The Pan-Pacific Entomologist

EDITORIAL BOARD

J. T. Sorensen, Editor

S. S. Shanks, Treasurer

J. T. Doyen J. E. Hafernik, Jr.

R. M. Bohart

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 Biology, University of San Francisco, San Francisco, CA 94117.

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

addressed to: Vincent F. Lee, Secretary, 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 $ 15.00 for regular members of the Society,

$7.50 for student members, or $20.00 for subscription only. Members of the

Society receive The Pan-Pacific Entomologist. Single copies of recent numbers

are $5.00 each or $20.00 per volume. Make checks payable to the Pacific Coast

Entomological Society.

Pacific Coast Entomological Society

OFFICERS FOR 1990

Robert V. Dowell, President Sandra S. Shanks, Treasurer

Leslie S. Saul, President-Elect Vincent F. Lee, 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 1 May 1990

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

66(1): 1-8, (1990)

THE PAN-PACIFIC ENTOMOLOGIST:

FORMAT INFORMATION FOR CONTRIBUTORS

The Pan-Pacific Entomologist is published quarterly by the Pacific Coast Entomological Society, in

cooperation with the California Academy of Sciences. The journal serves as a refereed publication

outlet and accepts original manuscripts on all aspects of the biosystematics of insects and closely

related arthropods, especially articles dealing with their taxonomy, biology, behavior, ecology, life

history, biogeography and distribution. Articles with either a natural, descriptive orientation or a

technical and analytical emphasis are welcome. Articles that deal with the strictly economic aspects

of insects, however, are inappropriate for submission to the Pan-Pacific Entomologist. Manuscripts

must be in English, but foreign language summaries are permitted.

As of 1 January 1990, beginning with volume 66, number 1, the Pan-Pacific Entomologist will

employ changes in format which will be consistently applied. The format changes will require con¬

tributing authors to closely observe these guidelines. Because of comments by peer reviewers, but

more particularly copy editing, very few manuscripts are ultimately accepted without being redrafted

to incorporate changes.

Format

All manuscripts must be typed or printed on one side of 8.5 x 11 in nonerasable, high quality paper.

Three (3) copies of each manuscript must be submitted: an editor’s copy (original or high quality

photocopy), and two high quality review copies each including reductions of any figures to an 8.5 x 11

in page. Manuscripts must either be double or triple spaced in a legible typeface (preferably a serif

font) that makes identification of letters and numbers distinct, especially characters such as the number

1, lower case L, and upper case I. All margins should be 1.5 in, all pages must be numbered and the

senior author’s name should appear in the upper right comer of each page following the title page.

Do not break words with hyphenation at the right margin of the text.

The pages must be ordered and numbered separately in sequence 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, original figures (unnumbered).

Title Page

The upper right comer must include the statement “Send galleys to:” and list the corresponding

author’s name, address including ZIP code, and phone number. Type the title of the manuscript in

all upper case letters in the center of the page. Use a title that is informative and specific, but as brief

as possible. Compound titles are permitted and are separated by a colon; do not include a serial

number in the title. The title should not include both the common name and scientific binomial of

an insect. Titles must list the order and family involved within parentheses and separated by a colon.

The name of each author should be typed in upper case letters several lines below the title. In no

case should the professional or academic title of authors be noted. Several lines below the author’s

name, the institutional affiliation should be typed in upper and lower case letters, as the sequence:

department, institution, city, state or province, postal ZIP code or equivalent, and country if different

from the United States or Canada. Abbreviations must not occur within affiliations or addresses, spell

out all words in the affiliation, but omit building and room numbers (i.e., Department of Entomological

Sciences, University of California, Berkeley, California 94720); do not use two letter postal abbre¬

viations for states. In institutional addresses use a street number (i.e., 2258 Howard Avenue) or

building (i.e., Wellman Hall) only if mail would not be likely to ultimately reach its destination without

that addition. If the address on an article is different from the author’s current correspondence address,

use a footnote number after the address associated with the title and supply the current correspondence

address on a separate footnote page.

If multiple authors are associated with different institutions or addresses, separate these addresses

with a semicolon; place a footnote marker after each author’s name, and before the respective address.

Abstract Page

The second page of the manuscript should contain only the abstract and key words. It is essential

that the abstract be concise, not exceeding 250 words. The abstract should be an informative digest

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

of the significant findings or main conclusions of the article and is often the only information made

available by the abstracting publications. The abstract must be in a left indented paragraph format

and begin with the word Abstract in upper and lower case letters, underlined, and followed by a period

and two hyphens (i.e., Abstract .—). Begin the abstract to the right of the hyphen. When a taxonomic

binomial is first mentioned in the abstract, the author (unabbreviated) of the name must be stated.

Reference citation is not permitted in the abstract.

The key words paragraph (again, a left indented) follows several lines below the abstract. Its lead

is treated similarly to the abstract (i.e., Key Words .—). This is followed by five to seven key words or

concise phrases, the first of which should be “Insecta” (or the class of related arthropod if applicable).

The key words are aids to abstracting services that will index the article and as such should be chosen

wisely. You may repeat words or phrases from the title.

Text Pages

The text should generally follow the guidelines established in a recent edition of the Council of

Biological Editors Style Manual (CBE Style Manual Committee. 1983. CBE style manual: a guide for

authors, editors, and publishers in the biological sciences. 5th ed. Council of Biological Editors,

Bethesda, Maryland.). In the introduction avoid statements such as “The purpose of this paper

is . . .” and use instead simply “This paper . . .” followed by what is accomplished. Throughout the

article use simple and concise phrasing.

Major sections are delimited by centered headings using upper and lower case letters, such as:

Methods and Materials, Results, Discussion, Results and Discussion, Acknowledgment, Literature

Cited. Do not use a heading for the introduction. In taxonomic manuscripts the major headings might

include: Taxonomy, Biology, Behavior, etc. Minor headings are delimited as are the abstract and key

words paragraphs. Minor headings begin as left indented paragraphs with the word(s) in upper and

lower case letters and underlined, followed by a period and two hyphens (i.e., Description of Adult

Female .—).

References in the Text. — Cite published works by a single author by using the author’s last name

and the date of publication, without intervening punctuation (i.e., Burton 1989). For two authors cite

both names with an intervening asmpersand, but no punctuation, followed by the date (i.e., Burton

& Bickham 1989). For multiple authors, cite the senior author’s last name followed by “et al.” and

the date (i.e., Burton et al. 1989); place a period after “al” and do not indicate italics. When citing

multiple papers by an author or authors in a single year, follow the year by a lower case letter (i.e.,

Burton 1979a, b or Hicks 1986c). Lower case letters must follow the dates of the appropriate citations

in the Literature Cited section. Sequences of references in the text must be cited chronologically (i.e.,

Nelsonl978, Nieukoop & Sutasurya 1983, Raff 19 8 7). In cases where sequential citations occur within

the text and note multiple papers by the same author, the dates of articles should be separated by

commas and the authors separated by semicolons (i.e., Weber 1932, 1936, 1941; Whitcomb 1950,

1952; Henderson 1978, 1979). Citations within the text that include reference to pages, figures or

tables should be cited as: (Smith 1983: 149-153, Price 1985: fig. 7a, Nothwith 1987: table 3). Note

that citations within the text of illustrations or tables from other works should begin with a lower case

letter (i.e., fig. 2, table 2), to distinguish them from the figures and tables of the manuscript which

begin with upper case letters. All (and only) articles cited in the text must occur within the Literature

Cited section.

Cite as “in press” (i.e., Hawksworth in press), any manuscript which has been formally accepted

for publication by a journal (not which is in preparation, or that merely has been submitted, acknowl¬

edged, or is in review by a journal!); do not estimate a date of publication in either the text reference

or Literature Cited listing for articles that are in press. Data which does not exist in press (in this strict

sense) or in publication and is not being presented in the manuscript under consideration can be cited

as unpublished. If the unpublished data are from the sole author or all authors of the manuscript

under consideration, then cite simply as “(unpublished data)” but if the data are from only one of

multiple authors of the manuscript under consideration cite by initials (i.e., JAC, unpublished data).

Cite as “unpublished data” any manuscript in preparation or which has been submitted for publication

but has not yet been formally accepted after review. If the unpublished data are from a source other

that the author(s) of the manuscript under consideration, cite the data as a personal communication.

Personal communications should be kept to an absolute minimum, but where necessary should be

cited without abbreviation as: (D. Hille Ris Lambers, personal communication). Personal commu¬

nications and unpublished data citations are not listed in the Literature Cited section.

1990

FORMAT INFORMATION FOR CONTRIBUTORS

Measurements.— All measurements must be in metric units. Measurements in U.S. equivalents are

permissible only within quoted data (as from the label of a holotype), or within parentheses after their

metric counterpart if such display has a practical value. All fractions must be in decimal format (i.e.,

0.2, 0.33, 0.67, 0.125); if such numbers are less than one, place a zero before the decimal (i.e., 0.05,

not .05). When using measurement or count data and citing its range, mean and standard deviation,

use the format: low-high (mean ± SD) (i.e., 337-388 [361 ± 16]).

Numbers.— Numerals are expressed as their word equivalents if between one (1) and nine (9), unless

part of a sequence including values of 10 or greater; values greater than 10 are expressible as their

numbers. Always begin a sentence with the word equivalent of a numeral (i.e., “Four specimens were

. . .”)• When a large number begins a sentence it is usually preferable to modify the beginning of the

sentence to avoid awkwardness (i.e., “We examined 175 taxa . . .” rather than “One hundred and

seventy-five taxa were . . .”). Usage can be confusing, and acceptable examples are: “Six genera were

examined which contained a total of 26 species 10 males among the seven groups could

be classified into three categories...,” “The number of synapomorphies associated with each respective

intemodal segment in the phyletic sequence is: 3, 5, 13, 11, and 8.” Values representing counts or

distances should use their numerical rather than word equivalents, even if under 10; for example, in

the Material Examined section of a taxonomic paper use the data format: “.. . 3 km E of Wilsonville,

hwy 17, 6 males, 10 females,

Comparative Ratios.— Use a slash (/) rather than “per” to indicate standard units in relation to

other standard units (i.e., mg/liter, not mg per liter). Use “per” to indicate counts in relation to

nonstandard units (i.e., 10 insects per leaf). Indicate ratios using a colon (i.e., 10 males: 1 female).

Use a lower case “x” to indicate “times” in comparative ratios (i.e., mesothoracic width 0.8 x width).

Dates and Time.— Dates must be expressed in the format: day month year, without punctuation;

not as: month day, year. Months are abbreviated as their first three letters in upper and lower case:

Jan, Feb, Mar, Apr, May, Jun, Jul, Aug, Sep, Oct, Nov, Dec. Do not shorten the notation for the year,

cite in full (i.e., 1989, not 89 or ’89). Examples of acceptable dates are: 4 Jul 1990, 17 May 1753.

Time is expressed in 24 hour format followed by h. If appropriate, time zones can be noted following

the time, as a standard, three letter, upper case abbreviation. An example is 06:30 h (PDT), representing

6:30 AM, Pacific Daylight Time.

Italics.— Italics are indicated by underlining words. It is preferable not to use actual italicized words

in a manuscript, even if the capability exists in word processing, but rather to use a single underline

to denote words to be italicized; this promotes clarity and reduces the chance of error during the

editing and type-setting stages of publication. Italics are used only for taxonomic names, for denoting

minor subheading sections in the text, for highlighting categorical descriptors or “flags” within a block

of text, and for denoting mathematical variables as in formulae. Italics may be used to denote a word

which should stand out in a block of text. Examples are: the word “Thorax” when used within text

comprising a taxonomic description; the name of a county (or equivalent) when used in a Material

Examined section; the word “In” denoting a contribution to a larger work, as used in a bibliographic

citation in the Literature Cited section. Do not use italics to emphasize a word for effect in a sentence

(i.e., “Only some of the insects were . . .”). Do not indicate italics for Latin abbreviations such as:

i.e., e.g., ca., sic., et al., etc.

Abbreviations.— Reference to tables should not be abbreviated in the text, whereas reference to

figures should be (i.e., Table 4, Figs. 1-3, Fig. 6A); capitalize such references when they are part of

the manuscript but not in citations from other articles (i.e., Hamilton 1983: fig. 2, Rolling 1975: table

2). Abbreviations for conventional units are in lower case letters and not followed by a period. Examples

are: h, min, sec, km, m, mm, /um, cc, ml, g, mg, ng, kg. Spell liter completely, do not abbreviate as

Avoid the use of ca. by using approximately instead. States should be spelled out in full, do not use

their two letter postal abbreviations (except possibly in tables where space is at a premium). Do not

abbreviate male(s), female(s), morphs, castes or life forms as their symbols. Do not include periods

after, or spaces between, the letters of abbreviations of institutions (i.e., use USDA, NMNH and

BM[NH] for U.S. Department of Agriculture, National Museum of Natural History and British

Museum [Natural History], respectively). Abbreviate percent as %. Abbreviate feet as ft, not ('), when

necessary in elevations, after altitude in meters. Trademarks and registered brand names are noted as

® and ®, respectively.

Compass directions are presented entirely in upper case without periods (i.e., N, S, E, W, NW, SE,

NEE, SSW). Direction from a source, as in data presented in the Material Examined section of

taxonomic papers, should have the word “of” between the direction and source to avoid potential

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

confusion in locality names (i.e., 3 km E of Palo Alto, instead of 3 km E Palo Alto, which should be

taken as East Palo Alto rather than East of Palo Alto).

Taxonomic Studies and Citations

Taxonomic works present special editorial problems in maintaining relative consistency within and

between articles on systematics within the journal. Because of these inherent problems, there is

necessarily less flexibility for style and format in taxonomic works in the journal than for articles on

solely biology.

Taxonomic Citations. — Upon the first mention of a species or lower level taxon in both the abstract

and text, the author of an animal taxon must be cited using the International Commission of Zoological

Nomenclature convention; botanical names must be so cited using the ICBN. Do not abbreviate the

generic name of a taxon upon first mention. The names of authors of taxa must not be abbreviated,

except for Linnaeus (as L.) and Fabricius (as Fabr.). If more than a single taxonomic author with the

same last name worked in the taxonomic group considered, the author’s initials should be used to

avoid confusion (i.e., Papilio bairdii W. H. Edwards, to avoid confusion with H. Edwards who was

also a lepidopterist). Multiple authorship for taxa should substitute an ampersand (&) for “and”

between the names (i.e., Gillette & Palmer).

When citing authors of taxa, citation of the year of description is optional; if used, however, the

year must be enclosed within parentheses and the citation must be considered a reference citation

within the article and be listed in the Literature Cited section (see below for an exception). Examples

are: Chaitophorus populellus Gillette & Palmer (1928) (where populellus was originally described in

Chaitophorus ), and Chaitophoruspopulifolii (Essig 1912) (where populifolii was not originally described

in Chaitophorus ). In these cases Gillette & Palmer (1928) and Essig (1912) must be listed in the

Literature Cited section. If no year citation is noted for a species when citing its author, a literature

citation is not invoked. The only exception to treating an author-year citation as a literature citation

is in checklists and synonymies, where year citations may be more appropriate than in the text, and

where the numbers of such citations would make literature citation unwieldy. In such cases, cite years

as C. populellus Gillette & Palmer, 1928, and C. populifolii (Essig), 1912.

All nomenclature erected in taxonomic studies must follow the rules established in the most recent

nomenclatural code by the ICZN. When mentioning the ICZN code, refer to it as such, rather that

just “the code.” At any mention of a particular article in the ICZN code, cite the publication date of

the code version used, to avoid confusion (i.e., “ . . . due to ICZN article 49 [ICZN 1985].”) and list

the code citation in the Literature Cited section (see the Literature Cited examples for citation of the

ICZN code).

New taxa or synonymies that are erected should be clearly and appropriately marked, or “flagged,”

in upper case letters after mention in the text (i.e., NEW GENUS, NEW SYNONYMY, NEW STATUS,

etc.), and notation should also occur in the abstract. New taxa must be listed with the name of the

describing author(s), even if the it is same as the manuscript author, after the binomial but before any

taxonomic “flag” (i.e., Diodontus retiolus Eighme, NEW SPECIES).

Descriptions and Diagnoses.— Taxa being described must have the following: a description (pref¬

erably with appropriate illustrations), and a separate diagnosis. The diagnosis must occur as a separate

paragraph delimited as “ Diagnosis .-” and must be concise but not in telegraphic style. Comment

briefly and only on those attributes required to separate the taxon from related taxa in the diagnosis.

The description must be delimited by a minor subheading, as is the diagnosis; it need, however,

not be labeled as “description” but rather can be labeled as a sex, life stage or form (i.e., male, egg,

larvae, viviparous apterae, etc.). Descriptions must be concise and be in telegraphic style. It is preferable

to separate major components with periods (i.e., legs), minor components with semicolons (i.e., tibiae,

tarsi) and the details of minor components with commas (i.e., color of tibiae and the forms of setae

on them). Avoid conjunctives as much as possible in telegraphic descriptions; also avoid “-ish”

adjective suffix forms of colors, such as “head yellow with brownish” using instead “head yellow with

slight brown”; avoid latin color descriptors (i.e., testaceous). An example is: “. . . Genitalia (Fig. 5A):

valve breadth greatest ventrally, filling entire vinicular area, bilobed and constricted near indentation,

caudal extension tapered, blunt terminally; saccus diminutive, lobate with rounded margins; aedeagus

robust, ceacum two-fifths aedeagal length.”

Types. —Descriptions and revisions also require comments on the types involved. Comments on

types are to be in a separate paragraph delimited as “ Types .—” that lists the type of type(s) (holotype,

paratype, etc.) erected, and their data and deposition. An example is: “Holotype, male (Figs. 2E, 5A)

1990

FORMAT INFORMATION FOR CONTRIBUTORS

deposited BM(NH), data: PERU. Cajamarca, 2800 m, Simons collection. Paratype, 1 male deposited

NMNH, (poor condition with tails broken off), data: same as holotype but 3800 m, O. T. Baron

collection, ex Hamilton collection 1919.” When citing data from the labels of types it is permissible

to quote within quotation marks. If not directly quoting from type labels, use the data format listed

below for material examined. The deposition of types must be noted appropriately and be in accord

with ICZN requirements. Deposition of types in private collections should be avoided for reasons of

“professionalism,” but when deposition is so designated, the ultimate intended deposition after the

death of the author(s) should be stated; neotypes require institutional deposition (ICZN 1985: article

75-d-6).

Keys.— Keys are not required in taxonomic works, but are highly recommended. When presented,

keys must be concise, clear, easy to follow and have reversibility provisions. Keys must also be in

adjacent couplet style, and each couplet should contain preferably more than a single, nonoverlapping

attribute. It must be clear to which life stage, sex, caste, morph, etc., the key pertains; keys requiring

more than a single such life form are discouraged, and it is recommended that separate keys be

provided in such cases.

Material Examined.— Data for material studied in taxonomic manuscripts must be listed under a

separate paragraph delimited as “ Material Examined .—This taxonomic section should have the

countries, as well as major (i.e., state, province) and minor (i.e., county or equivalent) political units

spelled out in upper case, with the minor political units also underlined for italics. Use the following

format, with modification as appropriate: USA. ARIZONA. APACHE Co.: 10 km N of Lupton, hwy

12, 2070 m, 11 Sep 1978, J. T. Sorensen (JTS 78118), P. ponderosa, 6 females. COCHISE Co.: nr

Rustler Park, Chiricahua Mts, 2500 m, 16 Sep 1978, J. T. Sorensen (JTS 78147), P. ponderosa, 12

females. COLORADO. ARCHULETA Co.: 25 km W of Pagosa Springs, hwy 160, 2140 m, 8 Aug

1978, J. T. Sorensen (JTS 78H50), P. ponderosa, 1 female. CANADA. BRITISH COLUMBIA.

Fairmont Hotsprings, hwy 93, 17 Jul 1978, J. T. Sorensen (JTS 78G91), P. ponderosa, 25 females.

Acknowledgment Page

Begin this section on a separate page and spell the heading as Acknowledgment, not as Acknowl¬

edgement, Acknowledgements or Acknowledgments. The acknowledgment should be concise, thanking

people first, institutions second where necessary, and grant or contract support third where appropriate.

Do not use the professional or academic titles of those being acknowledged. If the affiliations of those

acknowledged are included, do not abbreviate institutional names but do include their locations. Do

not abbreviate number as “No.” or as “#” in citing grant or contracts; rather, for example, cite as

“NSF grant BSR-8908456.”

Literature Cited Pages

Begin this section on a separate page, titled Literature Cited, not References, References Cited or

Bibliography. All paragraphs should be hanging format (left block first line and indented thereafter).

Do not list references which are not cited in the text. Do not list unpublished data, personal com¬

munications, or works in preparation in the Literature Cited section. Citations listed should be in

alphabetical order first and then in chronological order; if multiple citations bearing the same author(s)

and year are cited, they should be listed using lower case letters after the year and be in the sequence

in which they appear in the text. Do not use Ibid.

