these the combat was initiated by the female. On 7 Jul 1990, when a pair

was in the process of burying a brood ball with the male digging beneath the ball

to bury it, and the female on top of the ball, another female approached. The two

females grappled together, producing clearly audible scraping, while the male

continued digging and standing beside the ball. After approximately 1 min, the

146

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(2)

intruding female departed and the original pair continued to bury the brood ball.

At no time did the male enter the combat, although he emerged from beneath

the ball while the combat was still occurring.

The second instance was on 26 Jul 1990. A male was cutting and shaping a

dung ball at the dung source when he was approached by a female. A brief fight

ensued. Normally this would end quickly with pair-bond formation when the

male recognized the sex of the opposing beetle. In this case, however, the fight

ended with the female rolling the ball away by herself, the male remaining behind

atop the dung pat. The female buried the ball 15 min later, and the male burrowed

under the dung pat.

These observations, although anecdotal, indicate that the behavior of C. pilu-

larius may be even more complex than has been reported. Previous accounts have

shown that males will make a brood ball in the absence of females, apparently

choosing the riskier investment in reproduction rather than the more certain

feeding opportunity. My observation of a female making a brood ball suggests

that females may follow the same strategy. The observations of females engaged

in combat suggest that females will fight to defend their reproductive investment

if a male is not present, and that females may choose to feed rather than reproduce

by stealing a dung ball from a male rather than mating with him.

The unusual behaviors that I observed may be partly due to a more arid

environment with shorter summers than those in which previous observations

were made (e.g., Miller, A. 1954. Am. J. Trop. Med. Hyg., 3: 372-389; Matthews

1963). The Nunn, Colorado observation site is a semiarid shortgrass prairie near

the edge of the geographical range of C. pilularius\ winters are long and cold,

summers are dry and hot, and annual precipitation averages about 325 mm.

Different climatic conditions may impose different behavioral strategies upon the

beetles. Future observations on these beetles may reveal if the behaviors I observed

are repeatable or if they were chance aberrances.

Acknowledgment.— I thank T. Crist and J. Wiens for their helpful comments

on the manuscript. Funding was supplied by NSF grant BSR-8805829 and Grant

DE-FGO2-88ER60715 from the Ecological Research Division, Office of Health

and Environmental Research, U.S. Department of Energy.

David S. Guertin, Department of Biology, Colorado State University, Fort Col¬

lins, Colorado 80523.

Received 31 October 1990; accepted 9 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(2): 147-157, (1991)

PROCEEDINGS OF THE PACIFIC COAST

ENTOMOLOGICAL SOCIETY

Four Hundred and Fifty Eighth Meeting

The 458th meeting of the Pacific Coast Entomological Society was held on Friday, 15 Jan 1988, at

8:00 PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Alan I. Kaplan presiding. The minutes of the 457th meeting were read by the recording secretary and

accepted. Mr. Daniel Gross, membership chairman, proposed three new regular members and two

new student members, who were accepted as members. Several guests were introduced: Laurie Swiadon,

who recently received her masters from Ftebrew University of Jerusalem, Dr. John E. Lattke of the

University of Maracay, Venezuela, Mr. Albert Horn, David Zuckermann of the East Bay Regional

Parks and Margaret Kelley of the Tilden Nature Center. Darryl Ubick announced the spider workshop

would be postponed until February. Several notes were given. Kirby Brown presented three slides of

insect art and artifacts from Hong Kong, China, Singapore and Thailand. The slides depicted stamps,

coins, carvings and etchings. Dr. Ferguson presented a note on recovering a tagged monarch butterfly

at an overwintering site in Mexico, 80 miles from Mexico City. It is the longest migrant ever recovered.

It was tagged on the eastern shore of Georgia Bay in Canada. Contact Dr. Ferguson for detailed

direction to the overwintering site. Mr. Kaplan announced that Gail Grodhaus of the State Department

of Public Health passed away Christmas eve, 1987. The Gail Grodhaus Memorial Fund will be set

up for chironomid study. For tax deductible contributions to the fund in memory of Gail Grodhaus,

checks can be sent to the Chironomis Grodhaus Memorial Fund, % Dr. James Sublette, Life Science

Building, University of Southern Colorado, 2200 Bonfort Blvd., Pueblo, Colorado 81000.

President Kaplan introduced the speaker for the evening, John Lane, Education Director of the

Santa Cruz Museum. Mr. Lane presented an interesting lecture titled “Overwintering Monarch But¬

terflies in California: Past and Present.” The monarch butterfly is the only insect that has a long

distance, annual, two way migration. A survey of California overwintering sites of the monarch butterfly

was given. Historical documentation was presented along with a summary of the natural history of

the monarch and a list of the trees used for overwintering. The Natural Bridges State Park has from

40,000-50,000 butterflies overwintering and sometimes up to 200,000.

President Kaplan announced that the next meeting would be held on 19 Feb 1988 and the speaker

would be Leslie Saul. The title of the talk will be “Insect Zoos and their role in Public Education.”

The meeting was adjourned at 9:30 PM and a social hour was held in the entomology department of

the California Academy of Sciences. The following 43 persons were present: (27 members) P. H.

Amaud Jr., A. Balmy, L. G. Bezark, D. S. Brennan, T. S. Briggs, K. W. Brown, H. K. Court, K.

Dabney, L. Dong, S. S. Ferguson, W. E. Ferguson, D. F. Gross, A. I. Kaplan, B. Keh, C. Y. Kitayama,

V. F. Lee, P. A. Luft, S. S. Mead, J. A. Powell, E. S. Ross, L. S. Saul, W. E. Savary, H. I. Scudder,

C. Y. Takahashi, J. E. Tobler I, D. Ubick, and M. L. Utheim; (16 guests) M. M. Amaud, L. M. Bravo

I. Brown, J. E. Court, P. R. Craig, M. Davies, T. W. Davies, L. V. Dubay, D. Hayward, A. Horn, M.

Kelley, J. E. Lattke, W. C. Rauscher, L. Swiadon, B. Tobler, and D. Zuckermann.

Four Hundred and Fifty Ninth Meeting

The 459th meeting of the Pacific Coast Entomological Society was held on Friday, 19 Feb 1988, at

8:00 PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Alan I. Kaplan presiding. The minutes of the 458th meeting were read by the recording secretary and

accepted. Mr. Daniel Gross, membership chairman, proposed one new student member, three new

regular members, who were accepted as members. Tom Briggs and Darrell Ubick are working to get

threatened status for a species of phalangid at Edgewood Park whose habitat is the site of a proposed

golf course. Dr. Edward Smith presented a note on new thysanuran drawings by Dr. Kukalova-Peck.

Leslie Saul presented a note on several insects that have been used as ornamental jewelry.

President Kaplan introduced the speaker for the evening, Leslie Saul, Director of the Insect Zoo at

the San Francisco Zoological Gardens. Ms. Saul presented a slide lecture titled “Insect Zoos and their

Role in Public Education.” Ms. Saul gave an informative talk on the scope and mission of the Insect

Zoo’s public education program and detailed the criteria for choosing insects for public exhibit.

Methods of collecting, maintaining and transporting live insects for public exhibit were also illustrated.

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President Kaplan announced that the next meeting would be held on 18 Mar 1988 and the speaker

will be Dr. William Olkowski of the Bio-Integral Resource Center. The title of the talk will be “Recent

Developments in Urban Integrated Pest Management.” The meeting was adjourned at 9:30 PM in

memory of Dr. Donald Denning, a world renowned authority on caddisflies. Dr. Denning died on 7

Feb 1988. A social hour was held in the entomology department of the California Academy of Sciences.

The following 50 persons were present: (35 members) P. H. Amaud Jr., B. T. Berke, L. G. Bezark, F.

E. Cave, J. S. Chinn, J. R. Clopton, H. K. Court, T. D. Cuneo, D. K. Dabney, J. G. Edwards, R. Fall,

S. V. Fend, N. E. Gershenz, D. F. Gross, J. E. Hafemik Jr., A. Horn, A. I. Kaplan, B. Keh, R. L.

Langston, V. F. Lee, J. Leong, P. A. Luft, S. S. Mead, D. P. Pline, R. G. Robertson, L. S. Saul, W.

E. Savary, H. I. Scudder, E. L. Smith, G. S. Spicer, R. E. Stecker, C. Y. Takahashi, D. Ubick, M. A.

Wolf, and R. L. Zuparko; (15 guests) M. M. Amaud, L. W. Berke, J. E. Court, P. R. Craig, M. Davies,

T. W. Davies, T. Eager, P. Guard da Silva, K. Hobson, S. Kauffman, P. K. Kleintjes, G. Peterson, S.

Renkes, J. Robertson, and L. Swiadon.

Four Hundred and Sixtieth Meeting

The 460th meeting of the Pacific Coast Entomological Society was held on Friday, 18 Mar 1988,

at 8:00 PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with

president Alan I. Kaplan presiding. The minutes of the 459th meeting were read by recording secretary

Leslie Saul and accepted. Mr. Daniel Gross, membership chairman, proposed one new student member,

Mr. John W. Brown and two new regular members: Mr. James C. Milstead and Mr. John D. McCarty,

who were accepted as members. Several announcements were made. Ben Keh announced that Dr. Y.

Z. Erzinclioglu of the Department of Zoology, University of Cambridge will be giving a U.C. Berkeley

departmental seminar titled “Application of Entomology to Forensic Sciences” on 18 Apr, in 101

Moffitt Hall at 4:00 PM. Bill Olkowski will have a poster display with handouts after the meeting. A

memorial service was held for Dr. Donald Denning. President Kaplan announced that he was still

seeking speakers for April and May and called for any speaker suggestions from the members. It was

suggested that one evening could be devoted to member notes and exhibits.

President Kaplan introduced the speaker for the evening, Dr. William Olkowski, Technical Director

of the Bio-Integral Resource Center in Berkeley. Dr. Olkowski gave an interesting lecture on “Recent

Developments in Urban Integrated Pest Management.” The Bio-Integral Resource Center publishes

two journals: one for professionals, “The IPM Practitioner,” and one for non-professionals, “Common

Sense Quarterly.” Dr. Olkowski outlined the principles of integrated pest management and the various

approaches which can be used on shade trees, home vegetable gardens, structural and medical pest

problems. He discussed several programs that BIRC had designed and mentioned several new tech¬

niques for monitoring, detection and control.

President Kaplan announced that the next meeting would be held on 15 Apr 1988. The meeting

was adjourned at 9:20 PM. A social hour was held in the entomology department of the California

Academy of Sciences. The following 36 persons were present: (24 members) P. H. Amaud Jr., A. M.

Balmy, L. G. Bezark, T. S. Briggs, P. Buickerood, R. Buickerood, J. S. Chinn, J. R. Clopton, D. K.

Dabney, L. Davis, N. E. Gershenz, A. Horn, A. I. Kaplan, B. Keh, L. P. Kite, V. F. Lee, L. B. Mak,

W. W. Pitcher, L. S. Saul, W. E. Savary, H. I. Scudder, M. M. Tierney, D. Ubick, and R. L. Zuparko;

(12 guests) M. Anstett, M. M. Amaud, M. Brennan, N. Brownfield, S. Chinn, M. Furey, S. V. Garity,

M. Haines, C. Harvey, D. Mello, W. Olkowski, and W. C. Rauscher.

Four Hundred and Sixty First Meeting

The 461st meeting of the Pacific Coast Entomological Society was held on Friday, 15 Apr 1988, at

8:00 PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Alan I. Kaplan presiding. The minutes of the 460th meeting were read by recording secretary Leslie

Saul and accepted. Mr. Daniel Gross, membership chairman, proposed one new student member, Mr.

John K. Jackson and two new regular members: Mr. Don C. Force and Mr. Tony Harris, who were

accepted as members. Several announcements were made. Leslie Saul announced that the Insect Zoo’s

Annual open house would be held on 12 Jun 1988. Alan Kaplan thanked Albert Horn for making

arrangements for the speaker for April. He also thanked Pat Kite, Nancy Brownfield and Dr. Harvey

Scudder for their suggestions for future speakers for PCES.

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President Kaplan introduced the speaker for the evening, Dr. Fernando Agudelo Silva, Senior

Researcher at BIOSYS, a Palo Alto firm. Dr. Silva’s talk was titled “The Worm’s Turn: Developing

Nematodes for Insect Pest Management.” Dr. Silva gave a very interesting talk on the methods used

for mass culturing nematodes for use in biological control programs. These nematodes can be used

to control borers in oak and fig trees, in the greenhouse, in cranberry bogs and in apple and almond

orchards. In the future it may be possible to control cockroaches, black fly and fire ants. In 1979, the

use of nematodes for control took a big leap when an artificial media was developed by Australian

researchers. Infective stages are reared. These enter the host through the mouth, spiracles, anus, or

through the integument. Once inside, a bacteria is released which quickly kills the insect host. The

nematodes are nonpathogenic for people and nontarget organisms.

President Kaplan announced that the next meeting would be held on 20 May 1988. The meeting

was adjourned at 9:20 PM. A social hour was held in the entomology department of the California

Academy of Sciences. The following 22 persons were present: (17 members) P. H. Amaud Jr., T. S.

Briggs, P. Buickerood, R. Buickerood, L. Davis, W. A. Doolin, J. G. Edwards, A. Horn, A. I. Kaplan,

L. P. Kite, V. F. Lee, G. J. Mallick, S. W. Miller, L. S. Saul, C. Y. Takahashi, D. Ubick, and R. L.

Zuparko; (5 guests) F. Agudelo Silva, M. M. Amaud, P. R. Craig, S. Pollitt, and W. C. Rauscher.

Four Hundred and Sixty Second Meeting

The 462nd meeting of the Pacific Coast Entomological Society convened 20 May 1988, at 8:10 PM,

in the Morrison Auditorium of the California Academy of Sciences. PCES president Alan Kaplan

presided. The minutes of the 461st meeting were read by Mr. Warren Savary and accepted as read.

Mr. Daniel Gross, PCES membership chairman, proposed four individuals for student membership:

Mr. Gordon C. Snelling, Ms. Danielle Jahnke, Ms. Robin Jean Rathman and Mr. Martin Melia; three

individuals for regular membership: Dr. V. R. Vickery, Mr. John Steiner, Mr. Patrick R. Craig; and

one individual for sponsoring membership: Ms. Nancy T. Brownfield. The proposal was seconded

and these individuals were granted membership. Mr. Warren Savary announced that he had been

asked by the San Francisco Beekeepers Association to make available a petition that proposed making

the honey bee (Apis mellifera ) the national insect. Mr. Alan Kaplan introduced the guest speaker of

the evening, Mr. Glenn Conner of the Alameda County Mosquito Abatement District, who presented

an informative lecture entitled tl Aedes albopictus : Asian Tiger Mosquito, a New Threat to California’s

Health.” Mr. Conner detailed the spread of this potential disease vector which apparently entered the

United States at Memphis, Tennessee in 1983. By 1987, it had been reported from 77 counties in 17

states, and had reached California. This tree-hole breeding mosquito, which feeds aggressively through¬

out the day, has been shown to serve as vector of dengue fever. Although a lack of summer rains may

serve as a barrier to its establishment in California, its potential transport in used tires suggests our

need for continued vigilance. The meeting adjourned for refreshments at 9:45 PM. The following 27

persons were present: (23 members) P. H. Amaud Jr., L. G. Bezark, T. S. Briggs, P. Buickerood, R.

Buickerood, R. Fall, D. F. Gross, A. Horn, A. I. Kaplan, B. Keh, W. A. Maffei, G. J. Mallick, S. M.

Mcllraith, S. S. Mead, M. Melia, W. E. Savary, G. A. Sayre, H. I. Scudder, S. S. Shanks, R. E. Stecker,

C. Y. Takahashi, D. Ubick, and T. J. Zavortink; (4 guests) M. M. Amaud, G. E. Conner, W. C.

Rauscher, and S. Renkes.

Four Hundred and Sixty Third Meeting

The 463rd meeting of the Pacific Coast Entomological Society was held on 16 Sep 1988, at 8:10

PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Alan I. Kaplan presiding. The acting recording secretary read the minutes of the 462nd meeting.

The minutes were approved as read. Five persons were proposed by Daniel Gross and elected as new

regular members: Dr. Elizabeth A. Bemays, Dr. Ryutaro Iwata, Dr. Penelope F. Kukuk, Ms. Patricia

F. Neyman, and Dr. Richard S. Zack. The following guests were introduced: Dr. Janice Edgerly-Rook

and her husband, Edward; Dr. C. Riley Nelson, Tilton Fellow in the Department of Entomology,

California Academy of Sciences; and Mrs. Heidi E. M. Dobson.

Mr. Warren Savary showed some slides showing the orientation of first instar larvae in several

genera of vaejovid scorpions. Mr. Kaplan announced that some members from Chico will host a

collecting trip in their local area for the June, 1989, annual meeting of the American Association for

the Advancement of Science, Pacific Division, in Chico.

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Mr. Kaplan introduced the featured speaker, Dr. Gordon Frankie, who spoke on “Lomas Barbudal:

A New Biological Reserve in Costa Rica.” Dr. Frankie, president of Friends of Lomas Barbudal,

discussed the formation, goals and future plans for the Lomas Barbudal Reserva Biologica, a lowland

dry forest. He compared the preserve with the well known wet forests of La Selva and Monteverde

and showed that a dry forest can also harbor unique and distinct fauna and flora. He talked about

fire as a major problem in Costa Rica, where introduced vegetation gets established after a fire

outcompetes the native flora which is not adapted to it. He stressed the importance of educating local

people about their native fauna and flora. He also stressed that it is important for them to get involved

in conservation projects and decision making regarding the use of their land.

The meeting was adjourned and a social hour was held in the entomology department of the

California Academy of Sciences. A total of 52 persons attended the meeting: (30 members) P. H.

Amaud Jr., T. S. Briggs, K. W. Brown, J. S. Chinn, J. S. Cope, K. Dabney, L. H. Davis, H. E. M.

Dobson, J. G. Edwards, S. V. Fend, S. S. Ferguson, E. M. Fisher, G. W. Frankie, D. F. Gross, J. E.

Hafemik, Jr., A. I. Kaplan, B. Keh, L. P. Kite, R. L. Langston, Y. F. Lee, L. B. Mak, G. J. Mallick,

M. Melia, N. D. Penny, L. Saul, W. E. Savary, H. I. Scudder, S. S. Shanks, M. M. Tierney, and D.

Ubick; (22 guests) M. M. Amaud, B. Atkinson, D. De Raue, R. W. Douglas, C. W. Fox, J. Edgerly-

Rooks, M. Flexer, J. Frankie, J. B. Fraser, M. H. Fraser, M. Furey, J. Hirabayashi, R. Nelson, T. K.

Ohsumi, R. A. Raguso, S. Renkes, N. Robinson, E. Rooks, D. Schmidt, P. Sullivan, and A. Zoidis.

Four Hundred and Sixty Fourth Meeting

The 464th meeting of the Pacific Coast Entomological Society was held on Friday, 21 Oct 1988, at

8:10 PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Alan I. Kaplan presiding. The acting recording secretary read the minutes of the 463rd meeting and

they were accepted as read. Two new members were proposed and accepted. Several announcements

were made. President Kaplan announced the possibility of having a joint meeting with AAAS and

that Vincent Lee has suggested presenting papers; the deadline for the decision on participation is 15

Mar 1988. Mr. Kaplan also announced that a field day will be held in Chico, which will coincide with

the June AAAS meeting. Vincent Lee described the Academy’s new exhibit “Wild California” and

urged all members to make plans to see it. The following guests were introduced: Nora Welk and

Derham Giuliani. Mr. Guiliani has been donating specimens to the entomology department at the

California Academy for many years. Edward Smith showed a paper wasp nest that he collected near

his home. He noted that moths were beginning to emerge in great numbers. Larry Bezark announced

that he had brought along some arthropod specimens which had been collected in Papua New Guinea

by Larry Orsak. The specimens were available for examination during the social hour. Mr. Kaplan

announced that he was preparing to call on members to serve on various committees for the upcoming

year.

President Kaplan introduced the speaker for the evening, Mr. Patrick Craig, of Antioch West College

in San Francisco, who presented a slide lecture on “Amber and its Inclusions.” Mr. Craig began by

discussing the different types of amber collected in various parts of the world. His talk focused on the

amber mined in the Dominican Republic where amber is an important commodity. Mr. Craig discussed

the mining process, how the amber is fashioned and marketed. He pointed out that scientific collectors

could not financially compete with merchants that were willing to pay more than twenty thousand

dollars for single pieces with large inclusions. He showed photographs of animals preserved in amber

which included a fossilized slug, an entire gecko and a variety of arachnids, insects and other arthropods.

He noted that animals preserved in amber retained their genetic material which could still be analyzed.

He also noted that even though the arthropods in amber are mostly pre-Oligocene, they are for the

most part indistinguishable from the species living today in the same type of habitat, indicating the

stability of tropical ecosystems over time.

President Kaplan announced that the next speaker would be Dr. Vincent Resh of U. C. Berkeley

who would speak on “Geothermal Development and Endangered Species.” The meeting was adjourned

at 9:20 PM and a social hour was held in the entomology department of the California Academy of

Sciences. The following 60 persons were present: (38 members) P. H. Amaud Jr., L. G. Bezark, T. S.

Briggs, K. W. Brown, J. S. Chinn, J. R. Clopton, Patrick R. Craig, T. D. Cuneo, L. Dong, J. T. Doyen,

B. Ehreth, S. S. Ferguson, W. E. Ferguson, C. W. Fox, D. F. Gross, D. R. Herlocker, A. Horn, P. S.

Johnson, A. I. Kaplan, L. P. Kite, R. L. Langston, V. F. Lee, W. E. Maffei, L. B. Mak, G. J. Mallick,

M. Melia, N. D. Penny, D. P. Pline, K. J. Ribardo, R. G. Robertson, W. E. Savary, H. I. Scudder, S.

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S. Shanks, E. L. Smith, G. S. Spicer, C. Y. Takahashi, D. Ubick, and T. J. Zavortink; (22 guests) M.

M. Amaud, F. Baker, I. Baker, Y. Black, T. L. Davis IV, E. A. Doyen, D. Giuliani, B. Hall, B. I.

Hiler, T. F. Hlavac, D. Maffei, D. L. Mead, C. R. Nelson, B. Ohlanul, J. Ohlanul, W. C. Rauscher,

J. Ribardo, J. Robertson, L. S. Saul, M. A. Tenorio, R. Teys, and B. A. Wilson.

Four Hundred and Sixty Fifth Meeting

The 465th meeting of the Pacific Coast Entomological Society was held on Friday, 18 Nov 1988,

at 8:00 PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with

president Alan I. Kaplan presiding. The minutes of the 464th meeting were read and accepted by

Leslie Saul, recording secretary. Mr. Daniel Gross proposed two new members, Bernice Demarce and

C. Riley Nelson, and they were accepted as members. Several notes were given. Larry Bezark presented

a note on rain beetles in the genus Pleocoma stating that there were 25 species found throughout

California. Another note was presented on insect avoidance behavior in found in caribou where biting

flies occur. It was determined that 7-50% of caribou time was spent in insect avoidance behavior

depending on biting fly population density. Leslie Saul presented two slides of a species of Amblypigida

from the Jimba Caves, Kilifi district of Kenya, Africa collected by Dr. Robert Drewes in May 1988.

The first slide was of an adult female with 25-30 young on her back that hatched on 14 Nov 1988.

The second slide was a close-up of a one week old juvenile that was 6 mm in length. A third slide

was of an artistically decorated cockroach from a nonentomological conference to demonstrate the

increasing public interest in entomology. President Kaplan announced that PCES will have a con¬

tributed paper session at the AAAS meeting to be held in Chico in June. Detail will be included in

the January newsletter and AAAS newsletter. The Society of Vector Ecologists and the American

Ecological Society will also be participating. Committee members were announced. Vannoy Davis

will be a member of the audit committee; Stanley C. Williams, Larry Bezark, and Curtis Takahashi

will be members of the nominating committee. The nominating committee will make their recom¬

mendations next meeting. Mr. Kaplan announced that PCES dues are due.

President Kaplan introduced the speaker for the evening, Dr. Vincent Resh of U. C. Berkeley, whose

talk was titled “Hot Springs, Geothermal Development, and Rare and Endangered Species: The Search

for the Wilbur Spring Shore Bug.” Dr. Resh gave a brief history of geothermal energy in the west. Dr.

Resh detailed the natural history of the hot springs-inhabiting shore bug, Saldula usingeri which was

proposed for listing as an endangered species. Dr. Resh studied 119 springs discovering a range for

this saldid from Glenn to Contra Costa County. Saldula usingeri was discovered to be sluggish, a

behavioral advantage in hot spring habitats. It does not drink in highly mineralized springs but instead

gets its water by feeding on an ephydrid which is a good osmoregulator. It is thought that spider

predation keeps the shore bug from some habitats that it could otherwise occupy. Because of its wide

distribution, the Wilbur Springs Shore Bug was not listed as an endangered species.

President Kaplan announced that the next meeting would be held on 16 Dec 1988 and the speaker

would be Alan Kaplan. The title of the talk will be “Insects in the News.” The meeting was adjourned

at 9:35 PM and a social hour was held in the entomology department of the California Academy of

Sciences. A total of 46 persons attended the meeting: (35 members) P. H. Amaud Jr., L. G. Bezark,

T. S. Briggs, J. Burberry, J. R. Clopton, S. V. Fend, W. E. Ferguson, S. S. Ferguson, C. W. Fox, N.

E. Gershenz, D. F. Gross, D. A. Jensen, P. S. Johnson, A. I. Kaplan, L. P. Kite, R. L. Langston, V.

F. Lee, G. J. Mallick, S. S. Mead, M. Melia, C. R. Nelson, N. D. Penny, D. P. Pline, V. H. Resh, K.

J. Ribardo, L. S. Saul, W. E. Savary, S. S. Shanks, W. D. Shepard, R. E. Stecker, C. Takahashi, M.

M. Tierney, D. Ubick, T. J. Zavortink, R. L. Zuparko, and one unreadable; (11 guests) M. Amaud,

W. Cole, G. T. Coppe, B. I. Hiller, D. Mead, A. M. L. Penny, S. Renkes, J. M. Ribardo, L. Saul, S.

M. Villegas, and A. Zoidis.

Four Hundred and Sixty Sixth Meeting

The 466th meeting of the Pacific Coast Entomological Society was held on Friday, 16 Dec 1988,

at 8:20 PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with

president Alan I. Kaplan presiding. The minutes of the 465th meeting were read by recording secretary

Leslie Saul and accepted. Membership chairman Mr. Daniel Gross proposed two new student members,

Mr. Erik Johanson and Noel Caroline, and two new regular members, Mr. Scott Beckman and Ms.

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Irene Terry, who were accepted as members. Several announcements were made. Daniel Gross an¬

nounced that old schmidt boxes were available in the entomology department for a nominal donation.

A request was made for increased support of the refreshment committee. Larry Bezark announced

that Cornell press was offering a 20% book discount for a limited time. Two notes were given. Wesley

Maffei showed SEM slides of Morpho butterfly scales and discussed the possibility of their use taxo-

nomically. Slides of Morpho cypris, M. aega, M. deidamia, M. achilles, M. godarti, M. menelaus, M.

didius and M. amathonte were shown. Larry Bezark made two literary presentations: one covering a

robbery by men armed with a cicada; and another about a mouse-eating locust from 1915. President

Kaplan called for the annual committee reports. Membership chairman Daniel Gross reported that

in 1988, 40 new members were admitted: 25 regular, 14 student and 1 sponsoring, bringing the total

number of members to 467. Dr. Paul Amaud gave the Treasurer’s report for Dr. W. Pulawski. He

reported that the Society’s income was $28,880, expenses were $22,968, resulting in a balance of

$5,912. A thank you was extended to H. Vannoy Davis for his services on the auditing committee.

The Auditing Committee reported that all was in order and that a statement will be published in the

Annual Report. Dr. Paul Amaud reported for the Historical Committee that the reorganization of the

archives has been completed with 161 archive boxes. A special thanks was extended to Madeline

Amaud for her help with the archival material. Vincent Lee announced that the Society receives 63

serials from 51 institutions as exchanges. There have been six new exchanges this year. The Publications

Committee reported that the Society will be returning to Allen Press after the 3rd issue which should

be ready this week. Dr. Stanley Williams reported nominations for next year for the Nominating

Committee: President 1989, Dr. Thomas Zavortink; President-elect 1990, Dr. Robert Dowell; Trea¬

surer, Dr. W. Pulawski; Managing Secretary, Vincent Lee; and Recording Secretary, Leslie Saul.

Dr. Thomas Zavortink took charge of the meeting and introduced the speaker for the evening,

current President of the Society Alan Kaplan. Mr. Kaplan, who received his Masters from U. C.

Berkeley, is Naturalist at the Tilden Nature Center of the East Bay Regional Parks. President Kaplan

gave an entertaining and informative talk titled “Insects and Entomologists in the media.” Mr. Kaplan

presented a synthesis of entomology through the ages which was well researched. He also summarized

public attitude changes through time giving a survey of insect images in films.