Authors cited are listed as last name first followed by initials for sole or senior authors and initials

followed by last name for subsequent authors. Omit reference to number (issue) in citations after the

volume. Omit the number of total pages in books, separates, pamphlets, etc. Abbreviate journal titles

as listed in the International Serials Catalogue: Part I: Catalogue (International Council of Scientific

Unions Abstracting Board, 1978). Do not abbreviate single word journal names (i.e., Evolution,

Ecology). All citations must fully cite the authors, even when more that one article is present for any

author(s). When listing articles or books, all letters in the title should be lower case, except the first

letter of the title’s first word and any proper nouns. Examples of acceptable citation formats follow:

One author articles:

Arnold, R. A. 1983. Speyeria callippe (Lepidoptera: Nymphalidae): application of information-

theoretical and graph-clustering techniques to analyses of geographic variation and evalu¬

ation of classifications. Ann. Entomol. Soc. Am., 76: 929-941.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Two author articles:

Ferrari, J. A. & K. S. Rai. 1989. Phenotypic correlates of genome size variation in Aedes albopictus.

Evolution, 42: 895-899.

Articles with more than two authors:

Atchley, W. R., E. V. Nordheim, F. C. Gunsett & P. L. Crump. 1982. Geometric and probabilistic

aspects of statistical distance functions. Syst. Zool., 31: 445-460.

Manuscripts in press:

Sorensen, J. T. (in press). Three new species of Essigella (Homoptera: Aphididae). Pan-Pacif.

Entomol.

Books:

Anderson, T. W. 1984. An introduction to multivariate statistical analysis (2nd ed). John Wiley

& Sons, New York.

Parts of larger works:

Klecka, W. R. 1975. Discriminant analysis. Chapter 23. pp. 434-465. In Nie, N. H., C. H. Hull,

J. G. Jenkins, K. Steinbrenner & D. H. Brent. 1975. SPSS: statistical package for the social

sciences (2nd ed.). McGraw-Hill, 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 Czechoslo¬

vakia, Sept. 9-14, 1985. SPB Academic Publishing, The Hague, The Netherlands.

Proceedings of meetings:

Philbrick, R. N. (ed.). 1967. Proceedings of the symposium on the biology of the California

islands. Santa Barbara Botanic Garden, Santa Barbara, California.

Wilson, M. R. & L. R. Nault (eds.). 1987. Proceedings of the second international workshop on

leafhoppers and planthoppers of economic importance, held in Provo, Utah USA, 28th

July-1st August 1986. CAB International Institute of Entomology, London.

Governmental or Institutional publications:

Little, E. L. Jr., & W. B. Critchfield. 1969. Subdivisions of the genus Pinus (Pines). U.S. Dept.

Agric., Forest Serv. Misc. Publ., 1144.

Hafemik, J. E. Jr. 1982. Phenetics and ecology of hybridization in buckeye butterflies (Lepidop-

tera: Nymphalidae). Univ. Calif. Publ. Entomol., 96.

Anonymous Institutional or organizational publications:

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

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

California Department of Food & Agriculture. 1987. Environmental assessment of gypsy moth

and its eradication in California, 1987 program. California Department of Food & Agri¬

culture, Division of Plant Industry, Sacramento, California.

Theses and Dissertations:

Sorensen, J. T. 1983. Cladistic and phenetic analysis of Essigella aphids: systematics and phy¬

togeny in relation to their Pinaceae host plants (Homoptera: Aphididae, Lachninae). Ph.D.

Thesis, University of California, Berkeley.

Computer programs:

Felsenstein, J. 1984. PHYLIP—Phytogeny inference package (Version 2.5). (A phylogenetics

computer program package distributed by the author). J. Felsenstein, Dept, of Genetics,

University of Washington, Seattle, Washington.

1990

FORMAT INFORMATION FOR CONTRIBUTORS

Pimentel, R. A. & J. D. Smith. 1985. Biostat II. (A multivariate computer program package

distributed by the authors). Sigma Soft, Placentia, California.

Footnote Pages

All title and text footnotes must appear on a separate and numbered footnote page and be indicated

by consecutive superscript numbers. Author-line footnotes that denote a change of affiliation or current

address must also appear on the separate footnote page. Only multiple author and affiliation address

footnotes (as described under the title page section) should not be on the footnote page. Text footnotes

provide additional information and are usually discouraged; such information, if short, may preferably

be enclosed within parentheses in the text. Footnotes cannot not be used for acknowledgments, list

these in the Acknowledgment section.

Figure Caption Page

Figure captions should briefly interpret the figures but need not be complete sentences. Captions

should not unnecessarily repeat or explain information provided in the text. Be sure, however, to

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THE PAN-PACIFIC ENTOMOLOGIST

Yol. 66(1)

Papp, C. S. 1976. Manual of scientific illustration, with special chapters on photography, cover

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John T. Sorensen, Insect Taxonomy Laboratory, California Department of Food & Agriculture,

Sacramento, California 95814.

PAN-PACIFIC ENTOMOLOGIST

66(1): 9-12, (1990)

BEE POLLINATION OF CUPHEA (LYTHRACEAE)

SPECIES IN GREENHOUSE AND FIELD

Frank D. Parker 1 and Vincent J. Tepedino 2

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

Screwworm Research, American Embassy,

PSC Box 342, APO Miami, Florida 34020

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

Bee Biology & Systematics Laboratory,

Utah State University, Logan, Utah 84322-5310

Abstract. — Several species of native bees were examined as pollinators of three species of Cuphea,

a prospective oil-seed crop, in greenhouse and field trials, and records of flower visitation by

bees were assembled from literature and museum sources. The most successful pollinator in the

greenhouse was the gregarious megachilid, Osmia bruneri Cockerell, which constructed 106 nests

and produced 139 progeny using the pollen, nectar and leaves of Cuphea leptopoda Hemsley.

Two female Xylocopa californica Cresson also produced progeny in the greenhouse. In the field,

the only consistent visitors of Cuphea were three species of bumblebees ( Bombus fervidus (Fabr.),

B. huntii Greene and B. occidentalis Greene).

Key Words. — Insecta, Cuphea, Osmia, Xylocopa, Bombus, pollination, seed production, oil-seed

crop, brood

Cuphea is a large genus of plants (260 species) with potential agricultural im¬

portance because the seeds of some species contain oils that are rich in medium-

chain fatty acids (MCT). Such oils contain a high proportion of lauric acid whose

main source is coconut and palm kernel, and capric acid which is derived from

petrochemicals. These MCTs have important industrial applications (Hirsinger

& Knowles 1984, Thompson 1984, Hirsinger 1985).

Several federal and state research units are conducting intensive breeding pro¬

grams to develop Cuphea for commercial production. Because many of the species

being studied require animal vectors to transport pollen from one plant to another,

seed set is contingent upon providing effective pollinators. Although species in

the genus display a wide variety of pollination syndromes, including hummingbird

pollination (Feinsinger 1976), insects appear to be the most important pollinators

of species with commercial promise. The honeybee, Apis mellifera L., however,

does not visit flowers of many cross-pollinated species, even under duress (S. J.

Knapp, personal communication). Thus, this study was conducted to find man¬

ageable bee species that would visit and pollinate the flowers. In addition, we

assemble available bee-flower association records from various sources.

Methods and Materials

Three species of Cuphea were grown from seeds. Seedlings of Cuphea leptopoda

Hemsley were planted in a greenhouse (6 x 6 x 3 m) in February, 1986. Flowering

of the 450 plants began 1 May and continued until October. In June, two flats of

blooming C. lutea Rose in Koehne, an autogamous species, were introduced to

the same room. At the same time, about 100 plants each of C. laminuligera

Koehne and C. lutea were planted at Greenville Experimental Farm, North Logan,

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Table 1. Species of bees released in greenhouse containing Cuphea plants and numbers of cells

produced. Logan, Utah 1986.

Species

No. adults

Date

No. nests

Ashmeadiella aridula

1922, 466

2-7 May

Chelostoma minutum

2022, 20 66

20 Jun

Hoplitis fulgida

2622, 2066

24-25 Jun

Osmia bruneri

2422, 166

27-30 May

106

Osmia sanrafaelae

2122, 1266

27-30 May

1 +

Xylocopa arizonensis

222, 366

27 Apr

Utah, to provide flowers for field visitation by pollinating insects. These flowers

were observed several times each week and flower visitors were recorded.

The bee species tested as potential pollinators in the greenhouse were an ar¬

bitrary selection of those obtained from trap nests placed in Utah in 1985. In

addition, several adults (three males, two females) of Xylocopa californica Cresson

were also tested. These bees were found overwintering in a dead Yucca flower

stalk collected near St. George, Washington Co., Utah, in February. The bees and

stalks were held at 3° C until late April and then set among the blooming Cuphea

in the greenhouse.

Nesting units of several types were included for the megachilid bees. Some were

wood blocks with holes drilled therein. Two sizes of soda straws (5, 8.5 mm) were

alternately inserted in the holes. Other wood blocks contained 10 holes each of

5 diameters (2, 4, 6, 8, 10 mm) in Latin square design but without straw inserts.

Elderberry (Sambucus sp.) stems with 3 mm diameter holes drilled longitudinally

through the pith to a depth of 45 mm, and 3 mm holes drilled 1-2 mm into the

pith perpendicular to the longitudinal axis, were also provided. Finally, several

elderberry stems, each with 12 mm diameter holes drilled perpendicular to the

longitudinal axis and into the pith, were included as nest sites for X. californica.

Results and Discussion

Greenhouse Visitation. — Five species of megachilid bees and X. californica were

released in the greenhouse with Cuphea. The date and number of bees released,

and the number of nests recovered, are shown in Table 1. Only C. leptopoda was

consistently visited. The rare visits to C. lutea plants by Osmia and Xylocopa

were of brief duration. The bees clearly preferred C. leptopoda. The avoidance of

the flowers by half of the species is probably related to the presence of glandular

trichomes on the flower (Hirsinger & Knowles 1984), which frequently entangled

the mouthparts of foraging bees. Even large carpenter bees occasionally had dif¬

ficulty in freeing themselves from the viscous flowers.

Of the megachilids, only the metallic-blue Osmia bruneri Cockerell constructed

many nests and reared much brood on the pollen and nectar of Cuphea. Pollen

and nectar were collected from about 08:30-19:00 h. These bees produced 139

cells of which 80.5% contained live adults in diapause in September. Most nests

(98) and progeny (125) were produced in wood blocks in holes of 5-6 mm diameter

and irrespective of whether straws were present.

1990

PARKER & TEPEDINO: BEE POLLINATION OF CUPHEA

Female O. bruneri exclusively masticated pieces of Cuphea leaves to line and

partition their nests even though plants of other acceptable species were provided

for this purpose. This leaf selection behavior is noteworthy in that O. bruneri has

refused leaves of other crop plants such as red clover ( Trifolium pratense L.) in

other greenhouse studies (FDP, unpublished data). Thus, managing this bee for

Cuphea pollination in greenhouse or field is simplified because it is unnecessary

to provide another plant species as a leaf source.

Other attributes of O. bruneri make its use as a greenhouse pollinator of Cuphea

promising. Populations of this spring bee are abundant and can be obtained by

placing trap nests in mountainous locations in many western states. Its nesting

behavior has been studied in the greenhouse by Frohlich (1983) whose data and

that of Parker (FDP, unpublished data) show that several sizes of boring (10, 8,

6, 4 mm) are accepted for nesting. Osmia bruneri is univoltine and enters diapause

in the adult stage in the fall. Adults can then be stored overwinter at 3° C until

needed the following spring. At that time, nests with adults are incubated at 30°C

for a few days to break diapause. A small percentage of overwintering adults are

parsivoltine; i.e., diapause is not broken until the second spring (FDP, unpublished

data).

A single nest with a few cells was produced by a female Osmia sanrafaelae

Parker, a species whose life cycle and management requirements (Parker 1984)

are similar to O. bruneri. The female nested in a chamber formed by the meeting

of L-shaped metal joints that held the sides of the plant beds together. Other O.

sanrafaelae were observed collecting pollen and cutting and gathering pieces of

Cuphea leaves. These females probably also produced nests in obscure places but

those nests were not found.

Both X. californica females lived for approximately two months and used Cu¬

phea pollen and nectar to produce progeny. In contrast, adult males died about

a week after their introduction to the greenhouse. Although males were seen to

attempt copulation with females several times at the flowers, females produced

only a few male offspring each and, thus, may have remained unmated. Foraging

was limited to early morning and late evening; during the heat of the day, they

remained in their nests.

Should Cuphea be grown commercially in the desert southwest, X. californica

could be an important pollinator. Nests are commonly found in flower stalks of

Yucca and Agave in Arizona and New Mexico. It may be feasible to collect stalks

containing quiescent adults and transport them to agricultural plantings.

Field Visitation. — Only eight species of bees were recorded visiting Cuphea

plants in the field and of these, only the three species of bumblebees ( Bombus sp.)

were numerous. The following species were collected from C. laminuligera : B.

fervidus (Fabr.), B. huntii Greene, and B. occidentalis Greene. Three specimens

of Dialictus sp., one Agapostemon texanus Cresson and one Megachile rotundata

(Fabr.) were also taken from this same species. A few specimens of Anthophora

urbana Cresson and Calliopsis coloradensis Cresson were collected from C. lute a.

Both species produced abundant seed. Cuphea lutea, however, is known to be

autofertile (Hirsinger & Knowles 1984).

Collection Records.— The most intriguing information is from S. J. Knapp

(personal communication) who collected 6 bees from C. laminuligera in Mexico.

All specimens but one Bombus sonorus Say were Loxoptilus longifellator LaBerge

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

(Anthophoridae). This is the first flower association for Loxoptilus, a genus char¬

acterized by peculiar pads of curved hairs on the galea. Perhaps these hair pads

enable Loxoptilus females to more effectively negotiate the viscous trichomes of

Cuphea flowers.

We (FDP) recently (July 1987) collected four species of bees foraging on an

unknown species of Cuphea in the state of Chiapas, Mexico. Females of two new

species [Anthidium and Dianthidium (Mecanthidium)\ were collecting pollen, but

the others {Bombus mexicanus Cresson and Anthidium maculifrons Smith) were

collecting only nectar. L. A. Stange collected both sexes of an undescribed mega-

chilid, Hypanthidioides (Moureanthidium ), from flowers of Cuphea in Argentina.

Hurd (1979) listed Augochloropsis metallica metallica (Fabr.) and Melissodes te-

paneca Cresson visiting C. balsamona and Melissodes bimaculata bimaculata

(Lep.) from C. petiolata. Whether any of these bee-flower records represent as¬

sociations that are more than fortuitous remains to be studied.

Acknowledgment

We are most grateful to S. J. Knapp (Oregon State University, Corvallis) for

supplying seeds, bee specimens, unpublished information, and for reviewing the

manuscript. The nest of Xylocopa californica was found and donated by Utah

State University students D. Broemeling and N. Youssef. Technical assistance

was supplied by D. Veirs, M. E. Klomps, S. Jennings and J. Hennebold. The

manuscript also benefited from reviews by W. E. LaBerge and A. E. Thompson.

This is a contribution from the Utah Agricultural Experiment Station, Utah

State University, Logan, Utah 84322-4810, Journal Paper No. 3455, and USDA-

Agricultural Research Service-Bee Biology & Systematics Laboratory, Utah State

University, Logan, Utah 84322-5310.

Literature Cited

Feinsinger, P. 1976. Organization of a tropical guild of nectarivorus birds. Ecol. Monogr., 46: 257-

291.

Frohlich, D. R. 1983. On the nesting biology of Osmia ( Chenosmia ) bruneri. J. Kans. Entomol. Soc.,

56:123-130.

Hirsinger, F. 1985. Agronomic potential and seed composition of Cuphea, an annual crop for lauric

and capric seed oils. JAOCS, 62: 76-80.

Hirsinger, F. & P. F. Knowles. 1984. Morphological and agronomic description of selected Cuphea

germplasm. Econ. Bot., 38: 439-451.

Hurd, P. D., Jr. 1979. Apoidea. pp. 1741-2209. In Krombein, K. V., P. D. Hurd, Jr., D. R. Smith

& B. D. Burks (eds.). Catalog of Hymenoptera in America north of Mexico. Smithsonian

Institution Press, Washington, D.C.

Parker, F. D. 1984. A candidate legume pollinator, Osmia sanrafaelae Parker. J. Apic. Res., 24:

132-136.

Thompson, A. E. 1984. Cuphea—& potential new crop. Hort. Science, 19: 352-354.

Received 23 February 1988; accepted 3 April 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 13-18, (1990)

DESCRIPTION OF A NEW SPECIES OF METRIUS

(COLEOPTERA: CARABIDAE: PAUSSINI) FROM IDAHO

WITH COMMENTS ON THE TAXONOMIC STATUS OF THE

OTHER TAXA OF THE GENUS

Yves Bousquet and Henri Goulet

Biosystematics Research Center,

Agriculture Canada, Ottawa, Ontario K1A 0C6, Canada

Abstract. — A new species of Metrius Eschscholtz, M. explodens Bousquet & Goulet, is described

from Idaho. Specimens representing the three previously described Metrius taxa were examined,

but conclusions as to the taxonomic status of each of these taxa could not be drawn from the

material at hand.

Key Words. — Insecta, Carabidae, Metrius explodens, bionomics, interspecific variations

The genus Metrius Eschscholtz (1829) as recently discussed (Bousquet 1986),

represents the sister-group of the remaining Paussini (= ozaenines + paussines

of authors). The adults are odd, looking superficially more like tenebrionids than

carabids. The genus is confined to the Pacific Coast, ranging from southern British

Columbia to the Sierra Nevada in California. The beetles live along forest edges

and in woodlands under rocks, rotten logs and in leaf litter. Lindroth (1961)

recognized two species of Metrius, M. contractus Eschscholtz (1829) and M. ser-

iceus Rivers (1900) but Van Dyke (1925) suggested they may represent subspecies

of a single species. In the same paper, Van Dyke (1925) described a subspecies,

M. contractus planatus, from the alpine areas of the Lake Tahoe region.

This paper describes a new species of Metrius collected several years ago in

Idaho, compares it with Metrius, and briefly discuss the status of taxa within

Metrius.

Metrius explodens Bousquet & Goulet, New Species

Type Material. — Holotype (male) and paratype (male). IDAHO: “Ida: Idaho

Co. Hwy 12, 39 mi N.E. Lowell (2860') 30.VII.1968, R.E. Leech.” The holotype

is deposited in the California Academy of Sciences, San Francisco (CAS Type

No. 16494) and the paratype in the Canadian National Collection, Ottawa (CNC

No. 20010).

Description. —Coloration: body, including palpi and legs, uniformly brownish with head and first

four antennomeres slightly darker than remaining parts. Microsculpture: nearly entire body with

isodiametric, flat microsculpture. Head: frons with one supraorbital seta on each side; clypeus with

one long seta on each side and one to three smaller ones medially; labrum with 13-14 setae along

anterior margin; eyes small, flat; antennae pubescent from fifth segment, antennomere X about 2 x

as long as wide. Mentum with epilobes rounded anteriorly; tooth truncate; labial pits not apparent;

paraglossa fused to ligula; ligula with one pair of setae near anterior margin; last segments of both

labial and maxillary palpi slightly dilated in apical half; penultimate labial palpomere with two long

setae on anterior margin and usually a few additional smaller setae. Pronotum (Fig. 2): pronotum

rather flat, about 1.5 x wider than long; lateral margins barely sinuate in posterior half, with three or

four (only two unilaterally in one specimen) setae on each side; lateral bead thin, especially in anterior

half; anterior angles rather protruding, rounded; posterior angles sharp, distinctly protruding poste¬

riorly; base without lateral impression; basal margin strongly sinuate. Elytra: elytra oviform, convex,

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Figure 1. Metrius explodens Bousquet & Goulet.

1990

BOUSQUET & GOULET: A NEW METRIUS

4 5

Figures 2-5. Figure 2. Pronotum, M. explodens. Figure 3. Pronotum, M. contractus. Figure 4.

Aedeagus (left lateral view), M. explodens. Figure 5. Aedeagus (left lateral view), M. contractus.

coalescent, without discal or parascutellar setae; striae shallow; intervals slightly convex. Underside:

underside impunctate; prothoracic epipleura as wide anteriorly as at middle; prostemum and prostemal

apophysis with setae medially; apophysis not margined, pointed at apex; metastemum and metepi-

stemum short; metastemum with a few setae laterally; metepimera distinct; abdominal stemite III

with a patch of small setae at level of trochanter; stemites III, IV and V with two or three setae on

each side near posterior margin; stemite VI with a few setae medially and usually with two setae on

each side at posterior margin. Legs: tibiae smooth dorsally near apex; fore tibia with 8-10 clip setae;

all tarsomeres sparsely pubescent dorsally; last tarsomeres with rows of spiniform setae on ventral

side; first three fore tarsomeres of male with adhesive setae ventrally; middle tibia densely pubescent

in apical half; hind trochanter with two or three small setae dorsobasally and one longer seta near

posterior margin. Genitalia (Fig. 4): median lobe of aedeagus with apical part distinctly curved, nearly

perpendicular to remaining part, apex straight, not twisted in lateral view, rounded in dorsal view;

parameres asymmetric, left one without setigerous puncture, right one with brush of hair apically on

medial side. Body length: 11.0-11.5 mm.

Diagnosis .—The other Metrius differ most importantly from this species by

the following character states: microsculpture on body convex or beadlike; eyes

more convex; antennae proportionally shorter, antennomere X approximately

1.5 x longer than wide; epilobe of mentum longer and more oblique anteriorly;

tooth of mentum emarginate; lateral margins of pronotum (Fig. 3) more sinuate

in posterior half, with two setae on each side; lateral bead thick over entire length;

pronotum with anterior angles more protruding and more or less angulate, pos¬

terior angles less protruding posteriorly; elytral intervals flat; prothoracic epipleura

THE PAN-PACIFIC ENTOMOLOGIST Vol. 66(1)

M. contractus

ffc M. contractus

form planatus

(f) M. contractus

form sericeus

Figure 6. Distribution (including M. explodens [■]) and variation of the apex of median lobe

(dorsal view) of M. contractus (•), M. contractus form “ planatus ” (3) and M. contractus form “ sericeus ”

(O). Outside the map, M. contractus is also known from Vancouver, B.C.

wider anteriorly than at middle; prostemal apophysis rounded apically; abdominal

stemites III, IV and V usually with only one seta on each side at posterior margin;

tibiae more or less furrowed dorsally near apex; first two fore tarsomeres of male

with adhesive setae ventrally; metatrochanter without setae basally; median lobe

of aedeagus with apex twisted (Fig. 5), left paramere usually with few small setae

apically (Fig. 5).

Etymology. — The specific name explodens is derived from the Latin verb ex-

plodo, -ere (in the figurative expression to expel). It refers to the ability of the

species to expel the content of the pygidial glands like the bombardier beetles (H.

B. Leech, personal communication).

Distribution. — The species if known only from the type locality in Idaho.

Bionomics. — The two specimens of M. explodens were collected in a partly

logged pine-fir forest adjacent to a river (H. B. Leech, personal communication).

Leech (1971) reported collecting a female of the spider Callobius enus (Chamberlin

& Ivie) eating an adult of M. explodens. He also noted that the egg sac of the

spider was covered with remains of Metrius.

Taxonomic Status of Other Metrius Taxa. —In addition to the new species,

three Metrius taxa have been described previously: (1) M. contractus, occurring

BOUSQUET & GOULET: A NEW METRIUS

1990

Figures 7, 8. Figure 7. Microsculpture near the middle of the fifth elytral interval, M. contractus.

Figure 8. Same, M. contractus form “ sericeus

from SW British Columbia to southern California along the coast and in the Sierra

Nevada, usually at low elevation (< 600 m); (2) M. planatus, a montane form

recorded from near Lake Tahoe and from the Yosemite National Park, originally

described as a subspecies of contractus ; (3) M. sericeus, found in the southern

Sierra Nevada, from Kings Canyon and Sequoia National Parks to northern part

of Kern County.

Differentiation between these three forms is based on the shape of the apex of

the median lobe of the aedeagus and the type of microsculpture of the elytral disc.

These differences are summarized in Table 1.

Whether this complex consists of a single species or three taxa (subspecies or

species) is uncertain. The form planatus appears to be intermediate both geo¬

graphically and in the shape of the microsculpture. On the other hand, the dif¬

ference in the shape of the median lobe of these three forms, though slight, is

constant. Moreover, one typical male of contractus was found in proximity to the

other two forms that occur in the Sierra Nevada. We feel that better sampling of

Metrius from the Sierra Nevada is needed to resolve the complex. Therefore, we

currently consider all specimens to represent one species, M. contractus Esch-

scholtz, without recognition of subspecific status for the different forms.

Two of these three forms show no geographical variation. Adults of the form

contractus are uniform from southern British Columbia south to San Francisco,

California. From there southward, they tend to be smaller.

This study is based on 601 adults from the California Academy of Sciences,

Table 1. Distribution of character states between the three forms of M. contractus

Character states

Forms

Apex of median lobe (Fig. 6)

Elytral microsculpture

contractus

symetrically fanlike and sharp

laterally

sculpticells moderately convex and meshed

(Fig. 7)

“ planatus ”

symetrically fanlike and round

laterally

sculpticells convex and not meshed in spots

“ sericeus ”

asymetrically expanded

sculpticells very convex (wartlike) and not

meshed (Fig. 8)

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

San Francisco and from the Biosystematics Research Center, Ottawa. All speci¬

mens studied were labelled according to the form to which they belong to facilitate

future studies.

Acknowledgment

We thank H. B. Leech and D. Kavanaugh (California Academy of Sciences,

San Francisco) for the loan of specimens; A. Smetana and J. M. Campbell (Bio¬

systematics Research Center, Agriculture Canada, Ottawa) for reviewing the

manuscript; and G. Sato for preparing the habitus and inking the drawings.

Literature Cited

Bousquet, Y. 1986. Description of first-instar larva of Metrius contractus Eschscholtz (Coleoptera:

Carabidae) with remarks about phylogenetic relationships and ranking of the genus Metrius

Eschscholtz. Can. Entomol., 118: 373-388.

Eschscholtz, J. P. 1829. Zoologischer Atlas, enthaltend Abbildungen und Beschreibungen neuer

Thierarten, wahrend des Flottcapitains v. Kotzebue zweiter Reise um die Welt, auf der Russisch-

Kaiserlichen Kreigsschlupp Predpriaetie in den Jahren 1823-1826. Erstes Heft. G. Reimer,

Berlin.

Leech, R. E. 1971. Revision of the Amaurobiid spiders of the Nearctic Region (Arachnida: Araneida).

Ph.D. Thesis, University of Alberta.

Lindroth, C. H. 1961. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska.

Part 2. Opusc. Entomol. Suppl., 20: 1-200.

Rivers, J. J. 1900. A new Metrius from California. Entomol. News, 11: 389.

Van Dyke, E. C. 1925. Studies of western North American Carabinae (Coleoptera) with descriptions

of new species. Pan-Pacific Entomol., 1:111-125.

Received 9 February 1989; accepted 13 October 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 19-23, (1990)

FEMALE NESTING BEHAVIOR AND MALE

TERRITORIALITY IN

APHILANTHOPS SUBFRIGIDUS DUNNING

(HYMENOPTERA: SPHECIDAE)

Kevin M. O’Neill

Entomology Research Laboratory, Montana State University,

Bozeman, Montana 59717

Abstract. — The behavior of females and males of Aphilanthops subfrigidus Dunning was studied

in southcentral Montana in 1987, 1988, and 1989. As with the related species A. frigidus (F.

Smith), females of A. subfrigidus were found to specialize on alate queens of Formica (F. subpolita

Mayr) that they capture at mating swarms of this ant. Several aspects of female nesting behavior

are described, including their reaction to the presence of miltogrammine flies in the nesting area.

In addition, male territoriality and scent-marking, which are widespread within the Philanthinae,

are reported for the first time for this genus.

Key Words.— Insecta, Aphilanthops, Formica subpolita, nesting, territory

Aphilanthops is one of three North American genera of the sphecid subfamily

Philanthinae with females that provision their nests with ants (Bohart & Menke

1976). Females of Clypeadon and Listropygia prey upon workers of Pogonomyr-

mex spp., with each wasp species apparently specializing on a single species of

ant. Female Aphilanthops frigidus (Smith), restrict their prey to alate queens of

Formica (Peckham & Peckham 1905, Wheeler 1913, Evans 1962). Several prey

records obtained for A. subfrigidus Dunning suggest that it also specializes upon

queens of Formica spp. (Evans 1970). In contrast, Evans (1977) reports bees as

prey of Aphilanthops hispidus W. Fox, indicating that predation upon ants is not

a consistent feature of this genus. Our knowledge of the behavior of female Aphi¬

lanthops is incomplete. Furthermore, of the five genera of Philanthinae that are

relatively widespread in North America ( Aphilanthops, Cerceris, Clypeadon, Eu-

cerceris, and Philanthus), Aphilanthops is the only one for which male territoriality

and scent-marking have not been reported (Evans & O’Neill 1988). We report

observations of female nesting behavior and male territorial behavior in A. subfri¬

gidus.

Methods

Observations were made on Aphilanthops subfrigidus from 29 Jun-3 Jul 1987,

21 Jun-1 Jul 1988, and from 3-19 Jul 1989 at a site 13.5 km south of Three

Forks, Gallatin Co., Montana. Male territories and female nest sites were located

on the bottom and sides of several gullies, 0.5 to 1.5 km west of the Madison

River. Over a three day period in 1987, five hours of focal observations were

made on females at eight nests. Intermittent observations of both females and

males were made at other times. The grassland site was within the Stipa comata

Trinius & Ruprecht /Bouteloua gracilis (Humboldt, Bonpland, & Kunth) Lagasca

y Segura ex. Steudel habitat type (Mueggler & Stewart 1980). Also present were

three-leaf sumac ( Rhus trilobata Nuttall), junipers ( Juniperus spp.), and soapweed

(Yucca glauca Nuttall).

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 66(1)

Results and Discussion

Female Behavior.— In 1987, a nesting aggregation ultimately consisting of 11

nests within 4 m of one another, was discovered at the bottom of a gully 1 km

west of the Madison River. While no nests were found at this location in 1988,

a group of six nests was located in the same gully, approximately 100 m to the

east. Only three other nests, each well-separated from other nests, were located

during the three years of study. When present, the tumulus at each nest entrance

was less than 1 cm in diameter. The absence of a substantial tumulus was also

noted for A. frigidus (Evans 1962).

Eight of the nests were excavated, but the burrows were difficult to track, as

was reported for A. frigidus (Peckham & Peckham 1905, Wheeler 1913, Evans

1962). All of the nests occurred in fine silt interspersed with small rocks and fine

roots. The burrows entered the soil at angle of 30° to 45°. One burrow was followed

for 24 cm to a depth of 23 cm. The main burrow of the nest doubled back upon

itself, so the terminus was only 10 cm in horizontal distance from the entrance.

All prey found within nests were alate queens of Formica subpolita Mayr (For-

micidae). Alate queens of Formica neogates Emery were reported as prey of A.

subfrigidus from a single nest excavated in Jackson Hole, Wyoming (Evans 1970).

Two nest cells containing three and four prey were found at depths of 15 and 20

cm, each at a distance of 1 cm from the main burrow. Twelve additional prey

were found stored in the main burrows of four other nests (one to seven per nest),

all within 10 cm of the entrance. All seven prey found in the nest cells and two

of those found in burrows lacked wings. The wings of all but one of these prey

had apparently been chewed off by the provisioning female, rather than shed

because the fragmented bases of the wings were present. Ten of 12 ants found in

the burrows had wings.

At least some of the prey were captured at mating swarms of F. subpolita,

several of which were within 3 m of the 1988 nest site. A. subfrigidus were observed

at the mating swarms 13 times. Successful capture of an alate female at swarms

was observed once; three other females were seen leaving swarms carrying prey.

The female A. subfrigidus themselves were taken a prey by the robber fly Ejferia

staminea (Williston) (Asilidae) near the ant swarms ( n = 4).

Females began bringing prey into nests shortly after 09:00 h RMST and con¬

tinued foraging until between 10:17 and 11:05 h. This overlapped with the period

during which mating swarms of F. subpolita were observed on Rhus trilobata on

11 days in June and July, 1988 and 1989 (unpublished data). Seasonal synchrony

between predator and prey species was also observed. The first sightings of A.

subfrigidus in 1988 and 1989 occurred one and four days, respectively, after

swarms of the ants first appeared.

Storage of prey in the burrow allowed females to spend very little time in the

nest during the short daily period available for hunting. They spent 0.5 to 3.6

min (X = 1.8, n = 23) inside the nest between foraging flights 2.2 to 9.8 in duration

(X = 5.2, n = 20). During a 1 h observation period, three females brought 5, 6,

and 11 prey to their nests. Thus, if three or four prey per cell is typical for this

species, females were obtaining prey for more than a single cell in a morning. This

would seem advantageous because the mating swarms of Formica occurred for

only several hours per day, on only 11 of the 17 days on which A. subfrigidus was

1990 O’NEILL: SPHECID NESTING BEHAVIOR AND TERRITORIALITY

observed in 1988 and 1989. Winged females were only rarely observed on days

on which swarms did not take place.

The nest entrance was left open by the females during 86% of the foraging flights

observed (n = 37). However, seven females returning to nests after the final daily

foraging flight and five leaving the nests for prolonged non-foraging periods all

closed the nest with a loose plug of soil. Facultative temporary nest closures have

also observed in A. subfrigidus (Evans 1962) and in several species of Philanthus

(Evans & O’Neill 1988) and are thought to prevent entrance by cleptoparasites.

One final nest closure was observed. Over a period of 30 min, the female plugged

the top three cm of the burrow with soil before initiating a second nest two m

away.

Females approaching nests carrying prey sometimes behaved differently when

being pursued by cleptoparasitic Senotainia spp. (Sarcophagidae). In 22 of 32

return flights, the females with prey were not followed by the miltogrammine flies.

These 22 flew to the nest and either entered directly, or after dropping the prey

at the entrance, entered, turned, and pulled in the prey. However, of the 10 flights

observed when females were pursued by one (n = 5) or two (n = 5) flies, five

females continued their approach and entered the nest, whereas the other five

turned in flight and left the area. This apparently evasive action was observed in

A. frigidus (Wheeler 1913) and in many species of Philanthus (Evans & O’Neill

1988). Ristich (1956) and Evans (1962) found high levels of miltogrammine

parasitism in A. frigidus.

Male Behavior.— Groups of males were found in four locations from 21-29 Jun

1988 where they perched 1-2 m high on Rhus trilobata in three areas and on

grasses growing adjacent to Yucca in another. The types of behavior displayed by

males were generally identical to those of many other Philanthinae (Evans &

O’Neill 1988). From perches 0.5-2 m high, males pursued passing insects and

engaged in interactions with conspecific males. These interactions included: (1)

“swirling flights” when two to four males rapidly and repeatedly circled one

another in flight within a radius of less than 10 cm; (2) “butting” when males

made brief, but forceful and often repeated head-on contact in mid-air; and (3)

“grappling” when males grasped one another in mid-air and sometimes fell to

the surface below where they continued wrestling for several seconds. All three

of these behaviors have also been observed in the related genus Philanthus. In

that genus, specific butting and grappling episodes are sometimes best interpreted

as misdirected mating attempts. However, in Philanthus, interactions among males

often differ markedly from male-female encounters. They occur repeatedly be¬

tween two males over short intervals and commonly result in the perch occupant

being driven from the area and replaced by the intruder. Thus, in Philanthus and,

probably Aphilanthops as well, these behaviors, especially repeated bouts of but¬

ting and grappling, represent aggressive interactions associated with territorial

defense and exclusive occupation of areas immediately surrounding perches (Ev¬

ans & O’Neill 1988).

At three of the four aggregations, 7-14 territories were found. All territories in

each aggregation were within 5-20 m of one another, with perches of adjacent

males often less than a meter apart. Because the number of males at an aggregation

was at least twice the number of territories, interactions were common.

Territorial males also exhibited “abdomen dragging,” another behavior typical

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 66(1)

of male philanthines. Males walked up the stems of grasses with the hair brushes

of their clypeal area and posterior venter of their abdomen in contact with the

plant. On Rhus, they walked along the edges of the leaves. This behavior probably

serves to deposit a pheromone that attracts females (Evans & O’Neill 1988).

Schmidt et al. (1985) found that male Philanthus basilaris deposited volatile

substances onto plant from their mandibular glands while abdomen dragging. The

clypeal hair brushes of male Aphilanthops are shorter and less dense than those

of male Philanthus. The abdominal hair brushes are also sparser, although distinct

rows of short hairs (“fimbriae”) are present on abdominal sterna IV and V (Bohart

& Grissell 1975).

The territorial areas lacked flowering plants or honeydew sites likely to attract

conspecific females and although nests were nearby, nests did not occur in the

large aggregations sometimes found in philanthines (Evans & O’Neill 1988). Other

nests were scattered throughout the area that contained territories. Although ter¬

ritories of males at two of the areas were within swarms of Formica subpolita,

the latter were not observed at the other two territorial areas.

The behavior of female A. subfrigidus is similar to that of A. frigidus (Peckham

& Peckham 1905, Wheeler 1913, Evans 1962), particularly with regards to prey

use. Comparative data from other philanthines suggest the interactions observed

among male A. subfrigidus result from attempts of males to exclude conspecific

males from areas containing scent-marked plants. Although territoriality has been

reported in many Sphecidae, scent-marking of territories has been observed only

within five genera of the Philanthinae (Evans & O’Neill 1988). The presence of

the clypeal hair brushes in four of the six remaining undocumented genera (Bohart

& Menke 1976) suggests that territoriality and scent-marking maybe even more

widespread within the subfamily.

Acknowledgment

I thank Andre Francoeur for identifying the ants, Eric Fisher for identifying the

robber fly, Ruth O’Neill for assistance in the field, and Howard E. Evans, Daniel

Bartell, Stephen Harvey, Douglas Streett, and Saralee Visscher for reviewing the

manuscript. This publication is contribution 2326 of the Montana Agricultural

Experiment Station.

Literature Cited

Bohart, R. M. & E. E. Grissell. 1975. California wasps of the subfamily Philanthinae (Hymenoptera:

Sphecidae). Bull. Calif. Insect Surv., 19: 1-92.

Bohart, R. M. & A. S. Menke. 1976. Sphecid wasps of the world: a generic revision. University of

California Press, Berkeley.

Evans, H. E. 1962. A review of the nesting behaviour of digger wasps of the genus Aphilanthops,

with special attention to the mechanics of prey carriage. Behaviour, 19: 239-260.

Evans, H. E. 1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming. Bull.

Mus. Comp. Zool., 140: 451-511.

Evans, H. E. 1977. Aphilanthops hispidus as a predator of bees (Hymenoptera: Sphecidae). Pan-

Pacif. Entomol., 53: 123.

Evans, H. E. & K. M. O’Neill. 1988. The natural history and behavior of North American beewolves.

Comstock Publishing Associates, Ithaca, New York.

Mueggler, W. F. & W. L. Stewart. 1980. Grassland and shrubland habitat types of western Montana.

U.S. Dept. Agriculture, Forest Serv. General Technical Report, INT-66.

Peckham, G. W. & E. G. Peckham. 1905. Wasps social and solitary. Houghton Mifflin, Boston.

1990 O’NEILL: SPHECID NESTING BEHAVIOR AND TERRITORIALITY

Ristich, S. S. 1956. The host relationship of a miltogrammid fly Senotainia trilineata (VDW). Ohio

J. Sci., 56: 271-274.

Schmidt, J. O., K. M. O’Neill, H. M. Fales, C. A. McDaniel, & R. W. Howard. 1985. Volatiles from

mandibular glands of male beewolves (Hymenoptera: Sphecidae, Philanthus) and their possible

roles. J. Chem. Ecol., 11: 895-901.

Wheeler, W. M. 1913. A solitary wasp (Aphilanthops frigidus F. Smith) that provisions its nest with

queen ants. J. Anim. Behav., 3: 374-386.

Received 23 March 1989; accepted 27 October 1989.

PAN-PACIFIC ENTOMOLOGIST

66 ( 1 ): 24 - 32 , ( 1990 )

LIFE HISTORY OF EUTRETA DIANA (OSTEN SACKEN) ON

ARTEMISIA TRIDENT AT A NUTTALL IN

SOUTHERN CALIFORNIA (DIPTERA: TEPHRITIDAE)

Richard D. Goeden

Department of Entomology, University of California,

Riverside, California 92521

Abstract. —Eutreta diana (Osten Sacken) is univoltine on Artemisia tridentata Nuttall (Astera-

ceae), its only reported host plant in southern California. Eggs are inserted in axillary and terminal

buds on young branches in late spring. First instars diapause, and gall growth essentially ceases,

for the next six or seven months until larvae molt and then overwinter as second instars in very

small galls. Branch elongation and gall and larval growth resume in spring. Galls are described

in detail and pictured. Pupation and adult emergence occur mainly in April and early May.

Newly emerged females require two to three weeks to mature eggs, mate, and begin oviposition.

Principal natural enemies of E. diana include the parasitoids, Eurytoma sp. and Rileya sp.

(Eurytomidae), Pteromalus sp. (Pteromalidae), Tetrastichus sp. (Eulophidae), and Torymus ci-

tripes (Huber) (Torymidae), and avian bushtits, Psaltriparus sp., besides arthropod predators

and other mortality factors identified.