The meeting was adjourned at 9:25 PM. A social hour was held in the entomology department of

the California Academy of Sciences. The following 42 persons were present: (29 members) P. H.

Amaud Jr., L. G. Bezark, T. S. Briggs, K. W. Brown, P. Buickerood, J. S. Chinn, H. K. Court, J. G.

Edwards, S. V. Fend, N. E. Gershenz, D. F. Gross, K. S. Hagen, P. S. Johnson, A. I. Kaplan, B. Keh,

V. F. Lee, W. A. Maffei, L. B. Mak, C. R. Nelson, N. D. Penny, K. J. Ribardo, R. G. Robertson, L.

S. Saul, W. E. Savary, S. S. Shanks, C. Y. Takahashi, D. Ubick, S. C. Williams, and T. J. Zavortink;

(13 guests) M. M. Amaud, K. Blakwell, R. Buickerood, J. E. Court, D. W. Fletcher, M. Furey, M.

Hagen, A. M. L. Penny, W. C. Rauscher, S. Renkes, J. M. Ribardo, J. Robertson, and S. M. Villegas.

Four Hundred and Sixty Seventh Meeting

The 467th meeting of the Pacific Coast Entomological Society was held on 20 Jan 1989, at 8:00

PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, president Thomas

Zavortink presiding. The minutes of the 466th meeting were read by Leslie Saul, recording secretary,

and accepted. Membership chairman Daniel Gross proposed four new members which were accepted:

Mr. Donald Barwell, Mr. Randall Ridley, Dr. Richard Mason and Mr. Bmce Tilden. Several an¬

nouncements were made. PCES will participate in a contributed paper session at the AAAS Pacific

Division meeting being held in Chico, 11-15 Jun 1989. Rewards for student papers are being offered.

A Program Committee was formed. Dr. John Hafemik was appointed as chairman and Dr. Jerry

Powell will serve on the committee. Vincent Lee announced that money can be donated and designated

specifically to the entomology department of the California Academy of Sciences. Daniel Gross will

be retiring from the refreshment committee. Dr. J. W. Tilden passed away 30 Dec 1988, at the age

of 83. His specialty was nymphalid butterflies. He was the author of the books “A Field Guide to

Western Butterflies” and “California Butterflies.” Ben Keh showed slides of the Penang Butterfly

House and described its operation.

President Zavortink introduced the speaker for the evening, Dr. Robbin Thorp, professor of En¬

tomology at U. C. Davis. Dr. Thorpe gave an interesting slide lecture titled “Australian Bees and

Scenes,” on his research trips to Australia covering several visits there to study the impact of the

introduced honey bee on native bees and flora there.

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Dr. Zavortink announced that the next meeting would be held on 17 Feb 1989 and the speaker

would be Dr. C. Riley Nelson speaking on “Winter Stoneflies in California.” The meeting was ad¬

journed at 9:45 PM and a social hour was held in the entomology department of the California

Academy of Sciences. The following 44 persons were present: (31 members) P. H. Amaud Jr., F. L.

Blanc, T. S. Briggs, K. W. Brown, J. S. Chinn, K. Dabney, J. G. Edwards, E. M. Fisher, D. F. Gross,

A. Horn, B. Keh, R. L. Langston, V. F. Lee, M. Melia, N. D. Penny, D. P. Pline, K. J. Ribardo, R.

G. Robertson, E. S. Ross, L. S. Saul, W. E. Savary, H. I. Scudder, S. S. Shanks, J. A. Skinner, J. T.

Sorensen, R. E. Stecker, C. Y. Takahashi, R. W. Thorp, D. Ubick, S. P. Welles Jr., and T. J. Zavortink;

(13 guests) M. M. Amaud, F. Blanc, L. M. Bravo, L. V. Dubay, A. M. L. Penny, W. C. Rauscher, J.

M. Ribardo, J. Robertson, C. Thorp, J. Thorp, P. Thorp, S. M. Villegas, D. Welles, and P. Welles.

Four Hundred and Sixty Eighth Meeting

The 468th meeting of the Pacific Coast Entomological Society was held on 17 Feb 1989, at 8:00

PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Thomas Zavortink presiding. The minutes of the 467th meeting were read by Leslie Saul, recording

secretary, and accepted. Membership chairman Daniel Gross proposed one student member, Mr. Lee

H. Simons, and one sponsoring member, Mr. Pat Sullivan, which were accepted. Several announce¬

ments were made. Daniel Gross will be retiring from the refreshment committee. A worthy replacement

is still being sought. All members are encouraged to volunteer. Dr. Ross introduced guests Gene Hall

and Dr. George Ball. Leslie Saul announced that the Insect Zoo’s tenth anniversary celebration would

be coming up in June. All PCES members are invited to contribute any ideas for this event.

President Zavortink introduced the speaker for the evening, Dr. C. Riley Nelson, current Tilden

Fellow in the Department of Entomology at the California Academy of Sciences. Dr. Nelson gave an

informative talk titled “Winter Stoneflies in California.” Dr. Nelson began with an introduction and

overview of early work on Plecoptera with an emphasis on the genus Capnia. He noted that there

were 55 species found in North America exhibiting a high degree of endemism. Several habitats were

discussed including the Feather River, Navarro River, Alder Creek, Long Valley Creek, Lake Tahoe,

Dolores River and Lone Pine Creek. Capnia licustra from Lake Tahoe was noted to be aquatic in the

adult stage and found at depths of 300 ft.

Dr. Zavortink announced that the next meeting would be held on 17 Mar 1989 and the speaker

would be Dr. Edward Ross. The meeting was adjourned at 9:30 PM and a social hour was held in

the entomology department of the California Academy of Sciences. A total of 47 persons attended

the meeting: (26 members) L. G. Bezark, R. L. Bottoroff, P. Buickerood, J. S. Chinn, H. K. Court, T.

D. Cuneo, J. G. Edwards, S. V. Fend, W. E. Ferguson, S. S. Ferguson, D. F. Gross, P. Johnson, D.

H. Kavanaugh, B. Keh, G. Mallick, C. R. Nelson, N. D. Penny, W. W. Pitcher, K. J. Ribardo, R. G.

Robertson, L. S. Saul, S. S. Shanks, W. D. Shepard, G. C. Snelling, C. Takahashi, and T. J. Zavortink;

(21 guests) G. E. Ball, R. Buickerood, J. E. Court, M. Duffy, M. Gardner, T. Glenn, C. Greene, W.

E. Hall, B. A. Knight, A. W. Knight, M. Monsees, A. M. L. Penny, P. Pitcher, S. Renkes, J. M.

Ribardo, J. Robertson, D. Sipes, L. Snelling, J. Snelling, V. Snelling, and S. Wright.

Four Hundred and Sixty Ninth Meeting

The 469th meeting of the Pacific Coast Entomological Society was held on 17 Mar 1989, at 8:00

PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Thomas Zavortink presiding. The minutes of the 468th meeting were read by Leslie Saul, recording

secretary, and accepted. Membership chairman Daniel Gross proposed one student member, Mr.

Alvin Glot, and two regular members, Dr. Susan B. Halbert and Dr. Robert S. Martin, which were

accepted. Several announcements were made. PCES is still seeking a refreshment committee. Con¬

tributing papers for the AAAS meeting in Chico must be received by 31 Mar 1989. Student awards

will be presented. Journal Volume 64 number is at A-R Editions and number 1 and 2 of volume 65

have been approved for publication. Daniel Gross gave a short slide presentation on insects of pre¬

historic nature and their extinction and survival, based upon a new exhibit at the Academy.

President Zavortink introduced the speaker for the evening, Dr. Edward Ross, who gave a slide

lecture titled “Recent Entomological Experiences in Mexico.” Dr. Ross gave a well illustrated lecture

which focused on the embiids and various trips to Baja California spanning from 1931 to a recent

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Yol. 67(2)

adventure along the same route. He also discussed witnessing the monarch butterfly overwintering

sites in Mexico and mentioned Dr. Lincoln Brower’s conservation efforts there. Unique arthropods,

trees and angiosperms encountered were presented.

Dr. Zavortink announced that the next meeting would be held on 21 April and the speaker would

be Dr. Robert Dowell. His talk will be titled “Evolution of the Host Range of the Citrus Blackfly.”

The meeting was adjourned at 10:00 PM and a social hour was held in the entomology department

of the California Academy of Sciences. A total of 60 persons attended the meeting: (31 members) P.

H. Amaud Jr., B. T. Berke, F. L. Blanc, T. S. Briggs, K. W. Brown, D. J. Burdick, D. K. Dabney, L.

H. Davis, J. Chinn, J. R. Clopton, J. G. Edwards, W. E. Ferguson, S. S. Ferguson, D. F. Gross, L. P.

Kite, V. F. Lee, W. Maffei, J. D. McCarty, S. Mead, C. R. Nelson, W. W. Pitcher, K. J. Ribardo, R.

G. Robertson, E. S. Ross, W. E. Savary, L. E. Serpa, W. D. Shepard, C. Takahashi, M. M. Tierney,

J. E. Tobler I, D. Ubick, and T. J. Zavortink; (29 guests) M. Amaud, K. Attipaid, F. Blanc, Y. Burgess,

L. Disterheft, J. Edwards, T. Glenn, E. A. Goff, K. Haires, D. Holth, K. Horn, J. Johnton, R. Kolesen,

D. Maffei, D. Mead, M. Mistry, P. Pitcher, W. Rauscher, J. Ribardo, J. Robertson, N. Solberg, Ca.

Tobler, Ch. Tobler, N. Tobler, A. Tung, S. Villegas, and three unreadable.

Four Hundred and Seventieth Meeting

The 470th meeting of the Pacific Coast Entomological Society was held on 21 Apr 1989, at 8:10

PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Thomas Zavortink presiding. The minutes of the 469th meeting were read, corrected and accepted.

One new member was proposed, David C. Taylor, and accepted. Several announcements were made.

Five abstracts were submitted for presentation at the AAAS meeting to be held on 11 Jun 1989. A

refreshment chairman is needed to provide refreshments for the social hour after the meetings. The

program committee would welcome suggestions for speakers for upcoming meetings. Howell Daly

would like to step down as publication chairman of The Pan-Pacific Entomologist. Leslie Saul an¬

nounced that the zoo keeper association will be sponsoring a Bowl-A-Thon on 13 May 1989 at Rock

and Bowl and the proceeds will go to benefit habitat protection in Guanacaste National Park in Costa

Rica.

President Zavortink introduced the speaker for the evening, Dr. Robert Dowell of the California

Department of Food and Agriculture and president-elect of PCES, who gave an interesting talk titled

“Evolution and Host Range of the Citrus Black Fly.” The citms black fly is a whitefly in the Aleyrodidae

family, that is now found in Africa, the Caribbean, Mexico and Florida. It originated in Malaya where

citrus was the only host. This whitefly was first found in on Key West, Florida, in 1934 was eradicated

using fish oil spray. It reappeared in 1976 in Broward County Florida and eradication with sprays

failed. Two parasitic wasps were released reducing populations by over 95%. The pest is polyphagus,

feeding on 48 genera of plants and 35 different families. Dr. Dowell presented a detailed description

of the citrus black fly’s life history, morphology, oviposition behavior, dispersal behavior and foodplant

preference. Oviposition preference was studied and it was found that kumquat, mango and pink

trumpet were as attractive as citrus. Dr. Dowell also discussed the spread of this insect and its correlation

with the spread of different horticultural plants.

The meeting was adjourned at 9:35 PM and a social hour was held in the entomology department

of the California Academy of Sciences. The following 22 persons were present: (19 members) P. H.

Amaud Jr., T. S. Briggs, W. A. Doolin, R. V. Dowell, J. G. Edwards, A. Horn, P. S. Johnson, R. L.

Langston, V. F. Lee, W. A. Maffei, G. J. Mallick, C. R. Nelson, K. J. Ribardo, L. S. Saul, W. E.

Savary, S. S. Shanks, C. Y. Takahashi, S. P. Welles, and T. J. Zavortink; (3 guests) D. Maffei, W. C.

Rauscher, and J. M. Ribardo.

Four Hundred and Seventy First Meeting

The 471st meeting of the Pacific Coast Entomological Society was held on 19 May 1989, at 8:00

PM, in Morrison Auditorium of the California Academy of Sciences, San Francisco, with president

Thomas Zavortink presiding. The minutes of the 470th meeting were read by recording secretary

Leslie Saul, and accepted. Seven new regular members and one new student member were proposed

by Daniel Gross and elected: Russell C. Bingham, Dr. Lloyd M. Dosdall, Dr. Boris C. Kondratieff,

Dr. Robert E. Page Jr., Dr. Michael Pitcairn, James E. Wappes, Ms. Carrie J. S. Wong, as regular

members, and Mr. Dean F. Messer, as a student member.

1991

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Several announcements were made. Mr. Warren Savary announced that seven papers were to be

presented at the upcoming AAAS meeting. The Western Apiculturist Society will be meeting on the

week of 10 Jun 1989 at San Francisco State University and a short course on starting an apiary will

be given. Leslie Saul announced that the Insect Zoo’s tenth annual open house will be held on 10-11

June 1989. James Tobum of Santa Rosa has back issues of the journal which will be sold individually

or as a set for $85.00. There is a new printing of A Glossary of Entomology by J. R. de La Torre-

Bueno, which will be available at pre-publication price of $35.00; the publication date is 1 Jun 1989;

it will include an entomological spelling checker. Copies of the order forms will be available in the

entomology department of the Academy. Dr. Thomas Zavortink announced that Volume 65, issue 1

should be arriving soon. Dr. Edward Ross introduced two guests, Dr. P. Jolivet, a chrysomelid expert,

and J. Segress. Dr. Zavortink introduced the speaker for the evening, Dr. Kirby Brown of the San

Joaquin County Department of Food and Agriculture. His talk was titled “Gold Bugs and other

Numismatic Insects.” Dr. Brown gave an informative talk on the field of insect coins. There are over

2500-3000 different types of coins that depict insects or bee hives. The first coins date back to 650

BC. Dr. Brown showed a series of slides of various coins depicting insects from his collection and

studies from such places as Italy, Papua New Guinea, Panama, Tonga and Utah. Bees were the insect

most often represented.

The meeting was adjourned at 9:40 PM and a social hour was held in the entomology department

of the California Academy of Sciences. The following 35 persons were present: (23 members) P. H.

Amaud Jr., K. W. Brown, J. S. Chinn, H. K. Court, J. G. Edwards, A. Horn, A. S. Hunter, B. Keh,

R. L. Langston, V. F. Lee, W. A. Maffei, G. J. Mallick, M. Melia, N. D. Penny, K. J. Ribardo, R. G.

Robertson, L. S. Saul, W. E. Savary, R. E. Stecker, C. Y. Takahashi, J. E. Tobler, C. J. S. Wong, and

T. J. Zavortink; (12 guests) M. M. Amaud, J. E. Court, E. A. Goff, P. Jolivet, D. Maffei, D. S. Mayuoba,

A. M. Penny, J. M. Ribardo, J. Robertson, J. Segress, S. Villegas, and one illegible signature.

Four Hundred and Seventy Second Meeting

The 472nd Meeting of the Pacific Coast Entomological Society was held 17 Nov 1989, in the Goethe

Room of the California Academy of Sciences with president Dr. Thomas Zavortink presiding. The

meeting was called to order at 8:00 PM. Minutes of the 471st meeting were read, corrected and

accepted. Four new regular members and one new student member were proposed and accepted.

The October 1989 meeting of the Pacific Coast Entomological Society meeting was cancelled due

to the October 17th earthquake in San Francisco.

President Zavortink made several announcements. Dr. Zavortink read a letter from Howell Daly

stating that Dr. John Sorensen will assume the duties of Editor of The Pan-Pacific Entomologist. A

special thanks was extended to Dr. Chemsak for his effort to bring the journal, The Pan-Pacific

Entomologist, up to date. The publication is now back with Allen Press because of their ability to

produce a higher quality product. Thanks were also extended to Ken Cooper, Paul Amaud and

Wojciech Pulawski. The auditing committee members now include Helen Court, Vannoy Davis,

chairman of the committee and Dr. Paul Amaud. The publication committee needs one additional

member. The nominating committee includes Ron Stecker. Patty Pratt announced that five of her

sculptural works dealing with insects will be on display at the Insect Zoo through 15 Dec 1989. Leslie

Saul announced that Dr. Robert Pyle, compiler of the IUCN Invertebrate Red Data Book, will give

a lecture on “The Conservation of Monarch Butterflies” on 25 Jan 1990, in Morrison Auditorium at

the California Academy of Sciences. Dr. Edward Smith described the exciting development of the

new “Life through Time” exhibit at the California Academy of Sciences. The exhibit is scheduled to

open in April. Dr. Scudder announced that 50% of the insect paleontology group in the U.S. is retired,

but still going strong. Two volumes of “Genera of Fossil Insects” by Frank M. Carpenter from Harvard

are being published. Dr. Edward Smith announced that Dr. Alex Rasnitzen, foremost specialist on

fossil insects in the world, will be at the Academy in January. Wes Maffei and his wife have generously

offered to serve on the refreshments committee for one year. There will be a meeting of the executive

board of PCES on 15 Dec 1989. A notice will be sent. A special dedication for the R. L. Usinger Rare

Book Collection in the Entomology Library will be held on 15 Dec 1989, at 7:00 PM in the Department

of Entomology of the California Academy of Sciences. An invitation will appear in Bits & PCES.

Debra Meed, a graduate student at San Francisco State University working on Hygrotis, was introduced.

Charles Franklin, a graduate student at San Jose State University, was introduced.

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

Dr. Edward Smith gave a note describing the unusual life history of the primitive sawfly, Cimbex

americana. It squirts fluid from pores above its spiracles. A large infestation was found on Zorea

americana (Caprifoliacea). Slides picturing oviposition were shown. Eggs are laid in the leaf margin.

After the first feeding, the larvae turn green and retain that color. The first instar have a waxy bloom;

late instar larvae do not.

Dr. Zavortink introduced the speaker for the evening, Dr. Philip Ward from the Department of

Entomology at U.C. Davis. His talk was titled “Ants in Plants.” Dr. Ward discussed the debate about

the origin and evolution of the interactions between ants and plants. Different associations between

ants and plants were outlined: casual visitation to plants, detrimental to plant, beneficial to plant, and

co-evolved mutualism. Plants with domatiums and their ant associates were discussed in detail. Fifty

representative taxa were subjected to phylogenetic analysis and a consensus tree was drawn which

suggested that these associations evolved a number of different times, perhaps 12 times. The ecology

of various relationships was described.

The meeting was adjourned at 9:35 PM. Refreshment were served in the entomology department.

A total of 45 persons attended: (32 members) P. H. Amaud Jr., T. S. Briggs, R. Buickerood, P.

Buickerood, J. S. Chinn, D. K. Dabney, S. S. Ferguson, W. E. Ferguson, C. W. Fox, D. F. Gross, P.

S. Johnson, D. Kavanaugh, Y. F. Lee, W. E. Maffei, L. B. Mak, G. J. Mallick, D. L. Mead, N. D.

Penny, W. W. Pitcher, R. G. Robertson, L. S. Saul, W. E. Savary, H. I. Scudder, S. S. Shanks, E. L.

Smith, G. S. Spreer, R. E. Stecker, C. Y. Takahashi, P. Welles, S. C. Williams, and T. J. Zavortink;

(13 guests) M. M. Amaud, V. M. Barlow, C. D. Franklin, C. Greene, S. Haughes, L. Hiugich, D.

Maffei, M. Marcus, J. Osman, P. Pratt, W. Rauscher, A. A. Roberts, and P. Ward.

Four Hundred and Seventy Third Meeting

The 473rd meeting of the Pacific Coast Entomological Society was held 15 Dec 1989 in the Goethe

Room of the California Academy of Sciences at 8:10 PM with president Dr. Thomas Zavortink

presiding. The minutes of the 472nd meeting were read, corrected and accepted. Two new candidates

were proposed by Daniel Gross, chairman of the membership committee, and elected as new members:

Mr. Charles Franklin, graduate student at San Jose State University, was elected as a new student

member, and Mr. Gregory S. Sautter of Mishawaka, Indiana, was elected as a new regular member.

Several announcements were made. Vincent Lee turned in his letter of resignation as managing

secretary of the Society to be effective 1 Jan 1991. Robin Thorpe will assume the position of chairman

of the publications committee effective in January, 1990. Dr. Stanley Williams will join that committee.

There were 1400 oral presentations made at the San Antonio meetings of the Entomological Society

of America. The 1990 meetings of the ESA will be held in New Orleans, and the 1991 meetings will

be held in Reno. Dr. Zavortink extended his thanks to everyone for their support during the year of

his presidency. He announced that a PCES Board meeting was held on 15 Dec 1989 at 3:00 and

summarized the meeting as follows. There was a discussion of the Society’s financial situation. Data

will be collected from other organizations on their dues and subscription rates. Dues were raised to

$20.00 for regular members, $30.00 for subscribing members, $40.00 for sponsoring members, $21.00

for family members, $10.00 for student members, and $10.00 for retired members. The expenditure

of money to defray speakers travel costs to AAAS meeting symposium was discussed. The production

and mailing costs of Bits & PCES were also discussed.

Dr. Zavortink called for annual reports from the various committees. Daniel Gross, chair of the

membership committee reported that in 1989 the membership remained fairly stable. Twenty-five

new members were added (19 regular, 5 student and 1 sponsoring). In the past five years this was the

lowest number to apply. There were a total of 448 members in 1989: 352 regular members, 55 student

members, and 41 sponsoring members. Under the chairmanship of H. Vannoy Davis, Helen Court

reported that the audit committee reports that everything is in order. The full report will be published

in the proceedings. Lester Ehler and Ron Stecker asked Zavortink to report that the nominating

committee proposed the following slate of candidates for office in 1990: Robert Dowell as president,

Leslie Saul as president-elect, Sandra Shanks as treasurer, Leslie Saul as recording secretary, and

Vincent Lee as managing secretary.

The gavel was passed to president-elect Dr. Dowell. Dr. Dowell introduced Dr. Tom Zavortink,

of the University of San Francisco, who presented the presidential address. Dr. Zavortink presented

an excellent and informative talk on “The Biology of Mosquitos.” He related that there are 3146

species in the world, 167 species in North America and 47 species in California representing 38 genera.

1991

PROCEEDINGS

157

The classification is conservative. The majority of the species fall into three genera: 379 in Anopheles,

946 in Aedes and 767 in Culex. The morphological differences between the three main genera were

described. The largest mosquito in the world is in the genus Toxerinchides. This beautiful mosquito

is incapable of penetrating the skin and only sucks nectar. The biology of several groups was discussed

in detail focusing on oviposition behavior. Dr. Zavortink closed his lecture with a discussion of the

difficulties with drawing phylogenetic trees for mosquitos considering all stages. Dr. Dowell announced

the upcoming lecture in January “Grape Phylloxera in California: No Sex in the Vineyard” by Dr.

Jeffrey Granett. The meeting was adjourned at 9:45 PM. The following 42 persons were present: (35

members) P. H. Amaud Jr., L. G. Bezark, T. S. Briggs, P. Buickerood, R. Buickerood, J. S. Chinn,

H. K. Court, L. Dong, R. V. Dowell, J. G. Edwards, R. Fall, D. W. Gray, D. F. Gross, J. E. Hafemik

Jr., A. Horn, B. Keh, R. L. Langston, V. F. Lee, P. A. Luft, W. A. Maffei, L. B. Mak, D. L. Mead, S.

S. Mead, N. D. Penny, K. J. Ribardo, R. G. Robertson, L. S. Saul, W. E. Savary, J. T. Sorensen, P.

H. Sullivan, C. Y. Takahashi, D. Ubick, S. C. Williams, B. A. Wilson, and T. J. Zavortink; (7 guests)

M. M. Amaud, J. E. Court, C. D. Franklin, D. Maffei, A. M. L. Penny, W. C. Rauscher, and J. M.

Ribardo.

ANNOUNCEMENT

Beginning with the July, 1990, issue of the Pan-Pacific Entomologist, the Pacific

Coast Entomological Society began sending membership/subscription renewal

notices for the following year with its July (number 3) issue. Please check the July

issue of the journal for these renewal notices before discarding its shipping wrap¬

per. Only a final subscription notice will be sent, if PCES does not receive a

response to the July issue’s enclosure.

158

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(2)

ANNOUNCEMENT

PUBLICATIONS OF THE PACIFIC COAST

ENTOMOLOGICAL SOCIETY

PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.

Vol. 1 (16 numbers, 179 pages) and vol. 2 (9 numbers, 131 pages). 1901-1930.

Price $5.00 per volume.

THE PAN-PACIFIC ENTOMOLOGIST.

Vol. 1 (1924) to vol. 51 (1975), price $10.00 per volume of 4 numbers, or $2.50

per single issue. Vol. 52 (1976) to vol. 57 (1981), price $15.00 per volume, or

$3.75 per single issue, except for vol. 57, no. 1, $10.00. Vol. 58 (1982) and

subsequent issues, $20.00 per volume or $5.00 per single issue.

MEMOIRS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.

Volume 1. The Sucking Lice by G. F. Ferris. 320 pages. Published October

1951. Price $10.00 (plus $1.00 postage and handling).*

Volume 2. A Revision of the Spider Mite Family Tetranychidae by A. Earl

Pritchard and Edward W. Baker. 472 pages. Published July 1955. OUT-OF-

PRINT.

Volume 3. Revisionary Studies in the Nearctic Decticinae by David C. Rentz

and James D. Birchim. 173 pages. Published July 1968. Price $4.00 (plus

$0.75 postage and handling).*

Volume 4. Autobiography of an Entomologist by Robert L. Usinger. 343 pages.

Published August 1972. SPECIAL PRICE $5.00 (plus $ 1.00 tax, postage, and

handling for California orders, $0.70 postage and handling for non-California

U.S. orders, or $ 1.70 for foreign orders). No members discount at this special

price.

Volume 5. Revision of the Millipede Family Andrognathidae in the Nearctic

Region by Michael R. Gardner. 61 pages. Published January 21, 1975. Price

$3.00 (plus $0.75 postage and handling).*

Send orders to:

Pacific Coast Entomological Society, % California Academy of Sciences, Golden

Gate Park, San Francisco, California 94118-9961, U.S.A.

* (Add 6 % sales tax on all California orders (residents of Alameda, Contra Costa, San Francisco,

Santa Clara, and Santa Cruz counties add 6 1 / 2 %). Members of the Society will receive a 20% discount

on the price of the books.)

PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

See volume 66 (1): 1-8, January 1990, for detailed format information and the issues thereafter for examples. Manuscripts must be

in English, but foreign language summaries are permitted. Manuscripts not meeting the format guidelines may be returned. Please

maintain a copy of the article on a word-processor because revisions are usually necessary before acceptance, pending review and copy¬

editing.

Format. — Type manuscripts in a legible serif font IN DOUBLE OR TRIPLE SPACE with 1.5 in margins on one side of 8.5 x 11 in,

nonerasable, high quality paper. THREE (3) COPIES of each manuscript must be submitted, EACH INCLUDING REDUCTIONS

OF ANY FIGURES TO THE 8.5 x 11 IN PAGE. Number pages as: title page (page 1), abstract and key words page (page 2), text

pages (pages 3 + ), acknowledgment page, literature cited pages, footnote page, tables, figure caption page; place original figures last.

List the corresponding author’s name, address including ZIP code, and phone number on the title page in the upper right comer. The

title must include the taxon’s designation, where appropriate, as; (Order: Family). The ABSTRACT must not exceed 250 words; use

five to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.

Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases

underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet

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Literature Cited. — Format examples are:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley & Sons, New York.

Blackman, R. L., P. A. Brown & V. F. Eastop. 1987. Problems in pest aphid taxonomy: can chromosomes plus morphometries

provide some answers? pp. 233-238. In Holman, J., J. Pelikan, A. G. F. Dixon & L. Weismann (eds.). Population structure, genetics

and taxonomy of aphids and Thysanoptera. Proc. international symposium held at Smolenice Czechoslovakia, Sept. 9-14, 1985.

SPB Academic Publishing, The Hague, The Netherlands.

Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus. Evolution, 42: 895-899.

Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif. Entomol.