Key Words.— Insecta, Eutreta, Artemisia, gall, parasitoids, life history, predation, larval mor¬

phology

Live specimens of Eutreta diana (Osten Sacken), with their pale green eyes,

white-dotted black wings, and bright red abdomens, represent one of the more

beautiful species of fruit flies (Diptera: Tephritidae) in California. This aesthetic

consideration, plus my discovery of several field populations of E. diana, provided

me with ample reasons to study its life history.

Taxonomy. — First described as Trypeta diana by Osten Sacken (1877), Eutreta

diana and two separately described color variants of the adults were synonymized

by Stoltzfus (1977), who also illustrated the adults in his comprehensive revision

of the genus. Second and third instars and the puparium of E. diana were described

and illustrated by Steck & Wharton (1986).

Distribution, Hosts and Study Sites. —Stoltzfus (1977) mapped the North Amer¬

ican distribution of E. diana as California, Oregon, and Washington east into the

Dakotas, Nebraska, Colorado, and New Mexico, and adjacent parts of Canada

and Baja California, Mexico. Benbow & Foster (1982) also reported this fly from

northwestern Texas and called it “one of the most widely encountered gall-forming

tephritids in western North America.” Foote & Blanc (1963) and Stoltzfus (1977)

noted that it has been collected as far east as Missouri, well east of the distribution

actually mapped by the latter author, who also suggested that its southern distri¬

bution extends into Mexico, presumably meaning east of Baja California. Foote

& Blanc (1963) plotted the distribution of E. diana in northern California along

the length of the northern and central Sierra Nevada Mountains, and in southern

California, from the Palomar, Cuyamaca, and Laguna Mountains in Riverside

and San Diego Counties southwest to the Pacific Coast.

My field-study locations for flies and galls on Artemisia tridentata Nuttall (As-

teraceae) in the southern Sierra Nevada and San Bernardino Mountains serve to

1990

GOEDEN: LIFE HISTORY OF EUTRETA DIANA

connect the apparent northern and southern segments of its California distribu¬

tion: Horse Meadow, 2225 m, Mahogany Creek, and Taylor Creek, Sequoia Na¬

tional Forest (northern section), Tulare Co., 1985 and 1986, and Mormon Rocks,

Cajon Pass, San Bernardino National Forest, San Bernardino Co., 1986-1989.

Additional locations where the characteristic galls on A. tridentata were observed

or collected included: Rattlesnake Creek above S. Fork of Santa Ana River, San

Bernardino Mts., San Bernardino National Forest, San Bernardino Co., 9 Jun

1981; Mojave River Forks, 975 m, San Bernardino National Forest, San Ber¬

nardino Co., 22 Apr 1986. Adults were also swept as follows: road to Horseshoe

Meadow, 2750 m, Inyo National Forest, Inyo Co., 22 Jul 1987; White Mountain,

3150 m, Inyo National Forest, Inyo Co., 19 Aug 1987. Unless otherwise stated,

most of the observations were made at the Mormon Rocks site.

The recorded hosts of E. diana include six species and two varieties of Artemisia

representing tall and erect as well as low-growing shrubs (Fronk et al. 1964,

Stoltzfus 1977, Benbow & Foster 1982).

Egg. — No intact egg was found in nature because they are inserted for most of

their length tightly into buds, cannot readily be seen with the naked eye, and are

easily overlooked and destroyed even during dissection under a stereomicroscope.

Three ova dissected from a sexually mature, field-collected female were fusiform,

smooth, white, bluntly rounded at both ends, 0.55 mm long and 0.25 mm wide,

and at the cephalic end bore a 0.06 mm, peg-like pedicel. Within a bud, an egg

was surrounded by brown necrotic tissue.

Larva. — Eclosion was not studied, except to note that an embryo had rotated

180° before hatching. Upon hatching, first instars tunneled basipetally into the

pith but moved only 2 to 3 mm during the six to seven months following ovi-

position, which mainly took place in May 1987. The first instar is 1.1-1.2 mm

long and is easily distinguished by its red-brown cephalopharyngeal skeleton and

a black patch of verrucae on the abdominal ventor (Fig. la).

Examination of this instar by scanning electron microscopy (see Headrick &

Goeden in press for materials and methods) determined that the verrucae actually

ringed several abdominal segments (II-V dorsally, Fig. 2a; I-VII ventrally, Fig.

2b), but only the central portion of the ventral verrucae were pigmented (Figs.

2a, 2b). Two forms of verrucae were seen at higher magnification (Fig. 2c); the

larger morphs are less common, unevenly distributed, and have a central pore.

The dark ventral verrucae are less scattered than the lateral and dorsal verrucae,

being aligned in transverse rows (Fig. 2d).

The function of the segmental verrucae is uncertain, but an ambulatory role is

probable, although the larva could move little in its confined quarters. Like the

cephalopharyngeal skeleton, the patch appeared to darken as the first instar aged,

but was absent, or at least not prominent on the second and third instars. Fur¬

thermore, no such patch was mentioned by Steck & Wharton (1986) in their

detailed descriptions of the last two instars.

Thirty-one small and barely recognizable bud galls were collected on 9 Nov

1988, at the beginning of overwintering by E. diana. These young galls were

merely slightly clavoidal swellings of the distal parts of lateral and terminal veg¬

etative branches (Fig. lb). Twenty-one (68%) of the 31 galls contained live first

instars; seven had begun to molt, indicating the second instar served as the main

overwintering stage. The 21 galls with live larvae averaged 1.89 ± 0.06 (x ± SE)

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Figure 1. Life stages and galls of Eutreta diana on Artemisia tridentata : (a) first instar in small gall

(note black verrucae on ventor) (15 x); (b) dissected (left) and intact (right) small galls contained first

instars (5x); (c) overwintered second instar in gall (11 x); (d) puparium in central cavity with exit

tunnel and exit hole to right (note tunnels of agromyzid larvae in walls) (4x); (e) mature galls (0.8 x);

(f) female at rest (8 x); (g) puparia of E. diana and Liriomyza sp. in gall heavily mined by the latter

(5 x); (h) gall with hole through which bushtit extracted larva (4x).

1990

GOEDEN: LIFE HISTORY OF EUTRETA DIANA

Figure 2. First instar Eutreta diana : (a) lateral view; (b) ventral view; (c) detail of two morphs of

verrucae; (d) detail of ventral verrucae (pigmented).

(1.87-2.55) mm in diameter, measured at the level of the gall cavities, and 2.03

± 0.06 (1.50-2.57) mm at their maximal width slightly more distally. Benbow &

Foster (1982) reported that galls of E. diana on A. filifolia Torrey containing first

instars averaged 8 mm in diameter only a month or so after eclosion. The cavities

within the young galls from Mormon Rocks ended an average of 2.51 ± 0.17

mm from the gall apices (Fig. la) and were ellipsoidal, frass-free, open, and

smooth-surfaced; they averaged 0.97 ± 0.03 mm in diameter and were 1.56 ±

0.07 mm long. The thickness of gall walls varied from 0.2-0.6 mm. Ten of the

31 first instars examined were dead, and on one of these a parasitoid adult de¬

veloped (identified below).

On 21 May 1986, just after snow melt allowed access to the Horse Meadow

site (2225 m), 122 overwintered galls were collected, refrigerated, and dissected

in the laboratory during the next week. Only one gall showed little growth, mea¬

sured 6 mm long by 3 mm wide, and contained a live early second instar in a 2

mm long, 0.5 mm wide cell. This may typify the overwintering state. This gall

sample also contained 44 (36%) more advanced second instars (Fig. lc), 21 (17%)

of which had been killed by parasitoids or unknown causes. In the 21 galls con¬

taining dead larvae, the cavities had become wholly or partly filled with the

enlarged parenchymatous pith cells constituting most of the gall tissue, and which

the larvae rasp with their mouth-hooks while feeding. The 23 (19%) subspheroidal

galls that contained actively feeding or molting second instars averaged 8.7 ± 0.4

mm long by 7.5 ±0.3 mm wide. These lively larvae occupied open, approximately

1 mm wide, central-longitudinal feeding chambers which also contained a small

accumulation of whitish frass (Fig. lc). No solid feces littered the gall cavities.

After molting, the exuvium and cephalopharyngeal skeleton were discarded, usu¬

ally at the base of the feeding chamber. Even at this early date the remaining galls

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 66(1)

in this sample contained two live third instars and 22 puparia, so larval devel¬

opment progressed at different rates even in galls along the same branch.

A sample of 92 galls collected at Taylor Creek (2072 m) on 22 May 1986

contained only three (3%) second instars, 39 (42%) third instars, and 50 (54%)

puparia in contrast to the Horse Meadow sample. Earlier development by E.

diana at low elevations was further suggested by a sample of 55 galls collected

for dissection at Mormon Rocks (1017 m) on 18 May 1987. This sample contained

no larvae, only three (5%) intact puparia that probably were parasitized, and 18

(33%) empty puparia from which adults already had emerged. The remaining

galls bore evidence of bird predation or larval parasitism (see below).

Most larval growth takes place during the third stadium. Although Benbow &

Foster (1982) stated that gall growth ceased if the larva died before it reached the

third instar, my data, like their graphed mean gall diameters, suggested that gall

growth continued during the third stadium. One hundred mature galls that each

bore a puparium (Fig. Id) averaged 12.0 ± 0.4 (6-19) mm in length and 10.0 ±

0.2 (6-13) mm in maximal width. The central cavities within these mature galls

averaged 7.5 ± 0.2 (5-10) mm in length and 3.1 ±0.1 (2-4) mm in width. The

cavities usually were mostly frass-free, circular in cross-section, rounded basally,

and smooth-walled. The thickness of galls walls varied from 1.5-4 mm.

Externally, mature galls that contained live puparia usually were subspheroidal

(Figs. Id, le). This gall shape also resulted when larval feeding or exit hole con¬

struction destroyed the terminal bud at the apex of a gall. Upon death or removal

of a larva, or after emergence of the adult, apical growth and intemode elongation

usually resume and the gall assumes a spindle-shape. This causal relation for

branch-gall shape was reported for other tephritids ( Trupanea conjuncta [Adams]:

Goeden 1987, Tephritis stigmatica [Coquillett]: Goeden 1988), as well as for a

gallicolous moth ( Carolella beevorana Comstock, Cochylidae: Goeden & Ricker

1981). Though apparently fairly common among branch-gall-forming Tephriti-

dae, this relationship is not widely recognized (Freidberg 1984). Thus, the gall of

E. diana is a modified branch composed of shortened and inflated intemodes

(Fig. le). The galls are grey-green when young, sometimes purple-tinted, especially

when infested by inquilines and certain parasitoids (see below), but turn brown

and woody with age. They are covered with a closely appressed whitish tomentum

when young and are laterally marked by transverse, scale-like expanded leaf bases

bearing sessile leaves and axillary buds that may grow into one or more branches

adorning old empty galls. Empty galls of the current season end in clusters of

leaves or a terminal branch, depending on whether the apical bud has been killed.

Galls formed from axillary buds may be sessile or borne on pedicels of various

lengths, whereas galls of apical buds of main branches may have pedicels several

cm long. For example, 11 (37%) of 30 galls sampled at Mahogany Creek on 13

Jun 1985 were sessile but the remainder were borne on pedicels from 1-13 mm

long. Similarly, 19 (50%) of 38 galls from Taylor Creek on 12 Jun 1985 were

sessile although the remainder had pedicels which varied from 1-9 mm long.

Usually only a single larva of E. diana inhabits each gall. One gall, however,

collected at Mormon Rocks on 20 May 1987 contained two intact puparia formed

in separate chambers. These puparia did not yield flies or adult parasitoids when

caged at room temperature in a humidity chamber. This is a rare example of a

monothalmus tephritid producing a biloculate, polythalmous gall (Freidberg 1984).

1990

GOEDEN: LIFE HISTORY OF EUTRETA DIANA

Two dead second in stars found within a single chamber in a gall collected at

Taylor Creek on 12 Jun 1985 apparently wounded and killed each other, providing

another outcome to abnormal multiple oviposition in a single bud or branch tip.

Freidberg (1984) reported that the highest record for the number of tephritid

galls on a single plant probably was the 135 galls of Spathulina sicula Rodani he

counted in Israel. I removed and counted 147 current season galls of E. diana

from a single plant at Mormon Rocks on 27 May 1987. However, Dodson (1987)

noted “over 2000” galls of Aciurina bigeloviae (Cockerell) reported from aim

diameter Chrysothamnus nauseosus (Pallen) Britton.

Toward the end of their feeding period, the third instars usually extended their

feeding chambers distilaterally to form a curved exit tunnel. This narrowed to a

1.5 to 2 mm diameter, circular window formed by a thin layer of epidermis that

is usually not visible externally when intact (Fig. Id). A few exit tunnels projected

straight outward and destroyed the apical bud. One gall was observed in which

the exit tunnel had been formed basilaterally near the juncture with the pedicel.

Pupariation often occurs with the anterior end of the puparium projecting into

the mouth of the exit tunnel.

Pupa. — The puparium description of Steck & Wharton (1986) was based on

eight specimens from California. The 40 puparia I measured (Fig. Id) averaged

4.2 ±0.1 (3.3-5.0) mm in length and 1.7 ± 0.1 (1.0-2.5) mm in width. These

dimensions compare favorably with the 2.8-4.8 mm lengths and 1.0-1.8 mm

widths reported by Steck & Wharton (1986). The puparia I examined were trans¬

lucent and changed from mustard-yellow to ochrous tan with age. The abdomen

eventually appeared red-brown and the rest of the body dark brown through the

puparium just before adult emergence.

Adult.— Adults (Fig. If) were uncommon, but when found they were always on

gall-bearing A. tridentata. The ability of E. diana adults to feign death and drop

to the ground when disturbed reported by Steck (in Freidberg 1984) was not

observed. Both sexes exhibited the alternate and synchronous wing-waving be¬

havior described for other nonfrugivorous tephritids (e.g., Trupanea bisetosa [Co-

quillett]: Cavender & Goeden 1982, Neotephritis finalis [Loew]: Goeden et al.

1987, Paracantha gentilis Hering: Headrick & Goeden, unpublished data).

Adult emergence began approximately on 6 May 1987 as judged from emergence

records for flies reared from larvae and puparia dissected from galls collected at

Mormon Rocks on 30 Apr. Thirty-two males (63%) and 21 females were reared

from all galls sampled in 1985; 23 males (64%) and 13 females, from galls sampled

in 1986. Both sexes emerged over week-long emergence periods during both years.

Newly emerged females (n = 3) possessed immature ovaries. Weekly dissection

of three females, each held at room temperature (21° C) in separate cages and

supplied with water and honey, showed that they contained full size ova between

two and three weeks after emergence.

Gall-bearing plants are attacked year after year as reported by Benbow & Foster

(1982), although why a plant is attacked for the first time remains unanswered

(Freidberg 1984). Adults continued to frequent the plant from which I had re¬

moved all galls of the current season. The only occurrence in common that I

found was that galled plants always had access to a good supply of ground water

(e.g., in a wash at Mormon Rocks or other drainage at Mojave River Forks), were

on the margin of a meadow (Horse Meadow), or above a running stream (Ma-

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

hogany Creek, Taylor Creek). Plants with galls of E. diana also bore galls of one

or more species of Cecidomyiidae, as noted on A. tridentata in Wyoming (Fronk

et al. 1964). Adults of E. diana appeared early in the year when night temperatures

were low and days remained cool. Their morning activities mainly consisted of

sunning to gain heat with their black bodies and dark wing color. Flies rested with

their long axis perpendicular to, and dorsa facing, the sun and with their wings

held flat, motionless, and parted (Fig. If). Their characteristic dark wings no

doubt function in heat gain, raising body temperature to facilitate early daytime

and seasonal activities at higher altitudes.

Mating was observed once, at 08:15 h on 9 Jun 1987. Although the start and

finish of this single act of copulation was not observed, the flies were initially

located on the underside of a vertically oriented leaf. The male was positioned

over the posterior of the thorax and the abdomen of the female with his wings

motionless and straight back but overlapping and flat upon his dorsum. The wings

of the female were slightly parted and also motionless. The male was held off the

substrate and carried by the female as she alternately walked or rested, as they

pumped their mouthparts and remained in copula.

Oviposition by a single female was observed between 13:30-14:00 h on 27 May

1987. She backed into, and oviposited directly into, two axillary buds, while

probing other buds for 15-20 sec each without ovipositing. The two acts of

oviposition lasted 3:50 min and 3:45 min, and were subsequently confirmed by

observing the eggs in the excised buds after dissection under a stereomicroscope.

Phenology. — Adult emergence and activity coincided with flowering by Yucca

whipplei Torrey (Agavaceae) and Eriodictyon crassifolium Bentham (Hydrophyl-

laceae) at the Mormon Rocks site. The adults disappeared about the time that

Senecio douglasii deCandolle began to bloom there in 1986 and 1987.

Seasonal History. —Eutreta diana is univoltine on A. tridentata in southern

California, although it might be able to produce additional generations on one or

more alternate but as yet undetected hosts (Foote & Blanc 1963, Wasbauer 1972),

as predicted by host records from Wyoming (Fronk et al. 1964) and Texas (Benbow

& Foster 1982). Two of the four alternate hosts, A. arbuscula Nuttall and A. cana

Pursh, reported by Fronk et al. (1964) also occur in northern California (Munz

& Keck 1959), but not in southern California (Munz 1974) where these studies

were conducted. I have reared E. simplex Thomas from galls on a different species

of Artemisia (unpublished data), but not E. diana. My sweep records of live adults

from high elevations in July and August also indicate that alternate hosts are

present in California.

The biology and seasonal history of E. diana on A. filifolia in Texas (Benbow

& Foster 1982) is very different than on A. tridentata in southern California. Both

populations appear to be univoltine. The overwintering stage is the egg in Texas,

but mainly the second instar in southern California. First instars hatch in late

March to mid-April in Texas, where the first stadium lasts approximately a month,

in contrast to the six or seven months in southern California! Subsequent larval

and gall growth are continuous in Texas, without summer and fall diapause by

the first instar and concomitant arrested gall development that is now known

from southern California. Pupation occurs in late August to early September in

Texas and lasts about a month, whereas, in southern California it occurs in late

1990

GOEDEN: LIFE HISTORY OF EUTRETA DIANA

spring (April and May) and lasts about two weeks. Adults emerged in the fall to

oviposit the overwintering eggs in Texas, but emerged and oviposited in late

spring in southern California. Life history differences of this magnitude suggest

that Texas and California populations may be temporally, spatially, and etholog-

ically separated by their different host plant affinities and therefore possibly sibling

species which should undergo morphological re-examination. Stoltzfus (1977)

recognized great variability in the color of E. diana adults.

Natural Enemies and Inquilines. — The larvae were heavily parasitized by Hy-

menoptera including: Eupelmus sp. (Eupelmidae), a solitary, ectoparasitoid of the

first instar; Pteromalus sp. (Pteromalidae), a solitary, larval ectoparasitoid; Eur-

ytoma sp. and Rileya sp. (Eurytomidae), solitary, larval, endoparasitoids; Tet-

rastichus sp. (Eulophidae), a gregarious, larval-pupal endoparasitoid; and Torymus

citripes (Huber) (Torymidae), a solitary, larval, ectoparasitoid. These parasitoids

were reared from isolated parasitized hosts (humidity chamber). The latter species

killed and devoured the young larvae of E. diana, and then completed its devel¬

opment by tunneling in the walls of the gall. This same behavior was ascribed to

a Eurytoma sp. by Benbow & Foster (1982), who reared an additional five species

of parasitic Hymenoptera from galls of E. diana in Texas, including a Tetrastichus

sp. Fronk et al. (1964) also reported a braconid ( Dacnusa sp.) as a parasitoid of

E. diana.

Larvae of Apion sp. (Coleoptera: Apionidae) and an unidentified species of

Lepidoptera similarly fed inside the galls on the walls. They killed, and in a few

cases partly ate, the E. diana larvae encountered, thus accidentally functioning

as predators. The walls of the galls also were heavily mined and darkened by the

small larvae of an unidentified species oi Liriomyza (Diptera: Agromyzidae) (Figs.

Id, lg), which apparently did not harm E. diana (Fig. lg). Mining of gall walls

by the Apion sp., Liriomyza sp., the torymid, and lepidopteran hastened gall decay

and disintegration, in contrast to empty galls uninfested by inquilines that turned

woody and persisted for several years (Benbow & Foster 1982).

The larvae of an unidentified clerid beetle were observed as predators of E.

diana larvae and an adult was seen that had been freshly killed by a salticid spider.

However, the more common predators of E. diana larvae and puparia were birds,

which were especially active at the Mormon Rocks study site, where many galls

bore open holes pecked by social, insectivorous bushtits, Psaltriparus sp. (Fig.

lh). Stoltzfus (1977) suggested that woodpeckers opened galls presumably formed

by E. longicornis Snow on Artemisia cana.

Acknowledgment

My thanks to D. W. Ricker and D. H. Headrick for technical assistance, in¬

cluding the insect photography involved in Figs. 1 and 2, respectively. Thanks

also to F. L. Blanc, D. H. Headrick, J. A. McMurtry, and B. A. Stoltzfus for their

helpful comments on early drafts on the manuscript. The parasitoids were iden¬

tified by J. LaSalle and the insectivorous birds by A. C. Sanders (Department of

Entomology and Department of Botany and Plant Sciences, respectively, Uni¬

versity of California, Riverside). Apion sp. and Liriomyza sp. were identified

respectively by D. R. Whitehead and R. V. Peterson (Systematic Entomology

Laboratory, USDA, ARS, Beltsville, Maryland).

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Literature Cited

Benbow, S. M. & D. E. Foster. 1982. Biology of Eutreta diana Osten Sacken on sand sagebrush

Artemisia filifolia Torr. (Diptera: Tephritidae). Pan-Pacif. Entomol., 58: 19-24.

Cavender, G. L. & R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae)

on wild sunflower in southern California. Ann. Entomol. Soc. Am., 75: 400-406.

Dodson, G. 1987. Host-plant records and life history notes on New Mexico Tephritidae (Diptera).

Proc. Entomol. Soc. Wash., 89: 607-615.

Foote, R. H. & F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect

Survey, 7: 1-117.

Freidberg, A. 1984. Gall Tephritidae (Diptera). pp. 129-167. In Ananthakrishnan, T. N. (ed.). Biology

of gall insects. Edward Arnold, London.

Fronk, W. D., A. A. Beetle & D. G. Fullerton. 1964. Dipterous galls on the Artemisia tridentata

complex and insects associated with them. Ann. Entomol. Soc. Am., 57: 575-577.

Goeden, R. D. 1987. Life history of Trupanea conjuncta (Adams) on Trixus californica Kellog in

southern California (Diptera: Tephritidae). Pan-Pacif. Entomol., 63: 284-291.

Goeden, R. D. 1988. Gall formation by the capitulum-infesting fruit fly, Tephritis stigmatica (Diptera:

Tephritidae). Proc. Entomol. Soc. Wash., 91: 37-43.

Goeden, R. D. & D. W. Ricker. 1981. Life history of the gall-forming moth, Carollela beevorana

Comstock, on the ragweed, Ambrosia dumosa (Gray) Payne, in southern California (Lepidoptera:

Cochylidae). Pan-Pacif. Entomol., 57: 402-410.

Goeden, R. D., T. D. Cadatal & G. A. Cavender. 1987. Life history of Neotephritis finalis (Loew)

on native Asteraceae in southern California (Diptera: Tephritidae). Proc. Entomol. Soc. Wash.,

89: 522-558.

Headrick, D. & R. D. Goeden. (in press). Description of the immature stages of Paracantha gentilis

(Diptera: Tephritidae). Ann. Entomol. Soc. Am.

Munz, P. A. 1974. A flora of southern California. University of California Press, Berkeley.

Munz, P. A. & D. D. Keck. 1959. A California flora. University of California Press, Berkeley.

Osten Sacken, C. R. 1877. Western Diptera: descriptions of new genera and species of Diptera from

the region west of the Mississippi and especially from California. U. S. Geological and Geo¬

graphical Survey of the Territories Bull., 3: 189-354.

Steck, G. J. & R. A. Wharton. 1986. Descriptions of immature stages of Eutreta (Diptera: Tephrit¬

idae). J. Kansas Entomol. Soc., 59: 296-302.

Stoltzfus, W. B. 1977. The taxonomy and biology of Eutreta (Diptera: Tephritidae). Iowa State J.

Res., 51: 369-438.

Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of American north of Mexico

(Diptera: Tephritidae). California Dept. Agriculture, Bur. Entomol. Occas. Papers, 19: 1-172.

Received 5 April 1989; accepted 1 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 33-38, (1990)

LIFE HISTORY OF EUTRETA SIMPLEX THOMAS ON

ARTEMISIA L UDO VICIANA NUTTALL IN

SOUTHERN CALIFORNIA (DIPTERA: TEPHRITIDAE)

Richard D. Goeden

Department of Entomology, University of California,

Riverside, California 92521

Abstract. —Eutreta simplex Thomas is univoltine on Artemisia ludoviciana Nuttall (Asteraceae),

its only known host plant reported here for the first time. Eggs are inserted in terminal buds of

young emerging shoots in the fall by the long lived females that emerged from galls late during

the preceding spring. The second instars overwinter. Shoot, gall, and larval growth all resume

early in the next spring. Galls are described in detail and pictured. Pupation occurs in the gall.

Adults emerged from puparia after about three weeks during early July. Females convert their

fat body tissues to yolk, but postpone oviposition until fall; moreover, they apparently can resorb

these ovarian eggs, as a means of increasing fecundity and longevity.

Key Words.— Insecta, Eutreta, Artemisia, gall, life history, ovisorption

My discovery of a somewhat accessible field population as well as the host plant

of the heretofore little known, gall-forming fruit fly, Eutreta simplex Thomas

(Diperta: Tephritidae), allowed me to study those aspects of its life history reported

herein. The conduct of this field study and the interpretation of my findings on

this still rare and elusive tephritid were greatly aided by my concurrent, more

extensive field study of E. diana (Osten Sacken), reported separately (Goeden in

press).

Taxonomy.— Thomas (1914) described E. simplex from a single female. The

adults were illustrated and the male was described, again from a single specimen

by Stoltzfus (1977).

Distribution, Hosts and Study Sites.— The North American distribution of E.

simplex as recorded by Stoltzfus (1977) consists of two sweep records, one for the

holotype female from 2440 m at Sunset, Colorado, 19 Jul 1903, and the other

for a male and female from Big Meadow (probably on San Gorgonio Mountain,

San Bernardino National Forest), SW San Bernardino Co., California, 8 Jul 1950.

A female swept in Yosemite National Park in central California in 1940 by E. E.

Kenage was identified for the U.S. National Museum in 1978 by F. L. Blanc.

The three principal study areas in 1987 and 1988 where I collected galls on

Artemisia ludoviciana Nuttall (Asteraceae) from which I reared E. simplex adults

included: Near the summit (2330 m), at Santa Rosa Spring, and on the south¬

ernmost point along the truck trail (2110 m) on Santa Rosa Mountain of San

Bernardino National Forest, Riverside Co. Collections were also made at Coxey

Meadow (1770 m), San Bernardino Mts., San Bernardino National Forest, San

Bernardino Co., 26 Apr 1988.

Wasbauer (1972) and Stoltzfus (1977) listed no host information for E. simplex.

Whether this tephritid is monophagous on A. ludoviciana (which also occurs in

Colorado: Munz & Keck 1959), or forms galls on one or more other species of

Artemisia (like E. diana: Fronk et al. 1964, Stoltzfus 1977, Benbow & Foster

1982, Goeden in press), is unknown.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Egg. —No intact or newly hatched egg of E. simplex was found in nature;

however, like the egg of E. diana, it probably is inserted for most of its length

singly into a bud (Goeden in press). The terminal buds of succulent young shoots

formed in the fall, and arising from underground rhizomes (Fig. la), are the

probable oviposition sites of E. simplex. Only one empty, apparently infertile egg

was recovered from this position following week-long cagings of six mature lab¬

oratory-reared females with field-collected bouquets of these young shoots; the

origin of this egg was uncertain. No eggs or newly hatched first instars were

dissected from another 21 of these young shoots collected on 3 Nov 1988 from

the same immediate areas at Santa Rosa Springs where galls were harvested earlier

in the year.

Twenty eggs were dissected from a newly dead, 115 day old, laboratory-reared

female (Fig. lb). Like those of E. diana, these ova were fusiform, smooth, white,

rounded at both ends, with a peg-like 0.03 mm pedicel at the cephalic end. They

averaged 0.70 ± 0.006 mm (X ± SE) long and 0.25 ± 0.003 mm wide, slightly

longer on average but with shorter pedicels than those of E. diana (Goeden in

press).

Larva. — Despite repeated searches of specific areas previously yielding mid- to

full-size galls, no immature gall was found on current season shoots in October

and November, 1987 and 1988. The herbaceous galls of E. simplex on A. ludo-

viciana were observed to persist for at least six months after the adults had

emerged, but most decayed or became buried by the second year; moreover, each

emergent shoot is galled only once. Nevertheless, collection of several half-grown,

overwintered galls containing second instars (May 1987 and 1988, Santa Rosa

Mountain, Fig. lc) strongly indicates this is the overwintering stage in southern

California, as for E. diana (Goeden in press).

The fully developed, mature monothalmous galls of E. simplex consist of short¬

ened and thickened terminals of sessile or short-stalked vegetative shoots formed

at or just above the litter or soil surface (Fig. Id). Externally, they are subovoidal,

smooth, grey-green tomentose, and bear sessile leaves with expanded, transversely

oriented, fish scale-like leaf bases. The leaf blades elongate at the gall apices to

form characteristic, flattened rosettes 2-6 cm in diameter (Fig. le). The lengths

and widths of 38 galls (collected on Santa Rosa Mountain and containing fully

grown larvae or puparia, Fig. If, lg) averaged 12.0 ± 0.4 (8-20) mm and 9.0 ±

0.2 (6-11) mm, respectively, after leaves were removed. The largely buried ped¬

icels measured up to 7.6 cm in length between gall bases and their juncture with

the rhizome (Figs. If, lg). The central feeding chambers were clavoidal, open,

rounded basally, narrowing apically, and measured 7.0 ± 0.3 (6-9) mm long and

3.0 ± 0.1 (2-4) mm wide (n = 17). The walls of the galls were pitted and covered

with frass from larval feeding (Fig. If). The walls of mature galls were thickest

basally (4-7 mm), thinner laterally (2-3 mm), and thinnest apically (< 1 mm) at

the point of exit for the adult (Figs. If, lg).

Galls were found singly or in small groups of up to three beneath conifers on

steep (45°), shaded, north-facing slopes in thick litter (upper site, Santa Rosa

Mountain, 1987). They were scarce at this location in 1988. The first galls found

were on plants growing intermixed with Eriogonum wrightii Torrey ex Bentham

in an open, south-facing, gently sloping area (lowest site, Santa Rosa Mountain,

1987). None were found there in 1988. Since E. diana galls on A tridentata usually

1990

GOEDEN: LIFE HISTORY OF EUTRETA SIMPLEX

Figure 1. Life stages and galls of Eutreta simplex on Artemisia ludoviciana: (a) emergent and

emerging vegetative shoots (lx); (b) photomicrograph of an unresorbed ovum (~ 100x); (c) over¬

wintered second instar in gall (3.5 x); (d) lateral view of mature gall (2 x); (e) view of mature gall from

above (1.5 x); (f) saggital view of mature gall containing full-grown third instar (5.5 x); (g) puparium

in mature gall (4.4 x); (h) mating adults, lateral view (4.7 x); (i) mating adults, dorsal view (4.6 x).

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

were located in drainages or riparian habitats (Goeden in press), I searched for

galls at Santa Rosa Springs; most galls were found there in 1988 on plants growing

on steep, moist, north-facing slopes beneath conifers.

Pupa. — The puparia (Fig. lg) are formed with their anterior ends at or within

the central, apical meristem. This usually is destroyed by larval feeding or adult

emergence. The puparia are smooth, ellipsoidal, bluntly rounded posteriorly, and

slightly more tapered anteriorly. They are translucent mustard-yellow at first but

later turn dark orcherous brown at both ends. Puparia (n = 21) averaged 5.3 ±

0.08 (4.40-5.8) mm in length and 2.2 ± 0.04 (1.9-2.5) mm in maximal width.

Pupation lasted about three weeks (27 ± 2° C, humidity chamber). One dead

adult was observed that had become trapped in its gall upon emerging; it could

not break through the tissue layer (left uneaten as a larva) at the gall apex. Galled

shoots usually did not elongate apically after adult emergence, although if the

larva within a gall died the terminal bud growth resumed forming a more elongate,

narrower, fusiform gall. The shapes of E. diana galls on A. tridentata were similarly

influenced by larval mortality and terminal bud reactivation (Goeden in press).

Adult.— No adults of E. simplex were found. Ten males and 13 females were

reared from larvae and puparia dissected from galls collected in 1988, emerging

during early July. The four galls collected at Coxey Meadow in late April (same

year) yielded three females emerging in late May.

The adults were caged separately in screened-topped, clear-plastic, 850-ml cages

supplied with a water-wick and honey to study longevity. They remained relatively

inactive when isolated, usually resting on the side of the cage facing laboratory

windows. Adult males lived 51-121 days ( x = 85), and females (n = 11) lived

26-128 days (x = 83). Dissections indicated females were sexually immature at

emergence, but (like males) had substantial fat tissue. In females the fat bodies

subsequently were sequestered for yolk in ova (Fig. lb).

Mating occurred readily without elaborate courtship behavior when surviving

males and females were caged together in pairs about six weeks after emergence.

During premating behavior, flies faced each other at a distance of 1.5-2.0 cm and

slowly, alternately, waved their wings. The male then climbed atop the female

from the rear. While in copula the male was largely off the substrate (Fig. lh),

except for his hind tarsi, positioned above the posterior of the thorax and abdomen

of the female. His wings were held motionless, straight backward, overlapping,

slightly parted, and flat upon his dorsum (Fig. li). The male’s mouthparts touched

the female’s abdominal dorsum anteriorly, as both sexes pumped their mouthparts

rhythmically. The female’s wings were held perpendicularly and angled at 45°

from the body (Fig. li). The male’s foretarsi grasped the female’s abdomen an-

terolaterally, his mesotarsi grasped her abdomen mid-laterally, while his metatarsi

either intermittently stroked her oviscape, held it posteriorly or rested on the

substrate. Matings observed in cages (n = 5) were protracted (3.5-4 h), usually

beginning in the afternoon and ending at dusk (19:00). Twice matings were ex¬

tended under artificial light for 6 and 9 h. Stoltzfus (1977) observed a caged pair

of E. caliptera (Say) that remained in copula for about 4 h. Whether copulations

of such duration normally occur is questionable. One male E. simplex died shortly

after two protracted copulations on successive days. Stoltzfus (1977), however,

reported that a pair of E. caliptera copulated on each of seven successive days!

Ovaries of four females that lived 115, 116, 127, and 128 days were found to

1990

GOEDEN: LIFE HISTORY OF EUTRETA SIMPLEX

contain 66, 66, 58, and 48 full sized ova. The female that lived 127 days contained

white, intact, unresorbed ova, but the other three females also contained varying

proportions of nonwhite translucent ova whose contents had been partly resorbed.

Ovisorption has long been known in parasitic Hymenoptera as an adaptation for

extending the fecundity and longevity of synoovigenic females (c.f., Flanders 1935,

Legner & Gerling 1967, Legner & Thompson 1977). Ovisorption in long-lived

adult Tephritidae was first observed in Trupanea conjuncta (Adams) (Goeden

1987), and also occurs in long-lived adults of certain species of Procecidochares

(unpublished data).

Seasonal History. —Eutreta simplex is univoltine on A. ludoviciana in southern

California, as is E. diana on A. tridentata (Goeden in press); however, their

seasonal histories show several major differences. Both species: overwinter as

second instars in small, undeveloped bud galls; complete larval and gall devel¬

opment once vegetative growth resumes in the spring; pupariate in their galls;

and emerge as sexually immature adults. However, E. diana adults mature, mate,

and begin oviposition within three weeks. The larvae pass the summer and fall

in diapause as first instars, and molt to the second instar just before winter begins.

In contrast, E. simplex adults are long-lived and postpone their oviposition until

fall. Eutreta diana on A. filifolia Torrey, another herbaceous host, displays a

different seasonal history in Texas involving fall-emerging adults and overwin¬

tering eggs, but no summer and fall adult aestivation or larval diapause (Benbow

& Foster 1982).

Natural Enemies.— No parasitoids and only one parasite exuvium were re¬

covered from dissected galls or puparia of E. simplex. In contrast E. diana has a

well-developed complex of natural enemies (Fronk et al. 1964, Benbow & Foster

1982, Goeden in press). Thirty-two galls were collected at the highest study area

on Santa Rosa Mountain on 21 Oct 1987. Upon dissection: 19 contained empty

puparia indicative of fly emergence; eight had apical openings but were empty,

indicating predation; three were intact and small, containing carcasses of larvae

that died of unknown causes; one intact gall contained the dead adult that failed

to emerge; and only one had an exit hole and contained a larval carcass and

parasitoid exuvium. No symptoms of bird predation or gall inquilines were ob¬

served with E. simplex, as for galls of E. diana on A. tridentata (Goeden in press).

Acknowledgment

My thanks to D. W. Ricker for technical assistance, including the insect pho¬

tography involved in Fig. 1. Thanks also to F. L. Blanc, D. H. Headrick, and B.

A. Stoltzfus for their helpful comments on early drafts of the manuscript.

Literature Cited

Benbow, S. M. & D. E. Foster. 1982. Biology of Eutreta diana Osten Sacken on sand sagebrush

Artemisia filifolia Torr. (Diptera: Tephritidae). Pan-Pacif. Entomol., 58: 19-24.

Flanders, S. E. 1935. An apparent correlation between the feeding habits of certain pteromalids and

the condition of their ovarian follicles. Ann. Entomol. Soc. Am., 28: 438-444.

Fronk, W. D., A. A. Beetle & D. G. Fullerton. 1964. Dipterous galls on the Artemisia tridentata

complex and insects associated with them. Ann. Entomol. Soc. Am., 57: 575-577.

Goeden, R. D. 1987. Life history of Trupanea conjuncta (Adams) on Trixus californica Kellogg in

southern California (Diptera: Tephritidae). Pan-Pacif. Entomol., 63: 284-291.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Goeden, R. D. (in press). Life history of Eutreta diana (Osten Sacken) on Artemisia tridentata Nuttall

in southern California (Diptera: Tephritidae). Pan-Pacif. Entomol., 66: 24-32.

Legner, E. F. & D. Gerling. 1967. Host-feeding and oviposition on Musca domestica by Spalangia

cameroni, Nasonia vitripennis and Muscidifurax raptor (Hymenoptera: Pteromalidae) influences

their longevity and fecundity. Ann. Entomol. Soc. Am., 60: 678-691.

Legner, E. F. & S. N. Thompson. 1977. Effects of the parental host on host selection, reproductive

potential, survival and fecundity of the egg-larval parasitoid, Chelonus sp. near curvimaculatus

Cameron, reared on Pectinophora gossypiella (Saunders) and Phthorimaea operculella (Zeller).

Entomophaga, 22: 75-84.

Munz, P. A. & D. D. Keck. 1959. A California flora. University of California Press, Berkeley.

Stoltzfus, W. B. 1977. The taxonomy and biology of Eutreta (Diptera: Tephritidae), Iowa State J.

Res., 51: 369-438.

Thomas, F. L. 1914. Three new species of Trypetidae from Colorado. Can. Entomol., 46: 425-429.

Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico

(Diptera: Tephritidae). California Dept. Agriculture, Bur. Entomol. Occas. Papers, 19: 1-172.

Received 5 April 1989; accepted 1 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 39-42, (1990)

A MORPHOMETRIC ANALYSIS OF MELANOPLUS

FEMALES (ORTHOPTERA: ACRIDIDAE)

S. J. Guenther and W. Chapco

University of Regina, Regina, Saskatchewan S4S 0A2, Canada

Abstract. — Females of certain species-groups in the genus Melanoplus are difficult to distinguish

morphologically, especially for the novice. Three linear discriminant functions based upon mor¬

phometric traits are defined that allow separation of females of M. sanguinipes Fabr. from each

of M. femurrubrum De Geer, M. confusus Scudder, and M. gladstoni Scudder.

Key Words. — Insecta, Orthoptera, Acrididae, Melanoplus, discriminant function

Species of Melanoplus are conventionally identified by genital characters of the

male. Females of certain species-groups, however, are difficult to distinguish,

especially for the novice. One practice is to associate co-occurring females with

identified males; this is probably satisfactory if males of no other Melanoplus

species are found. Nevertheless, congeneric species do often appear in the same

location. It is not uncommon to find M. sanguinipes Fabr. associated with M.

femurrubrum De Geer, M. confusus Scudder, and M. gladstoni Scudder. Features

used to distinguish M. sanguinipes from these species are unreliable. For instance,

the red-legged trait of M. femurrubrum also occurs polymorphically in M. san¬

guinipes (Chapco 1983). The early spring emergence of M. confusus can aid in its

separation from M. sanguinipes, but by late June adults of both species are found.

The more robust M. gladstoni is less difficult to distinguish from M. sanguinipes.

Pale laterodorsal lines which form a diamond-back marking are common in the

former, but M. sanguinipes also has this trait polymorphically (Bidochka 1984).

Brooks’ (1958) monograph on Acridoidea of the Canadian Prairies states that

there are differences in the dorsal angle of ovipositor among Melanoplus species,

but we have found considerable variation for the trait; moreover, its measurement

is awkward. We demonstrate that discriminant functions based on a few easily

made measurements can, with reasonable confidence, help separate M. sangui¬

nipes from M. femurrubrum, M. confusus and M. gladstoni.