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Volume 67

THE PAN-PACIFIC ENTOMOLOGIST

April 1991

Number 2

Contents

CHAPCO, W. & P. W. Riegert—Maternal effects and egg hatchability in Melanoplus (Orthop-

tera: Acrididae). 81

GOEDEN, R. D. & D. H. HEADRICK—Life history and descriptions of immature stages of

Tephritis baccharis (Coquillett) on Baccharis salicifolia (Ruis & Pavon) Persoon in south¬

ern California (Diptera: Tephritidae). 86

DAANE, K. M., M. S. BARZMAN, C. E. KENNETT & L. E. CALTAGIRONE-Parasitoids

of black scale in California: establishment of Prococcophagus probus Annecke & Myn-

hardt and Coccophagus rusti Compere (Hymenoptera: Aphelinidae) in olive orchards .. 99

BROWN, R. L. & J. A. POWELL—Description of a new species of Epiblema (Lepidoptera:

Tortricidae: Olethreutinae) from coastal redwood forests in California with an analysis

of the forewing pattern . 107

GRISWOLD, T. E. —A review of the genus Microthurge (Hymenoptera: Megachilidae). 115

POLHEMUS, J. T.—A new and primitive genus of Cryphocricinae (Heteroptera: Naucoridae) 119

WELCH, J. L. & B. C. KONDRATIEFF—The Mydidae (Diptera) of Costa Rica. 124

BARRENTINE, C. D. — Post-egestive survival of Sphenophorus phoeniciensis Chittenden (Co-

leoptera: Curculionidae) egested by western toads. 135

BALLING, S. S. & V. H. RESH —Seasonal patterns in a San Francisco bay, California, salt

marsh arthropod community. 138

SCIENTIFIC NOTE

GUERTIN, D. S. — Atypical sex role behavior in the ball-rolling dung beetle, Canthon pilularius

L. (Coleoptera: Scarabaeidae). 145

Pacific Coast Entomological Society, Proceeding for 1988 . 147

Announcement—notification of subscription renewals . 157

Announcement —publications of the Pacific Coast Entomological Society. 158

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 67

July 1991

Number 3

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

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PAN-PACIFIC ENTOMOLOGIST

67(3): 159-170, (1991)

THE ACANTHOSOMATIDAE (HETEROPTERA) OF

NORTH AMERICA

Donald B. Thomas

U.S. Department of Agriculture, Agricultural Research Service,

Screwworm Research Laboratory,

Apartado Postal 544, Tuxtla Gutierrez, Chiapas, Mexico 1

Abstract. — The heteropteran family Acanthosomatidae (Pentatomoidea) is represented in North

America by six species, one new, in two genera. The species are diagnosed and figures of the

male and female genitalia and notes on the biology and distribution are given. Keys for the

identification of genera and species are also provided.

Key Words.— Insecta, Heteroptera, Acanthosomatidae, taxonomy, stinkbugs

Acanthosomatids are stinkbugs (Pentatomoidea) that have two-segmented tarsi

and a large, exposed, eighth abdominal segment in males. In other pentatomoids

the eighth segment is reduced to a narrow ring at the base of the male genital cup

(Schaeffer 1977) and is not visible externally. In most, but not all, genera of

acanthosomatids, the female possesses a pair of abdominal structures known as

Pendergrast’s organs. These organs function during oviposition when the female

transfers a secretion from the gland to the newly laid egg mass using her hind legs

(Pendergrast 1953). Interestingly, in Elasmucha, one of the few genera lacking

Pendergrast’s organs, maternal brooding of the egg clutch and early instar nymphs

is a characteristic behavior (Frost & Haber 1944, Kudo et al. 1989). Although

Pendergrast’s organs are unique to the acanthosomatids, paired external organs

of unknown homology occur on the abdomens of scutellerids (Carayon 1984),

asopine pentatomids (Aldrich 1988), cyrtocorids, plataspids and lestoniids (Miller

1971), although usually on males, not females. In male acanthosomatids incon¬

spicuous pheromone secreting dermal glands are found in patches mainly on

abdominal stemites V and VI (Staddon 1990).

Kirkaldy (1909) considered the Acanthosomini to be a tribe within the nominate

subfamily of Pentatomidae. American workers Van Duzee (1916) and Torre-

Bueno (1939a) placed them at the level of subfamily within the Pentatomidae.

Subsequently, Kumar (1974) accorded the group family status, and most American

workers since then (e.g., Rolston & McDonald 1979, Froeschner 1988) have

treated the group as a full family. Kumar (1974) divided the acanthosomatids

into three subfamilies and provided diagnoses for the world genera. Rolston &

Kumar (1974) provided a key to the genera of the Western Hemisphere.

Acanthosomatids are most abundant in boreal or high latitude temperate regions

or in subtropical regions at high elevation. In North America, the family is rep¬

resented by six species in two genera. Until early in this century, North American

species were placed in the genus Acanthosoma Curtis. Bergroth (1907) transferred

the North American species to the genera Elasmostethus Fieber and Elasmucha

St&l. Elasmostethus was revised by Torre-Bueno (1939b) who resolved the dispute

1 Current Address: USDA-AG/SEA (TUXTLA), P.O. Box 3149, Laredo, Texas 78041.

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

over synonymies between Barber (1932) and Van Duzee (1935). Torre-Bueno

(1939b) recognized three species and provided a key for their separation, but did

not describe the genitalic structures on which definitive diagnoses depend. In his

initial report of Elasmostethus interstinctus (L.) in North America, Barber (1932)

included a figure of the male genitalia. Subsequently, McDonald (1966) described

and figured the genitalia of males and females of Elasmostethus cruciatus and

Elasmucha lateralis. Kumar (1974) described the male genitalia of Elasmostethus

dentatus (DeGeer), a synonym of the Holarctic E. interstinctus (L.). Kumar also

examined the female type of the Mexican Acanthosoma flammatum Distant, 1893,

and removed it to Elasmucha St&l. Until recently, Elasmucha flammatum was

known only from this single female type. However, I have been able to collect

this species in Chiapas, Mexico, and subsequently numerous specimens have been

located. In addition, a closely related but separate species was discovered in the

Sierra Madres of Mexico with specimens occurring as far north as New Mexico

in the United States. I have compared these specimens to Distant’s type of Acan¬

thosoma flammatum in the British Museum (Natural History). I have also been

able to obtain specimens of Elasmostethus interstinctus (L.) from Alaska and have

compared these to European specimens determined as either interstinctus or its

synonym dentatus. I, therefore, have been able to study the genitalic structures

of all five previously described species reported from North America and confirm

their correct identity. In the present article I provide a description and comparison

of the external morphology, including the male and female genitalia, for all six

North American species.

Key to the North American Genera of Acanthosomatidae

1. Scent gland ruga auriculate, short, extending only half-way to metapleural

margin; females lack glandular structures on abdominal segments VI and

VII. Elasmucha

- Scent gland ruga elongate, extending three-quarters of the distance to meta¬

pleural margin; females with glandular structures, visible as darkened,

ovoid depressions on abdominal segments VI and VII .... Elasmostethus

Elasmucha StAl

Clinocoris Hahn 1834:70 [not Fallen 1829]. (Type-species: Cimexferrugatus Fabr.)

Elasmucha St&l 1864:54.

Meadorus Mulsant & Rey 1866:238, synonymized by Kumar (1974). Type-spe¬

cies: Cimex griseus L., designated by Kirkaldy (1909).

Pseudostollia Breddin 1901:65. (Type-species: Acanthosoma delicatula Walker,

by monotypy), synonymized by Kumar (1974).

Galasastra Breddin 1903:205. (Type-species: Galasastra salebrosa Breddin, by

monotypy), synonymized by Kumar (1974).

Type-species. — Cimex ferrugatus Fabr. designated by Kirkaldy (1909).

Diagnosis. —Elasmucha can be separated from other North American genera

of Acanthosomatidae by the following character combinations: the anteclypeus

is slightly longer than paraclypei; the first antennal segment reaches and slightly

surpasses the apex of the paraclypeus; the mesostemal carina are elevated, pro¬

jecting over the prostemum, with the carina received by a groove in the proster-

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THOMAS: NORTH AMERICAN ACANTHOSOMATIDAE

161

num; the margins of the prostemal groove are elevated and carinate; the scent

gland ruga is auriculate and broad, reaching half-way to the metapleural margin;

the base of the abdomen has an anteriorly projecting spine that reaches to the

mesocoxae. Females lack Pendergrast’s organs on abdomen.

Key to North American Species of Elasmucha

1. Anterolateral pronotal margin carinate, weakly reflexed; abdominal ster-

nites with dark spot on each side (mainly Mexico) . 2

- Anterolateral pronotal margin obtuse; abdominal stemites without dark

spot laterally (USA & Canada). lateralis

2(1). Rostrum of moderate length, in repose attaining spine-bearing abdom¬

inal segment (second visible); male paramere with an angular tooth

laterally (Fig. 3) . cordillera

Rostrum long, in repose attaining at least third visible abdominal seg¬

ment; male paramere subcapitate and bent (Fig. 1). flammatum

Elasmucha lateralis (Say)

(Figs. 2, 7)

Edessa lateralis Say 1831:3.

Edessa nebulosa Kirby 1837:277, synonymized by Uhler (1878).

Acanthosoma affinis Westwood 1837:30, synonymized by Distant (1900).

Acanthosoma nebulosum: Dallas 1851:307.

Acanthosoma later ale: Distant 1881:101.

Meadorus lateralis: Van Duzee 1904:73.

Elasmucha lateralis : Bergroth 1907:49.

Clinocoris lateralis: Van Duzee 1908:109.

Types. — Not examined. Say collection largely destroyed (Beidleman 1986).

Redescription.—Male. Dorsal color yellow with tan or brown mottling; connexivum alternately

yellow medially and red-brown near sutures; legs immaculate. Head, pronotum and scutellum with

coarse dark brown punctures; punctures on hemelytral corium smaller. Pleura densely brown punctate;

abdominal venter with few punctures. Length 6-9 mm. Antennal segment I thickest and longest,

segment II almost as long as I, V shortest, III and IV subequal. Rostrum attains second or third visible

abdominal stemite in repose. Mesostemum with elevated carina projecting over prostemum. Base of

abdomen with anteriorly projecting spine which projects over metastemum reaching mesocoxae, its

apex acute and lying aside mesostemal carina. Pendergrast’s organs absent. Dorsoposterior margin of

proctiger with reflexed carina bearing erect spine mesially. Pygophore with fringe of setae on ventro-

posterior margin (Fig. 2); on each side, lateral margin at middle with sclerotized cusp; dorsoposterior

margin deeply emarginate, emargination flanked on each side by setose lobes. Parameres elongate,

undulate.

Female. — Basal gonocoxites longer than wide (Fig. 7). Eighth paratergite with deep, U-shaped mesial

emargination; spiracles present.

Diagnosis. — See the key to species of Elasmucha.

Biology.— The primary hosts are birch and beech trees (Jones & McPherson

1980). A good account of the biology is given by McPherson (1982). The brooding

behavior was studied by Frost & Haber (1944). I have collected E. lateralis in

numbers on beech on the margins of lakes in Ontario and Manitoba.

Distribution.— Maine to British Columbia, south to Tennessee in the east, to

Nevada in the west. Froeschner & Halpin (1981) report it from Alaska. I have

seen one specimen from Dallas, Texas, which I consider probably mislabeled.

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Figures 1-6. Male genital capsule, caudal view. Figure 1. Elasmucha flammatum. Figure 2. Elas-

mucha lateralis. Figure 3. Elasmucha cordillera. Figure 4. Elasmostethus interstinctus. Figure 5. Elas-

mostethus cruciatus. Figure 6. Elasmostethus atricornis.

Material Examined.— USA. CALIFORNIA. MARIPOSA Co.: Yosemite Natl Park, 4 Sep 1974,

A.J. Gilbert, ex Soldidago, 1 female. MICHIGAN. IRON Co.: 29 Jul 1975, C. Buskirk, 1 female.

MARQUETTE Co.: Champion, 14 Jun 1977, D. Flynn & J. Mahar, 1 male. MINNESOTA. BECKER

Co.: Itasca St Park, 28 Jun 1984, N. Downie & J. Wappes, 1 male. CROW WING Co.: 4 Jul 1983,

D.L. Caldwell, 1 female. ITASCA Co.: Deer Lake, 15-21 Jun 1986, D.A. Rider, 5 males, 3 females.

NEW JERSEY. MONMOUTH Co.: Lincroft, 30 Jul 1979, A. Hook, 1 female. NEW YORK. ESSEX

Co.: Whiteface Mtn, 16 Jul 1990, J. Huether, 2 males, 3 females. OREGON. LINCOLN Co.: Agate

Beach, 15 Jun 1976, R.L. Westcott, in beach wash, 1 female. PENNSYLVANIA. ALLEGHENY Co.:

Pittsburgh, 14 Jun 1982, D. Colterrah, on birch, 2 males, 1 female. TEXAS. DALLAS Co.: Dallas, 5

Apr 1981, J. Wellso, 1 male.VIRGINIA. WASHINGTON Co.: Clinch Mtn, Hidden Valley, 24 May

1990, B. KondratiefF, J.L. Welch, R.F. Kirchner, 1 male. WASHINGTON. KING Co.: Hwy 1-90 &

S Fork of Snoqualmie River, 15 May 1988, M.E. Rice, 1 female. CANADA. BRITISH COLUMBIA:

Lions Bay, 7 Jul 1988, D.B. Thomas, 3 males, 6 females; Spahats Creek Prov Park, Spahats Creek,

17 Aug 1986, P.H. Amaud, 2 males, 2 females. ONTARIO: Quetoco Prov Park, 8-9 Aug 1978, D.

Legg, 2 males, 1 female.

Elasmucha flammatum (Distant)

(Figs. 1, 8)

Acanthosoma flammatum Distant 1893:458.

Elasmucha flammatum : Kumar 1974:49.

Types.—Holotype, female, data: MEXICO. GUERRERO : Omilteme, July, H.H.

Smith; deposited in the British Museum (Natural History), London.

Redescription. —Male. Dorsal color yellow-tan with dense red-brown punctation. Connexivum al¬

ternately yellow mesially, dark brown near sutures. Pleura punctate; legs immaculate. Length 7-11

mm. Antennal segment II longest, I shortest, III, IV and V subequal. Rostrum long, attaining third

or fourth visible abdominal stemite in repose. Anterolateral pronotal margin weakly reflexed, carinate.

Abdominal stemites with shagreened, dark spot on each side just mesad of trichobothria. Pendergrast’s

organs absent. Proctiger with low, carinate fold mesially on dorsal surface. Ventroposterior lip of

pygophore bearded with double row of setae: ental row short and dense, ectal row long, paler, both

rows interrupted mesially (Fig. 1). Lateral margin of pygophore with sclerotized cusp on each side at

middle. Dorsoposterior margin of pygophore not deeply emarginate. Parameres asymetrically clavate.

Female. Similar to male except distinctly larger and paler. Basal gonocoxites about equally long as

wide (Fig. 8). Eighth paratergite with deep U-shaped emargination mesially; spiracles present.

Diagnosis. — See the key to the species of Elasmucha.

1991

THOMAS: NORTH AMERICAN ACANTHOSOMATIDAE

163

Figures 7-12. Female genital plates, ventral view. Figure 7. Elasmucha lateralis. Figure 8. Elas-

mucha flammatum. Figure 9. Elasmostethus atricornis. Figure 10. Elasmostethus cruciatus. Figure

11. Elasmucha cordillera. Figure 12. Elasmostethus interstinctus.

Biology. — I collected two specimens at light at two different localities in Chiapas.

One of these locations, El Chorreadero, is a tropical deciduous forest; the other,

near Cuxtepeques, is a pine-oak forest. Both collection localities were montane

riparian habitats.

Distribution. —Mexico and Guatemala.

Material Examined. —GUATEMALA. Bananera, Apr 1930, JJ. White, 1 male. MEXICO. CHIA¬

PAS'. Cuxtepeques, 15 Jun 1987, D.B. Thomas & A.M. Mendoza, 1 male; Chorreadero Cyn, 9 km E

of Chiapa de Corzo, 18 Sep 1985, D.B. Thomas, B.C. RatclifFe & C. Messenger, 1 female; Chorreadero,

26 May 1987, D.A. Rider, D.B. Thomas, E.G. & T.J. Riley, 1 female; Municipio Pantepec, road from

Rayon between Pantepec and Tapalapa, 1707 m, 22 Sep 1981, D.E. Breedlove, 1 female. NUEVO

LEON\ 29 km W of Linares, 23 Jul 1976, Peigler, Gruetzmacher, Murray & SchafFner, 3 males, 4

females; Monterrey, 4 Nov 1978, L.M. Koenig, 1 male. VERACRUZ : Fortin de las Flores, Planta de

la Cerveceria Moctezuma, 17-18 May 1965, D. Rabago, 1 male, 1 female.

Elamucha cordillera Thomas, NEW SPECIES

(Figs. 3, 11, 13)

Types.— Holotype, male, data: NEW MEXICO. CATRON Co.: Whitewater

Forest Camp, 9 km (5 mi) NE of Glenwood, 21 Aug 52, H.B. Leech & J.W.

Green; deposited, California Academy of Sciences, San Francisco. Paratypes: 6

males and 6 females. One male, data: NEW MEXICO. SANTA FE Co.: 10 Jul

1940, D.J. & J.N. Knull; deposited in the personal collection of D.B. Thomas.

One male, data: MEXICO. OAXACA: 10 km N of Oaxaca, 1900 m, 13 Sep 1986,

R. Baranowski, sifting litter at small stream, trop. mont. forest; deposited, private

collection of D.A. Rider. Two males, 5 females, data: MEXICO. “2390,” C.F.

Baker; deposited, United States National Museum of Natural History. One male,

data: MEXICO. San Antonio, “No. 114,” 1 May 1945, with leaf mold; deposited,

United States National Museum of Natural History. One female, data: MEXICO.

MEXICO D.F.: L. Conradt; deposited, United States National Museum of Natural

History. One male, data: GUATEMALA. Bananera, Apr 1930, J.J. White; de¬

posited, United States National Museum of Natural History.

Description. —Male. Elongate-oval (Fig. 13), depressed dorsally, convex ventrally. Color yellow with

dense, irregularly dispersed, red-brown punctations. Length from tip of tylus to apex of abdomen: 6.5

mm. Head', elongate; tylus gradually widened apically and slightly surpassing juga. Antennal segments

variably black or darkened; segment I shortest, II longest, III, IV and V subequal. Rostrum in repose

attaining base of abdominal stemite III (second visible). Thorax : anterolateral pronotal margin with

obtuse reflexure; weakly concave in dorsal view. Humeri obtuse, not produced. Corium with lateral

164

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Figure 13. Elasmucha cordillera, dorsal habitus.

1991

THOMAS: NORTH AMERICAN ACANTHOSOMATIDAE

165

punctures larger and denser. Scent gland auricle reaching almost one-half distance to metapleural

margin. Pleura strongly punctate. Legs yellow, tibia terete, except protibiae prismatic distally. Abdo¬

men: venter yellow, impunctate; each stemite with round dark spot on each side just mesad of spiracle.

Connexivum appearing alternate with apex of each segment having a broad nearly square dark spot.

Genitalia: ventral margin of pygophore bearded with thick fringe of setae each side of middle; lateral

margin without tooth or spinous process (Fig. 3). Proctiger with small triangular tooth at middle on

posterior margin. Parameres thick, compressed; ectal margin with small acute angular tooth-like

projection (Fig. 3).

Female.— Length 8.0 mm. Basal gonocoxites with strong longitudinal wrinkles near and parallel to

mesial margin. Posterior margin of ninth tergite with U-shaped mesial emargination; spiracles present

(Fig. 11).

Diagnosis. —Definitive separation of this species from the others in the genus

requires examination of the male genitalia. The male paramere has an angular

tooth on the lateral margin that is lacking in the others.

Remarks. — The last mentioned paratype, from Bananera, Guatemala, may be

mislabeled because it bears a collection label that is identical with a specimen of

flammatum. Because both specimens are males, they are easily differentiated.

Many of the paratypes bear no other locality information than “Mex.” The range

of this species is, therefore, open to question. Reliably labeled males are from

New Mexico, in the United States, and Oaxaca, in Mexico.

Material Examined.—see types.

Elasmostethus Fieber

Elasmostethus Fieber 1860:78.

Cypostethus Fieber 1860:78. (Type-species, Cimex tristriatus Fabr. by monotypy),

synonymized by Kumar (1974).

Oxydalus Mulsant & Rey 1866:247. (Type-species: Cimex dentatus DeGeer, by

monotypy), synonymized by St&l (1868).

Elasmatostethus Marshall 1868:281. [invalid emendation].

Stictocarenus St&l 1871:638. (Type-species: Rhynchoris ligatus Erichson, by

monotypy), synonymized by Kumar (1974).

Dichobothrium Breddin 1903:207. (Type-species, Dichobothrium sastragaloides

Breddin), designated by Kirkaldy (1909), synonymized by Kumar (1974).

Ditaenius Bergroth 1912:361. (Type-species, Cimex emeritus Bergroth, by mono-

typy), synonymized by Kumar (1974).

Type-species.—Cimex dentatus DeGeer [=Cimex interstinctus L.], by mono¬

typy.

Diagnosis. — The following characters and states separate Elasmostethus from

other New World genera of Acanthosomatidae: The anteclypeus is longer than

paraclypei; the apex of antennal segment I clearly surpasses the anteclypeus; the

mesostemum has an elevated carina projecting over the prostemum which is

received by a groove in prostemum; the margins of the prostemal groove are

elevated and subcarinate; the base of abdomen is produced forward as an elongate

spine lying along side of the mesostemal carina and projects over the metastemum

to the mesocoxae; the scent gland ruga are straight and elongate, about five times

longer than wide, and reaching three-quarters of the distance to metapleural mar¬

gin. Pendergrast’s organs are present on the abdomen of females. These are glan¬

dular structures appearing as small, darkened, ovoid depressions on each side of

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the abdomen, near the posterior border of stemite VI and the anterior border of

stemite VII.

Key to North American Species of Elasmostethus

1. Pronotal disc and head with many dark punctures; first antennal segment

pale. 2

Pronotal disc and head without dark punctures; first antennal segment

dark . atricornis

2(1). Abdominal venter with row of dark spots on each side . interstinctus

- Abdominal venter immaculate. cruciatus

Elasmostethus interstinctus (L.)

(Figs. 4, 12)

Cimex interstinctus L. 1758:445.

Cimex dentatus DeGeer 1773:260. Synonymized by Kirkaldy (1909).

Cimex haemagaster Schranck 1781:270. Synonymized by Kirkaldy (1909).

Cimex bidens Gmelin 1789:2149 [not Linnaeus 1758]. Synonymized by Kirkaldy

(1909).

Cimex arboreus Gmelin 1789:2160. Synonymized by Kirkaldy (1909).

Cimex collaris Fabricius 1803:170 [homonym]. Synonymized by St&l (1868).

Pentatoma liturata Latreille 1804:192. Synonymized by Kirkaldy (1909).

Pentatoma stolli Le Pelletier & Serville 1825:53. Synonymized by Kirkaldy (1909).

Acanthosoma haematogaster (sic): Burmeister 1835:360.

Meadorus interstinctus Mulsant & Rey 1866:238.

Acanthosoma dentatum : Saunders 1892:39. Synonymized by Horvath (1899).

Elasmostethus interstinctus : Horvath 1899:366.

Types. — Cimex dentatus DeGeer [= Cimex interstinctus L.] designated by Stcil

(1864). Not examined.

Redescription. —Male. Dorsal color green-yellow with variable red tinge. Connexivum uniformly

yellow. Dense black punctations on head, pronotum, scutellum and corium. Venter yellow, impunctate.

Legs immaculate. Abdominal stemites with shagreened, dark, oval spot on each side just mesad of

trichobothria. Length 8-10 mm. Antennal segment IV longest, I and III shortest, II and V subequal,

longer than I and III. Rostrum in repose attaining metacoxae. Anterolateral pronotal margin calloused.

Mesosternal carina elevated, projecting over prostemum and received by prostemal groove, margins

of groove elevated, subcarinate. Scent gland ruga elongate, attaining three-fourths distance to meta-

pleural margin. Posterior margin of proctiger arcuate, reflexed. Ventro-posterior margin of pygophore

bearded with fringe of setae, interrupted mesially; margin of pygophore either side of fringe bearing

sharp, sclerotized tooth (Fig. 4). Paramere elongate, compressed, with asymmetrical quadrate head.

Female. — Basal gonocoxites slightly wider than long (Fig. 12). Eighth paratergite undulately emar-

ginate in caudal view; spiracles present.

Diagnosis.— See the key to the species of Elasmostethus.

Biology. —This bug is known as the “birch bug” in England. Kirkaldy (1909)

also lists alders, beech and aspen as hosts. Staddon (1990) has studied the dis¬

tribution of pheromone glands and the chemistry of their secretions.

Distribution. — Holarctic: Alaska, Northwest Territories of Canada, northern

Europe and Asia.

1991

THOMAS: NORTH AMERICAN ACANTHOSOMATIDAE

167

Material Examined. —USA. ALASKA. Delta Junction, Fourth judicial district, Aug 1981, C. Papp,

3 males, 2 females. GREAT BRITAIN. ENGLAND. NORFOLK : Thetford, 16 Sep 1977, A.J. & M.E.

Gilbert, 1 male; Cambridge, Wicken Fen, 16 Sep 1977, A.J. & M.E. Gilbert, 1 female.

Elasmostethus cruciatus (Say)

(Figs. 5, 10)

E dess a cruciata Say 1831:2.

Acanthosoma borealis Westwood 1837:30. Synonymized by Distant (1900).

Acanthosoma cruciata : Uhler 1861:23.

Acanthosoma cruciata var. cooleyi Van Duzee 1904:74.

Elasmostethus cruciatus: Bergroth 1907:49.

Elasmostethus cooleyi : Bergroth 1907:49. Synonymized by Torre-Bueno (1939b).

Types.—Not examined. Say collection reportedly destroyed (Beidleman 1986).

Redescription. —Male. Dorsal color yellow-tan with red band on ental margin of hemelytral corium.

Red-brown punctures dense on pronotum, scutellum and most of corium, sparse on head and large

spot on endocorium. Connexivum uniformly yellow. Venter yellow impunctate, immaculate; apices

of last abdominal stemite red. Legs immaculate. Length 8-11 mm. Antennal segment III shortest, II,

IV and V subequal, longer than I. Rostrum attains metacoxae in repose. Anterolateral pronotal margin

calloused. Posterior margin of proctiger bluntly acuminate, broadly reflexed. Ventroposterior margin

of pygophore bearded with dense fringe of setae, interrupted mesially; margin either side of fringe

with a sharp, sclerotized tooth (Fig. 5). Paramere elongate, compressed, with ovoid head which has

outer, apical margin deflexed.

Female .—Basal gonocoxites about as long as wide, posterior margin arcuate (Fig. 10). Posterior

margin of eighth paratergite broadly V-shaped in ventral view with weak mesial emargination; spiracles

present.

Diagnosis.— See the key to the species of Elasmostethus.

Biology. — Jones & McPherson (1980) report that this species feeds and repro¬

duces on alders. McPherson (1982) provides notes on collection records, distri¬

bution and parasites. I have collected this species in numbers on alders in Nevada

and in British Columbia.

Distribution.— Labrador to British Columbia in Canada, south to California,

Texas and South Carolina.

Material Examined.—USA. CALIFORNIA. SHASTA Co.: Burney, 20 Aug 1969, R.P. Allen, 3

males, 3 females. COLORADO. LARIMER Co.: Cameron Pass, 19 Aug 1948, D.G. Denning, 1

female. NEVADA, DOUGLAS Co.: Carson City, Spooner Ridge, 6 Jul 1978, on Alnus tenuifolia, 3

males, 2 females. NEW YORK. ESSEX Co.: Whiteface Mt, 11 Jul 1990, J. Heuther, 2 males, 3

females. WASHINGTON. THURSTON Co.: Millersylvania St Park, nr Maystown, 53 m, 11 Jul

1988, E.L. Sleeper, 1 male. WYOMING. ALBANY Co.: Laramie, Pole Mtn, 7 Jul 1948, D.G. Denning,

1 male, 1 female. CANADA. BRITISH COLUMBIA: Lions Bay, 7 Jul 1988, D.B. Thomas, 1 male,

1 female; Vancouver, UBC campus, S.E. Woods, 2 Jul 1988, E.L. Sleeper, 1 male, 1 female. MAN¬

ITOBA. Whiteshell Prov Park, South Cross Lake, 13 Aug 1983, D.B. Thomas & J.E. Pasek, 1 female.

Elasmostethus atricornis (Van Duzee)

(Figs. 6, 9)

Acanthosoma atricornis Van Duzee 1904:75.

Elasmostethus atricornis: Kirkaldy 1909:177.

Types.— Not examined.

Redescription.— Male.— Dorsal color yellow with humeral angles of pronotum and antennae black;

scutellum basally and broad ental margin of hemelytral corium infuscated. Venter yellow with apices

168

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

of last abdominal segment orange. Anterior pronotum and head with colorless punctures; base of

pronotum, corium and scutellum with scattered brown punctures. Venter and legs immaculate. Length

8-10 mm. Antennal segments II and IV longest, subequal, III and V shortest, subequal; rostrum

attaining mesocoxae in repose. Anterolateral pronotal margins obtuse. Ventroposterior border of

pygophore bearded with double row of setae, ental row short dense, ectal row longer, interrupted

mesially; margin of pygophore lacking teeth or cusps (Fig. 6). Proctiger broad, obtusely acuminate

distally, apex broadly reflexed. Head of paramere angulate, capitate, feebly bilobate.