Materials and Methods

Eleven morphometric traits were determined for roughly 30 specimens per

species: prozona length (PZL), metazona length (MZL), pronotum width (PRW),

head width (HW), eye height (EH), eye width (EW), femur length (FL), femur

width (FW), tibia length (TL), interoccular distance (ID), and suboccular suture

length (SSL). Traits can be easily measured to the nearest 0.5 mm with calipers.

Melanoplus sanguinipes, M. femurrubrum, and M. confusus were collected within

a 20 km radius of Regina, Saskatchewan; M. gladstoni were collected at the Last

Mountain House Historical site, about 45 km NW of Regina.

Results and Discussion

Table 1 shows mean values for each trait and species; pooled within species

mean square errors (MSE) provide measures of variability. Because analysis of

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

variance (not shown) revealed significant interspecific variation for all traits, a

Duncan’s multiple range test (a = 0.01) was used to determine homogeneous

groupings. Uniform sets are indicated by common letters (a, b, .. .) in the table.

Although Melanoplus femurrubrum appears as the smaller species in Table 1,

there is no consistent ranking among the other species. Many of the shape indices

(ratios of various traits such as MZL/PZL, TL/FL, etc., see Eades 1970) custom¬

arily used in morphometric investigations of acridids were calculated, but inter¬

estingly, with a few exceptions (see below), these showed considerable overlap

among species. Allometric similarity for body proportions may, therefore, explain

why some Melanoplus groups are difficult to identify.

Discriminant functions (DF) between M. sanguinipes and each of the other

three species were arrived at by applying stepwise discriminant analysis (Wilks)

to all variables and to subsets of variables and derived ratios (e.g., MZL/PZL,

PZL/FW, etc.). Those sets with the largest discriminatory power (percentage of

correctly classified specimens) were retained.

Resultant discriminant functions and classification rules follow:

1. M. sanguinipes vs. M. femurrubrum :

DF = (-11.0366) + (0.8882 x (TL))

- (9.3904 x (PZL/FW)) + (0.7142 x (FL))

If DF < 0.0787, classify as M. femurrubrum.

If DF > 0.0787, classify as M. sanguinipes.

96.6% correct classification

2. M. sanguinipes vs. M. confusus :

DF = (—10.2199) + (3.1389 x (FW)) - (4.0698 x (PZL))

+ (2.0752 x (HW)) - (1.9843 x (SSL)) + (0.9190 x (FL))

- (1.0061 x (TL) + (1.1778 x (EH))

If DF ^ 0, classify specimen as Af. confusus.

If DF 0, classify specimen as AT. sanguinipes.

96.4% correct classification.

3. M. sanguinipes vs. M. gladstonv.

DF = (—13.3864) - (1.8198 x (EW)) + (2.1149 x (FL))

- (1.7867 x (PZL + MZL))

If DF < 0.0984, classify specimen as M. gladstoni.

If DF > 0.0984, classify specimen as M. sanguinipes.

98.3% correct classification.

Traits are not difficult to measure although the inclusion of seven variables in the

separation of M. sanguinipes and M. confusus may be bothersome to some in¬

vestigators. An alternative discriminant function based on five variables yielded

92.7% correct classification and may be more attractive. This is given by

DF = (-0.6759) - (1.2193 x (FL)) + (1.0714 x (TL))

- (1.6382 x (EH)) + (12.0694 x (PZL/FW)).

To classify specimens, their character measurement values are substituted into

1990

GUENTHER & CHAPCO: ANALYSIS OF MELANOPLUS

Table 1. Mean values for 11 morphometric traits for four Melanoplus species. Letters following

values indicate homogeneous subsets of species. MSE = mean square error with df = 111. See text for

meaning of other symbols.

Trait

fern

sang

conf

glad

MSE

PZL

2.10a

2.29b

2.58c

2.32b

0.054

MZL

2.33a

2.77c

2.63bc

2.53b

0.060

PRW

3.20a

3.71b

3.78bc

3.92c

0.060

3.70a

4.28b

4.15b

4.17b

0.064

2.22a

2.43b

2.25a

2.43b

0.047

1.52a

1.71b

1.52a

1.83c

0.034

11.32a

13.02b

12.69b

11.17a

0.451

2.98a

3.55d

3.22b

3.38c

0.049

9.18a

10.59b

10.74b

9.33a

0.312

IND

0.93a

l.OOab

1.05b

0.013

SSL

1.50a

1.57a

1.52a

1.80b

0.052

Sample n

lll(df)

the appropriate discriminant function and their DF scores are assessed accord¬

ingly. For example, one specimen suspected to be M. femurrubrum had the fol¬

lowing measurements: TL = 9.0, PZL = 2.0, FW = 3.0, FL = 10.5. Substitution

into the first discriminant function yielded a discriminant score DF = (—11.0366)

+ (0.8882 x (9.0)) - (9.3904 x (2.0/3.0)) + (0.7142 x (10.5)) = -1.80. Because

-1.80 < 0.0787, the specimen is identified as M. femurrubrum.

It is not known whether these discriminant functions can be applied to speci¬

mens outside the geographic range of the present collection; this is currently being

investigated. Because the samples used to establish the DFs were the same as

those used to test the procedure, quoted discriminatory powers are likely over¬

estimates of true values. A more appropriate test of discriminatory power could

be achieved by randomly splitting samples into parts and using one part for DF

construction and the other part for assessing power. Although sample sizes larger

than those used here are usually required, the procedure was tried using the same

variables as above. Discriminatory powers then dropped to 84.0%, 66.7% and

90.6% for “unknown” samples of M. sanguinipes and each of M. femurrubrum,

M. confusus, and M. gladstoni, respectively. For the alternative DF involving M.

confusus, the percentage of correctly classified specimens remained high at 92.3%.

Despite shortcomings our procedure illustrates that a high degree of discrimination

among morphologically similar females of different species is possible using rel¬

atively few variables; researchers in other locations could define similar functions.

Acknowledgment

We thank the Natural Sciences and Engineering Research Council of Canada

(Grant A-0485 to WC) for financial support.

Literature Cited

Bidochka, M. J. 1984. Genetic variation in natural populations of the migratory grasshopper, Me¬

lanoplus sanguinipes. M.Sc. Thesis, University of Regina.

Brooks, A. R. 1958. Acridoidea of southern Alberta, Saskatchewan and Manitoba (Orthoptera). Can.

Entomol., Supl. 9.

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Vol. 66(1)

Chapco, W. 1983. The genes, T R , Ost, Pro R and L in natural populations of Melanoplus sanguinipes.

Can. J. Genet. Cytol., 25: 88-92.

Eades, D. C. 1970. Theoretical and procedural aspects of numerical phyletics. Syst. Zool., 19: 142-

171.

Received 10 April 1989; accepted 7 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 43-54, (1990)

THE INTRODUCTION OF SIPHONINUS PHILLYREAE

(HALIDAY) (HOMOPTERA: ALEYRODIDAE) INTO

NORTH AMERICA: NICHE COMPETITION,

EVOLUTION OF HOST PLANT ACCEPTANCE,

AND A PREDICTION OF ITS POTENTIAL RANGE

IN THE NEARCTIC

John T. Sorensen, 1 Raymond J. Gill, 1

Robert V. Dowell 2 and Rosser W. Garrison 3

insect Taxonomy Laboratory and 2 Pest Detection/Emergency Projects,

California Department of Food & Agriculture, Sacramento, California 95814

3 Los Angeles County Agricultural Commissioner’s Office,

El Monte, California 91732

Abstract.— Siphoninus phillyreae (Haliday), the ash whitefly, was initially collected in North

America in Los Angeles, California, August 1988 and underwent a vast population explosion

shortly afterwards. We document the current status of its expanding distribution in North Amer¬

ica and note over 25 host plant species/varieties and five plant families that are used by this

whitefly in California but unreported as hosts in its native Palaearctic range. We predict a potential

ultimate range in the Nearctic for the whitefly, based upon isotherm data with reference to its

Old World distribution. We list the diagnostic features of the species, and comment on the

taxonomic status of S. phillyreae, noting that pupae from California seem closer in appearance

to material from Egypt than from southern Europe. We suggest that acceptance of new host

plants in California may be a mutation driven phenomenon with the increased host acceptance

mutations due to its extreme and unchecked population size. We comment upon the potential

host plant niches that this whitefly will occupy in the Nearctic, and the taxa that could be its

most serious ecological competitors in these niches.

Key Words. — Insecta, Aleyrodidae, Siphoninus phillyreae, ash whitefly, Nearctic distribution,

host plant evolution, niche competition

On 18 Aug 1988, Siphoninus phillyreae (Haliday), the ash whitefly, was collected

for the first time in North America by R. Orsbum in Los Angeles, California.

This species’ previous distribution was palaearctic, from Ireland, Morocco and

Cameroun in the west to India in the east. Shortly after its collection, an extraor¬

dinary population explosion of S. phillyreae occurred throughout the Los Angeles

basin which was attributed to a lack of natural enemies. Populations grew rapidly

with the flying adults described as appearing similar to a light snow flurry. Within

a year the species distribution within California (Fig. 1) had spread along the

southern California coast from Santa Barbara to San Diego, and ranged inland

to Riverside (Riverside Co.), Victorville (San Bernardino Co.) and Lancaster (Los

Angeles Co.) in the desert; by December 1989 it had expanded through California’s

central valley to Sacramento, and to San Jose in the San Francisco Bay Area,

probably through the movements of infested plant material. In December 1989,

S. phillyreae was found infesting nursery stock arriving in Los Angeles from

Phoenix, Arizona; the whitefly had been found in Phoenix two months before.

The severe infestation found in the Los Angeles basin has caused honeydew

and sooty mold problems, as is common with other whiteflies but seldom noticed

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

because of their smaller populations. The whitefly is not dependent upon fresh,

flush growth on its hosts as are several other aleyrodids (e.g., Parabemisia myricae

[Kuwana], Walker & Aitken 1985) which have been recently introduced into

California. Therefore, the entire shrub or tree canopy is subject to infestation

during the season. Locally occurring aleyrodid natural enemies have not been

found associated with the S. phillyreae populations. Although this whitefly has

been reported to have two to three generations per year in Egypt (Priesner &

Hosny 1932), it is thought to have considerably more in southern California with

a potential generation time of 25 days (T. Bellows, personal communcation).

This explosive, invasive occurrence has revealed several interesting biological

1990 SORENSEN ET AL.: SIPHONINUS PHILLYREAE IN NORTH AMERICA 45

Figure 2. Pupae of Siphoninusphillyreae (after Priesner & Hosney 1932). Smaller pupae (described

as S. granati by Priesner & Hosney) and larger pupae (figured as S. phillyreae by Priesner & Hosney).

Note size related occurrence of siphon tubes on dorsomedial abdominal segments.

aspects about this whitefly which are atypical of most others that have become

established in California. We comment upon some of these aspects and their

evolutionary significance in the expanding allopatric distribution of this species.

Taxonomy

Siphoninus phillyreae has a unique phena in its pupal stage. The living pupa is

pale with a melanic stripe dorsomedially; its dorsum is covered with ‘siphon

tubes’ that are somewhat similar in appearance to the siphunculi (cornicles) of

aphids. Each siphon produces a droplet of wax that causes the entire structure to

appear as a glassy club; there are 40-50 such clubs on each pupa. The pupae also

have a dorsomedial series of tufts composed of white fibrous wax which form a

dorsomedial line obscuring medially located siphon tubes.

Slide mounted pupae show these numerous siphon tubes and a dorsomedial

melanic stripe that fades in the middle of the otherwise pale body (Fig. 2). Live

adults are undistinguished, but slide mounted males have a single posteromedially

directed tooth that is immediately anteriad of the terminal process (apical tooth)

on each clasper (Fig. 3); this tooth is apparently homologous with the subapical

tooth of the clasper (Gill in press) exhibited by other aleyrodids.

Mound & Halsey (1978) consider S. phillyreae to be a single, but variable

species, noting that the number and placement of the dorsal siphons on its pupal

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Operculum

Terminal

Abdominal

Segment

Clasper

(Paramere)

Subapical Tooth

Lingula

Aedeagus

Inflatable Sac

Terminal Process

(Apical Tooth)

Figure 3. Male genitalia of Siphoninus phillyreae, dorsal view, anterior to top.

cases is variable and is related in part to the overall size of the pupal case (Goux

1949); see Fig. 2 for size-related variability in siphon numbers, where larger

specimens reputedly show more siphons in dorsomedial series on the mesal por¬

tion of the abdominal dorsum. Mound & Halsey’s (1978) assessment of S. phil¬

lyreae as a single (taxonomic) entity has considerable importance to its successful

biological control because if a species-complex or series of biotypes were involved,

it could complicate efforts to find compatible controlling agents.

Material in California thus far examined shows a characteristic heavy dorso¬

medial band of wax. However, CDFA efforts in August, 1989, to secure natural

enemies in northern Italy and adjacent France found that S. phillyreae pupae in

that area lack such a conspicuous wax band (L. Bezark, personal communication).

The California material appears similar to Egyptian material of S. granati Priesner

& Hosny (1932: plate 1) described from pomegranate. That species nomen, as

well as inaequalis Gautier, dubiosa Haupt, dubious Heeger, phylliceae Bouche,

finitimus Silvestri, and multitubulatus Goux, have been synonymized under phil¬

lyreae Haliday by Mound & Halsey (1978). It will be interesting to find out if

imported and released biological control agents “recognize” such taxonomic treat¬

ment.

Biology

Potential Nearctic Range.—We suspect that S. phillyreae, if it is a single and

uniform biological entity with respect to temperature tolerances, will continue to

1990 SORENSEN ET AL.: SIPHONINUS PHILLYREAE IN NORTH AMERICA 47

Figure 4. Isotherm map of Europe for average daily temperature in January. Siphoninus phillyreae

occurs throughout the area shown, north to approximately the -7° C (20° F) isotherm, except in

southern Norway and Sweden.

expand its range well into temperate North America. This is because unlike many

more tropical aleyrodids that have been introduced into California, such as Aleu-

rothrixus floccosus (Maskell) or Dialeurodes citrifolii (Morgan) (Dowell & Gill

1989), S. phillyreae occurs well north into Europe in Germany, Poland and the

western and southern U.S.S.R. Its Palaearctic distribution appears limited to south

of isotherms for an average January daily temperature of between — 7° C (20° F)

and — 1° C (30° F) (Fig. 4), although it has not been reported in southern Norway

or Sweden (Hulden 1986) which also fall into this zone. If these same isotherms

are also limiting in North America, S. phillyreae could potentially expand its range

well into the Nearctic (Fig. 5) to southwestern British Columbia, western Montana,

southern South Dakota, extreme southern Minnesota, and eastward across south¬

ern portions of Wisconsin, Michigan, and Ontario, to New England.

Host Associations. — This whitefly is polyphagous on relatively hard-leaved shrubs

and small trees in the Palaearctic, where it shows host associations in the Oleaceae

(Fraxinus, Olea, Phillyrea ) and the Rosaceae ( Crataegus, Cydonia, Mespilus, Pru-

nus, Pyrus ), but is also recorded from the Leguminosae ( Afzelia ), Punicaceae

(Punica), Rhamnaceae ( Rhamnus ) (Mound & Halsey 1978) and Rutaceae ( Citrus)

(Khan et al. 1985). In California, S. phillyreae has been found on many hosts

(Table 1) including over 25 species (and varieties) and five families not recorded

in the Palaearctic. (This list has been updated while in press to reflect additional

hosts reported by Bellows et al. [1990] and subsequent data, these have not [yet]

been verified by CDFA.)

Although S. phillyreae has been found to use many new plants in California,

it shows some differentiation in its acceptance of hosts. The preferred hosts in

California have thus far been evergreen ash ( Fraxinus uhdei [Wenzig] Lingelsheim)

and evergreen (flowering) pear {Pyrus kawakamii Hayata), which seem particularly

susceptible with large populations causing partial defoliation (Gill 1989). In con-

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Figure 5. Isotherm map of North America for average daily temperature in January. As judged

by its distribution in Europe with relation to climate, Siphoninus phillyreae should be able to tolerate

those areas in North America north to the -7° C (20° F) or at least the -1° C (30° F) isotherm line.

trast, this whitefly seems to use Rhaphiolepis (K. Arakawa, personal communi¬

cation) and Citrus mostly during winter months in southern California, when

preferred deciduous hosts are barren.

Siphoninus phillyreae appears to survive the winter in all stages in southern

California on some of its nondeciduous hosts (e.g., Citrus, Rhaphiolepis, Hetero-

meles). This facultative use of “overwintering” hosts in such climates has no

doubt aided S. phillyreae in building up the dramatic numbers that are reported

in the Los Angeles basin. By using evergreen overwintering hosts in California,

populations of this whitefly need not crash in the fall, when their preferred de¬

ciduous hosts are unavailable. This permits a large number of surviving individ¬

uals in the spring to allow a greater population growth during the next season, as

seen in the expression: population growth = r m N, where r m is the species’ innate

capacity for increase (Andrewartha & Birch 1954) and N is the surviving popu¬

lation in the spring. As S. phillyreae moves into more continental climates in the

Nearctic, we expect that it will cease to have year-around breeding populations

and that it will necessarily exhibit the distinct seasonality seen in northern Europe.

Evolution of Host Acceptance. —The prediction of ultimate range potential in

the Nearctic assumes that compatible hosts are available and that the source

population for the California introduction has an unrestricted genetic tolerance

1990 SORENSEN ET AL.: SIPHONINUS PHILLYREAE IN NORTH AMERICA 49

for cold with respect to the overall genome of S. phillyreae. Since “founder”

populations are considered to classically show restricted genetic variance with

respect to entire parental populations across their geographic ranges (Lewontin

1974), it is doubtful the last assumption is true. Alternatively, however, California

populations of walnut husk fly, Rhagoletis completa Cresson, derived from intro¬

duced founders from parental populations in eastern North America, have been

found (Berlocher 1984) to have developed both differing patterns of isozymic

variation and greater isozymic variation around individual locii, than occurs in

the populations in eastern North America. Hence, evolutionary mechanisms ap¬

parently exist in some cases to increase genetic variance and heterozygosity in

populations derived from genetically restricted founder groups. Such newly dis¬

covered and poorly understood evolutionary mechanisms make potential and

rapid evolutionary change under allopatric conditions over short time periods

increasingly feasible and probable; this is because the genetic variance of daughter

populations may be increased, allowing them to meet selection pressures of their

new environment that their parental populations were not subjected to.

One potential result of such an evolutionary mechanism in the case of S.

phillyreae might be the increase in acceptable host plants (both as numbers of

species and particularly families) that we report as used in California in comparison

to the Palaearctic. The increase in acceptable hosts is no doubt a result of either

favorable allelic (re)combinations or mutations allowing exploitation of a new

host. Such host plant utilization models based upon genetic variance and com¬

binations have been proposed in tephretids previously (Bush 1969, 1975). Under

these models, the genetic variance in some Rhagoletis species has been considered

great enough to create discrete behavioral traits for host selection; because mating

occurs on hosts this has led to differentiation of mating sites and therefore to

potential sympatric speciation between individuals accepting different hosts. It is

doubtful, however, that simply new allelic combinations alone (barring mutations)

are responsible for the increased level of host acceptance seen in California for

S. phillyreae, because surely such recombination would have occurred previously

in the species’ palaearctic range. Rather it is likely that mutations have caused

the observed increase in host acceptance.

Ordinarily, mutations are usually estimated to occur at one mutation per 10 -5

or 10 -6 individuals in the natural environment for organisms in general (Mettler

& Gregg 1969). This rate has been reported to be as high as more than 1 per 10 -4

for some traits in Drosophila (Dobzhansky 1970). Clearly, at these higher rates

(> 1 mutant individual per 10,000) the explosive population growth exhibited

by S. phillyreae in California (or other new agricultural pests in new environments

free of natural enemies) would provide adequate numbers from which new ac¬

ceptable host plant mutants might occur. Normally, the number of mutations in

populations shows a balance between those mutations imparting an increased

selective advantage and those that are lethal or sublethal. When populations

suddenly begin to grow rapidly, however, the ascendancy of advantageous genes

do not necessarily have to occur at the expense of normally lethal genes; the latter

would thus functionally be eliminated more slowly (Emlen 1973). This is because

if lethal genes were initially in equilibrium in a population, any rapid population

increase would lessen the selection pressures upon them (in terms of population

rather than individual response), allowing their frequency to increase, and ulti-

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Table 1. Hosts recorded for Siphoninus phillyreae in California from which adults have emerged.