Female.— Basal gonocoxites about as long as broad; posterior margin of eighth paratergite sym¬

metrically biarcuate in ventral view; spiracles present (Fig. 9).

Diagnosis.— See the key to the species of Elasmostethus.

Biology.— This species breeds on Aralia racemosa L. (Blatchley 1926, Jones &

McPherson 1980).

Distribution. —Quebec to Montana, south to South Carolina (McPherson 1982).

Material Examined. -USA. NEW YORK. TOMPKINS Co.: Ithaca, 13 Sep 1952, W.D. Stockton,

1 male, 2 females. CANADA. QUEBEC: 20 km SE of Kazabazua, 29 Jul 1989, B.D. Gill, 3 males,

3 females.

Acknowledgment

I thank David A. Rider (Louisiana State University, Baton Rouge), Joseph C.

Schaffner (Texas A&M University, College Station), Richard C. Froeschner (Unit¬

ed States National Museum of Natural History, Washington D.C.) and Paul H.

Amaud (California Academy of Sciences, San Francisco) for the loan of specimens;

and William R. Dolling (British Museum [Natural History], London) for access

to the type of Acanthosoma flammatum and European material of Elasmostethus

interstinctus in his care. I especially thank L. H. Rolston (Louisiana State Uni¬

versity, Baton Rouge) for a critical review of the manuscript. The habitus drawing

of Elasmucha lateralis was rendered by Richard Ashley of Tuxtla Gutierrez,

Chiapas.

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Received 26 November 1990; accepted 16 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 171-176, (1991)

NEST PROVISION AND POLLEN FORAGING IN THREE

MEXICAN SPECIES OF SOLITARY BEES

(HYMENOPTERA: APOIDEA)

Stephen H. Bullock, 1 Ricardo Ayala, 1

Gregorio Rodriguez-GonzAlez, 1 Rodolfo Palacios-ChAvez, 2

Belfina Ramos-Zamora, 2 D. Leonor Quiroz-Garcia 2 and

Maria de la Luz Arreguin-SAnchez 2

^stacion de Biologia Chamela, Universidad Nacional Autonoma de Mexico,

Apartado Postal 21, 48980 San Patricio, Jalisco, Mexico;

2 Laboratorio de Palinologia, Departamento de Botanica,

Escuela Nacional de Ciencias Biologicas, Instituto Politecnico Nacional,

Prol. Carpio y Plan de Ayala, 11340 Mexico, D.F., Mexico

Abstract. — Pollen selection was determined by two methods for three ground-nesting bees: anal¬

ysis of nest cell contents (provisions or feces), and of female pollen foraging records. Ancyloscelis

wheeleri Cockerell and Melitoma marginella Cresson (Anthophoridae) were oligolectic on dif¬

ferent genera of Convolvulaceae.The two methods differed in emphasis of particular hosts.

Mesoxaea nigerrima Friese (Oxaeidae) had unrelated hosts in different months. As many as nine

pollen species were found in a single cell, but the dominant pollen comprised at least 95% in

most cells for all three bee species. Not all minor pollen species were clearly contaminants.

Resumen.— La selection de polen fue determinada por dos metodos para abejas que anidan en

el suelo: el analisis del contenido de las celdas de los nidos (provision o excremento), y la

observation de hembras en pecoreo de polen. Ancyloscelis wheeleri Cockerell y Melitoma mar¬

ginella Cresson (Anthophoridae), fueron oligolecticos sobre diferentes generos de Convolvula-

ceae. Los dos metodos enfatizaron diferentes hospederos. Mesoxaea nigerrima Friese (Oxaeidae)

tenia plantas hospederas no emparentadas en los diferentes meses. Se encontraron hasta nueve

especies de polen por celda, pero el polen dominante comprendio al menos un 95% en la mayoria

de las celdas para las tres especies de abejas. No todas las minoritarias parecieron ser conta-

minantes.

Key Words. — Insecta, feeding specialization, tropical deciduous forest, Convolvulaceae, Legu-

minosae, Tiliaceae

Pollen is a major nutritional source for most larval bees. The eusocial Apidae

in general are highly polylectic (Kleinert-Giovannini & Imperatriz-Fonseca 1987,

Lobreau-Callen et al. 1986) and their selection of pollen may be greatly affected

by its abundance (Roubik et al. 1986). However, pollen from various plant species

may vary demonstrably in nutritional value, as shown for Apis mellifera L. (Her¬

bert et al. 1970) and for the solitary bee Osmia lignaria Say (Levin & Haydack

1957) .The vast majority of bee species are solitary (Linsley 1958), and their host

selection evolves quite distinctly from that of eusocial bees (Lobreau-Callen &

Coutin 1987). In solitary species the repetition of particular host relations across

isolated generations may represent a physiological link. For example, Colletes

fulgidus longiplumosus Stephen females show a preference for the surface oils of

the pollen on which they were reared (Dobson 1987).

The diversity of hosts varies greatly between species of solitary bees (Linsley

1958) , and to a lesser extent among their populations (Linsley et al. 1963, Tasei

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

1976), among individuals (Michener & Rettermeyer 1956), and during or between

seasons (Rust 1990). These variations are particularly difficult to quantify in

habitats with low visibility of nesting areas, flight patterns and hosts. Hosts for

pollen are usually described from observation of foraging females, and not from

direct quantitative assessments of pollen types in the nest cells; the latter is an

important tool for studying feeding specialization (Cripps & Rust 1989). In this

paper, we apply both of these approaches to evaluate the pollen selectivity of

populations of three solitary species from a single locality in neotropical deciduous

forest.

Materials and Methods

The site of this study was the Estacion de Biologia Chamela (19°30' N, 105°03'

W) in the Pacific lowlands (10-150 m) of Mexico, where the rainy season is

typically four months long (Bullock 1986) and the mean annual rainfall is 714

mm (1977-1990). The vegetation at this site is principally tropical deciduous

forest, with a floristically distinct semideciduous forest along the larger seasonal

watercourses (Lott et al. 1987). General collection of bees on flowers has been

made sporadically since 1980 by resident and visiting scientists (principally SHB

and RA). Tree phenology was reported by Bullock & Solis (1990); other plants

were observed while collecting the flora and fauna.

The bees studied were Ancyloscelis wheeled Cockerell and Melitoma marginella

Cresson (Anthophoridae), and Mesoxaea nigerrima Friese (Oxaeidae). Closed nest

cells of each species were collected in November 1986. Ancyloscelis and Melitoma

were nesting in dense aggregations, so the individual cells collected may have

been from various nests. The Mesoxaea nests were solitary. Ancyloscelis and

Mesoxaea nests were active but Melitoma nests were already closed; the respective

samples were 17 cells, five cells from two nests, and 11 cells. In October 1988,

15 cells were collected from closed nests of Melitoma and one from a closed nest

of Mesoxaea.

The pollen samples were prepared by standard acetolysis methods. Percentage

composition was determined from counts of 1000 grains from three mounts

(Ramalho & Kleinert-Giovannini 1986). Volumetric measurements are not a clear

solution to the related problem of nutritional values.

Results

Ancyloscelis wheeled. — Pollen of five plant species was found in nest provisions,

including three species of Convolvulaceae. Merremia quinquefolia (L.) Hallier f.

contributed 99.67%, and there were small amounts of an Ipomoea sp. (0.28%)

and of Jacquemontia nodiflora (Desrousseaux) G. Don (0.05%). The other two

pollen types were Heliocarpuspallidus Rose (Tiliaceae), 0.09%, and an unidentified

Compositae, 0.02%. The latter two may have come from exploratory or nectar

foraging, or may have been introduced as contaminants by the bees because these

forms are commonly airborne (RPC & SHB, unpublished data).

The nesting and principal flight period is October and November, but we have

found occasional individuals as late as April. Pollen-foraging females were col¬

lected on only three plant species: M. quinquefolia, J. nodiflora and Ipomoea

crinicalyx Moore. Nectar-foraging individuals of one or both sexes were collected

on five Convolvulaceae, one Acanthaceae and one Boraginaceae species.

1991

BULLOCK ET AL.: POLLEN FORAGING IN BEES

173

Melitoma marginella. —The 1986 nest samples all showed only one form of

pollen, most closely resembling reference material of Ipomoea pedicellaris Ben-

tham.

In the 1988 sample, three species of Ipomoea were predominant: I. pedicellaris,

I. clavata (G. Don) Oostroom and I. wolcottiana Rose. The most abundant species

comprised 99% to 100% in eight cells (monospecific in Ipomoea), 87% to 97% in

four cells, and 53.6% to 70.5% in three cells. As many as three Ipomoea species

were present in only three cells. Ipomoea nil (L.) occurred only once (2. l%).Non-

Ipomoea constituents in four cells were probably all contaminants introduced by

the bees, including Poeppigia procera Presl (Leguminosae), an unidentified Po-

aceae, and two species not present in the area of the Chamela station, Alnus sp.

(Betulaceae) and Pteris sp. (Adiantaceae), which are distributed by wind. In all

the samples of fecal material, the pollen grains were broken into many pieces.

The flight season was September and October, and again February and March;

the life cycle remains problematic (GRG, unpublished data). Pollen-foraging fe¬

males were collected on Ipomoea muricata Cavanillas, I. nil and I. pedicellaris.

Nectar-foraging males were collected on I. muricata, I. trifida (HBK) G. Don,

and on I. wolcottiana (Bullock et al. 1987).

Mesoxaea nigerrima.—A single pollen type dominated the November 1986

nest samples: Heliocarpus pallidus comprised 99.9%, although trace amounts of

Phaseolus sp. and Apoplanesia paniculata Presl (both Leguminosae) appeared also.

The 1988 sample was collected before Heliocarpus began to flower: the contents

were dominated by Schrankia diffusa Rose (Leguminosae, 95.6%), which is a

common vine and the most frequently observed host of foraging Mesoxaea fe¬

males. Eight other species were present in this single cell provision: Poeppigia

procera (1%), Cassia sp. (0.8%), and Senna sp. (0.8%) (all Leguminosae), Doyerea

emetocathartica Grosourdy (Cucurbitaceae, 0.7%), Serjania brachycarpa A. Gray

(Sapindaceae, 0.6%) and trace amounts of Psidium sartorianum (Berg) Niedenzu

(Myrtaceae, 0.2%), Cleome sp. (Capparidaceae, 0.1%), and Pedilanthus calcaratus

Schlechtendal (Euphorbiaceae, 0.1%).

The flight season extended from July through November. Mesoxaea was col¬

lected foraging on 11 species from four families (Leguminosae, Malpighiaceae,

Sapindaceae and Tiliaceae). Females were observed collecting pollen from four

mimosoid and one caesalpinoid legume species {Mimosa brandegei B. L. Rob¬

inson, M. leptocarpa Rose, Piptadenia obliqua (Persoon) MacBride, Schrankia

diffusa, and Apoplanesia paniculata ) and from Heliocarpus pallidus.

Discussion

Hurd (1979: 2118, 2123) stated that all species of Ancyloscelis and Melitoma

visit flowers of Ipomoea for pollen and nectar, but also visit a variety of other

flowers for nectar. Our results confirm the Melitoma-Ipomoea association and

extend its generality in terms of species, habitat and geography. Ancyloscelis whee-

leri, however, is a specialist on Convolvulaceae but not on Ipomoea at Chamela,

as far as our samples are representative. Both fit the restricted definition of oli-

golecty—“species that confine their pollen foraging to species within a genus”—

suggested by detailed comparative study of provisions in seven Osmia species of

a montane meadow (Cripps & Rust 1989: 136).

Intermingled nests ofM. marginella and A wheeled (Hurd 1979) have not been

174

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

observed at Chamela, although large aggregations of each are known. Our finding

that virtually all grains in the feces of M. marginella are highly fragmented is

apparently unique.

Both these anthophorids have relatively brief flight seasons which may not

exceed the flowering seasons of single host species. Yet their host specificity is

rather remarkable because of the range of closely related plants available simul¬

taneously. October and November are the months of peak flowering of Convol-

vulaceae at Chamela, typically including six genera and 23 species, with 16 species

of Ipomoea (as currently revised).

Pollen hosts of oxaeids are reported from Solanaceae, Zygophyllaceae and Le-

guminosae, especially Cassia (Hurd & Linsley 1976, Camargo et al. 1984). Flowers

in the former two families are common at Chamela during the flight period of

Mesoxaea nigerrima, but there are no foraging or provision records. Foraging has

not been observed on Cassia or Senna but these were minor species in one cell

provision. The rarity of visits to buzz-pollinated plants at Chamela is conspicuous;

the common pollen hosts were mimosoid legumes. Typical of Oxaeidae, M. ni¬

gerrima is a polylectic species, and several plant families may be present in any

cell; however, one species is heavily dominant.

Mesoxaea change principal host within an individual female’s (?) flight season.

Schrankia flowers for several months starting about the same time as the flight

season of Mesoxaea , but Heliocarpus has only a brief abundant flowering in

October or November. The flower visitation records and our observations on

other plant species clearly show that Mesoxaea nigerrima discriminates and is

not simply dependent on flower-frequency in its host selection.

Little is known about the means of pollen host recognition in newly emerged

solitary bees. Experiments with Colletes fulgidus showed that bees can recognize

their host plant by the odor of its pollen, suggesting that larvae may be conditioned

to volatile oils of pollen in their provision (Dobson 1987). This hypothesis might

be appropriate in M. marginella and A. wheeleri. However, our data suggest that

some M. nigerrima individuals begin collecting pollen on plants that are unrelated

to their larval host.

The actual host range in a bee population at any one time may be significantly

greater than we could sample. Certainly the data indicate that generally only one

host species is used over several to many foraging trips, over periods of days, and

among neighboring females in Ancyloscelis . Any one larva of any of these species

is unlikely to experience more than a one percent admixture of a second pollen

type in its provision, and the combined percentage of secondary species exceeded

five percent only in some cells of M. marginella.

Both flower visitation records and palynological analysis of nest contents have

limitations in terms of optimum sampling—particularly in forests and for ground¬

nesting bees. In the present case the contribution of pollen analysis is in the

quantification of the purity, or lack of it, in the provisions for individual bees.

Observation of foraging females had not missed any major hosts. But in a forest

habitat, following an individual bee throughout a single foraging trip, or during

several trips or days, is virtually impossible.

The presence of pollen species other than the principal hosts may be attributable

to several causes. Of considerable importance are the opportunities for contam¬

ination of the bee’s pollen load. Foreign grains may be added to flowers of a given

1991

BULLOCK ET AL.: POLLEN FORAGING IN BEES

175

species through the visits of more polylectic insects, somewhat analogous to intra¬

hive inter-bee transfer of pollen (DeGrandi-Hoffman et al. 1986). Also, airborne

pollen is ubiquitous and remarkably diverse (Palacios 1985; RPC & SHB, un¬

published data) which will also contact the bees via soil, leaves, etc. However,

the most important causes are probably the admixture of pollen from plants visited

for nectar, and from exploratory visits by bees to potential alternate resources

(Heinrich 1979, Cripps & Rust 1989).

Acknowledgment

We are grateful to H. E. M. Dobson, F. D. Parker, and anonymous reviewers

for comments on the manuscript.The ENCB-IPN group was supported in part by

COFAA and by the Direccion de Estudios de Posgrado e Investigation (grant

870285).

Literature Cited

Bullock, S. H. 1986. Climate of Chamela, Jalisco, and trends in the south coastal region of Mexico.

Arch. Met. Geoph. Biocl. Ser. B, 36: 297-316.

Bullock, S. H., R. Ayala, I. Baker & H. G. Baker. 1987. Reproductive biology of the tree Ipomoea

wolcottiana (Convolvulaceae). Madrono, 34: 304-314.

Bullock, S. H. & J. A. Solis-Magallanes. 1990. Phenology of canopy trees of a tropical deciduous

forest in Mexico. Biotropica, 22: 22-35.

Camargo, J. M. F., G. Gottsberger & I. Silberbauer-Gottsberger. On the phenology and flower visiting

behavior of Oxaea flavescens (Klug) (Oxaeinae, Andrenidae, Hymenoptera) in Sao Paulo, Brazil.

Beitr. Biol. Pflanzen, 59: 159-179.

Cripps, C. & R. W. Rust. 1989. Pollen preferences of seven Osmia species (Hymenoptera: Mega-

chilidae). Environ. Entomol., 18: 133-138.

DeGrandi-Hoffman, G., R. Hoopingamer & K. Baker. 1986. Influence of honey bee (Hymenoptera:

Apidae) in-hive pollen transfer on cross-pollination and fruit set in apple. Environ. Entomol.,

15: 723-725.

Dobson, H. E. M. 1987. Role of flower and pollen aromas in host-plant recognition by solitary bees.

Oecologia, 72: 618-623.

Heinrich, B. 1979. “Majoring” and “minoring” by foraging bumblebees, Bombus vagans: an ex¬

perimental analysis. Ecology, 60: 245-255.

Herbert, E. W., W. E. Bickley & H. Shimanuki. 1970. The brood-rearing capability of caged bees

fed dandelion and mixed pollen diets. J. Econ. Entomol., 63: 215-218.

Hurd, P. D., Jr. 1979. Superfamily Apoidea. pp. 1741-2209. In Krombein, K. V., P. D. Hurd, Jr.,

D. R. Smith & B. D. Burks (eds.). Catalog of the Hymenoptera in America north of Mexico.

Smithsonian Institution Press, Washington, D.C.

Hurd, P. D., Jr. & E. G. Linsley. 1976. The bee family Oxaeidae with a revision of the North

American species (Hymenoptera: Apoidea). Smiths. Contrib. Zool., 220: 1-75.

Kleinert-Giovannini, A. & V. L. Imperatriz-Fonseca. 1987. Aspects of the trophic niche of Melipona

marginata marginata Lepeletier (Apidae, Meliponinae). Apidologie, 18: 69-100.

Levin, M. D. & M. H. Haydack. 1957. Comparative value of different pollens in the nutrition of

Osmia lignaria. Bee World, 38: 221-226.

Linsley, E. G. 1958. The ecology of solitary bees. Hilgardia, 27: 543-599.

Linsley, E. G., J. W. MacSwain & P. H. Raven. 1963. Comparative behavior of bees and Onagraceae.

II. Oenothera bees of the Great Basin. Univ. Calif. Publ. Entomol., 33: 25-50.

Lobreau-Callen, D. & R. Coutin. 1987. Ressources florales exploitees par quelques Apoiodes des

zones cultivees en savane arboree senegalaise durant la saison des pluies. Agronomie, 7: 231-

246.

Lobreau-Callen, D., R. Darchen & A. Le Thomas. 1986. Apport de la palynologie a la connaissance

des relations abeilles/plantes en savanes arborees du Togo et du Benin. Apidologie, 17: 279-

306.

Lott, E. J., S. H. Bullock & J. A. Solis-Magallanes. 1987. Floristic diversity and structure of upland

and arroyo forests in coastal Jalisco. Biotropica, 19: 228-235.

176

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Vol. 67(3)

Michener, C. D. & C. W. Rettenmeyer. 1956. The ethology of Andrena erythronii with comparative

data on other species. Univ. Kansas Sci. Bull., 37: 645-684.

Palacios-Chavez, R. 1985. Liuvia de polen modemo en el bosque tropical caducifolio de la Estacion

de Biologia Chamela, Jal. (Mexico). An. Esc. Nac. Ciencias Biol., Mex., 29: 43-55.

Ramalho, M. & A. Kleinert-Giovannini. 1986. Some aspects of the utilization of pollen analysis in

ecological research. Apidologie, 17: 159-174.

Roubik, D. W., J. E. Moreno, C. Vergara & D. Wittmann. 1986. Sporadic food competition with

the African honey bee: projected impact on neotropical social bees. J. Trop. Ecol., 2: 97-111.

Rust, R. W. 1990. Spatial and temporal heterogeneity of pollen foraging in Osmia lignaria propinqua

(Hymenoptera: Megachilidae). Environ. Entomol., 19: 332-338.

Tasei, J. N. 1976. Recolte des pollens et approvisionnement du nid chez Osmia coerulescens L.

Apidologie, 7: 277-300.

Received 20 September 1990; accepted 15 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 177-180, (1991)

FORAGING OF STENOPOGON INQUINATUS (LOEW)

(DIPTERA: ASILIDAE) ON MONTANA RANGELAND SITES

Kevin M. O’Neill 1 and William P. Kemp 12

Entomology Research Laboratory, Montana State University,

Bozeman, Montana 59717

2 USDA, Agricultural Research Service, Rangeland Insect Laboratory,

Bozeman, Montana 59717

Abstract.— The robber fly Stenopogon inquinatus Loew preyed upon a wide variety of insects at

rangeland sites in Montana. As is typical of Stenopogon, prey use varied with local insect

abundance. Stenopogon inquinatus tended to take alate male Formica subpolita Mayr near ant

mating swarms, honey bees {Apis mellifera L.) near flowers, and dung beetles (Scarabaeidae) on

open range. These three groups, and other robber flies, comprised 90% of the prey records for

S. inquinatus.

Key Words. — Insecta, Diptera, Asilidae, Stenopogon, foraging, prey

Stenopogon is a wide-ranging genus of robber flies in western North America,

where more than 50 species occur (Wilcox 1971). Several studies of the behavior

and ecology of Stenopogon have been undertaken (Cole 1958, Powell & Stage

1962, Lavigne & Holland 1969, Dennis & Lavigne 1975, O’Neill & Kemp 1990,

O’Neill in press). Foraging Stenopogon make short flights in pursuit of prey from

perches on or near the ground and, as a group, prey on a variety of insects. Local

populations tend to take a relatively narrow range of prey, but differences among

species and populations of Stenopogon may be due to spatial and temporal vari¬

ation in the abundance of potential prey, rather than to variation in prey preference

(Powell & Stage 1962, Dennis & Lavigne 1975).

Stenopogon inquinatus Loew is a large species, 20 to 37 mm in length (Dennis

& Lavigne 1975), that is widely distributed in North America (Wilcox 1971).

Here, we report observations on prey use of S. inquinatus in Montana that sup¬

plement earlier studies from Wyoming (Dennis & Lavigne 1975) and provide

evidence that prey use in this species reflects local insect abundance.

Methods

Stenopogon inquinatus was studied at two sites, 14 km S of Three Forks and 5

km NE of Logan, Gallatin County, Montana, during the summers 1987-1990.

The species was typically present from early June through early August and was

easy to distinguish from other asilid species because of its large size. The sites are

classified within the Stipa comata Trinius & Ruprecht /Bouteloua gracilis (Hum¬

boldt et al.) Lagasca y Segura ex. Steudel habitat type (Kemp et al. 1990). Prey

and observations of predation events were sampled ad libitum. Body lengths of

prey and head widths of robber flies were measured to the nearest 0.1 mm, using

a micrometer accurate to 0.05 mm.

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Results

Foraging behavior for S. inquinatus followed the patterns observed by Dennis

& Lavigne (1975) in Wyoming. Foraging flies perched on the soil surface or on

plants up to 1 m in height and pursued insects flying within about 1 m of the

perch. The 170 prey observed at our site included members of five orders, 16

families, and at least 21 species of insects.

The prey specimens that we collected were identified as (numbers of specimens

are in parentheses): Hemiptera: Coreidae, Chelinidea vittiger Uhler (1); Penta-

tomidae, Chlorochroa sp. (2); Reduviidae, Apiomerus sp. (1); Coleoptera: Scar-

abaeidae, Onthophagus nuchicornis (L.) (1), Aphodius fossor L. (12), Dichelonyx

sp. (1); Cicindellidae, Cicindella punctata Olivier (1), Cicindella purpurea Casey

(1), Meloidae, Epicauta sp. (1); Lepidoptera: Noctuidae (2); Pyralidae (2); Diptera:

Tipulidae (1), Calliphoridae (1); Stratiomyidae (1); Tabanidae (1); Asilidae, Efferia

staminea (Williston) (13), Megaphorus willistoni (Williston) (1), Machimus oc¬

cidentals (Hine) (2), Stenopogon inquinatus (11); Hymenoptera: Formicidae, For¬

mica subpolita Mayr (55 alates), Formica sp. (4 alates); Halictidae (2), Apidae,

Apis mellifera (12). Forty-one other prey were identified in the field, but not

collected: one Efferia staminea, one Machimus occidentals, and 39 Formica

alates.

The 129 prey collected had a mean length of 12.6 mm (SE = 0.50). Although

the S. inquinatus collected at this site varied widely in body size (i.e., from 150

to 320 mg wet mass, O’Neill in press), there was no correlation between robber

fly head width and prey body length for the 83 pairs for which both values were

known (r = 0.06, P = 0.59). However, some constraints from predator size are

evident. There was a significant correlation between predator head width and prey

body length when only the largest prey for each predator size class (i.e., 0.1 mm

increment) was used in the analysis ( r = 0.61, n = 11, P = 0.04). Furthermore,

all of the asilids taken as prey by S. inquinatus were smaller than the predator

(O’Neill in press).

Although S. inquinatus had a relatively broad diet breadth, the type of prey

used reflected local abundance of insects. For example, 93% of the prey (n = 70)

collected on 10 days within 5 m of active mating swarms of the ant Formica

subpolita were winged ants. On eight occasions, we observed S. inquinatus fly

into swarms of F. subpolita on and above Rhus trilobata Nuttall and capture an

ant. We also collected prey before and after Formica swarms on the same 10 days

and on six other days when swarms did not appear (but during weeks when swarms

appeared on other days). In combined records for these nonswarm periods, only

25% of the prey were winged Formica {n = 28, %i = 40.2, P < 0.0001).

By switching to winged Formica subpolita during swarms, S. inquinatus ap¬

peared to have a greater frequency of successful foraging flights. On 4 Jul 1990,

17 of the 20 S. inquinatus observed during ant swarms had prey, all but one of

which were winged Formica. Four days later at the same location, when the ants

were not swarming, only one of 55 S. inquinatus observed had prey (%? = 31.71,

P < 0.0001). By taking Formica, they used smaller prey than at other times. The

mean length of winged Formica prey was 9.1 mm (SE = 0.11, n = 59), although

that of all other prey was 15.5 mm (SE = 0.76, n = 70; Mann-Whitney test, P <

0.0001). There was a limit to which S. inquinatus would switch to locally abundant

insects. They did not prey upon winged Feptothorax sp. (Hymenoptera: Formic-

1991

O’NEILL & KEMP: STENOPOGON FORAGING

179

idae) that swarmed at the same location as the F. subpolita on several days in

1988. This ant, which ranged in length from 2.2 to 4.4 mm {x = 3.1 mm, SE =

0.23, n = 13), was well below the size range (6.6 to 29.5 mm) of prey used by S.

inquinatus.

Other examples of site-specific prey use are evident in the prey records. Most

honey bee {Apis mellifera) prey were collected at patches of leafy spurge {Euphorbia

esula L.) and Snowberry {Symphoricarpos spp.) on which the bees were foraging.

There was an apiary about 1 km E of the study site. The presence of cattle in the

area may account for dung beetles (Scarabaeidae) being commonly taken as prey

on open range.

Discussion

Stenopogon inquinatus, in the Montana population that we observed, used a great

variety of insects as prey. However, we found site-specific prey use near ant swarms

and flowers. Increased predation on alate ants near mating swarms was also

observed in the robber flies Heteropogon wilcoxi (James) (Lavigne & Holland

1969) and Efferia staminea (Williston) (unpublished data). Site-specific predation

by Stenopogon has also been observed in studies conducted near apiaries, where

68% of the prey of S. inquinatus (Dennis & Lavigne 1975: Riverton, Wyoming

prey samples) and 89% prey of -S. rufibarbis Bromley (Cole 1958) were honey

bees. Bromley (1942, 1948) also documents that honey bees are common com¬

ponents of robber fly prey records near flowers and apiaries, but does not compare

these to records in other locations. Powell & Stage (1962) noted a difference in

prey types of S. engelhardti Bromley between dry and moist habitats, with prey

records in the latter being dominated by Tipula coloradensis (Doane) (Diptera:

Tipulidae).

Given such site specificity, it is perhaps surprising that our prey records and

those from Wyoming (Dennis & Lavigne 1975) are similar. Honey bees, winged

ants, scarab beetles, and robber flies comprised 82% of the prey in Wyoming and

90% in Montana, although our value may be somewhat inflated due to our frequent

observations near ant swarms. The Wyoming prey records do not include He-

miptera, but do include insects from two orders, Homoptera and Orthoptera, not

found in our samples. However, these orders comprised just a small proportion

of all prey in both our samples and those of Dennis and Lavigne. The size range

of prey from the Wyoming populations (Dennis & Lavigne 1975: range = 5.0 to

29.5 mm, x = 14.5 mm, n = 127) was also generally similar to that at our site

(range = 6.6 to 29.5 mm, x = 12.5 mm). The correspondence of prey records

between areas is probably a reflection of similar grassland insect communities at

the two sites.

The prey records presented cannot be used to draw strong conclusions regarding

prey preference in an opportunist such as S. inquinatus. Simple lists of prey, even

from a relatively restricted array of habitats, are liable to reflect several unrelated

correlates of prey use. The contents of prey records are not only a function of

local insect abundance, but of variation in visibility and catchability among prey

species, as well as evolved preferences of robber flies for nutritionally superior or

non-toxic insects (Shelly 1984). Our observations and other published reports on

Stenopogon suggest that both local abundance and catchability (as influenced by

size and flight speed) influence prey use. Slow flying insects that are probably easy

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Vol. 67(3)

to capture are common in Stenopogon prey records (e.g., the use of alate Formica

by S. inquinatus and crane flies by S. engelhardti, Powell & Stage 1962). However,

the flies also include several strong flying species in their diets, such as honey bees

and robber flies. Although honey bees are strong fliers, their tendency to hover

and approach flowers slowly probably makes them susceptible to robber flies

while foraging. In all cases of S. inquinatus predation on robber flies, the predator

was larger than its prey, suggesting that predator size constrains its ability to take

relatively formidable and strong flying prey (O’Neill in press). Thus, the upper

limit of S. inquinatus prey size (represented by conspecifics) is apparently deter¬

mined by catchability. The lower limit may be defined by the lower visibility or

perceived unprofitability of small insects, such as Leptothorax alates that may be

locally abundant, but which do not appear in prey records.

Acknowledgments

We thank Ruth O’Neill, Kathleen Johnson, and James Cummins for assistance

with the field work and Andre Francoeur (Universite du Quebec), C. Riley Nelson

(University of Texas), and M. Ivie and T. Keith Philips (Montana State University)

for providing determinations. Howard Evans, Michael Ivie, Robert Nowierski,

and Douglas Streett kindly provided comments on the manuscript. This work

was supported by USDA Agricultural Research Service, USD A Animal and Plant

Health Inspection Service Plant Protection and Quarantine, and the Montana

Agricultural Experiment Station. Contribution J-2631 from the Montana Agri¬

cultural Experiment Station.

Literature Cited

Bromley, S. W. 1942. Bee-killing asilids in New England. Psyche, 49: 81-83.

Bromley, S. W. 1948. Honey-bee predators. J. New York Entomol. Soc., 56: 195-199.

Cole, F. R. 1958. Prey of robber flies of the genus Stenopogon (Diptera: Asilidae). Pan-Pacif. En¬

tomol., 34: 176-178.

Dennis, D. S. & R. J. Lavigne. 1975. Comparative behavior of Wyoming robber flies. II. (Diptera:

Asilidae). Univ. Wyoming, Agric. Exp. Sta., Sci. Monogr., 30.

Lavigne, R. J. & F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming robber

flies (Diptera: Asilidae). Univ. Wyoming, Agric. Exp. Sta., Sci. Monogr., 18.

Kemp, W. P., S. J. Harvey & K. M. O’Neill. 1990. Patterns of vegetation and grasshopper community

composition. Oecologia, 83: 299-308.

O’Neill, K. M. (in press). Body size asymmetries in predatory interactions among robber flies (Diptera:

Asilidae). Ann. Entomol. Soc. Am.

O’Neill, K. M. & W. P. Kemp. 1990. Behavioral responses of the robber fly Stenopogon inquinatus

Loew (Diptera: Asilidae) to variation in the thermal environment. Environ. Entomol., 19: 459-

464.

Powell, J. A. & G. I. Stage. 1962. Prey selection by robber flies of the genus Stenopogon, with

particular observations on S. engelhardti Bromley (Diptera: Asilidae). Wasmann J. Biol., 20:

139-157.

Shelly, T. E. 1984. Prey selection by the Neotropical robber fly, Atractia marginata (Diptera: Asi¬

lidae). Proc. Entomol. Soc. Wash., 86: 120-126.

Wilcox, J. 1971. The genera Stenopogon Loew and Scleropogon Loew in America north of Mexico

(Diptera: Asilidae). Occ. Papers, Calif. Acad. Sci., 89: 1-134.

Received 31 December 1990; accepted 22 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 181-188, (1991)

A NEW GENUS AND SPECIES OF TRICHIINI FROM

MEXICO (COLEOPTERA: MELOLONTHIDAE)

Leonardo Delgado-Castillo and Miguel Angel Moron

Instituto de Ecologia, A.C., Apdo. Postal 63,

91000, Xalapa, Veracruz, Mexico

Abstract. — The adult and larva of Iridisoma acahuizotlensis NEW GENUS, NEW SPECIES from

the State of Guerrero, Mexico, are described and illustrated. Relationships with Paragnorimus,

Dialithus and Giesbertiolus are discussed. A modification of Howden’s key to the North and

Central American genera of Trichiini is included.

Resumen. — Se describen e ilustran el adulto y la larva de Iridisoma acahuizotlensis nov. gen.,

nov. sp. colectados en el estado de Guerrero, Mexico. El unico ejemplar conocido se obtuvo a

partir de la cria de una larva de tercer estadio encontrada dentro de un tronco podrido. Se

discuten sus relaciones con Paragnorimus, Dialithus y Giesbertiolus, comentando su distribution

geografica-ecologica. Tambien se incluye una modification a la clave para los generos Norte y

Centroamericanos de Trichiini.

Key Words. — Insecta, Coleoptera, Melolonthidae, Iridisoma acahuizotlensis

The North and Central American Trichiini are represented by 37 species in¬

cluded in eight genera (Howden 1968, 1970, 1971, 1972, 1988). All these taxa

are apparently related to Old World genera, and the distribution of most of their

species is relictual. The relationships between the American genera are vague,

and at present they appear as distant taxa (Howden 1968). Only the larva of eight

species of American Trichiini are known; these are in Trigonopeltastes, Gnorimella

and Trichiotinus (Ritcher 1966, Moron 1983).

During summer, 1986, one cetoniid-like larva was collected from a rotten log

in a disturbed subdeciduous tropical forest near Acahuizotla, Guerrero, Mexico.

The larva was reared in the laboratory for seven months, emerging as a perfect

adult male that showed characters that permitted easy separation from all other

known genera of American Trichiini. This paper describes the new genus and

species, based on the adult male, and the third instar larva, based upon its cast

skin.

Iridisoma Delgado-Castillo & Moron, NEW GENUS

Type Species.—Iridisoma acahuizotlensis Delgado-Castillo & Moron, NEW

SPECIES.

Description. — Body oval-elongate, with conspicuous dorsal and ventral setae, without scales or

cretaceous marks. Pronotum, scutellum, pygidium and venter iridescent green; elytra dull orange.

Clypeus nearly as long as wide, with short erect setae, anterior border thickened, moderately emar-

ginated. Mandibles membranous, maxillae with long, apical setae, mentum weakly widened, deeply

emarginated. Frons and vertex with dense erect setae. Anteocular nocht present. Antennae 10-seg-

mented. Pronotum longer than wide, semihexagonal, posterolateral borders parallel to and longer than

convergent anterolateral margins; posterior angles well marked; dorsal surface with iridescent, rugose

striae and numerous short, erected setae; margins well defined. Scutellum iridescent setose. Elytra

longer than wide, with nonelevated interstriae and striae nearly obsolete; surface setose; lateral elytral

declivity separated from elytral disc by a strong carina running from humeral to apical callus. Ventral

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Vol. 67(3)

region with long setae, especially on metastemum. Middle coxae contiguous; tarsi longer than tibiae,

tarsal claws entire, equal in length.

Male. — Pygidium wider than long, strongly convex apically, with abundant setae; fore tibiae uni-

dentate; middle tibiae short, recurved inward, with inner apex projected as a hook-like structure and

with an articulated spur.

Diagnosis.—Iridisoma is related with Paragnorimus, Dialithus and Giesber-

tiolus, but can be distinguished by the following combination of characters: pro-

notum, scutellum, pygidium and venter completely iridescent, unique pronotal

sculpture and elytral carina. The shape of the pronotum and elytra give a distinct

appearance to this trichiine.

Distribution. — Known only from the type locality.

Etymology. —Greek: “ iridos a rainbow, “soma” the body, in relation with

the iridescent color of all of the body.

Discussion. —Because this genus is described only with the male characters, it

is possible that many of the female structures may be different, especially the

shape of the clypeus, the pronotal convexity, the shape of pygidium and abdomen,

the tarsal length, and the structure of the fore and middle tibiae.

Key to the North and Central American Genera of Trichiini

Modified from Howden (1968)

1. Anterior border of clypeus rounded, truncate or slightly emarginated.

Color variable but not iridescent . 2

1Anterior border of clypeus moderate or strongly emarginated. Color at

least partially iridescent . 7

2(1). Pronotal disc without complete “V”-shaped or triangular impressions,

uniformly convex or with slight longitudinal or transverse depressions

. 3

2'. Pronotal disc with complete “V”-shaped or triangular impressions,

these marks usually cretaceous . Trigonopeltastes Burmeister

3(2). Pronotal disc with at least some fine, erect setae; in some cases with

transverse cretaceous stripes or spots. Body ovate . 4

3'. Pronotal disc without setae or impressions, slightly and uniformly con¬

vex. Body elongate. Apeltastes Howden

4(3) Elytral disc pubescent . 5

4'. Elytral disc without setae . Gnorimella Casey

5(4). Elytra dull, without shiny elevations . 6

5'. Elytra shiny, sometimes with two shiny elevations ... Trichiotinus Casey

6(5). Pronotal disc with an incomplete cretaceous “V”-shaped or middle

transverse line. Peltotrichius Howden

6'. Pronotal disc without cretaceous lines or dots. Paragnorimus Becker

7(1'). Dorsal surface glabrous or with sparse setae. Pronotum, pygidium and

venter partially iridescent; elytra with iridescent spots. Pronotum

with simple punctures and lines or with longitudinal sulci. Elytra

convex, with lateral declivity sparsely marked . 8

7' Dorsal surface densely setose. Pronotum, pygidium and venter com¬

pletely iridescent, elytra dull. Pronotum with dense, transversely,

rugose striae. Elytra with a strong carinae between the calla .

... Iridisoma NEW GENUS

1991

DELGADO-CASTILLO & MORON: NEW GENUS OF TRICHIINI

183

8(7). Clypeus longer than wide, bilobed. Pronotum with three central, lon¬

gitudinal sulci. Fore tibiae of male unidentated . Dialithus Parry

8'. Clypeus slightly wider than long and moderately emarginated. Prono¬

tum with a single middle longitudinal sulcus. Fore tibiae of male

bidentated. Giesbertiolus Howden

Iridisoma acahuizotlensis Delgado-Castillo & Moron, NEW SPECIES

(Figs. 1-14)

Type.— Holotype male, deposited in M.A. Moron collection (Xalapa, Mexico),

final deposition in Instituto de Ecologia, A.C. (Mexico); data: “MEXICO: Guer¬

rero, Mochitlan, Acahuizotla, Canada de Rio Escondido. Altitud 650 m. Bosque

tropical mediano subperennifolio.” “Ex larva colectada l-IX-86 en albura de

Cedrela sp. C.Deloya y L.Delgado, cols. Prepupa 7-III-87. Pupa 14-III-87. Adulto

3-IV-87.”

Male.— Holotype (Fig. 1). Total length (from the apex of clypeus to the pygidium): 15.1 mm.

Maximum elytral width: 6.4 mm. Clypeus. Nearly as long as wide (0.9:1.0), wide and strongly emar¬

ginated, but not bilobed; anterior border thickened, slightly elevated; lateral borders keeled from the

anteocular notch towards apical third; clypea dull orange, darkened in the emarginated area; regular

or fine punctulate with short, sparse, semierect setae. Frons. Black, strongly rugose toward the apex,

with long, erect, dense setae. Antenna. Club slightly shorter than the funicle. Pronotum. Slightly longer

than wide (1.0:0.9), anterior angles scarcely marked, middle angles rounded, and posterior angles acute,

directed ventrally. Margins black. Covered with dense, semierect setae. Scutellum. Semiovate, slightly

concave, colored as pronotum, with some semierect setae. Elytra. Longer than wide (1.0:0.84); dull

orange, lateral margins and sutura black; area contiguous to lateral margin darkened; elytral carina

brilliant orange. Surface finely rugose (best seen with counterlight), with nearly obsolete striae, except

sutural striae slightly elevated near posterior third; short, semierect setae best seen at base and center.

Venter. Iridescent, more intense than on pronotum. Metastemum with long, dense, decumbent yellow

setae and a central arrow-shaped mark on the flat semitriangular disc. Abdomen. Slightly concave.

Stemites I-IV equal in length, stemite V as long as III+IV, with the posterior border broad, slightly

emarginated. Anal plate as long as the stemite V, posterior border nearly straight. Setae on the stemites

long and decumbent. Pygidium. Wider than long (1.0:0.72), same color as stemites, with only narrow

mesial stripe and apex dark red; rugose, with long, abundant, semierect setae distributed evenly. Legs.

Coxae iridescent; femora brilliant orange with iridescent green shine on the anterior face and near the

coxae. Fore tibiae spur shorter than basal tarsomere. Fore tibiae orange ventrally and dorsally, dor¬

solateral face black. Fore tarsi black, almost one-third longer than fore tibiae; tarsomeres I-IV short

tuft of ventral setae, denser towards the tibiae. Middle tibiae dark orange, bent at apical one-half;

mesotibial spur shorter than basal tarsomere (Fig. 2). Middle tarsi 2.0 x longer than middle tibiae.

Hind tibiae black, long and fine, scarcely thickened towards the apex, with a short, dorsal, transverse

carina and two elongated, apical spurs. Hind tarsi nearly twice as long as tibiae. Middle and hind tarsi

with long, stout setae on the ventral and apical surfaces (except the last tarsomere). All the tarsal claws

nearly as long as tarsus V. Genitalia. Basal piece and tectum strongly convex; paramera directed

downward in the basal part, but abruptly bent distally, with two long and fine processes widened

apically; the apices are rounded and mesially directed (Figs. 3-4).

Third Instar Larva.—{ Figs. 5-14). Head. Maximum width of head capsule: 3.5 mm. Surface of

cranium slightly rugose, light yellow. Frons laterally bearing one minute posterior seta, one minute

exterior seta, and two minute anterior setae; remaining cranial surface glabrous, except one dorsal

epicraneal seta and two paraocellar setae laterally. Clypeus laterally bearing two long anterior setae

and one minute exterior seta. Labrum semiovate, with two central setae and two lateral setae. Ocelli

present (Fig. 5). Epipharynx (Fig. 6), without clithra and zygum, with only nine erected setae on

haptomera; right chaetoparia with 38—40 spine-like setae and few minute fine setae; left chaetoparia

with 20-22 fine setae; dexiotorma, laeotorma and sense cone well developed, acanthoparia with four

or five curved setae. Scissorial area of left mandible with three teeth, well separated by two notches;

molar area with one lobe, rounded acia, very prominent, acute ventral process, dense brustia (Fig. 7).

Scissorial area of right mandible with two teeth; molar area with three acute lobes, ventral process

184

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Figure 1. Dorsal view of the holotype of Iridisoma acahuizotlensis NEW GENUS, NEW SPECIES.

Scale line 5 mm.

1991

DELGADO-CASTILLO & MORON: NEW GENUS OF TRICHIINI

185

Figures 2-4. Iridisoma acahuizotlensis. Figure 2. Distolateral view of the apex of left middle tibia.

Figure 3. Distal view of the male genital capsule. Figure 4. Lateral view of the male genital capsule.

Scale lines are 1 mm.

less acute and prominent pointed calx and short brustia. Stridulatory area of each mandible formed

by 13 fine striae. Galea with sharply pointed conical uncus; lacinia with two fused large conical unci;

maxillary stridulatory area with only three sharp, anteriorly directed, recurved or angulated teeth, and

a prominent distal, rounded process (Figs. 8-9). Labium short, with scarce small setae; hypopharyngeal

sclerome strongly developed, prominent and heavily sclerotized (Fig. 10). Dorsal surface of last an-

tennomere with three sensory spots (Fig. 11). Thorax. Thoracic spiracles 0.45 mm long and 0.30 mm

wide. Respiratory plate with a maximum of 18-20 irregular elongate “holes” along any diameter, but

not in definite rows. Arms of respiratory plate nearly contiguous, separated by a distance much less

than the dorsoventral diameter of the bulla. Prothoracic sclerome light yellow. Dorsal and lateral setae

long, regularly distributed. Abdomen. Dorsa of abdominal segments covered with numerous fine, long

setae, regularly distributed. Spiracles I-VIII similar in size, 0.40 mm long and 0.30 mm wide. Distance

between the two lobes of the respiratory plate much less than the dorsoventral diameter of the bulla

(Fig. 12). “Holes” of the plate with irregular, elongate shape, similar to those on thorax (Fig. 13).

Upper anal lip with a great number of short and long mixed setae. Septula and palidia absent. Lower

anal lip with numerous long scattered setae. Legs. Claws falcate, sharp pointed, each bearing two

proximal setae (Fig. 14). Body length approximately 23 mm.

Diagnosis .—The species presents many unusual characters and is remotely

related to other Central American species. One diagnostic feature is the male’s

genitalia capsule, with the parameres long, narrow, strongly curved and widened

186

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Figures 5-9. Iridisoma acahuizotlensis third-instar larva. Figure 5. Frontal view of head. Figure

6. Epipharynx. Figure 7. Inner view of left mandible. Figure 8. Inner view of right maxilla. Figure 9.

Detail of right maxillary stridulatory area. Scale lines are 1 mm.

apically, not short, wide, straight or slightly curved and narrowed apically as in

Giesbertiolus and Dialithus.

Etymology. — From the Mexican (Nahuatl) word “ Acahuizotlan ,” translated as

“site of spiny reed grass” or “southern site of reed grass,” with the Latin suffix

“ensis,” belonging to Acahuizotla.

Discussion. — The existence of this species in the Sierra Madre del Sur reinforces

the relictual pattern of the montane southern Pacific distribution of these genera

of Trichiini. We expect many other surprises in this poorly collected area, es-

1991 DELGADO-CASTILLO & MORON: NEW GENUS OF TRICHIINI

187

C^> CT7

Figures 10-14. Iridisoma acahuizotlensis third-instar larva. Figure 10. Inner view of hypopharynx.

Figure 11. Dorsal view of last antennal article. Figure 12. Third abdominal spiracle. Figure 13. Detail

of “holes” in the respiratory plate. Figure 14. Right hind tarsal claw, external view. Scale lines are 0.3

mm.

pecially in small humid, tropical or subtropical canyons, which support remnants

of tropical subdeciduous forests. It is interesting that during three years of col¬

lecting, we only obtained the type specimen of Iridisoma; during that period, only

11 specimens of Trigonopeltastes discrepans Howden, T. s. sallaei Bates, T. ar-

quimedes Schaum, T. frontalis Bates and Apeltastes elongata Howden were col¬

lected, always associated with the lower parts of the vegetation. Perhaps adults

of Iridisoma are associated with the canopy of the subdeciduous tropical forest,

represented in that locality, with trees 18m high. The larval description is based

on a single cast skin of a third instar larva reared to the adult stage.

According with the key to the larvae of American Trichiinae proposed by Moron

(1983), Iridisoma must be placed near Inca, with reference to the absence of

clithra, the number of dorsal sensory spots of the antennae, and the haptomerum

without mesal tooth-like process. Also it is interesting that Iridisoma appears

more related to Trichiotinus than Trigonopeltastes, based upon the mandibular

and maxillar structures. It is necessary to know the larval morphology of other

Central American genera of Trichiinae to understand their precise relationships.

Material Examined.— Type, reared from third instar larva; data: MEXICO. GUERRERO : Canada

de Rio Escondido, Acahuizotla, Mochitlan, 650 m, 1 Sep 1986, C. Deloya and L. Delgado, ex Cedrela

sp. (Meliacea) rotten log.

188

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Vol. 67(3)

Acknowledgment

We thank Cuauhtemoc Deloya (Instituto de Ecologia, Mexico) for his collab¬

oration during the field work, and Henry F. Howden (Ottawa, Ontario) for the

donation of literature and specimens. This research is a contribution of the Biosys-

tematics and Ecology of Insects Program of the Instituto de Ecologia (Xalapa,

Ver.) and is paper number 13 of the “Studies of the Coleoptera Lamellicomia

from Sierra Madre del Sur” (P220CCOR-880061) project supported by CONA-

CYT, Mexico.

Literature Cited

Howden, H. F. 1968. A review of the Trichiinae of North and Central America (Coleoptera: Scar-

abaeidae). Mem. Entomol. Soc. Can., 54.

Howden, H. F. 1970. The genus Paragnorimus, with descriptions of two new species (Coleoptera:

Scarabaeidae). Can. Entomol., 102: 1385-1389.

Howden, H. F. 1971. A second species of Peltotrichius (Coleoptera: Scarabaeidae). Can. Entomol.,

103: 104-106.

Howden, H. F. 1972. New species of Dialithus Parry and a new synonym of Pantodinus Burmeister

(Coleoptera: Scarabaeidae: Trichiinae). Can. Entomol., 104: 647-654.

Howden, H. F. 1988. A new genus and four species of New World Trichiinae (Coleoptera:Scara¬

baeidae). Coleopts Bull., 42: 241-250.

Moron, M. A. 1983. Los estados inmaduros de Inca clathrata sommeri Westwood (Coleoptera,

Melolonthidae, Trichiinae); con observaciones sobre el crecimiento alometrico del imago. Folia

Entomol. Mex., 56: 31-51.

Ritcher, P. O. 1966. White grubs and their allies. University of Oregon Press, Corvallis.

Received 7 September 1990; accepted 4 February 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 189-194, (1991)

DEVELOPMENT AND SURVIVORSHIP OF

CYDIA POMONELLA (L.) (LEPIDOPTERA: TORTRICIDAE)

AT TEN CONSTANT TEMPERATURES

Michael J. Pitcairn, 1 Carolyn Pickel, 2 Louis A. Falcon, and Frank G. Zalom 1

Department of Entomological Sciences, University of California,

Berkeley, California 94720

Abstract.— Development and survivorship of Cyclia pomonella (L.) immature stages were ob¬

served at 10 constant temperatures between 8.9° and 34.4° C. All stages failed to complete

development at temperatures below 12.2° C. Fourteen percent of the eggs and no larvae or pupae

survived at 34.4° C. The lower developmental thresholds, estimated by regressing developmental

rate (1/days) against temperature, ranged between 10.5°-12.5° C. A nonlinear model based on

Logan’s function was fit to the observed data and provides a more complete description of the

developmental rate-temperature relationship than a linear model.

Key Words. — Insecta, Cydia pomonella, developmental rates, threshold temperatures, Logan’s

model

Cydia pomonella (L.), codling moth, is the most destructive insect pest of

walnuts, apples, pears and plums in California. Damage is caused by larvae feeding

directly on fruit resulting in early fruit drop or unmarketable mature fruit (Uni¬

versity of California in press). In the absence of effective biological and cultural

controls, damage is prevented by properly timed insecticidal treatments. Spray

timing is usually achieved using a degree-day model and pheromone trap catch

data (Riedl et al. 1976; University of California 1987, in press).

The relationship between temperature and development of C. pomonella has

been the subject of several studies (Glenn 1922, Shelford 1927, Hagley 1972,

Williams & McDonald 1982, Rock & Shaffer 1983). Early studies were performed

in an outdoor insectary under ambient air temperatures; later studies used constant

temperature cabinets. However, all these studies investigated temperatures in the

middle of the developmental rate curve, a portion that can be reasonably ap¬

proximated by a straight line. For example, Rock & Shaffer (1983) reared larvae

and pupae of moths collected in North Carolina at 16°, 22°, 28°, and 32° C. The

lower thermal threshold (9.9° C) was estimated by regressing developmental rate

against temperature; the upper thermal threshold was suggested to occur above

32° C.

Studies have shown that the relationship between codling moth developmental

rate and temperature is significantly nonlinear (Rock & Shaffer 1983, Gold et al.

1987). Recent field studies in California showed that development under high

summer temperatures (>30°C) was slower than predicted using a linear degree-

day model (MJP & FGZ, unpublished data). We suggest that a nonlinear model

may better represent moth development, especially when temperatures routinely

exceed the upper thermal threshold. Here, we examine the rate of immature

1 Statewide IPM Project, University of California, Davis, California 95616.

2 University of California Cooperative Extension, 142A Garden Highway, Yuba City, California

95991.

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THE PAN-PACIFIC ENTOMOLOGIST

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development of C. pomonella at 10 constant temperatures to provide a more

complete description of its developmental rate curve and fit a nonlinear model

to the data.

Materials and Methods

Cydia pomonella larvae, pupae, and adults were selected from a laboratory

colony maintained on artificial diet (modified after Vanderzant & Davich 1958)

at about 23° C and natural photoperiod. The colony was initiated with adults

collected in apple orchards near Placerville, California. Eggs were obtained by

placing 20 adult pairs in a metal rotating oviposition cage (Batiste & Olson 1973)

2-3 h before sunset; eggs were collected 4 h later and put into a vegetable crisper

at 100% RH and immediately placed into a temperature cabinet. Sample size

varied from 42-89 eggs.

Developing larvae were maintained individually on artificial diet (approxi¬

mately 10 ml) in 30 ml plastic cups. The end of the larval stage was when larvae

began to spin a cocoon. Pupal development was examined using individuals that

started cocoons on the same day.

Ten constant temperature regimes were used: 8.9°, 10.0°, 11.1°, 12.2°, 15.6°,

25.6°, 27.8°, 30.0°, and 34.4° C. At 15.6° C and above, egg hatch was checked

every 24 h; at 12.2° C and below, egg hatch was checked at 1-3 day intervals.

Larvae and pupae were examined for completion of their developmental stage

every 1-2 days at 15.6° C and above, and at 1-3 day intervals at temperatures

below 15.6° C. The end of a stage was recorded as the midpoint between two

observations.

The relationship between temperature and developmental rate (rate = 1/days)

was described by linear and nonlinear regression models. Parameter estimates of

the linear model were calculated by regressing observations at 15.6°-32.2° C (the

X-intercept method of Arnold 1959). Parameter estimates of the nonlinear model

were calculated using Marquardt’s method of least-squares (SAS Institute 1985).

Results and Discussion

The survivorship and mean duration of development for the egg, larval, and

pupal stages are presented in Table 1. Development was not observed in any

stages at 8.9° and 10.0° C but was noted in all stages at 11.1° and 12.2° C. Eggs

maintained at 8.9° and 10.0° C remained in the opaque white stage (Richardson

et al. 1982), but those at 11.1° and 12.2° C developed to the blackhead stage and

then died. Hagley (1972) studied the development of C. pomonella eggs at several

constant temperatures, and reported that while all eggs failed to hatch at 10.0° C,

27% developed to the blackhead stage before dying. Successful egg hatch, however,

did occur at 11° C. In our study, egg survival was highest between 15.6°-27.8° C,

was lower at 30.0°-32.2° C and substantially lower at 34.4° C. These results

corroborate Hagley’s (1972) observations of reduced egg hatch at temperatures

above 30° C. Larval survivorship in our study was >80% between 27.8°-32.2° C;

pupal survivorship was > 80% between 27.8°-30.0° C. No larvae or pupae survived

at 34.4° C.

The number of days required to complete embryogenesis declined between

15.6°-27.8° C then increased with increasing temperature. For the larval stage,

there was a general increase in developmental rate with increasing temperatures

Table 1. Survivorship and mean days required for development of immature stages of Cydia pomonella at different constant temperatures.

Stage

Egg

Larva

Pupa

Total

Temperature

(°C)

n a

S b

(%)

Development

(mean days ± SE)

Range

(days)

(%)

Development

(mean days ± SE)

Range

(days)

(%)

Development

(mean days ± SE)

Range

(days)

Development

(mean days)

8.9

0.0

10.0

0.0

—

0.0

—

0.0

—

11.1

0.0

—

0.0

—

0.0

—

12.2

0.0

—

0.0

—

0.0

—

15.6

77.5

19.26 ± 0.55

18.0-20.5

53.3

62.21 ± 2.75

54-82

25.0

68.18 ± 1.36

66-75

149.65

25.6

78.7

6.47 ± 0.42

5.0-7.0

76.7

17.14 ± 0.53

15-23

61.1

17.32 ± 0.56

15-21

40.93

27.8

79.7

4.21 ± 0.00

—

83.3

13.42 ± 0.22

11-17

89.5

10.10 ± 0.34

9-14

27.73

30.0

58.7

4.27 ± 0.33

3.5-5.5

87.0

14.68 ± 0.41

12-19

80.0

10.85 ± 0.58

7-15

29.80

32.2

49.3

4.56 ± 0.23

4.0-5.0

80.0

11.27 ± 0.25

11-14

22.2

10.71 ± 1.34

8-14

26.54

34.4

14.3

5.40 ± 0.57

5.0-6.0

0.0

—

0.0

—

a Sample size.

b Survivorship.