Family

Species

Common name

Apocynaceae: a ’ c

Plumeria rubra L. a c

plumeria

Bignoniaceae: 3

Catalpa sp. 3 ’ b

catalpa (hybrid)

Leguminosae:

Cercis occidentalis Torrey 3

western redbud

Cercis siliquastrum L. a - C

Judas tree

Lythracaceae: 3

Lagerstroemia indica L. a

crape myrtle

Magnoliaceae: 3

Liriodendron tulipifera L. a

tulip tree

Magnolia koba stellata Maximowicz 3

star magnolia

Oleaceae:

Fraxinus latifolia Bcntham 33

Oregon ash

Fraxinus uhdei (Wenzig) Lingelsheim

Shamel ash

Fraxinus uhdei (Wenzig) Lingelsheim

‘Tomlinson’ 3 ’ 3

Tomlinson ash

Fraxinus velutina Torrey ‘Modesto’ 3 - 3

Modesto ash

Fraxinus velutina var. glabra Rehder 3

Arizona ash

Fraxinus velutina var. coriacea (Watson)

western (or leather-

Rehder 3

leal) ash

Li gust rum sp. 3

privet

Phillyrea latifolia L.

phillyrea

Syringa x hyacinthifolia (Hort. Lemoine) Rehder 3

excel lilac (hybrid)

Syringa laciniata Miller 3 ' 3

cut-leaf lilac

Syringa vulgaris L. 3

lilac (or common lilac)

Punicaceae:

Punica granetum L.

pomegranate

Rosaceae:

Amelanchier sp. 3 ' d

serviceberry

Chaenomeles speciosa Nakai 3

flowering quince

Eriobotrya deflexa (Hemsley) Nakai 3

golden loquat

Eriobotrya japonica (Thunberg) Lindley 3 ’ 3

loquat

Heteromeles arbutifolia (Aiton) M. Roemer 3

toyon

Malus floribunda Siebold 3

Japanese flowering

crabapple

Malus fusea (Rafinesque) C. K. Schneid 3

Oregon crabapple

Malus scheidecker Zabel 3

Scheider crabapple

Malus pumila P. Miller 3

apple

Malus sp. ‘Hopa’ 3 - 3

crabapple

Malus sp. ‘Red Jade’ 3 3

crabapple

Prunus armeniaca L. 3

apricot

Prunus x blireiana Andre 3

blue plum (hybrid)

Prunus salicina Lindley ‘Santa Rosa’ 3

Santa Rosa plum

Prunus virginiana var. melanocarpa (A. Nelson)

Sargent 3

chokecherry

Pyracantha sp. 3

firethom

Pyrus calleryana Decaisne

ornamental pear

Pyrus communis L.

pear

Pyrus kawakamii Hayata 3

evergreen pear

Pyrus pyrifolia Nakai

Japanese sand pear

Rhaphiolepis indica Lindley 3

Indian hawthorn

Rubiaceae: 3

Cephalanthus occidentalis var. californicus

Bentham 3

buttonbush

Rutaceae:

Citrus aurantifolia Swingle 3

lime

Citrus limon (L.) Burman f. ‘Meyer’ 3

Meyer lemon

Citrus reticulata Blanco

tangerine

Citrus sinensis Osbeck ‘Valencia’ 3 ’ 1-

Valencia orange

Fortunella sp. 3 ’ 3

kumquat

a Host associations recorded from California only.

b Hybrid species reported as “ x Chiopa .”

1990 SORENSEN ET AL.: SIPHONINUS PHILLYREAE IN NORTH AMERICA 51

mately allowing the incorporation of these normally lethal genetic factors into the

genetic variance in ways which may occasionally prove to be beneficial innovations

(Emlen 1973). (For instance, a mutation allowing use of a new host plant might

be functionally lethal or sublethal to a species with a requirement for sexual

[recombinant] reproduction, if a mutated female on the new host could not find

mates which occur on the normal host[s].) The result is that more rapidly fluc¬

tuating populations generally display a greater increase in changes for selected

traits than do stable populations (Ford 1971).

An alternative explanation for S. phillyreae’s increase in host acceptance in

California over that of the Palaearctic is that previous host reporting in the Old

World was inadequate. This is unlikely, however, because the whitefly is a serious

agricultural pest and has been examined by many workers (for a bibliography see

Mound & Halsey 1978). Furthermore, it is not uncommon for other exotic white-

flies to show similar increases in host acceptance during the phase of unrestrained

population growth that initially follows their introduction into a new environment

(RVD, unpublished data). Models currently under development treat expansion

of host plant acceptance in species invading new environments, with particular

reference to agricultural pests (RVD, unpublished data).

Niche Competition.—In addition to lacking natural enemies, S. phillyreae is

almost devoid of competitors in California (especially rosaceous plants). Species

of Citrus are the only hosts of S. phillyreae, either previously reported or new

(Table 1), that are attacked regularly by a complement of whiteflies in California

(RJG, unpublished data). However, S. phillyreae uses Citrus primarily as an

overwintering host in California. Interestingly, Citrus are the chief evergreen hosts

of S. phillyreae in California with a substantial but facultative fauna of polypha-

gous aphids (Kono & Papp 1977).

Because many genera of aphids in a “Rosaceous series” within the Aphidinae

have evolved using plants in the Rosaceae as a primary (overwintering) host (Hille

Ris Lambers 1950, 1979; Blackman & Eastop 1984; Dixon 1987), aphids are

perhaps the most likely Homoptera to seriously compete with S. phillyreae for

the niche of feeding on deciduous leaves in the Rosaceae. Several aphid groups

have radiated using the Rosaceae in this way (e.g., the Rhopalosiphini: Hyalop-

terous, Hysteroneura, Melanaphis, Rhopalosiphum, Schizaphis\ and the Macro-

siphini: Anuraphis through Macrosiphum; among others). These holocyclic aphids

have evolved using a life cycle employing woody plants in the Rosaceae as over¬

wintering hosts, upon which to deposit their eggs in the fall. They move from the

Rosaceae after completing a few generations in the early spring, having taken

advantage of the high amino-nitrogen content of the rapidly growing leaves during

spring foliation (Dixon 1970, 1973, 1977). When the amino-nitrogen content of

c Host added while in press to reflect an update after Bellows et al. (1990) and subsequent data.

Identification not (yet) verified by CDFA.

d Reported as Amelanchier “'dentiolata” for which we can find no species reference in taxonomic

works.

e Reported as M. domestica, which is crabapple, but assumed here to be common apple due to the

common name association of “apple” on the record (Bellows et al. 1990 lists this record as M.

domestica ).

f Also recorded from navel oranges but without reference to a particular cultivated variety.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

the maturing leaves on these woody rosaceous hosts drop, the aphids customarily

emigrate to rapidly growing herbaceous plants during the summer, again because

of the higher available amino-nitrogen during their summer growth season. In

the fall, as the herbaceous plants decline, the summer populations of these hol-

ocyclic aphids immigrate back to their woody Rosaceae overwintering hosts, where

the amino-nitrogen content of the senescent leaves increases as leaf proteins are

broken down. An advantage of this heteroecious life mode for holocyclic aphids

is that favorable amino-nitrogen sources can be tracked throughout the year, and

that foliating woody hosts in the early spring provide a dependable nutrient source

when the aphid’s eggs hatch into poorly mobile first instar nymphs that require

immediate and adjacent high-quality food.

An ecological strategy of heteroecy and holocycly for aphidine aphids has many

exceptions, however. In warmer areas, such as California, many species of these

otherwise heteroecious aphid genera have adapted an anholocyclic strategy and

have become secondarily monoecious by remaining continuously parthenogenetic

on the ever-present growth of herbaceous plants. Many aphids considered serious

agricultural pests (e.g., Myzus persicae [Sulzer], Aphis gossypii Glover) are poly-

phagous and anholocyclic, taking advantage of whatever happens to be the best

amino-nitrogen source at the moment. Other entire aphid genera in rosaceous

evolutionary series have similarly adopted a secondarily monoecious strategy (e.g.,

Sitobion) and have moved permanently to herbaceous plants in noncontinental

climate areas. In contrast, some holocyclic species of Aphidinae (e.g., Aphis pomi

DeGeer) remain on their rosaceous hosts continuously. The abandoning of mid¬

summer teneral leaves with lower amino-nitrogen contents is not trivial for aphids;

some more primitive aphid groups that are monoeciously restricted to trees (e.g.,

drepanosiphines and some phyllaphidines) must cope with lower available nu¬

trients during the summer by ceasing reproduction and estivating as nymphs until

amino-nitrogen raises in the fall (Hille Ris Lambers 1966).

Nonaphid homopterans do not have such a strong and definite evolutionary

link with the Rosaceae, and either do not regularly feed on the deciduous leaves

of plants in the Rosaceae (e.g., Coccoidea, Aleyrodoidea), or appear to occupy

that niche as a relatively modem and derived “host capture” (e.g., several ty-

phlocybine leafhopper genera, Cacopsylla [Psyllidae] on Pyrus, Parthenolecanium

corni [Bouche] [Coccidae] nymphs on leaves during the summer).

Among S. phillyreae’s favored nonrosaceous and deciduous hosts, aphids again

appear as potential competitors. For example, Fraxinus is commonly used by

Prociphilus americanus (Walker) and P. fraxinifolii (Riley) as an overwintering

host (Kono & Papp 1977); these aphids, however, are also holocyclic, using co¬

nifers (Smith 1969) as secondary hosts during the summer.

In contrast to aphids, A. phillyreae prefers mature (teneral) foliage and is present

during the hot summer months on its deciduous hosts; this largely eliminates or

minimizes the synchronic use of these plants by this whitefly and aphids. Thus,

S. phillyreae appears to occupy a reasonably distinct and somewhat unoccupied

or less competitive niche that involves the summer use of teneral leaves among

plants in the Rosaceae and Fraxinus. With no natural enemies or serious com¬

petitors, the population of S. phillyreae has rapidly increased in California and

the whitefly has expanded its host plant range onto previously unused plants.

Economic Potential as a Pest. — Lacking competitors S. phillyreae will become

1990 SORENSEN ET AL.: SIPHONINUS PHILLYREAE IN NORTH AMERICA 53

a major summer feeding pest of stone and pome fruits and numerous ornamental

trees in California and elsewhere in North America. As such it could seriously

disrupt existing management programs on these plants as has occurred with other

whiteflies on Citrus (Dowell in press). In addition, management sprays for other

pests such as codling moth, Cydia pomonella (L.) and oriental fruit moth, Gra-

pholitha molesta (Busck) could interfere with biological control efforts (Dowell in

press) aimed at S. phillyreae.

The apparent requirement for an overwintering host in California will limit the

impact of S. phillyreae to those deciduous hosts near stands of overwintering

hosts. Such situations for S. phillyreae commonly exist throughout California in

urban and agricultural settings with Citrus, Rhaphiolepis and Heteromeles ; par¬

ticularly the latter in foothill areas near orchards. A similar situation exists for

Trialeurodes vittata (Quaintance) which is a pest in grape vineyards near stands

of its overwintering host, Rhamnus californica Eschscholtz (Joos 1981).

The extent of genetic variance in Californian S. phillyreae and its ultimate

expression and consequences remain unknown. We suggest that studies of iso-

zymic variation in the introduced Californian population, as it expands, in com¬

parison with similar studies of variation for the Palaearctic may be of significant

importance to both pragmatic economic estimates for the species as well as to an

increased knowledge of evolutionary parameters and mechanisms in general. We

suspect that eventually when successful natural enemies are introduced to control

S. phillyreae populations in California, the variety of host plants which it occurs

on will collapse to only those it does best on physiologically, ecologically and

competitively. We also suspect, however, that this species will be a significant

aleyrodid pest to some deciduous plants throughout its eventual Nearctic range.

Acknowledgment

We thank D. Barbe for botanical consultations, L. Bezark, T. Bellows and K.

Arakawa for information on S. phillyreae, and T. Kono for reviewing the manu¬

script. Many of the hosts listed as new within California were reported to the

California Dept, of Food & Agriculture by K. Arakawa, which verified the S .

phillyreae identification.

Literature Cited

Andrewartha, H. G. & L. C. Birch. 1954. The distribution and abundance of animals. The University

of Chicago Press, Chicago, Illinois.

Bellows, T., T. D. Paine, K. Y. Arakawa, C. Meisenbacher, P. Leddy & J. Kabashima. 1990. Biological

control sought for ash whitefly. California Agriculture, 44 (1, January-February): 4-6.

Berlocher, S. H. 1984. Genetic changes coinciding with colonization of California by the walnut

husk fly, Rhagoletis completa. Evolution, 38: 906-918.

Blackman, R. L. & V. F. Eastop. 1984. Aphids on the world’s crops, an identification guide. John

Wiley & Sons, New York.

Bush, G. L. 1969. Sympatric host race formation and speciation in frugivorus flies of the genus

Rhagoletis (Diptera, Tephretidae). Evolution, 23: 237-251.

Bush, G. L. 1975. Modes of animal speciation. Annual Rev. Ecol. & Syst., 6: 339-364.

Dixon, A. G. F. 1970. Quality and availability of food for sycamore aphid population, pp. 271-

287. In Watson, A. (ed.). Animal populations in relation to their food resources. Blackwell,

Oxford.

Dixon, A. G. F. 1973. Biology of aphids. Edward Arnold, London.

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Dixon, A. G. F. 1977. Aphid ecology: life cycles, polymorphism, and population regulation. Annual

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Dixon, A. G. F. 1987. Chapter 4. Biology. 4.1. The way of life of aphids: host specificity, speciation

and distribution, pp. 197-207. In Minks, A. K., & P. Harrewijn (eds.). Aphids, their biology,

natural enemies and control. Vol. 2A. World Crop Pests. Elsevier, Amsterdam.

Dobzhansky, T. 1970. Genetics of the evolutionary process. Columbia University Press, New York.

Dowell, R. V. (in press). Chapter 13. Integrating biological control of whiteflies into existing pest

management systems. In Gerling, D. (ed.). Whiteflies: their bionomics, pest status and man¬

agement. Intercept, Dorset, England.

Dowell, R. V. & R. J. Gill. 1989. Exotic invertebrates and their effects on California. Pan-Pacif.

Entomol., 65: 132-145.

Emlen, J. M. 1973. Ecology: an evolutionary approach. Addison-Wesley, London.

Ford, E. B. 1971. Ecological genetics (3rd ed). Chapman & Hall, London.

Gill, R. J. 1989. The ash whitefly in California, history, economic potential, biology, current status.

Analysis & Identification Branch, California Dept. Food & Agriculture, Sacramento, California.

Gill, R. J. (in press). Chapter 2. The morphology of whiteflies. In Gerling, D. (ed.). Whiteflies: their

bionomics, pest status and management. Intercept, Dorset, England.

Goux, L. 1949. Contribution a l’etude des aleurodes (Hem. Aleyrodoidea) de la France. VII. Le

genre Siphoninus Silvestri. Bull. mens. Soc. linn. Lyon N. S., 18: 7-12.

Hille Ris Lambers, D. 1950. Host plants and aphid classification, pp. 141-148. Proc. 8th Intemat’l.

Cong. Entomol., Stockholm, 1948.

Hille Ris Lambers, D. 1966. Polymorphism in Aphididae. Annual Rev. Entomol., 11: 47-78.

Hille Ris Lambers, D. 1979. Aphids as botanists? Symbolae Botanicae Upsaliensis, 22: 114-119.

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Division of Agricultural Sciences, University of California, Publication 4105. Agricultural Sci¬

ences Publications, University of California, Berkeley.

Khan, A. G., A. A. Goraya, A. I. Mohyuddin & C. Inayatullah. 1985. The whiteflies (Homoptera:

Aleyrodidae) of Pakistan. Pakistan J. Zool., 17: 29-33.

Kono, T. & C. S. Papp. 1977. Handbook of agricultural pests—aphids, thrips, mites, snails and

slugs. California Dept. Food & Agriculture, Sacramento.

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Mettler, L. E. & T. G. Gregg. 1969. Population genetics and evolution. Foundations of modem

genetics series. Prentice-Hall, Englewood Cliffs, New Jersey.

Mound, L. A. & S. H. Halsey. 1978. Whitefly of the world, a systematic catalogue of the Aleyrodidae

(Homoptera) with host plant and natural enemy data. British Museum (Natural History) and

John Wiley & Sons, New York.

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Aphididae). Ann. Entomol. Soc. Am., 62: 1128-1152.

Priesner, H. & M. Hosney. 1932. Contributions to a knowledge of the white flies (Aleurodidae) of

Egypt (I.). Ministry of Agric., Egypt, Tech, and Sci. Serv., Bull., 121.

Walker, G. P. & D. C. G. Aitken. 1985. Oviposition and survival of bayberry whitefly, Parabemisia

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14: 254-257.

Received 20 November 1989; accepted 4 December 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 55-61, (1990)

PODAGRITUS CORA (CAMERON) AND

P. ALBIPES (F. SMITH)

(HYMENOPTERA: SPHECIDAE: CRABRONINAE)

PREYING ON EPHEMEROPTERA AND TRICHOPTERA

Anthony C. Harris

Otago Museum, Dunedin, New Zealand

Abstract.— Podagritus albipes (F. Smith) and P. cora (Cameron) are recorded preying almost

exclusively on a Deleatidium sp. taxonomically close to D. myzobranchia Phillips, a Deleatidium

sp. close to D. lillii Eaton (Ephemeroptera) and Pycnocentrodes aureola (McLachlan) (Trichop-

tera). Multitudes of individuals of these carbronines, together with predatory muscids ( Spilogona

Schnabl), occupy large stones in the Blue Stream (Mt. Cook National Park, New Zealand) that

protrude above waterlevel. When Ephemeroptera nymphs eclose as subimagines and emerge

from the water onto the stones, they are often captured either by Podagritus wasps or by Spilogona

muscids. The wasps take the prey to nests in patches of sand on either side of the stream. Forty

(of 41) excavated Podagritus nests contained only Ephemeroptera and Trichoptera. This asso¬

ciation reoccurs every summer. The Podagritus populations generally do not feed upon the

Diptera, prey usually associated with this genus.

Key Words.— Insecta, Sphecidae, Podagritus cora, Podagritus albipes, Diptera, Spilogona,

Ephemeroptera, Deleatidium, Trichoptera, Pycnocentrodes aureola

Podagritus Spinola comprises 53 described species distributed in South America

(28), Australia (19) and New Zealand (6); New Zealand has an additional two

undescribed species. Species of this genus provision their nest cells exclusively

with Diptera (Bohart & Menke 1976). The present observation is unusual because

it shows that nesting aggregates along the entire course of a stream prey almost

entirely on aquatic insects, mostly Ephemeroptera and Trichoptera. The vast

majority of provisioned nest cells in this locality were completely devoid of Dip¬

tera.

A brook, the Blue Stream, originates from Tasman Glacier, Mt. Cook National

Park, New Zealand, emerging from the southern side approximately 8500 m

upstream of the terminal face of the glacier. Blue Stream runs for approximately

2 km along the southern wall of the Tasman Valley before joining the Tasman

River. The stream flows over outwash plains of boulders, gravel, shingle and sand;

large stones line its course and many boulders protrude above the surface of the

water.

Observations

There are large numbers of Ephemeroptera (principally Deleatidium myzo-

branchia- group, D. lillii- group) and Trichoptera (Pycnocentrodes aureola [Mc¬

Lachlan]), besides lesser numbers of miscellaneous immature aquatic insects on

the undersurfaces of the stones. During November-February between 11:00-

16:00 h, ultimate instar mayfly nymphs crawl rapidly from the undersides of

boulders, and emerge from the water. They stop (about 2 min after first appearing

underwater on the stone) and eclose to a subimago, that is free in 2-8 min. Teneral

caddisflies also emerge along the sides of the stones, pausing above the water

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

level. Large numbers of crabronine wasps, mostly Podagritus corn (Cameron) and

P. albipes (F. Smith), together with an undescribed muscid fly {Spilogona Schnabl,

formerly Limnophora Robineau-Desvoidy) are also on the stones. In January

there are up to 24 crabronines and five muscids per boulder, but more usually

there are two to six crabronines and two muscids per stone. On warm sunny days

between mid October and February, every available stone in the stream may be

occupied thusly, and the flies and wasps wait for mayflies and caddisflies. The

predators frequently observe the submerged prey and move closer to their emer¬

gence points. During the peak emergence of mayflies (often 13:00-15:00 h) cra¬

bronines fly from the tops of the boulders to the eclosing mayfly nymphs (Fig. 1)

and, as the integument splits, pull the subimago out, sting and paralyze it, and

fly rapidly about 15 m to sand patches on the sides of the stream where they nest.

Most of the nest-cells throughout a 1.5 km area on both sides of the Blue Stream

are provisioned solely with mayflies, although about 10% of nests contain a com¬

bination of mayflies and caddisflies.

The muscids are capable of defending their prey against relatively large cra¬

bronines such as P. corn with which they compete for mayflies (Fig. 2), despite

that in other parts of the country this wasp frequently provisions its nests with

large sarcophagids and calliphorids. Examination of 40 crabronid nests at the

Blue Stream indicated that Spilogona were not used as prey; no sphecids were

observed to take Spilogona.