1991 PITCAIRN ET AL.: DEVELOPMENT OF CYDIA POMONELLA

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THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Table 2. Linear regression equations and estimates of lower developmental thresholds using mean

developmental rates for immature stages of Cydia pomonella.

Stage

Regression equation 3

SE of regression

coefficient

r 2

Lower threshold 1 *

(°C)

Egg

Y =

-0.121 + 0.0115*X

0.0024

0.88

10.56

Larva

Y =

-0.048 + 0.0042*X

0.0005

0.96

11.54

Pupa

Y =

-0.065 + 0.0052*X

0.0011

0.89

12.49

Egg-to-Adult

Y =

-0.023 + 0.0019*X

0.0003

0.94

11.85

a Regression equation is Y = a + bX, where Y is the developmental rate (1/days) and X is temperature

(°C).

b X-intercept.

above 15.6° C, the fastest rate occurring at 32.2° C. Little change in the rate of

pupal development was observed at 27.8°-32.2° C. These results suggest that the

upper developmental thresholds are near 27.8° C for the egg and pupal stages,

and 32.2° C for the larval stage.

The estimated lower thermal thresholds and associated linear regression equa¬

tions are presented in Table 2. The coefficient of determination, r 2 , was high for

all stages. Estimates of the lower thermal threshold ranged between 10.5°-12.5°

C and are similar (or slightly lower) to the minimum temperatures for which

development was observed. The average number of degree-days (° D) above the

estimated threshold of 11.85° C for total development (egg to adult) is 529.45° D.

Temperature (°C)

Figure 1. Rate of total (egg to adult) development as a function of temperature for Cydia pomonella

(a = linear regression for the temperature range of 15.6-32.2° C; b = nonlinear model after Logan et

al. 1976; means are given with range of observed values; parameter estimates are given in Tables 2

and 3).

1991

PITCAIRN ET AL.: DEVELOPMENT OF CYDIA POMONELLA

193

Table 3. Parameter estimates of Logan’s model to describe the relationship between Cydia po-

monella developmental rate and temperature. In the model, rate = a{exp[b*x] — exp[b«c — (c — x)/d]}.

Parameter estimates

df a

Life stage

Eggs

0.0767

0.1921

36.1601

5.1617

265

Larvae

0.0036

0.1067

34.4463

1.2581

129

Pupae

0.0013

0.2443

34.4441

3.8968

Total development

0.0006

0.1490

34.4493

2.1796

a Degrees of freedom.

The most commonly reported lower thermal threshold in the literature for C.

pomonella is 10° C. Using this value for comparison, the average number of

degree-days required to complete development of the egg, larval, and pupal stages

are 93.87° D, 280.62° D, and 256.93° D, respectively; the mean duration of total

development is 631.42° D. Glenn (1922) presented data showing that an average

of 645.00° D (base 10° C) were required for total development. In our study, the

average number of degree-days from newly emerged larvae to adults was 537.55°

D (base 10° C), a value similar to other studies (all base 10° C): 552.78° D (Glenn

1922), 494° D (Williams & McDonald 1982), 514° D (Rock & Shaffer 1983).

The average number of days for the total developmental period was estimated

as the sum of the means for the egg, larval, and pupal stages. These data are

plotted in Fig. 1 along with the range of observed values. Low or no survival at

34.4° C indicates that high temperature can inhibit C. pomonella developmental

rate. A nonlinear function based on Logan’s model (Logan et al. 1976) was used

to describe the rate-temperature relationship (Fig. 1). The parameter estimates

for each life stage are given in Table 3. For total development, the calculated

lethal maximum temperature with 100% mortality was 34.45° C; the fastest rate

of development was calculated to occur at 30.82° C. In contrast with the linear

model, Logan’s model exhibits a rapid decline at temperatures >31° C and also

fits the data well in the intermediate temperature range.

Linear models of development have become widely used in practice because

they have worked well for a wide variety of insects, they are easily modified (e.g.,

adding an upper thermal threshold), and their parameters are easily estimated

from developmental data. Our study is significant because not only are the in¬

termediate temperatures examined, but also temperatures at the upper and lower

thermal limits of development. When a nonlinear function is fit to our data, a

more complete description of the developmental rate curve of C. pomonella is

presented. Air temperatures exceed 31° C almost daily during July through Sep¬

tember in most regions of California where C. pomonella is found (University of

California 1983) and use of this nonlinear model should improve prediction of

immature development during these summer months.

Literature Cited

Arnold, C. Y. 1959. The determination and significance of the base temperature in a linear heat unit

system. Proc. Amer. Soc. Hort. Sci., 74: 430^145.

Batiste, W. C. & W. H. Olson. 1973. Codling moth: mass production in controlled environment

rearing units. J. Econ. Entomol., 66: 383-388.

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Vol. 67(3)

Glenn, P. A. 1922. Relation of temperature to development of the codling-moth. J. Econ. Entomol.,

15: 193-199.

Gold, H. J., W. L. Kendall & P. L. Shaffer. 1987. Nonlinearity and the effects of microclimatic

variability on a codling moth population (Cydia pomonella ): a sensitivity simulation. Ecol.

Modelling, 37: 139-154.

Hagley, E. A. C. 1972. Observations on codling moth longevity and egg hatchability. Environ.

Entomol., 1: 123-125.

Logan, J. A., D. J. Wollkind, S. C. Hoyt & L. K. Tanigoshi. 1976. An analytic model for description

of temperature dependent rate phenomena in arthropods. Environ. Entomol., 5: 1133-1140.

Richardson, J. C., C. E. Jorgensen & B. A. Croft. 1982. Embryogensis of the codling moth, Laspeyresia

pomonella : use in validating phenology models. Ann. Entomol. Soc. Am., 75: 201-209.

Riedl, H., B. A. Croft & A. J. Howitt. 1976. Forecasting codling moth phenology based on pheromone

trap catches and physiological time models. Can. Entomol., 108: 449-460.

Rock, G. C. & P. L. Shaffer. 1983. Developmental rates of codling moth (Lepidoptera; Olethreutidae)

reared on apple at four constant temperatures. Environ. Entomol., 12: 831-834.

SAS Institute Inc. 1985. SAS user’s guide: statistics, version 5 edition. SAS Institute Inc., Cary,

North Carolina.

Shelford, V. E. 1927. An experimental investigation of the relationships of the codling moth to

weather climate. Ill. Nat. Hist. Surv. Bull., 16: 311-440.

University of California. 1983. Average Daily Air Temperatures and Precipitation in California.

Univ. of Calif., Div. of Agric. Sci., Spec. Publ., 3285.

University of California. 1987. Integrated pest management for walnuts (2nd ed.). Univ. of Calif.,

Div. of Agric. and Nat. Res. Publ., 3270.

University of California. (In press). Integrated pest management for apples and pears. Univ. of Calif.,

Div. of Agric. and Nat. Res. Publ., 3340.

Vanderzant, E. S. & T. B. Davich. 1958. Laboratory rearing of the boll weevil: a satisfactory larval

diet and oviposition studies. J. Econ. Entomol., 51: 288-291.

Williams, D. G. & G. McDonald. 1982. The duration and number of immature stages of codling

moth Cydia pomonella (L.) (Tortricidae: Lepidoptera). J. Aust. Entomol. Soc., 21: 1-4.

Received 3 December 1990; accepted 12 February 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 195-199, (1991)

DESCRIPTION OF THE LARVA OF ATRACTELMIS

(COLEOPTERA: ELMIDAE) AND NEW

INFORMATION ON THE MORPHOLOGY,

DISTRIBUTION, AND HABITAT OF

ATRACTELMIS WAW ON A CHANDLER

William D. Shepard and Cheryl B. Barr

Department of Biological Sciences,

California State University, Sacramento,

Sacramento, California 95819

Abstract .—Collection of additional specimens of Atractelmis wawona Chandler has allowed

descriptions of the female and larva. The species’ known geographic distribution is from northern

California to southern Oregon and Idaho. Submerged aquatic mosses are the preferred micro¬

habitat.

Key Words. — Insecta, Coleoptera, Elmidae, Atractelmis, descriptions, distribution, microhabitat

Atractelmis wawona Chandler for many years has been the most elusive riffle

beetle in the Nearctic region. This monospecific genus was described from two

specimens by Chandler in 1954. Despite serious attempts, it was not collected

again until 1969, when Harley P. Brown took a single specimen at Wawona,

California, the type locality. Sixteen years later, Brown collected a few more

specimens from a new locality near Wildwood, California. We spent several years

unsuccessfully searching for the species.

In 1988, a re-examination of the morphology of a specimen showed similarities

to Promoresia, an eastern Nearctic genus that usually inhabits aquatic mosses.

Careful examination of mosses at locations where Atractelmis wawona had been

previously collected revealed this species to be present, and demonstrated its

strong preference for inhabiting aquatic mosses. We now describe the female and

larva, both of which were unavailable to Chandler.

Atractelmis Chandler

Larva .—Body elongate (Fig. 6), hemicylindrical in cross section; dark brown; sclerites densely

tuberculate, intersegmental membranes lightly sclerotized, tan; head prognathous; posterior border of

thoracic and abdominal terga I-VIII with numerous tubercles bearing elongate stout plumose setae

(giving intersegmental areas a fringed look). Mature length 4.8-5.0 mm. Head highly sclerotized, dark;

dorsum very tuberculate; epicranial suture U-shaped with very short stem, arms ending near antennal

bases; stemmata distinct; edge of frons straight, smooth. Each antenna three-segmented, each segment

with apical setae; segment one as wide as long; segment two 3 x as long as wide; segment three 3 x

as long as wide but one-third the size of segment two. Labrum transversely rectangular, with anterior

border of several stout setae. Each mandible bluntly trifid; molar brush present. Each maxilla with

galea and lacinia distinct, armed with a brush of stout medially-curved setae; palpus three-segmented,

segment one as long as wide, segments two and three longer than wide, segment three with peg-like

setae apically. Labium 2 x as long as wide, apex truncate, one stout seta present on each side just

anterior to middle; each palpus short, two-segmented, segments each as wide as long and having blunt

apical setae. Cervical area lightly sclerotized; accessory sclerites (= cervical sclerites) greatly elongate,

diverging anteriorly away from a single area (V-shaped) (Fig. 7). Thoracic terga with middorsal suture

(or line); prothorax with a transverse triangular depression in anterior one-half on each side. Prostemum

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Vol. 67(3)

Figures 1-5. Adult Atractelmis wawona. Figure 1. Habitus. Figure 2. Normal color pattern. Figure

3. Color pattern variant. Figure 4. Male genitalia in dorsal, lateral and ventral views. Figure 5. Female

genitalia.

1991

SHEPARD & BARR: ATRACTELMIS BIOLOGY

197

Figures 6-9. Larval Atractelmis. Figure 6. Habitus. Figure 7. Ventral thoracic structures. Figure

8. Left thoracic pleurites. Figure 9. Abdominal segment IX (ventral view) and operculum.

reduced to triangular sclerite between and anterior to procoxae; meso- and metastemum very trans¬

verse, lightly tuberculate. Pro-, meso-, and metathoracic pleurites divided into two sclerites (Fig. 8);

prothoracic prepleurites with medial extensions that almost meet anterior to procoxae. All legs similar

(Fig. 7), all coxal cavities open; each coxa conical, anterolaterally with several stout setae; trochanter

well developed, subequal to femur; femur and tibia 2 x as long as wide, with numerous setae; tarsun-

gulus bluntly pointed, having one stout seta on medial border. Abdomen nine-segmented; segments

198

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

I-VI with separate tergites, pleurites and stemites; segments VII-IX with all sclerites united into a

ring. Middorsal line present on segments I-VIII. Segment IX 2 x as long as segment VIII, slightly

truncate apically, dorsally slightly sinuate (lateral view), having an elongate pentagonal operculum

(Fig. 9). Operculum with two posteroventrally directed claws on inner surface, and covering three tufts

of perianal tracheal gills each with 7-10 gill filaments. Gill filaments furcate basally (most bifurcate,

some trifurcate). Spiracles present on mature larva on sides of mesonotum and abdominal segments

I-VIII.

Atractelmis wawona Chandler

Adult Morphology. — The pronotal sculpturing is quite distinct from that of any

other Nearctic elmid (Fig. 1), particularly the bulbous disk and the transverse

posterior sulcus. The color pattern exhibited by this species is remarkably invar¬

iable. Each humerus has a broad comma-shaped macula with the tail near the

elytral suture (Fig. 2). The subapical maculae are elongate and directed medially

posteriorly. The single variant color pattern we have found has the spots somewhat

reduced in size (Fig. 3).

Chandler (1954) described the adult male well, except for one minor detail of

the genitalia: the median piece has an elongately sinuate, Y-shaped sclerite on the

ventral side (Fig. 4). Chandler’s drawing has detail in this area that resembles

sutures or folds in the membranes. We have found no external difference between

the sexes. The female genitalia are typical for elmids (Fig. 5).

Habitat and Microhabitat

Larvae and adults of A. wawona co-occur. They are usually found in cool, small-

to medium-sized, mountain streams; in these sites they are most numerous in

aquatic mosses. The greatest number of specimens we have collected at once came

from the moss Platyhypnidium riparioides (Hedwig) Dixon. However, we have

also collected specimens from submerged roots of Indian rhubarb, Peltiphyllum

peltatum (Torrey) Engler, and of riparian trees.

Geographical Distribution

Atractelmis wawona occurs throughout the northern half of California and parts

of southern Oregon and Idaho. Although it is probably even more widely dis¬

tributed, its restricted microhabitat has hidden it from those few who have searched

for it, as noted by Chandler (1954).

Distribution Records.— CALIFORNIA. HUMBOLDT Co.: 1.6 km (1 mi) west of Dinsmore, Van

Duzen River, 13 Jul 1988, W. D. Shepard, 2 adults and 1 larva. MARIPOSA Co.: nr Wawona, South

fork of Merced River, 17 Jul 1946, H. P. Chandler, 1 adult (holotype); Wawona, South Fork of Merced

River, 18 Jul 1969, H. P. Brown, 1 adult. PLUMAS Co.: 1.6 km (1 mi) NE of Indian Falls, Indian

Creek, 19 Aug 1989, W. D. Shepard and C. B. Barr, 2 larvae. SHASTA Co.: Middle Fork of Cottonwood

Creek, 24 Jun 1952, H. P. Chandler, 1 adult (paratype). SHASTA-TEHAMA Co. Border: Beegum,

Beegum Creek, 12 Jul 1988, W. D. Shepard, 10 adults and 25+ larvae; same but 4 Sep 1988, W. D.

Shepard and C. B. Barr, 5 adults and 31 larvae; 10.5 km (6.5 mi) S of Platina, North Fork of Beegum

Creek, 17 Jul 1990, W. D. Shepard and C. B. Barr, 15 larvae. SISKIYOU Co.: 4.8 km (3 mi) SE of

Cecilville, South Fork of Salmon River, 21 Jul 1990, W. D. Shepard and C. B. Barr, 11 adults and 1

larva. TRINITY Co.: 6.4 km (4 mi) W of Wildwood, Hayfork Creek, 12 Jul 1988, W. D. Shepard, 4

adults and 11 larvae; 3.2 km (2 mi) N of Wildwood, Hayfork Creek, 4 Sep 1988, W. D. Shepard and

C. B. Barr, 62 adults and 25+ larvae; same but 17 Jul 1990, W. D. Shepard and C. B. Barr, 24 adults;

below Wildwood, Gemmill Gulch, 23 Jun 1985, H. P. Brown, 3 adults; Mad River (town), (on the)

Mad River, 13 Jul 1988, W. D. Shepard, 1 larva. IDAHO. ADAMS Co.: 19.3 km (12 mi) N of New

Meadows, Little Salmon River, 17 Aug 1989, W. D. Shepard and C. B. Barr, 3 adults and 11 larvae.

1991

SHEPARD & BARR: ATRACTELMIS BIOLOGY

199

OREGON. CROOK Co.: 4.8 km W of Forest Service Headquarters, Meehan, Ochoco Creek, 26 Jul

1974, R. E. Miller, 1 adult.

Discussion

Brown’s (1972:62) key to the genera of aquatic dryopoid larvae of the United

States can be modified to include Atractelmis as follows (we retain Brown’s [1972]

style for the key couplets below; the figure numbers listed refer to that work):

26 (25) Body robust, broad, subtriangular in cross section; with spatulate

spines along lateral margins and mid-dorsal line (Figs. 169, 170):

Ampumixis

Body long and slender, hemicylindrical; without prominent clusters

of spines. 26A

26 A (26) Abdominal segments 7 and 8 with no longitudinal sutures, segments

1-6 with pleurites; posterior border of nota and terga with tu¬

bercles having elongate setae well sclerotized: Atractelmis

Abdominal segments 7 and/or 8 divided by tergostemal sutures,

pleurites on segments 1-6 or 1-7; posterior border of tubercles

with setae not or poorly sclerotized: Cleptelmis

Because Atractelmis was the last elmid genus in the United States for which larvae

were unknown, we now can concentrate on separating the species in the larval

stage.

Adult morphological characters indicate that Atractelmis is closely related to

Cleptelmis, Rhizelmis and Ampumixis. Externally, adult Atractelmis look most

similar to adult Rhizelmis that are quadrimaculate. This morph of Rhizelmis,

however, is equally as rare as Atractelmis. Fortunately, the two genera do not

occur in the same microhabitat. Larval morphology links Atractelmis very strongly

with Cleptelmis, and less strongly with Ampumixis. Larval synapomorphies shared

with Cleptelmis include the number of abdominal pleurites, and the posterior

fringe of tubercular setae on thoracic and abdominal segments (weakly developed

in Cleptelmis). Habitat and/or microhabitat preferences of Atractelmis also show

similarities to those of Rhizelmis, Ampumixis and, especially, Cleptelmis. In fact,

the collection of any of these genera suggests that the others may be present also.

Acknowledgment

We thank Harley P. Brown (University of Oklahoma, Norman) for providing

specimens for examination and information on his collection sites. We also greatly

appreciate the support and encouragement he has extended through the years.

Additionally, we thank William R. Buck (New York Botanical Garden, Bronx)

for identifying our samples of aquatic moss.

Literature Cited

Brown, H. P. 1972. Biota of freshwater ecosystems identification manual No. 6. Aquatic dryopoid

beetles (Coleoptera) of the United States. Water Pollution Control Research Series, U.S. En¬

vironmental Protection Agency, Washington, D.C.

Chandler, H. P. 1954. New genera and species of Elmidae (Coleoptera) from California. Pan-Pacif.

Entomol., 30: 125-131.

Received 17 December 1990; accepted 9 March 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 200-205, (1991)

A NEW SPECIES OF FOSSIL DASYMUTILLA

(HYMENOPTERA: MUTILLIDAE) FROM

DOMINICAN AMBER

Donald G. Manley 1 and George O. Poinar, Jr . 2

department of Entomology, Clemson University,

Pee Dee Research and Education Center,

Florence, South Carolina 29501

department of Entomological Sciences, University of California,

Berkeley, California 94720

Abstract.—Dasymutilla dominica, a new species of mutillid wasp, is described from a fossil

specimen embedded in a piece of Dominican amber estimated to be from 25 to 40 million years

old. Very few fossil mutillids have been recorded, and this represents the first description of a

fossil mutillid from the New World. This male belongs to the bioculata species-group, and can

be distinguished from closely related species by its totally black integument.

Key Words. — Insecta, Dasymutilla dominica, Mutillidae, new species, fossil, Dominican amber

Fossil velvet ants, or mutillids, are extremely rare and thus far have been limited

to Palaearctic forms. Menge (1856) mentioned finding six specimens of this family

in Baltic amber and Brischke (1886) cites another three in the same deposits. In

addition, Larsson (1978) mentions an additional three in the Copenhagen collec¬

tion of Baltic amber insects. To our knowledge, none of the above have been

described. Scudder (1891) also makes reference to an undescribed Mutilla from

the mid-Oligocene beds (not amber) located in Brunstatt, Alsatia (now Alsace),

in northeastern France.

Bischoff (1915) described seven species of fossil mutillids from Baltic amber,

placing them in a new genus, Protomutilla. Although Krombein (1979) suggests

that at least one of those species may not be correctly assigned to the family

Mutillidae, that does, nevertheless, appear to be the first description of a fossil

mutillid.

Sharov (1957) described another new species of aculeate Hymenoptera from

the Cretaceous of Siberia, which was ultimately assigned to Mutillidae. That fossil

specimen, estimated to be about 80 million years old, was called Cretavus sibiricus

Sharov, for which the author created a new family, Cretavidae. However, Ras-

nitsyn (1975) determined that the specimen was justified in being placed in the

family Mutillidae.

We describe here a mutillid from a piece of Dominican Republic amber. This

represents the first record and description of a fossil mutillid from the New World.

Methods and Materials

The piece of amber containing the mutillid was lent to GOP for study by Aldo

and Didi Costa of Puerto Plata, Dominican Republic. The original piece had to

be reshaped in order to better obtain a closer examination of the specimen. It is

presently semi-hemispherical, measuring 30 mm in greatest length, 16 mm in

1991

MANLEY & POINAR: A NEW FOSSIL MUTILLID

201

greatest width, 12 mm in greatest depth and weighing 3.5 g. The amber is golden

in color with a red tinge on one surface.

The piece of amber originated from mines located in the Cordillera Septentri¬

onal mountain ranges in the northern Dominican Republic. An analysis of amber

from seven Dominican mines by nuclear magnetic resonance spectroscopy sug¬

gests that amber from that area ranges in age from 25 to 40 million years (Lambert

et al. 1985).

Dasymutilla dominica Manley & Poinar, NEW SPECIES

(Figs. 1-3)

Type. — Holotype male, data: DOMINICAN REPUBLIC. Cordillera Septen¬

trional mountains; deposited in the Museum of Dominican Amber, Puerto Plata,

Dominican Republic.

Description. —Male. Integument black; pubescence sparse, mostly pale, erect or semierect; length

about 8 mm. Head black, rounded, sparse pale pubescence throughout; head slightly narrower than

thorax. Eyes and ocelli normal; eyes large, not bulging; ocelli small; ocello-ocular distance about 4.0 x

greatest width of ocelli. Relatively large, well-separated punctures throughout. Mandibles bidentate,

black throughout. Clypeus bidentate (but cannot be seen clearly). Antennae black; flagellomeres sub¬

equal in length; scape bicarinate beneath, coarsely punctate; antennal scrobes not carinate (but not

clearly visible). Thorax black, longer than broad, predominantly with sparse pale pubescence, with

moderately large, well-separated punctures. Pronotum with humeral angles rounded; anterior margin

apparently not emarginate (consistent with evenly rounded posterior margin of head). Mesonotum

without laterally expanded lobes. Propodeum coarsely reticulate. Tegulae subhemispherical, with

moderately large punctures; anterior margin with black pubescence. Pleura coarsely punctate through¬

out. Prothorax, scutellum, propodeum, and pleura with sparse pale pubescence; mesothorax with black

pubescence. Legs black, with sparse pale pubescence; calcaria pale; apices of middle and hind femora

rounded; posterior trochanters normal, not toothed; hind tibiae cylindrical, not flattened on inner

margin. Wings (badly damaged, clipped off near base) with basal third (right wing) hyaline, dark veins.

Abdomen black throughout. Terga and sterna I and II with coarse punctures; apical one-half of

remaining segments with smaller, shallower punctures. Tergite I with sparse, pale pubescence; con¬

spicuously lacking an apical fringe of hairs. Stemite I with a longitudinal carina about nine-tenths the

length of the segment. Tergite II with anterior two-thirds having sparse, pale pubescence; posterior

third with sparse, black pubescence; apical fringe pale. Terga III and IV with sparse, pale pubescence

with pale apical fringes of hairs. Terga V and VI with black pubescence throughout, including apical

fringes. Felt lines with black pubescence. Sterna I-IV with sparse, pale pubescence; II-IV with apical

fringes of pale hairs. Sterna V-VI with black pubescence. Stemite II with a round, median pit filled

with hairs (Fig. 3). Pygidium not readily visible, but appears to have an apical fringe of hairs. Hy-

popygium black, punctation not readily visible; posterolateral angles of last stemite appear to be

rounded, not dentate.

Diagnosis.—Dasymutilla dominica is placed in the bioculata species group. It

is easily distinguished from other, closely related members of the bioculata species-

group by the color of the second abdominal segment. In D. dominica, the integ¬

ument of the second abdominal segment is entirely black. In D. bioculata (Cresson),

the apical one-half of the second abdominal tergite is ferruginous. Dasymutilla

chiron (Blake) has a pair of large yellow spots on the posterior one-half of the

tergite. In D. hersilia Mickel, the second stemite is entirely, and the anterior,

posterior, and lateral margins of the second tergite are black, although the re¬

mainder is ferruginous; additionally, the integument of the second abdominal

segment is entirely ferruginous for D. lepeletierii (Fox), D. pyrrhus (Fox), and D.

praegrandis Mickel.

202

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Etymology. — The specific name is a Latinized version of the name of the re¬

public from which the specimen was collected.

Distribution.— Known only from the type specimen.

Material Examined. —Type.

Discussion

The genus Dasymutilla was first proposed by Ashmead (1899). The characters

that he attributed to the genus included the first abdominal segment being petiolate

with the second; rounded or hemispherical, highly polished eyes; males with wings;

and the body usually very hairy or pubescent. Inconsistencies in Ashmead’s clas¬

sification were pointed out by subsequent authors. Mickel’s (1928) list of diag¬

nostic characters for the group has been generally accepted since its publication.

Although his characters included many of the same as Ashmead’s, Mickel noted

that not all Dasymutilla are densely pubescent. Except for characters of the wings

(which cannot be seen), the type of D. dominica exhibits the characteristics at¬

tributed to Dasymutilla. The median pit filled with hairs on the second abdominal

stemite is an important character, but one that is present only in some male

Dasymutilla. However, the only other genus in which this character is present is

Traumatomutilla, and that genus is restricted in distribution to South America.

Thus, it seems apparent that, despite its age, this specimen belongs to Dasymutilla.

Mickel (1928) points out that many of the characters present in Dasymutilla

are superspecific; that is, present in more than one species. He uses the term

“species group,” with members of each group being closely related. Using Mickel’s

(1928) key to the species, D. dominica keys to couplet 54, with species in couplets

54-56 belonging to the bioculata species-group. Characters shared by males of

this group include the presence of a median pit filled with hairs on the second

abdominal stemite, the presence of an apical fringe of hairs on the last tergite,

the absence of a prominent tooth on the posterior trochanters, normal size ocelli

and eyes, cylindrical posterior tibiae, and rounded apices on the middle and hind

femora.

The subfamily Sphaeropthalminae, which contains two tribes, is cosmopolitan

in distribution. The tribe Dasylabrini is found primarily in the Old World, and

the tribe Sphaeropthalmini occurs primarily in the New World. The New World

distribution of the Sphearopthalmini may have resulted from a single introduction

of an ancestral sphearopthalmine from Africa into South America at about the

time that these two continents split apart about 110 million years ago (Brothers

1975). After becoming established in South America, the tribe apparently radiated

extensively and rapidly, establishing two basic lines, or subtribes, the Pseudome-

thocina and Sphaeropthalmina (to which Dasymutilla belongs).

The origin of Dasymutilla is not known. However, based upon its present

distribution, it may have radiated from an ancestral Sphaeropthalmina in the

Central American region after the two continents were reunited during the Cre¬

taceous period (Dietz & Holden 1970), with its distribution being mostly north¬

ward into North America. There are at least three extant species of Dasymutilla

that are found in the Dominican Republic. At least one other species is found in

Cuba. There is very little known about how the Caribbean region was formed.

However, the land forms presently in the Caribbean, including the Dominican

1991

MANLEY & POINAR: A NEW FOSSIL MUTILLID

203

Figures 1-3. Holotype of Dasymutilla dominica Manley & Poinar. Figure 1. Dorsal view. Figure

2. Lateral view. Figure 3. Ventral view of abdomen showing median pit filled with hairs on second

stemite (arrow).

204

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Republic, may have separated from the Central American region after the evo¬

lution of Dasymutilla and prior to their moving to their present location about

120 to 70 million years ago (LePichon 1968). Dasymutilla is the most widely

distributed and highly radiated (with about 150 species) group of mutillids in

North America (at least of the diurnal mutillids). The new Dasymutilla we describe

here is a relatively highly derived species, because of the pit on the second ab¬

dominal stemite. Thus, it seems likely that the genus radiated from a more prim¬

itive Sphaeropthalmina well over 40 million years ago.

Acknowledgment

We thank Aldo and Didi Costa for loan of the piece of amber containing the

specimen from which the description is made. We also thank Frank M. Carpenter

for informing the authors of the current status of Cretavus sibiricus. Technical

contribution no. 3099 of the South Carolina Agricultural Experiment Station,

Clemson University.

Literature Cited

Ashmead, W. H. 1899. Super-families in the Hymenoptera and generic synopses of the families

Thynnidae, Myrmosidae and Mutillidae. J. New York Entomol. Soc., 7: 45-60.

Bischolf, H. 1915. Bemsteinhymenopteren. Schrift. Phys.-Oekon. Ges., Konigsberg, 56: 139-144.

Brischke, C. G. A. 1886. Die Hymenopteren des Bernsteins. Schr. Nat. Ges. Danzig, N.F., 6: 278-

279.

Brothers, D. J. 1975. Phylogeny and classification of the aculeate Hymenoptera, with special reference

to Mutillidae. Univ. Kansas Sci. Bull., 50: 483-648.

Dietz, R. S. & J. C. Holden. 1970. The breakup of Pangaea, pp. 126-137. In Wilson, J. T. 1976.

Continents adrift and continents aground. W. H. Freeman, San Francisco.

1991

MANLEY & POINAR: A NEW FOSSIL MUTILLID

205

Krombein, K. Y. 1979. Biosystematic studies of Ceylonese wasps, IV. Kudakrumiinae, a new sub¬

family of primitive wasps (Hymenoptera: Mutillidae). Trans. Am. Entomol. Soc., 105: 67-83.

Lambert, J. B., J. S. Frye & G. O. Poinar, Jr. 1985. Amber from the Dominican Republic: analysis

by nuclear magnetic resonance spectroscopy. Archaemetry, 27: 43-51.

Larsson, S. G. 1978. Baltic amber—a palaeobiological study. Entomonograph Vol. 1. Scandinavian

Science Press, Klampenborg, Denmark.

LePichon, X. 1968. Sea-floor spreading and continental drift, pp. 89-121. In Cox, A. 1973. Plate

tectonics and geomagnetic reversals. W. H. Freeman, San Francisco.

Menge, A. 1856. Lebenszeichen vorweltlicher, im Bernstein eingeschlossener Tiere. Programm Pe-

trischule, Danzig, 1-32.

Mickel, C. E. 1928. Biological and taxonomic investigations on the mutillid wasps. U.S. Nat. Mus.

Bull., 143: 1-351.

Rasnitsyn, A. P. 1975. Hymenoptera Apocrita of the Mesozoic. Trudy Paleontol. Inst. Akad. Nauk

SSSR, 147: 1-140.

Scudder, S. H. 1891. Index to the known fossil insects of the world including myriapods and arachnids.

Bull. U.S. Geol. Survey, 71.

Sharov, A. P. 1957. First discovery of a Cretaceous aculeate hymenopteron. Dokl. Akad. Nauk

SSSR, 112: 943-944.

Received 15 November 1990; accepted 21 March 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 206-215, (1991)

A BIOGEOGRAPHICALLY BASED ASSESSMENT OF THE

POTENTIAL MAYFLY FAUNA OF NEVADA

Richard K. Allen 1 and Chad M. Murvosh 2

4048 La Sombra Drive, Lake San Marcos, California 92069

department of Biological Sciences, University of Nevada,

Las Vegas, Nevada 89154

Abstract. — A faunal assessment of the mayflies of Nevada is in progress. We hypothesize the

number and composition of species and genera expected for Nevada based on the known bio¬

graphical distribution of western species. Sixteen species and 12 genera are presently known from

Nevada, but we hypothesize the total number of species and genera will be 74 and 32, respectively.

Biogeographical patterns reveal where in the state species should be found.

Key Words. — Insecta, mayflies, Nevada, geographic distribution, biogeography, Ephemeroptera

The mayflies (Ephemeroptera) of North and Central America are now well

known and the species composition of only a few areas on the continent are poorly

reported. The most poorly known region of the Nearctic is Nevada, where only

15 species have been reported. Mayflies of most states surrounding Nevada (Ar¬

izona, California, Idaho, Oregon and Utah) had been thoroughly studied. Much

of this literature is scattered in many individual papers, but some summary

references include Allen & Edmunds (1956) for Oregon, Day (1956) for California,

Edmunds (1954) for Utah, Edmunds et al. (1976) and Edmunds (1984) for North

America, Jensen (1966) for Idaho and Kilgore & Allen (1973) for Arizona. We

predict the potential mayfly fauna for Nevada based upon the biogeographic

distribution of species in western North America. Allen (1990) has published the

distribution patterns of North and Central American mayflies; we refer to his

distribution patterns in this paper.

Collections of Nevada mayflies are now in progress to corroborate our hypoth¬

esis and also to extend the distributional range and limits of western North Amer¬

ican species.

Physiography of Nevada. — Nevada is a large state which has been poorly col¬

lected. The state is about 790 km long and 512 km wide. Drainage from the

southeastern comer of Nevada is to the Gulf of California via the Colorado River,

but most of the remainder of the state lies within The Great Basin. This is a

geographic/hydrologic region where no surface water leaves except by evaporation.

The usage of the term “Great Basin” has been confused at times by biologists

and anthropologists (Fiero 1986). The Great Basin lies in the northern part of a

much larger geologic province called the Basin and Range. A bird’s eye view of

Nevada would see many uplifted blocks (horsts) forming the mountain ranges

separated by broad elongated valleys or basins formed by down dropped blocks

(grabens). The long axes of the valleys and ranges generally trend north-south.

The valley floors are quite high, generally between 1200 and 1500 m above sea

level. The mountains may be 3900 m high (Fiero 1986).

Lentic habitats dominated the landscape during former pluvial periods (Hubbs

& Miller 1948), but these ecosystems are relatively uncommon today compared

1991

ALLEN & MURVOSH: MAYFLIES OF NEVADA

207

to the many lotic systems present. Two important river systems include the

Humboldt River in northern Nevada and the Colorado River in southern Nevada

but the latter has been largely transformed into a lentic habitat (Lake Mead).

There are some smaller rivers, but these are overshadowed in number by the

numerous small streams that flow down the mountain canyons. Few of these can

be seen from the basin highways and many are difficult to get to, often requiring

high clearance vehicles and/or four wheel drive. As a result, it may be a long time

before we have an accurate picture of the aquatic insect fauna of the state.

Mayfly Distribution Patterns. — The distributions of mayflies suggest that North

and Central America can be divided into five biogeographic subdivisions with

regard to Ephemeroptera (Allen 1990): Arctic, Northern American, Western North

American, Eastern North American, and Mesoamerican. The “Arctic Subdivi¬

sion” includes species that are distributed about 58° North Latitude. The “North

American Subdivision” includes species that are widely distributed in all of North

America. The “Eastern North American Subdivision” includes species that are

restricted to western or eastern North America in Mexico, the United States, and

Canada. The “Mesoamerican Subdivision” includes species that are distributed

mainly in tropical Mexico and Central America, but may occur as far north as

the southern U.S.

North American Subdivision

This subdivision includes two distinct distribution patterns, the “Pancontinen-

tal” and the “Widespread North American.” Both patterns include species known

in Nevada, and species expected there.

Pancontinental Distribution Pattern. — Ten species occur across North America,

except in the southwest deserts of the U.S. Stenonema terminatum (Walsh) is

now known from Elko Co., Caenis latipennis Banks, which is presently known

from only western Oregon and Washington, is not expected to occur in Nevada.

Baetis hageni Eaton and B. tricaudatus Dodds eventually should be found widely

distributed in the state. Attenella margarita (Needham), Caenis simulans Mc-

Donnough, Ephemera simulans Walker, and Hexagenia limbata (Serville) should

be found in the northern parts of the state. Ephemerella aurivillii Bengtsson and

Paraleptophlebia debilis (Walker) should be found in the western and northern

parts of Nevada.

Widespread North American Distribution Pattern.— Four species are widely

distributed in the United States and Canada, but have narrower distributional

limits than the pancontinental species. These species are also absent from the

southwest deserts. Ephoron album (Say) and Heptagenia elegantula (Eaton) are

presently known in Nevada. Baetis quilleri Dodds, which has been reported from

California, Arizona, Utah and Idaho, should be found in Nevada. Pseudiron

centralis McDunnough is known from only southern Canada, eastern Utah and

Wyoming, and the species is not expected in Nevada.

Western North American Subdivision

This subdivision includes nine distribution patterns which include species that

are presently known to occur in Nevada, and/or expected to occur in the state.

Widespread Western Distribution Pattern. — This pattern (Fig. 1) includes seven

species, of which six are known in Nevada: Drunella ( Myllonella ) coloradensis

208

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(3)

southwestern Alberta, Washington, most of Oregon, eastern California except for southern desert,

Idaho, western Montana, Wyoming, western Colorado, and New Mexico, eastern Arizona, northeastern

and northwestern Nevada, and most of Utah), Western United States (Washington, Oregon, and

California, south to near Los Angeles, northern Nevada, most of Idaho, western Montana, Wyoming,

Colorado and New Mexico, eastern Arizona and most of Utah, southern British Columbia and south¬

western Alberta), Northwest and Coast Ranges (Washington, Oregon, most of California south to about

1991

ALLEN & MURVOSH: MAYFLIES OF NEVADA

209

(Dodds), D. (Eatonella) doddsi Needham, D. ( Drunella) grandis (Eaton), Ephem-

erella inermis Eaton, Iron longimanus (Eaton), and Tricorythodes ( Tricorythodes)

minutus Traver. The distribution of Baetis bicaudatus Dodds (California, Idaho,

Oregon, and Utah) suggests it will eventually be found in Nevada.

Western United States Distribution Pattern. — This (Fig. 1) is composed of eight

species with four known from Nevada: Rhithrogena hageni Eaton, R. morrisoni

(Banks), Serratella tibialis (McDunnough), and Timpanoga hecuba Eaton, are

presently known from Nevada. Baetis insignificans McDunnough and Paralep-

tophlebia memorialis (Eaton) occur in all of the western states surrounding Nevada,

and Ephemerella infrequens McDunnough and Iron albertae McDunnough occur

in all of the surrounding states except Arizona. Eventually, all of these species are

expected to be found in Nevada.

Northwest Distribution Pattern. — This (Fig. 2) includes 71 species, none of which

are known in Nevada. As the name suggests, all species are distributed in the

northern reaches of western North America. Four are known to occur in southern

Idaho and Oregon and should occur in Nevada: Choroterpes albiannulatus

McDunnough, Paraleptophlebia heteronea (McDunnough), Rhithrogena futilis

(McDunnough), and Dactylobaetis cepheus Traver & Edmunds.

Southwest Distribution Pattern.— This (Fig. 2) includes 42 species from the

southwest deserts to west-central Mexico. It is impossible to hypothesize which

of these species will eventually be found in Nevada. Only three species of Lep-

tohyphes ( L. apache Allen, L. packeri Allen, and L. quercus Allen) are possible

contenders, because they occur in central Arizona. Leptohyphes apache also occurs

in the southwest comer of Utah.

Coast Ranges Distribution Pattern. — This (Fig. 3) includes 46 described species,

of which two, Baetis parallelus Banks and Ephemerella rama Allen, were described

from Nevada. Fourteen of the other 44 species, all of which are presently known

from the Sierra Nevada Mountains in California, are expected to be found, even¬

tually, in western Nevada: Ameletus amador Mayo, Attenella delantala (Mayo),

Baetis palisadi Mayo, Cinygmula tioga Mayo, Edmundsius agilis Day, Eury-

lophella lodi (Mayo), Iron dulciana (McDunnough), I. lepidus (Traver), Para-

cloeodes abditus Day, Paraleptophlebia placeri Mayo, Serratella levis (Day), S.

sequoia (Allen & Collins), S. velmae (Allen & Edmunds), and Siphlonurus spec-

tabilis Traver. The remaining species are not expected to occur in Nevada and

are known only from the coast ranges of the Pacific.

Rocky Mountain Distribution Pattern. — This (Fig. 3) is composed of 11 species,

and only Heptagenia solitaria (McDunnough) is expected in Nevada. This species

is known as far west as southern Oregon. The other species are widely distributed

in the Rocky Mountains from New Mexico to Canada, but do not occur west of

eastern Utah.

half of Baja California, Idaho, western Montana, northwestern Wyoming, southeastern British Co¬

lumbia and southwestern Alberta), and the Northwest and Rocky Mountain (western Northwest Territories

south, through eastern British Columbia and western Alberta, Oregon, northern California and eastern

Washington, Idaho, western Montana, Wyoming, Colorado and New Mexico, very northern Mexico,

eastern Arizona and northern and eastern Utah).

210

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Figure 2. Composite patterns for the following mayfly distributions: Northwest (Washington, Or¬

egon, Idaho, western Montana and Wyoming, northern Nevada, Utah, and northwestern Colorado,

British Columbia, western Alberta and Northwest Territories), Southwest (Southern California and

Nevada, Arizona, southern New Mexico, Baja Mexico and northwestern Mexico), Widespread Meso-

american (southeastern Arizona, western New Mexico, western Mexico, most of Central America

to South America, eastern Mexico into south central Texas).

1991

ALLEN & MURVOSH: MAYFLIES OF NEVADA

211

Figure 3. Composite patterns for the following mayfly distributions: Coast Ranges (western Wash¬

ington and Oregon and California south to near Los Angeles, part of central western tip of Nevada,

southwestern British Columbia), Rocky Mountain (western Northwest Territories, Alberta, Montana,

Wyoming, Colorado, and New Mexico, eastern British Columbia, Washington, Idaho, Utah and

Arizona), and Disjunct Western (southwestern Washington and Oregon, California south to most of

Baja California, parts of central and southeastern Arizona, central New Mexico, most of Colorado,

Wyoming and parts of central Montana into southwestern Alberta).

212

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(3)

Table 1. List of the mayflies currently reported from Nevada.

Family

Species

Locality

Baetidae

Baetis parallelus Banks

Reno (Type Locality)

Ephemerellidae

Drunella ( Myllonella ) coloradensis (Dodds)

Elko Co., Washoe Co.

Drunella ( Eatonella ) doddsi Needham

Elko Co., Washoe Co.

Drunella ( Drunella) grandis (Eaton)

Elko Co., Washoe Co.

Ephemerella inermis Eaton

Elko Co.

Ephemerella rama Allen

Reno (Type Locality)

Serratella tibialis (McDunnough)

Elko Co.

Timpanoga hecuba (Needham)

Carson River

Heptageniidae

Heptagenia elegantula (Eaton)

Reno

Iron longimanus (Eaton)

Washoe Co.

Rhithrogena hageni Eaton

Reno

Rhithrogena morrisoni (Banks)

Reno

Stenonema terminatum (Walsh)

Elko Co.

Oligoneuridae

Isonychia intermedia

Clark Co. Moapa River

(unpublished)

Palingeniidae

Ephoron album (Say)

Elko Co.

Tricorythidae

Tricorythodes (Tricorythodes) minutus

Traver

Elko Co.

Northwest and Coast Ranges Distribution Pattern. — This (Fig. 1) also includes

11 species, and two, Ametropus ammophilus Allen & Edmunds, and Centroptilum

conturbatum McDunnough, are not expected in Nevada; their known distribution

is from the coast ranges in the U.S. and Canada. The following species are known

to occur in the Sierra Nevada of California and should eventually be found in the

western part of Nevada: Ameletus validus (McDunnough), Caudatella heterocau-

data (McDunnough), C. hystrix (Traver), Drunella ( Myllonella ) flavilinea

(McDunnough), Drunella {Drunella) spinifera (Needham), Ironodes nitidus (Ea¬

ton), Ironopsis grandis (McDunnough), Serratella teresa (Traver), and Siphlonurus

columbianus McDunnough.

Northwest and Rocky Mountain Pattern. — This (Fig. 1) includes eight species

distributed within its geographical limits, and all of them are expected in Nevada.

Five species, Callibaetis coloradensis Banks, Iron deceptivus (McDunnough), Nixe

( Akkarrion ) criddlei (McDunnough), Rhithrogena undulata (McDunnough), and

Siphlonurus occidentalis Eaton, occur in southern Idaho and southern Oregon.

Callibaetis nigritus Banks is known from southern Idaho, but not from Oregon,

and Nixe (Akkarion) simplicioides (McDunnough) occurs in southern Idaho and

northern Oregon. Traverella albertana (McDunnough) occurs in southern Idaho

and also north central Arizona, and southeastern Utah. Siphlonurus occidentalis

also occurs in northern California, Utah and central Arizona.

Disjunct Western Distribution Pattern. — This (Fig. 3) presently includes six

species, of which four, Cinygmula par (Eaton), C. mimus (Eaton), Rhithrogena

flavianula (McDunnough) and Serratella micheneri (Traver), occur in the Sierra

Nevada of California. These species should eventually be found in western Ne¬

vada. Baetis adonis Traver, known from New Mexico and southern California,

and Tricorythodes {Homoleptohyphes ) dimorphus Allen, known from eastern Ar¬

izona and southern California, are not expected in Nevada.

1991

ALLEN & MURVOSH: MAYFLIES OF NEVADA

213

Table 2. List of the mayflies predicted to be collected in Nevada.

Family

Species

Area of State

Baetidae

Baetis bicaudatus Dodds

Widespread

Baetis hageni Eaton

Widespread

Baetis insignijicans McDunnough

Widespread

Baetis palisadi Mayo

Western

Baetis quilleri Dodds

Widespread

Baetis tricaudatus Dodds

Widespread

Callibaetis coloradensis Banks

Northern

Callibaetis nigritus Banks

Northern

Dactylobaetis cepheus Traver & Edmunds

Northern

Paracloeodes abditus Day

Western

Caenidae

Caenis simulans McDunnough

Northern

Ephemerellidae

Attenella delantela (Mayo)

Western

Attenella margarita (Needham)

Northern

Caudatella heterocaudata (McDonnough)

Western

Caudatella hystrix (Traver)

Western

Drunella ( Myllonella ) fiavilinea (McDunnough)

Western

Drunella ( Drunella ) spinifera Needham

Western

Ephemerella aurivillii Bengtsson

Northern & Western

Ephemerella infrequens McDunnough

Widespread

Eurylophella lodi (Mayo)

Western

Serratella levis (Day)

Western

Serratella micheneri (Traver)

Western

Serratella sequoia (Allen & Collins)

Western

Serratella teresa (Traver)

Western

Serratella velmae (Allen & Edmunds)

Western

Ephemeridae

Ephemera simulans McDunnough

Northern

Hexagenia limbata (Serville)

Northern

Heptageniidae

Cinygmula mimus (Eaton)

Western

Cinygmula par (Eaton)

Western

Cinygmula tioga Mayo

Western

Heptagenia solitaria (McDunnough)

Northern

Iron albertae (McDunnough)

Widespread

Iron deceptivus (McDunnough)

Northern

Iron dulciana (McDunnough)

Western

Iron lepidus (Traver)

Western

Ironodes nitidus (Eaton)

Western

Ironopsis grandis (McDunnough)

Western

Nixe (A kkarrion) criddlei (McDunnough)

Northern

Nixe ( Akkarrion ) simplicioides (McDonnough)

Widespread

Rhithrogena flavianula (McDunnnough)

Western

Rhithrogena futilus McDunnough

Northern

Rhithrogena undulata Banks

Northern

Leptophlebiidae

Choroterpes albiannulatus McDunnough

Northern

Choroterpes intermedia McDunnough

Southern

Paraleptophlebia debilis (Walker)

Northern & Western

Paraleptophlebia heteronea (McDunnough)

Northern

Paraleptophlebia memorialis (Eaton)

Widespread

Paraleptophlebia placeri Mayo

Western

Traverella albert ana (McDunnough)

Widespread

Siphlonuridae

Ameletus amador Mayo

Western

Ameletus validus McDunnough

Western

Edmundsius agilis Day

Western

Siphlonurus columbianus McDunnough

Northern

214

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Table 2. Continued.

Family

Species

Area of State

Siphlonurus occidentalis Eaton

Widespread

Siphlonurus spectabilis Traver

Western

Tricorythidae

Leptohyphes apache Allen

Southern

Leptohyphes packeri Allen

Southern

Leptohyphes quercus Allen

Southern

Mesoamerican Subdivision

This subdivision includes two patterns the “Mesoamerican” and the “Wide¬

spread Mesoamerican,” the latter may include species that eventually will be

found in Nevada.

Widespread Mesoamerican Distribution Pattern.— This (Fig. 2) includes nine

species that are distributed from tropical Mexico and Central America to the

southern United States. Two of these, the boreal Choroterpes intermedia Mc-

Dunnough and the austral Leptohyphes packeri Allen, are expected to be found,

eventually, in Nevada. The former species is known from western Arizona to

Mexico. The latter is the most widely distributed species in the Americas, from

Honduras to southwestern Arizona. The other species that are distributed in this

pattern do not occur north of southern Arizona and New Mexico and have not

been found north of their presently known distributional limits.

Conclusions

The mayfly fauna of Nevada (Table 1) presently includes 16 species in 12 genera.

We postulate that an additional 20 genera and 58 species will be found in the

state (Table 2) based upon biogeographic assessments of surrounding states. The

order of listing in Tables 1 and 2 are alphabetical, not phylogenetic. The total

potential number of genera and species should be 32 and 74, respectively. We

further hypothesize that 26 species will be found in western Nevada; 15 in northern

Nevada; four in the southern part of the state; two in the north and west; and 11

species will be found to be widespread. Future collections are expected to cor¬

roborate these predictions confirming the use of biogeographic information in

assessing the state’s fauna as a whole, and that of its partitioned regions.

Species of different genera and subgenera can be expected to be found in the

following habitats: Ameletus, Heptagenia, Iron, and Rhithrogena in small rapid

streams; Edmundsius in slow flowing, shallow streams; Caenis, Callibaetis, and

Siphlonurus in lentic situations; Attenella, Choroterpes, Ephemerella, Caudatella,

Drunella, Eurylophella, Serratella, Ephemera, and Hexagenia in a variety of lotic

and lentic habitats; Dactylobaetis, Cinygmula, Paraleptophlebia, and Leptohyphes

in a variety of lotic habitats; Paracloeodes in large streams; Baetis species in

streams that range from slow flowing to torrential. Detailed habitat descriptions

can be found in Edmunds et al. (1976) and Edmunds (1984).

Acknowledgment

We thank Dick Baumann for all the help he has given us in this study. The

comments of two anonymous reviewers were useful and improved the manuscript

in several ways.

1991

ALLEN & MURVOSH: MAYFLIES OF NEVADA

215

Literature Cited

Allen, R. K. & G. F. Edmunds Jr. 1956. A list of the mayflies of Oregon. Proc. Utah Acad. Sciences,

Arts and Letters, 33: 85-87.

Allen, R. K. 1990. Distribution patterns of North and Central American mayflies (Ephemeroptera).

pp. 155-167. In Campbell, I. C. (ed.). Mayflies and stoneflies. Life histories and biology. Kluwer

Academic Publishers, Dordrecht, Netherlands.

Day, W. C. 1956. Ephemeroptera. pp. 79-105. In Usinger, R. L. (ed.). Aquatic insects of California.

University of California Press, Berkeley.

Edmunds, G. F. Jr. 1954. The mayflies of Utah. Proc. Utah Acad. Science, Arts and Letters, 31:

64-66.

Edmunds, G. F. Jr. 1984. Ephemeroptera. pp. 94-125. In Merritt, R. W. & K. W. Cummins (eds.).

An introduction to the aquatic insects of North America (2nd ed.). Kendall Hunt, Dubuque,

Iowa.

Edmunds, G. F. Jr., S. L. Jensen & L. Berner. 1976. The mayflies of North and Central America.

University of Minnesota Press, Minneapolis.

Fiero, B. 1986. Geology of the Great Basin. University of Nevada Press, Reno.

Hubbs, C. L. & R. R. Miller. 1948. The zoological evidence: correlation between fish distribution

and hydrographic history in the desert basins of western United States. In The Great Basin

with emphasis on glacial and postglacial times. Bull. Univ. Utah (Biol. Ser. 10), 38: 17-166.

Jensen, S. 1966. The mayflies of Idaho. Master’s thesis, University of Utah.

Kilgore, J. I. & R. K. Allen. 1973. Mayflies of the Southwest: new species descriptions, and records

(Ephemeroptera). Ann. Entomol. Soc. Am., 66: 321-332.

Received 31 December 1990; accepted 22 March 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 216-221, (1991)

A NEW SPECIES OF UROLEUCON ( UROLEUCON) AND

COMMENTS ON UROLEUCON (UROLEUCON) TUCUMANL

(ESSIG) (HOMOPTERA: APHIDIDAE ) 1

Miguel Angel Delfino

Instituto Superior de Entomologia, Facultad de Ciencias Naturales,

Universidad Nacional de Tucuman,

Miguel Lillo 205, 4000-Tucuman, Argentina

Abstract. — Uroleucon ( Uroleucon ) garnicai NEW SPECIES on Eupatorium buniifolium Hooker

& Arnold from Argentina is described. Morphological and biological aspects were analyzed

together with those of its closely related species Uroleucon ( Uroleucon) tucumani (Essig). The

study of the type specimens of U. tucumani shows that they could belong to two different species;

those collected in Tucuman are true U. tucumani.

Key Words. — Insecta, Uroleucon garnicai, Uroleucon tucumani, Aphididae, morphometries, dis¬

criminant analysis

Uroleucon live on plants in the Asteraceae (= Compositae), and are broadly

distributed throughout the world with some almost cosmopolitan species. Smith

& Cermeli (1979) list 23 species in the Neotropical region, seven of which were

described from Argentina. Of these, Uroleucon (Uroleucon ) tucumani (Essig) was

described by Essig (1953) as a Macrosip hum from specimens collected on Bac-

charis sp. in Huanta (Guanta), Chile and on Baccharis sp. (?) in Tucuman province,

Argentina by Michelbacher.

All the aphids collected on Baccharis coridifolia De Candolle in Cordoba prov¬

ince, Argentina, as well as those collected on Eupatorium buniifolium Hooker &

Arnold in Cordoba and Tucuman provinces, had been previously considered to

be U. tucumani. Essig’s description is very brief and generalized, making it im¬

possible to recognize accurately this, and related, species.

To solve this problem, a detailed study of the Uroleucon collected on B. cori¬

difolia and E. buniifolium was undertaken, and showed two species exist with one

described as new here.

Materials and Methods

Aphids belonging to the genus Uroleucon were collected on E. buniifolium and

B. coridifolia in Tafi del Valle (Tucuman, Argentina) and Cabana (Cordoba, Ar¬

gentina). Adult aphids were mounted on slides in Hoyer’s mountant. Nineteen

body structures (Table 1) were measured from 60 apterae (30 of each host plant).

The data were analyzed on a VAX-11 computer using the discriminant analysis

program in the BMDP statistical package, at the University of California.

During comparative studies, the types of U. tucumani (Essig) were analyzed.

Their data follow: “Type,” ARGENTINA. TUCUMAN: 11 Feb 1951, Michel-

backer, Baccharis sp.?, 1 alate viviparous female. Paratypes: same data as type,

4 apterous viviparous females; CHILE. HUATA (GUANTA): 4 Dec 1950, Mich-

elbacker, Baccharis sp., 4 apterous viviparous females, 3 alate viviparous females.

1 Published with a grant from the C. P. Alexander Fund, Pacific Coast Entomological Society.

1991

DELFINO: NEW SPECIES OF UROLEUCON

111

Table 1. Morphometric data for U. garnicai and U. tucumani. All measurements are in mm as

range, followed by the mean in parentheses.

Character

U. garnicai

U. tucumani

Body length

1.92-2.67 (2.30)

1.86-2.59 (2.14)

Metatibia length

1.30-1.68 (1.51)

1.13-1.70 (1.41)

Metafemor length

0.75-0.85 (0.82)

0.65-0.93 (0.77)

Siphunculus length

0.49-0.62 (0.55)

0.44-0.60 (0.54)

Cauda length

0.31-0.38 (0.34)

0.24-0.31 (0.13)

Decimal proportion of reticulated

part of the siphunculus

0.21-0.27 (0.24)

0.20-0.29 (0.24)

Length of ant. seg. Ill

0.55-0.73 (0.64)

0.51-0.68 (0.61)

Length of ant. , seg. IV

0.50-0.62 (0.58)

0.33-0.53 (0.45)

Length of ant. seg. V

0.44-0.59 (0.50)

0.30-0.44 (0.40)

Length of base of ant. seg. VI

0.14-0.18 (0.17)

0.12-0.17 (0.14)

Length of processus terminalis

0.71-0.87 (0.80)

0.54-0.76 (0.67)

Length of ultimate rostral segment

0.13-0.15 (0.14)

Length of hind distitarsus

0.14-0.17 (0.16)

0.12-0.14 (0.13)

Diameter of ant. seg. Ill

0.032-0.035 (0.034)

0.030-0.037 (0.033)

Length of setae on ant. seg. Ill

0.030-0.040 (0.034)

0.017-0.027 (0.023)

Number of sensoria on ant. seg. Ill

4-13 (9)

7-15 (11)

Number of rostral setae

6-10 (8)

7-13 (9)

Number of caudal setae

9-12 (11)

8-11 (10)

Uroleucon ( Uroleucon) garnicai Delfino, NEW SPECIES

Types. — Holotype: apterous viviparous female; data: ARGENTINA. COR¬

DOBA PROVINCE: Cabana, 27 Jan 1988, M. A. Delfino, Eupatorium buniifoli-

um; deposited: Instituto Miguel Lillo, Tucuman. Paratypes: same data as holotype,

3 apterous viviparous females, 2 alate viviparous females; same data as holotype

except 28 Jan 1985, 17 apterous viviparous females. ARGENTINA. TUCUMAN

PROVINCE: Tafi del Valle, 2 Mar 1985, M. A. Delfino, Eupatorium buniifolium,

9 apterous viviparous females. All paratypes deposited: British Museum (Natural

History), London.

Apterous Viviparous Female.— {n = 30) Color: Living specimens with entire body brown. Cleared

specimens with dorsum colorless, bearing distinct separate brown sclerites; weakly sclerotized speci¬

mens with the sclerites poorly defined; head, antennal segments I, II and base of III brown; antennae

flagellum, rostrum, distal halves of the femora, distal fourth part of the tibiae, tarsi and siphunculi

dark brown to nearly black; cauda pale to slightly smoky. Morphological characters: Body length 1.92-

2.67 mm (Fig. 1A). Head smooth with antennal tubercles well developed, diverging; median frontal

tubercle inconspicuous (Fig. IE). Antennae with processus terminalis 4.28-5.31 x the base of segment

VI; flagellum almost smooth in the basal one-half of segment III and imbricate distally. Antennae

hairs on segment III 0.030-0.040 mm long, about 0.86-1.14 x as long as the basal diameter of segment.

Secondary rhinaria 6-13, distributed on nearly the whole length of antennal segment III (Fig. IB).

Rostrum reaching the hind coxae; ultimate rostral segment 0.13-0.15 mm; 0.81-1.00 x as long as the

hind distitarsus (Fig. 1H), with 7-10 additional hairs. Hind tibiae and femur 0.56-0.76 and 0.31-

0.40 x body length, respectively. First tarsal chaetotaxy usually 5:5:5. Dorsal hairs of abdomen on

small sclerites (Fig. ID); tergite VIII with 4 hairs. Postsiphuncular sclerites distinct, sometimes very

small; antesiphuncular sclerites not developed. Frequently small marginal tubercles, irregularly present

on segments II-IV. Siphuncular length 0.22-0.27 x body length (Fig. 1G), distinctly shorter than

antennal segment III, reticulated on distal 0.21-0.29, remainder imbricate. Cauda with distal two-

thirds rather slender, acuminate, bearing 9-12 hairs; about 0.56-0.68 x as long as the siphuncular

length. Genital plate with 2-4 (usually 2) hairs on anterior half and 8-12 hairs on the hind margin.

Measurements: See Table 1.

218

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

Figure 1. Uroleucon garnicai, NEW SPECIES. A. Body dorsum, right half (aptera). B. Secondary

rhinaria on antennal segment III (aptera), scale bar = 98 n. C. Secondary rhinaria on antennal segment

III (alate). D. Dorsal hair on abdomen with sclerite base (aptera), scale bar = 24 n. E. Frontal area of

head showing median frontal tubercle (aptera), scale bar = 98 n. F. Cauda (alate), scale bar = 40 n.

G. Siphunculus (aptera), scale bar = 40 yu- H. Hind distitarsus (aptera), scale bar = 24 n.

1991

DELFINO: NEW SPECIES OF UROLEUCON

219

0 00

0 0 0 0 0 0000

■A- ■ . 1——-1 >

0 0

o 0

00 OOOO 0 0 0 0

-ft—

®e> ©©©©«• oo

x— ft ft i

Figure 2. Standardized discriminant function showing canonical variable distributions for U. tucu¬

mani and U. garnicai. On this function, the unstandardized discriminant index, L OJ of 3.33 equals 0

on this standardized function. Black = U. garnicai, white = U. tucumani.

Alate Viviparous Female.—(n = 2) Color: Living specimens with head, antennae, thorax and legs

dark brown, abdomen brown. Cleared specimens with head and thorax brown; antennal segment I,

II and base of III concolorous with the head, remainder dark brown; legs with basal one-half of femora

pale, distal one-half dark brown; tibiae dark brown except basally slightly pale; tarsi concolorous with

apical tibiae; marginal sclerites small, on poorly pigmented specimens; siphunculi uniformly dark

brown; cauda, genital, and anal plates dusky, or slightly darker than abdomen. Morphological char¬

acters: Body length 2.39-2.48 mm. Antennae with processus terminalis 4.14-4.61 x the base of segment

VI. Hairs of antennal segment III 0.030-0.032 mm long, about 0.94-1.07 x the length of the basal

diameter of segment. Secondary rhinaria on segment III about 8-14 in number covering almost its

entire length (Fig. 1C). Ultimate rostral segment 0.14 mm; 0.78-0.86 x the length of the hind distitarsus,

with 7-8 additional hairs. Hind tibiae and femur 0.62-0.73 and 0.32-0.37 x body length, respectively.

Dorsal hairs of abdomen on small sclerites poorly pigmented. Siphunculi 0.22-0.26 x body length,

reticulated over apical 0.23-0.24, remainder imbricate. Cauda about 0.53-0.60 x siphuncular length

(Fig. IF). Otherwise like apterous viviparous females.

Diagnosis.— Because U. garnicai overlaps morphologically with U. tucumani,

a discriminant function must be calculated to separate these species reliably. See

Table 2, the discussion for character overlap and the diagnostic function.

Etymology. — This species is named after Rafael Gamica, Universidad de Leon,

Spain, who taught me so much on our field trips in that country.

Material Examined. —See types.

Discussion

Morphology. — The correlation matrices for both U. garnicai and U. tucumani

show that the lengths of the metafemur, antennal segment III and cornicle (si-

phunculus) are highly correlated; therefore, I consider only one of these characters

for the morphological analysis of the specimens. The use of discriminant analysis

between the two species shows that the lengths of the cauda, antennal segment

III, antennal segment IV, setae on antennal segment III and the base of antennal

segment VI are five characters that could differentiate the two species.

Table 2. Differences between U. garnicai and U. tucumani. Measurements are in mm as range

followed by the mean within parentheses.

Character

U. garnicai

U. tucumani

Color in life

brown

black

Cauda length

0.31-0.38 (0.34)

0.24-0.31 (0.28)

Processus terminalis length

0.71-0.87 (0.80)

0.54-0.76 (0.67)

Hind distitarsus length

0.14-0.17 (0.16)

0.12-0.14 (0.13)

Setae length of ant. seg. Ill

0.030-0.040 (0.034)

0.017-0.027(0.023)

Host plant

E. buniifolium

B. coridifolia

Linear discriminant function (L 0 3.33)

(see discussion section).

L g = 3.96

L t = 2.69

Ratio (see Fig. 3)

13.23-18.82

8.50-12.96

220

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

• U. garnicai

o U. tucumani

mm)

• »• •• •*

• •••

• •• •

0 0

• •

°0

£ .30

0 o

0 0 0 o 00

< .25

0 0

2 3 4 5 6

Ratio of setae on antennal III vs. distitarsus length

Figure 3. Regression of caudal length plotted against a ratio of the number of setae on antennal

segment III versus distitarsal length. White = U. tucumani, black = U. garnicai.

A linear discriminant function (L) distinguishes these species; for U. garnicai,

L g = 3.96; for U. tucumani, L t = 2.69; with discriminant index (L 0 ) of 3.33.

Starting from these values specimens belonging to these species could be correctly

classified according to the following function, Li.

Li = (7.4883 x X x ) - (4.2548 x X 2 ) + (1.8261 x X 3 )

+ (7.9576 x X 4 ) + (53.5169 x X 5 )

where: X x = cauda length; X 2 = length of antennal segment III; X 3 = length of

antennal segment IV; X 4 = length of base of antennal segment VI; X 5 = setae

length of antennal segment III. (Lengths are in mm.) The specimens are U. garnicai

if Li > 3.33, as L 0 , or are U. tucumani if Li < 3.33.

The histogram of the canonical variates (Fig. 2) shows the separation between

these two species. The setae length on antennal segment III x length of hind

distitarsus I also obtained values that discriminate these species; that value in U.

garnicai ranges from 4.8-6.4 and in U. tucumani it is 2.0-3.9. The ratio: (ant.

seg. Ill) x (hind distitarsus)/(cauda length), is 13.23-18.82 for U. garnicai and

8.50-12.96 for U. tucumani. These values are plotted in Fig. 3.

The length of hind distitarsus (Table 1) also seems to be a useful character to

separate these species univariately. However, three alatae and four apterae from

Chile (“Paratype” of U. tucumani) have tarsal lengths of 0.16-0.18 mm, similar

to U. garnicai-, however, one alatae from Tucuman (“Type” of U. tucumani) and

four apterae also from Tucuman (“Paratype” of U. tucumani) have tarsal lengths

of 0.11-0.13 mm, as do the specimens that I collected on B. coridifolia. Because

of this, although Essig’s “Type” and “Paratype” from Tucuman are necessarily

1991

DELFINO: NEW SPECIES OF UROLEUCON

221

“true” U. tucumani, as I collected on B. coridifolia, Essig’s “Paratype” from Chile

could be another very closely related species.

Furthermore, the apterae from Chile have a few secondary rhinariae (2-6) on

the antennal segment III and the processus terminalis is short in relation at the

base of antennal segment VI. Curiously, Essig drew two antennal segments III of

apterae and alatae when he described U. tucumani, one set with long setae and

the other with shorter ones. He obviously observed some differences in his material

that he did not explain in the description.

Biology. — Uroleucon garnicai colonizes the young shoots and stems of the in¬

florescences of E. buniifolium, a shrub widely distributed in Argentina. In Tafi

del Valle (1972 m el) this species forms crowded colonies that often cover several

stems; whereas, in Cabana (500-600 m el) it forms disperse and small groups of

1-3 adult aphids per group. This suggests that there could be different ecotypes

which are more or less suitable to U. garnicai, according to the geographic dis¬

tribution of E. buniifolium.

No sexuals of U. garnicai were found on this plant. Alate males were captured

in Moericke traps placed in Tafi del Valle; morphologically these males are fairly

similar to U. garnicai females. If the males are actually U. garnicai, considering

the abundance on E. buniifolium there, the possible origin of U. garnicai could

be in the highlands (e.g., Tafi del Valle) where its host plant is common. Uroleucon

garnicai seems to be completely monophagous on this single host species year-

round. It may be a holocyclical and monoecious species on E. buniifolium.

Uroleucon tucumani colonizes twigs and shoots on B. coridifolia, a shrub also

widely distributed in Argentina. In Cabana, its colonies contain only a few adults,

which fall readily when disturbed.

Both species are very closely related but are easily distinguished in nature by

their body color and host plant; U. garnicai is brown and lives on E. buniifolium,

whereas U. tucumani is black and lives on B. coridifolia.

Conclusions. —Uroleucon ( Uroleucon ) garnicai is a new species closely related

to Uroleucon ( Uroleucon ) tucumani (Essig). Both may be distinguished by bio¬

logical and morphological characters. The U. tucumani paratypes labelled by Essig

with specimens collected in Argentina and Chile could belong to two different

species. The specimens collected in Tucuman are the true U. tucumani.

Acknowledgment

The author is indebted to Lie. M. I. Torres de Plaza and Lie. L. M. Benitez de

Parra for statistical advice, and to P. H. Amaud, Jr. (California Academy of

Sciences, San Francisco) for sending the types of U. tucumani. Funding for pub¬

lication came from the C. P. Alexander Fund of the Pacific Coast Entomological

Society.

Literature Cited

Dixon, W. J. 1985. BMDP Statistical Software. University of California Press, Berkeley.

Essig, E. O. 1953. Some new and noteworthy Aphidae from western and southern South America.

Calif. Acad. Sci. Proc., 28: 59-164.

Smith, C. F. & M. M. Cermeli. 1979. An annotated list of Aphididae (Homoptera) of the Caribbean

Islands and South and Central America. N. C. Agr. Exp. Sta. Tech. Bull., 259: 1-131.

Received 6 May 1990; accepted 2 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 222-224, (1991)

Scientific Note

NOCTURNAL CLUSTERING OF THE SOLITARY WASP,

CHALYBION CALIFORNICUM (SAUSSURE)

(HYMENOPTERA: SPHECIDAE)

Chalybion californicum (Saussure) is a solitary wasp that provisions its nest

with spiders. It appears to make its nest in preexisting cavities, especially in

abandoned nests of the sphecid mud-daubers, Sceliphron. After provisioning the

nests, females close the entrance with mud (Bohart, R. M. & A. S. Menke. 1976.

Sphecid wasps of the world, a generic revision. University of California Press,

Berkeley). Chalybion californicum occurs throughout most of the United States,

and also has been reported from southern Canada, northern Mexico, Hawaii and

Bermuda (Bohart, R. M. & A. S. Menke. 1963. Univ. Calif. Publ. Entomol., 30:

91-182). This note confirms previous reports (Weiss, H. B. 1944. Entomol. News,

55: 270-271, Ward, G. L. 1972. Proc. Indiana Acad. Sci., 81: 177-181) that C.

californicum aggregates nightly in the same location in successive seasons.

We observed a dense cluster or “sleeping aggregation” (Evans, H. E. & E. G.

Linsley. 1960. Bull. So. Calif. Acad. Sci., 59: 30-37) of C. californicum that

appeared nightly beneath an overhanging eave of a ground-level apartment in

Riverside, California. The aggregation was first noticed in mid-June 1989 and

the number of individuals in the aggregation during the summer was estimated

to be about 50. This number declined noticeably in September, and the last

individual was seen on 13 Oct 1989. In late April 1990, the aggregation site was

examined nightly; no wasps were seen until 8 May when approximately 12 in¬

dividuals aggregated. The synchronous reappearance of the wasps after the long

winter absence was striking. The number of individuals fluctuated between three

and 16, during 8-29 May. On 2 June the number increased abruptly to 27, then

to 45-50 on 3 June. Through the remainder of June and July, the number of

individuals in the aggregation was estimated as 65-70. The number began to

decrease gradually in August; the rate of decrease was more rapid later in the

month than earlier. Between late September and mid October, the number of

individuals was down to about 20 and fluctuated up and down nightly. Between

the evenings of 14 and 15 Oct, the number dropped sharply from approximately

15 to seven. On the morning of 16 Oct, we marked the remaining wasps with

paint to see if they would overwinter and return again next year. This was the

last that the wasps were seen in 1990; however, it is uncertain if the disturbance

that we caused the wasps when we marked them hastened their departure from

the aggregation site.

Each year the aggregation site was the exact same spot on a 15 x 15 cm wooden

pillar where it joined with an eave approximately 2.4 m above the ground (Fig.

1). The eave was on the east side of the apartment and the wasps aggregated on

the north side of the pillar. In late afternoon, the wasps flew around the aggregation

site and some landed. Gradually more landed until all were settled one to several

hours before dark. In cooler weather (e.g., 8 May-1 June 1990, with daily max-

1991

SCIENTIFIC NOTE

223

Figure 1. Aggregating Chalybion californicum.

imum temperatures between 19 and 30° C), they generally aggregated earlier

(approximately 16:00-16:30 h); in hotter weather (e.g., 2-4 June 1990, maximum

temperatures between 35 and 38° C), they aggregated later, about 18:00-18:30 h.

During the period 8 May-4 June, the morning dispersal was not observed; how¬

ever, on cooler days they usually were not dispersed before 07:30 h, although on

hot days most had dispersed by 07:30.

At 22:00 h, 7 Sep 1989, the aggregation contained roughly 40-50 individuals

and we collected 10 for sex determination; of these, six were male and four were

female, demonstrating that both sexes were present. We did not observe any

mating behavior; however, because our observations were not continuous, we

cannot say for certain that mating did not occur.

Weiss (1944) initially documented the reappearance of C. californicum at the

same aggregation site in New Jersey in five successive summer seasons with

intervening winters when the wasps were absent. Weiss (1944) suggested that a

chemical aggregation cue (pheromone) persisted at the site from autumn to spring.

However, this does not seem likely because the pheromone would have to persist

throughout the winter and remain potent enough to attract wasps from a distance

in the spring. Nonchemical aggregation cues, such as visual cues, would be more

likely to persist from one season to the next. Alternatively, we suggest that the

aggregation in the spring may consist of the same individuals that were present

in the fall and that the overwintered wasps remembered the aggregation site in

the same manner that wasps learn and remember the location of nesting sites

(Evans, H. E. & M. J. W. Eberhard. 1970. The wasps. University of Michigan

Press, Ann Arbor). This may provide a more likely explanation for the synchro¬

nous reappearance of 12 wasps on 8 May 1990, as opposed to a scenario where

newly emerged wasps were searching for a nocturnal resting site and 12 wasps

were attracted to the same physical cue for the first time on the same night.

However, a weakness in our hypothesis is that C. californicum is reported to

overwinter as a prepupa (Rau, P. & N. Rau. 1916a. J. Anim. Behavior, 6: 27-

224

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 67(3)

63). To test our hypothesis, we marked some individuals in autumn 1990 and

will watch for their reappearance in the spring of 1991.

Nocturnal aggregations of several species of Chalybion have been reported: C.

californicum (= C. caeruleum [L.]) (Rau, P. & N. Rau. 1916b. Ann. Entomol.

Soc. Am., 9: 227-274; Rau, P. 1938. Ann. Entomol. Soc. Am., 31: 540-556; Weiss

1944; Ward 1972), C. zimmermanni Dahlbom (Rau, P. 1942. Can. Entomol.,

74: 196), C. bengalense (Dahlbom) (= C. violaceum [Fabr.]) (Williams, F. X.

1928. Hawaiian Sugar Planters’ Assoc., Entomol. Ser. Bull., 19: 1-179; Jayakar,

S. D. & R. S. Mangipudi. 1965. J. Bombay Nat. Hist. Soc., 61: 708-711), and C.

japonicum (Gribodo) (= Sceliphron inflexion Sickmann) (Iwata, K. 1963. Trans.

Shikoku Entomol. Soc., 7: 114-118). All four species are in the subgenus Chal¬

ybion; C. californicum and C. zimmermanni are New World species, and C.

bengalense and C. japonicum are Old World species (Bohart & Menke 1976). The

aggregations of these four species consisted of both sexes although no mating

behavior was observed by Rau & Rau (1916b) or Ward (19 72) or by us. Therefore,

the aggregations do not appear to facilitate mating. By day, Chalybion spp. are

solitary wasps; why they aggregate at night has remained a mystery since Rau &

Rau first described this behavior in 1916.

Acknowledgment. — We thank R. M. Bohart for his identification; a voucher

specimen from this study is deposited in the UCR Entomological Teaching and

Research Collection, University of California, Riverside.

T. R. Prentice and G. P. Walker, Department of Entomology, University of

California, Riverside, California 92521.

Received 20 July 1990; accepted 7 February 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 224-226, (1991)

Scientific Note

OCCURRENCE OF COCCINELLA SEPTEMPUNCTATA (L.)

(COLEOPTERA: COCCINELLIDAE) IN CENTRAL

BRITISH COLUMBIA

Coccinella septempunctata (L.), the seven spotted lady beetle (C7), is a Pa-

laearctic predator intentionally introduced into North America numerous times

since 1956 for the biological control of aphids (Angalet, G. W., J. M. Tropp &

A. N. Eggert. 1979. Environ. Entomol., 896-901; Schaefer, P. W., R. J. Dysart

&H. B. Specht. 1987. Environ. Entomol., 16: 368-373). During 1973 it was found

to be established in both northern New Jersey and Quebec (Angalet, G. W. & R.

L. Jacques. 1975. USDA Coop. Econ. Insect Rep., 25: 883-884; Larochelle, A.

1991

SCIENTIFIC NOTE

225

1979. Bull. Invert. Ins. Quebec, 1: 68-73). Since then, its range in North America

has rapidly expanded, both as a result of natural dispersal and redistribution by

biocontrol workers (Schaefer et al. 1987). This article reports a significant expan¬

sion of the known range of C. septempunctata to central British Columbia.

In early August, 1990, large numbers of beetles were noted on bareroot spruce

(Picea ) seedlings, caragana ( Caragana arborescens Lamarck) windbreaks, and weeds

alongside a greenhouse at a commercial conifer seedling nursery located at Ness

Lake near Prince George, British Columbia. Because of a concern for potential

damage to the seedlings, adults were collected and submitted to Pacific Forestry

Centre, Forestry Canada, for identification. Seven of these adults were C. septem¬

punctata. This identification was later confirmed by J. McNamara of Biosystemat-

ics Research Centre, Agriculture Canada, Ottawa, who also determined the re¬

maining beetles in the collection (2 females) to be Hippodamia sp. prob.

quinquesignata (Kirby). Voucher specimens of C7 (7 adults) and Hippodamia sp.

(2 adults) are deposited in the Forestry Canada, Forest Insect and Disease Survey

insect reference collection, Victoria, British Columbia.

The origin of this C7 population is enigmatic. As there have been no intentional

releases of C7 in central British Columbia, its presence at Ness Lake could have

resulted from either an adventive introduction or natural disperal. Although an

adventive introduction cannot be ruled out, other isolated recoveries of C7 in

North America indicate that natural dispersal is a credible mechanism to account

for its presence in the Prince George area. Coccinella septempunctata is an active

flier capable of long distance dispersal from release sites. It has been found on

Sable Island, Nova Scotia, more than 175 km from the nearest land (Schaefer et

al. 1987). As well, recent surveys in Saskatchewan and Manitoba have shown it

to be present in most agricultural areas of both provinces, with dispersal as far

north as Ile-a-La-Crosse (55°30'N) in the boreal forest of Saskatchewan, more

than 360 km from the nearest release site (Tumock, W. J., B. Timlick, J. F. Doane

& J. Soroka. 1990. Ag. Can. Res. Br. Biocontrol News, 3: 25-30). Further surveys

should be conducted in British Columbia to determine the distribution of C7.

Because releases of C. septempunctata in the Pacific Northwest have been con¬

fined to the Vancouver area of British Columbia, more than 520 km to the south

(Schaefer et al. 1987; B. D. Fraser, personal communication), and in agricultural

areas of Washington and Oregon, at least 750 km to the SSE (Schaefer et al. 1987;

Olkowski, W., A. Zhang & P. Thiers. 1990. IPM Practitioner, 12: 1-12) surveys

along the major drainages to the south (e.g., Fraser, Thompson, Okanagan and

Columbia rivers) would indicate if C7 has spread from more southerly popula¬

tions. Although dispersal from more southerly populations is more likely, the

recovery of C7 from a more northerly latitude in the boreal forest of Saskatchewan

indicates that dispersal into British Columbia from the northeast through the

Peace River drainage system cannot be discounted.

Material Examined. —Coccinella septempunctata: BRITISH COLUMBIA. Ness Lake (54°N,

123°05'W), about 25 km NW of Prince George, 16 Aug 1990, F. Donnelly, ex Picea and Caragana

arborescens, FIDS 90-1305-01.

Acknowledgment.— The cooperation of the staff of Ness Lake Nursery, Indus¬

trial Forest Services, Prince George and the assistance of Mr. R. Tumquist, For¬

estry Canada, Forest Insect and Disease Survey, Victoria, B.C., in surveying the

226

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 67(3)

site and Mr. R. W. Duncan and Drs. B. E. Callan and H. Moeck, Forestry Canada

for reviewing the draft manuscript are gratefully acknowledged.

L. M. Humble, Forestry Canada, Pacific Forestry Centre, 506 West Burnside

Road, Victoria, British Columbia, Canada V8Z 1M5.

Received 17 January 1991; accepted 31 January 1991.

PAN-PACIFIC ENTOMOLOGIST

67(3): 226-227, (1991)

Scientific Note

NOTES ON TWO PACIFIC COASTAL SPECIES OF

TENEBRIONIDAE (COLEOPTERA)

Scaphidema pictum Horn was transferred (Triplehom, C. A. 1961. Coleopt.

Bull., 15: 125-127) into Phaleromela, receiving the new name variegata Triple¬

hom, because pictum was preoccupied. At the time Phaleromela variegata was

known only from Oregon. Shortly thereafter, its range was reported as south¬

western British Columbia to Oregon, in sandy areas along river banks (Hatch, M.

1965. Beetles of the Pacific Northwest. Part IV. University of Washington Press,

Seattle). Subsequent collection records show that the range is more extensive,

including Idaho and California. The Idaho collections are from the western edge

(Idaho Co.) and the south central part (Blaine Co.) of that state (data below).

Probably, the beetles occur throughout the Snake River drainage, including the

western parts of Wyoming and Montana.

In California, several collections have been made (data below), where specimens

were taken from fine, dry, partially shaded sand near rivers or streams. Although

the beetles and their larvae inhabit dry sand, they are never more than a few

inches from moist sand and do not live long in the laboratory if water is withheld.

The sand bars and banks where they occur are often flooded during winter. It is

unknown how they survive this period. Collection localities include both the

coastal mountains and the Sierra Nevada, at elevations near sea level to almost

2000 m.

Phaleromela variegata (as Scaphidema pictum) and P. globosa (LeConte) (as

Phaleria ) were included in a list of Tenebrionidae (Papp, C. S. & W. D. Pierce.

1960. J. Kansas Entomol. Soc., 33: 154-156) found to be associated with stored

animal food in the Antelope Valley on the western edge of the Mojave Desert.

Phaleromela variegata appears to be strictly limited to the montane riparian

habitat described above and has not been recorded in barnyard situations. More¬

over, the closest geographic records are about 500 miles north. Phaleromela glo¬

bosa occurs strictly on maritime sand dunes from British Columbia to approxi-

1991

SCIENTIFIC NOTE

227

mately San Luis Obispo in central California, with no other records from interior

localities. Thus, both the occurrence of these species in the Mojave Desert and

their association with stored products need verification.

Diaperis californica Blaisdell, described in 1929, has remained rare in collec¬

tions. In a revisionary treatment (Triplehom, C. A. 1965. Proc. U.S. Nat. Mus.,

117: 349-458) only 14 specimens were recorded from four localities in Calaveras,

Placer, Madera and Fresno Counties. We provide additional locality records (see

below), with localities from the San Bernardino Mountains being outside the Sierra

Nevada and about 300 miles south of the previously known range.

Records. — Phaleromela pictum: CALIFORNIA. MENDOCINO Co.: Eel River Ranger Station, nr

Covelo, 9 Jun 1972; 4.8 km (3 mi) N of Branscomb, 18 May 1984. PLUMAS Co.: Humbug Creek,

4.8 km (3 mi) NW of Portola, 1584 m (5200 ft), 18 May 1982; 1.6 km (1 mi) S of Meadow Valley,

1173 m (3850 ft), 14 May 1982. SISKIYOU Co.: 14.4 km (9 mi) E of McCloud, 1067 m (3500 ft),

10 Jun 1974. TRINITY Co.: Hayfork Ranger Station, nr Hayfork, 23 May 1973; Forest Glen, 5 Jun

1988. IDAHO. BLAINE Co.: Big Woods River, 32 km (20 mi) N of Shoshone, 1 Jul 1966. IDAHO

Co.: Stites, 31 May 1975.

Diaperis californica: CALIFORNIA. CALAVERAS Co.: 6.4 km (4 mi) E of Murphy’s, 914 m (3000

ft), 12 Jul 1963. EL DORADO Co.: Blodgett Experimental Forest, 20.8 km (13 mi) E of Georgetown,

21 Jun 1981, blacklight. MARIPOSA Co.: El Portal, 16 Jun 1961, ex Laetoporis sulphureus (Bulliard

ex Fries). Yosemite National Park, Tenaya Canyon, 960 m (3150 ft), 17 May 1928. SAN BERNAR¬

DINO Co.: Crestline, 1494 m (4900 ft), 5 Jul 1968; Lake Arrowhead, 2 Jul 1949. TUOLUMNE Co.:

Twain Harte, 21 Jun 1961.

Acknowledgment.—The single fungal association listed was provided by J. F.

Lawrence (CSIRO, Canberra, Australia).

John T. Doyen 1 and F. T. Hovore, 2 1 Department of Entomological Sciences,

University of California, Berkeley, California 94720; 2 Placerita Canyon Nature

Center, Newhall, California 91321.

Received 20 January 1991; accepted 11 March 1991.

PAN-PACIFIC ENTOMOLOGIST

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five to seven words or concise phrases as KEY WORDS. Number FOOTNOTES sequentially and list on a separate page.

Text. — Demarcate MAJOR HEADINGS as centered headings and MINOR HEADINGS as left indented paragraphs with lead phrases

underlined and followed by a period and two hypens. CITATION FORMATS are: Coswell (1986), (Asher 1987a, Franks & Ebbet

1988, Dorly et al. 1989), (Burton in press) and (R. F. Tray, personal communication). For multiple papers by the same author use:

(Weber 1932, 1936, 1941; Sebb 1950, 1952). For more detailed reference use: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith

1987: table 3).

Taxonomy. — Systematics manuscripts have special requirements outlined in volume 66 (1): 1-8, including SEPARATE PARAGRAPHS