On overcast days when the crabronines remained underground in their burrows

the muscids were present on the boulders; the flies were also present on the

boulders for longer periods during sunny weather, arriving earlier and leaving

later than the wasps. They frequently captured and consumed aquatic insects on

the boulders.

The crabronines were best able to capture emerging mayflies on still, warm,

sunny days. When a small wind developed, the wasp and its prey were frequently

blown into the stream and swept away with the current. Mayfly subimagines

frequently fell into the stream by themselves and drifted away; some of these,

however, were plucked from the surface of the water by crabronines, stung in

mid-air, and transported to the nests.

Both P. com and P. albipes would attempt to take struggling insects held by the

surface tension of the water, but neither would accept as prey anything other than

Deleatidium subimagines or Pycnocentrodes aureola. On six occasions, P. albipes

females flew to a struggling Syrphus novozealandiae (Diptera: Syrphidae), a pro¬

visioned prey in its nests elsewhere, but these flies were ignored once identified

by the wasps at Blue Stream.

The association between Podagritus, Spilogona, Deleatidium and Pycnocen¬

trodes species at Blue Stream has occurred every summer since at least 1975,

beginning in November and ending at the end of February.

Although the Deleatidium ultimate instar nymphs taken by Podagritus wasps

at Blue Stream key to the myzobranchia- group and the //////-group using Winter¬

bourne et al. (1981), and the subimagines key to D. myzobranchia Phillips and

D. lillii Eaton using Phillips (1930), the two species taken from crabronine wasps

are both undescribed (M. Winterbourne, personal communication).

Nests.—Podagritus albipes nests were made in level sand. The 3.0 mm wide

entrance typically opened in the middle of a 40 mm diameter mound and was

left open during provisioning. The 3 mm wide main burrow sloped at 75°-77°

1990

HARRIS: SPHECID PREDATION ON AQUATIC INSECTS

Figures 1, 2. Figure 1. Three Podagritus cora females competing for a Deleatidium sp. subimago

(undescribed), that is emerging from its nymphal exuviae. One P. cora has grasped the subimago,

pulling it out. Other mayfly exuviae are visible. Figure 2. A Spilogona sp. (undescribed) has beaten a

Podagritus cora to a Deleatidium sp. subimago (undescribed) that started emerging from its last

nymphal exuviae.

and the first cell was 10-68 mm (typically 60 mm) below surface level. Nests

contained either one or two cells. The cells contained four to six mayflies ( De¬

leatidium myzobranchia-growp, D. //////-group), and less frequently a combination

of mayflies and caddisflies (Pycnocentrodes aureola). Of 15 nests excavated, 11

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Figure 3. Typical nest of Podagritus cor a at Blue Stream, a. Mound, b. Entrance, c. Main burrow,

d. Inner closure, e. Provisioned cell. f. Partly provisioned cell. g. Spur.

were one-celled and four were two-celled. Total prey comprised Deleatidium

myzobranchia-group and D. lillii- group (n = 89) and Pycnocentrodes aureola {n

= 22). All prey were positioned supine (venter up) facing the ends of the cells.

After an egg was laid on the prey, the cell-burrow, but not the main burrow was

closed.

Podagritus cor a nests were made in level sand. The entrance was a 3.5-3.9 mm

wide round hole in the center of a 30-50 mm wide mound. Nests were sometimes

dug beside stones. The main burrow was 3.5-3.9 mm wide and the first cell was

65-80 mm below the surface. The cells were on average 4.6 mm wide (greatest

diameter) and 9.2 mm long. Fifteen nests were excavated, and of these nine were

single-celled and six had two cells. Total prey comprised 139 Deleatidium my-

1990

HARRIS: SPHECID PREDATION ON AQUATIC INSECTS

Figures 4, 5. Figure 4. Position of egg of Podagritus cor a on Calliphora quadrimaculata. Figure 5.

Position of egg of P. albipes on Deleatidium sp. subimago (Blue Stream). When preying on Diptera

in other areas, eggs are positioned in the neck region, as in Fig. 4. (Both figures are tracings from

photographs.)

zobranchia-growp and D. lillii- group and 23 Pycnocentrodes aureola. The cells

contained seven to eight mayflies or eight mayflies and one caddisfly. Eggs laid

on mayflies were usually placed on the third and fourth abdominal sterna (Fig.

5). These same species place eggs on the neck region when utilizing Diptera as

hosts. All prey were placed in the cells supine with the head facing the end of the

cell. Main burrows were left open during provisioning.

Both P. albipes and P. cora held the mayfly upside-down, grasped by the meso-

thoracic legs, with the wings of the caddisfly projecting. The wasp would descend

slowly towards the nest and fly directly in without alighting on the sand.

On 24 Jan 1989 a solitary P. cora (initally assumed to be P. swalei until captured)

spent much time at the entrance to a rabbit burrow, preying on large flies. Its nest

was made in the sand, beside the burrows of its peers, and was single-celled. The

nest contained two Calliphora quadrimaculata (Swederus) and one undetermined

sarcophagid of similar size. The egg was placed between the head and thorax of

the fly.

The nests of Podagritus species were invaded by larvae of an undescribed

Anabarynchus (Diptera: Therevidae) which burrowed through the sand and de¬

voured the provisioned food as well as the developing immature wasps. The adult

stage of this therevid had a slate-grey body and white, black-veined wings; it

frequents the sandy patches during summer.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Discussion

Throughout its range in the Southern Hemisphere, Podagritus has been recorded

as preying only on Diptera (Bohart & Menke 1976). Solitary wasps capture prey

in habitats different than and often distant from their parental nest or where they

eventually build their own nests (Evans 1966, Evans & Eberhard 1970). Therefore,

the wasp must undertake initial hunting flights involving random flight before it

locates a source of appropriate prey. Sphecid wasps apparently learn certain sources

of prey, and return to them until the source is exhausted; they do not normally

make mistakes with respect to prey.

New Zealand Podagritus prey mostly on Diptera, but frequently take other

insects occasionally. Podagritus albipes and P. cora stock their nest-cells with a

combination of Diptera and Ephemeroptera from streams in the Tararua Range,

the South Island West Coast, and elsewhere (unpublished data), and P. albipes

uses predominantly Diptera but also Ephemeroptera at the Hutt River (D. Willis,

personal communication). Podagritusparrotti frequently stocks its nests with small

beetles ( Cyphon spp. [Scirtidae], Asilis tumidus Broun [Cantharidae], Luperus

vulgaris Broun [Chrysomelidae]) captured from flowering shrubs (unpublished

data; J. Charles, personal communication). Notwithstanding this evidence, such

instances usually appear almost as exceptions to the rule that Podagritus in New

Zealand are specialists on Diptera.

Despite such frequent break-downs in prey specificity among New Zealand

Podagritus, the situation at the Blue Stream is unusual. Throughout almost the

entire length of the Blue Stream, P. albipes and P. cora prey almost exclusively

on mayflies and, to a lesser extent, caddisflies and appear to reject seemingly more

appropriate prey. On six occasions, after careful examination, P. cora and P.

albipes females rejected syrphid flies (their usual prey). The association with

mayflies and caddisflies probably represents a population in evolutionary tran¬

sition, as noted in Australia involving the widespread genus Bembix Fabr. (Evans

& Matthews 1973). Bembix is best represented in North America, where Evans

& Matthews (1973) suggest that Bembix evolved, and later spread to other regions.

Evans accumulated thousands of prey records for Bembix in North America,

without finding a single record outside Diptera (Evans & Matthews 1973). Bembix

is also well represented in Australia, and Evans & Matthews (1973) found 14

species that preyed exclusively on Diptera, one which preyed on both flies and

damselflies (Odonata: Anisoptera), one that preyed on both flies and wasps, and

six that were each host specific for Hymenoptera, Odonata and Neuroptera. They

regarded the former as “species in transition,” and assumed that the six Australian

Bembix species that prey on single orders of insects exclusive of Diptera passed

through similar transitional stages. Wheeler & Dow (1933) suggested that if the

(then unidentified) Bembix that Wheeler observed capturing damselflies in Aus¬

tralia was taking its normal prey, it represented a “comparatively recent devel¬

opment in evolution.”

The situation at Blue Stream seems similar and may indicate how sphecids

which characteristically exhibit host specificity at the ordinal level change their

host preference to entirely new orders. Alternatively, the New Zealand Podagritus

might not be specialist hunters of Diptera, but rather prey generally on Diptera,

taking other orders if a good source of prey is located.

1990

HARRIS: SPHECID PREDATION ON AQUATIC INSECTS

Acknowledgment

I am grateful to J. Ward, J. Leader and M. Winterbourne for confirming my

identifications of Trichoptera and Ephemeroptera, to R. Harrison and L. Lyneborg

for their identifications of Diptera, and C. Lalas for the photographs.

Literature Cited

Bohart, R. M. & A. D. Menke. 1976. Sphecid wasps of the world: a generic revision. University of

California Press, Berkeley.

Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps. Harvard University

Press, Cambridge, Massachusetts.

Evans, H. E. & M. J. W. Eberhard. 1970. The wasps. University of Michigan Press, Ann Arbor.

Evans, H. E. & R. W. Matthews. 1973. Systematics and nesting behavior of Australian Bembix sand

wasps (Hymenoptera: Sphecidae). Mem. Am. Entomol. Inst. 20.

Phillips, J. S. 1930. A revision of New Zealand Ephemeroptera. Trans. New Zealand Inst., 61: 271—

390.

Wheeler, W. M. & R. Dow. 1933. Unusual prey of Bembix. Psyche, 40: 57-59.

Winterbourne, M. J., K. L. D. Gregson & D. R. Townes. 1981. Ephemeroptera (mayflies). Key to

larvae. In Winterbourne, M. J. & K. L. D. Gregson (eds.). Guide to the aquatic insects of New

Zealand. Bull. Entomol. Soc. New Zealand, 5: 14-18.

Received 7 June 1989; accepted 17 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 62-65, (1990)

INFLUENCE OF HOST PLANT ON FECUNDITY OF

ALEUROCANTHUS WOGLUMI ASHBY

(HOMOPTERA: ALEYRODIDAE)

Robert V. Dowell 1 and Bryan Steinberg 2

4681 Pebblewood Drive, Sacramento, California 95833

2 University of Florida, Research and Education Center,

Ft. Lauderdale, Florida 33314

Abstract. — The ovaries of citrus blackfly females are nonfunctional at eclosion but become

functional within 24-36 h thereafter. Egg development is synchronous and monotene, with all

available eggs deposited in each egg spiral. Maximum number of eggs laid was 118. There were

significant differences in the fecundity of females reared on different host plants with the greatest

from Citrus spp. and the least from Eriobotya japonica Lindley. Variation in fecundity is due

to host plant and not to differences in female survival or the percentage of females laying eggs.

Fecundity is significantly correlated with nymphal survival among the host plants indicating that

both are influenced by the same host plant factors.

Key Words.— Insecta, Aleyrodidae, Aleurocanthus woglumi, fecundity, hostplant, Citrus

Citrus blackfly (CBF), Aleurocanthus woglumi Ashby, is an aleyrodid of Asian

origin that is currently found in Florida, Texas and Mexico. Female CBF deposit

eggs in spirals on the underside of leaves, most notably on Citrus spp. (Dietz &

Zetek 1920). The host plant influences the number of eggs deposited on it and the

subsequent survival of nymphs. Oviposition is influenced by the levels of foliar

nitrogen, sugars, cysteine and methonine (RVD, unpublished data) but nymphal

survival is affected by the density of living tissue and the presence of allelochem-

icals (Dowell 1989, Dowell & Steinberg in press).

This study determined whether host plants of CBF nymphs influenced their

subsequent fecundity, and how ovary and oocyte maturation patterns influence

egg deposition.

Materials and Methods

Females were taken from potted citrus trees 4 h (n = 12) or 24-36 h (n = 6)

after eclosion and dissected to determine the condition of their ovaries. Females

reared on Citrus spp. were dissected after depositing an egg spiral to determine

whether any eggs were left in the ovaries or oviducts.

Six species of plants (Table 2) were exposed to CBF infested citrus trees in

October, 1980. The plants then were held in a screenroom until adult CBF began

to emerge. After emergence, single females were removed from the plant upon

which they developed and placed in a 30 dram vial with a Citrus x paradisi

Macfadyen leaf and one or two males. The vials were held at 24-27° C under

artificial light (16 h photophase) until the female died; she was dissected and the

number of eggs was noted. The leaf was then removed and the egg spirals and

eggs were counted. The leaf petiole was held in water until egg hatch to determine

fertility.

Longevity and fecundity data were transformed to log 10 (x + 1) prior to Analysis

of Variance. Transformed means were separated using Duncan’s Multiple Range

1990 DOWELL & STEINBERG: FECUNDITY OF CITRUS BLACKFLY

Table 1. Relationship between number of egg spirals and average fecundity and age of females at

death.

Number of spirals

Average number eggs per female (nf

Average age of female

at death (days) b

Average increase in

number eggs per female

2.5 ± 1.0 [a]

—

37.7 ± 9.7 (37) [a]

3.3 ± 1.0 [a]

37.7

55.8 ± 8.5 (11) [b]

7.4 ± 2.1 [b]

18.1

67.0 ± 24.1 (4) [c]

9.0 ± 1.0 [c]

11.2

4+ c

94.5 ± 33.2 (2) [d]

9.0 ± 0 [c]

8.8

a Mean ± SD, with means followed by different letters [a] differing at P < 0.05. ANOVA values F

= 15.99; df = 3, 51; P < 0.01.

b As a: ANOVA values F = 26.42; df = 3, 67; P < 0.01.

c Includes females laying 4 and 5 egg spirals.

Test (Little & Hills 1978). The relationship between host plant specific fecundity

and 34 chemical and physical attributes of the host plants (RVD, unpublished

data) or nymphal survival (Dowell & Steinberg in press) was determined using

regression analysis (Little & Hills 1978).

Results and Discussion

The ovaries of CBF are not functional at eclosion but elongate within 24-36 h

so that oocytes are visible. Oocyte development is synchronous and generally

monotene with a single oocyte developing in each ovariole. Each egg spiral consists

of all available mature eggs; no eggs were found in the ovaries or oviducts of

females that had just deposited an egg spiral. Both the number of eggs laid per

female and the age of the females at death significantly increase with the number

of egg spirals laid. In contrast, the average increase in the number of eggs laid per

female decreases with age (Table 1) indicating that fewer ovarioles produce eggs

as the female ages.

The greatest number of eggs laid was 118 by a female reared on Citrus spp.

(Table 2). The greatest longevity was 17 days. The average number of eggs per

spiral from Citrus spp. females was 31.5 (Table 2). These data agree closely with

previous studies where females laid 20-40 eggs per spiral and egg laying began

1-4 days after eclosion (Dietz & Zetek 1920). The data confirm the conjecture of

Dietz & Zetek (1920) that females can lay over 100 eggs.

There were significant differences in the fecundity of females from the six test

plants; the highest fecundity was on Citrus spp. and the lowest on Eriobotyra

japonica Lindley. There were significant differences in the average number of eggs

per spiral among the test plants; females bred on Ardisia solanacea Roxburg and

Citrus spp. laid the largest egg spirals and those from Eugenia uniflora L. the

smallest. The maximum number of eggs from a female and the percentage of

females laying eggs followed the trend of fecundity; the greatest values were for

Citrus spp. and the lowest for E. japonica. There were no significant differences

in female longevity (F = 1.80, df = 5, P > 0.05) or in percentage egg hatch (x 2

= 4.77, df = 5, P > 0.05) among the test plants (Table 2).

If fecundity was calculated using only those females that laid eggs significant

differences were evident among the plants with the highest values on A. solanacea

(50 ± 22) and Citrus spp. (48.4 ± 19) followed by Mangifera indica L. (39.6 ±

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Table 2. Host plant specific fecundity.

Plant

Average number eggs

per female (n)‘

Average number eggs

per spiral (n) b

Maximum

number eggs

Percentage

females laying

Citrus spp.

39.6 ± 25 (44) [a]

31.5 ± 13(53) [ab]

118

Ardisia solanacea

28.6 ±31 (7) [b]

33.3 ± 11 (6) [a]

Mangifera indica

17.4 ± 22 (25) [c]

27.3 ±7 (16) [b]

Gardenia thunbergi c

14.1 ± 20(13) [d]

26.1 ± 7 (7) [b]

Eugenia uniflora

10.2 ± 25 (6) [e]

15.0 ± 2 (4) [c]

Eriobotrya japonica

8.3 ±11 (12) [f]

19.8 ± 7 (5) [d]

a Mean ± SD, with means followed by different letters [a] differing at P < 0.05. ANOVA values F

= 5.69; df = 5, 102; P < 0.01.

b As a: ANOYA values F = 3.44; df = 5, 86; P < 0.01.

c Nymphal survival assumed to be 1.4% for regression analysis.

14), Gardenia thunbergi L. (36.6 ± 12) and E. japonica (19.8 ± 7) (F = 6.96; df

= 4, 57; P < 0.01). The average number of eggs in the ovaries of females that

died before ovipositing did not differ from the average number of eggs laid by

females ovipositing on Citrus spp. (38.0 ± 5.4 versus 48.4, t = 1.24, df = 28, P

> 0.05), on E. japonica (18.3 ± 5.6 versus 19.8, t = 0.61, df = 7, P > 0.05) or

on M. indica (31.8 ± 5.4 versus 39.6, t = 1.62, df = 19, P > 0.05). Thus, the

differences seen in Table 2 are due to the host plant and not to differences in

female longevity or the percentage of females laying eggs. Host plant mediated

differences in whitefly fecundity have been reported for Aleyrodes proletella (L.)

on crucifers (Ihaegwam 1980) and Trialeurodes vaporariorum Westwood on an

array of glasshouse vegetables (van Boxtel et al. 1978, van Sas et al. 1978).

There were no significant correlations between fecundity and 34 chemical or

physical attributes (RVD, unpublished data) of the test plants either singly or in

combination (r < 0.6, F < 2.4, P > 0.1). There was a significant correlation

between fecundity and the survival of CBF nymphs on plants (r=0.95;F=35.6;

df = 1, 4; P = 0.004) indicating that the same factors that influence survival

(believed to be allelochemicals: Dowell 1989, Dowell & Steinberg in press) are

affecting the fecundity of the resulting females.

Reduced survival of immature CBF on noncitrus hosts (Dowell & Steinberg in

press), combined with reduced fecundity of the resulting females, are primary

forces inhibiting the utilization of such hosts by CBF.

Literature Cited

Dietz, H. F. & J. Zetek. 1920. The blackfly of citrus and other subtropical plants. U.S. Dept.

Agriculture Bull., 885.

Dowell, R. V. 1989. Toxicity of water extracts of Murray a paniculata Jack leaves to immature citrus

blackfly, Aleurocanthus woglumi Ashby (Homoptera: Aleyrodidae). Pan-Pacif. Entomol., 65:

163-165.

Dowell, R. V. & B. Steinberg, (in press). Influence of host plant characteristics and nitrogen fertil¬

ization on development and survival of immature citrus blackfly (Hompt.: Aleyrodidae). J.

Appl. Entomol.

Ihaegwam, E. U. 1980. Influence of host plant ( Brassica species) and temperature on population

increase of the cabbage whitefly Aleyrodes brassicae. Ann. Appl. Biol., 95: 273-278.

Little, T. M. & F. J. Hills. 1978. Agricultural experimentation. John Wiley & Sons, New York,

van Boxtel, W., J. Woets & J. C. van Lenteren. 1978. Determination of host-plant quality of eggplant

{Solarium melongena L.), cucumber ( Cucumis sativus L.), tomato {Lycopersicum esculentum

1990 DOWELL & STEINBERG: FECUNDITY OF CITRUS BLACKFLY

L.) and paprika ( Capsicum annuum L.) for the greenhouse whitefly ( Trialeurodes vaporariorum

(Westwood)) (Homoptera: Aleyrodidae). Med. Fac. Landbouww. Rijksuniv. Gent, 43: 397-

408.

van Sas, J., J. Woets & J. C. van Lenteren. 1978. Determination of host-plant quality of gherkin

(Cucumis sativis L.), melon ( Cucumis melo L.) and gerbera ( Gerbera jamesonii Hook) for the

greenhouse whitefly, Trialeurodes varorariorum (Westwood) (Homoptera: Aleyrodidae). Med.

Fac. Landbouww. Rijksuniv. Gent, 43: 409^120.

Received 14 August 1989; accepted 7 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 66-70, (1990)

BIOLOGICAL OBSERVATIONS OF GLASSY CUTWORM

(LEPIDOPTERA: NOCTUIDAE) IN WESTERN OREGON

J. A. Kamm

National Forage Seed Production Research Center,

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

Corvallis, Oregon 97331-7102

Abstract. — The glassy cutworm, Crymodes devastator Brace, is a sporadic subterranean pest of

grasses grown for seed in the Pacific Northwest. Light trap captures indicate that adult flight

occurs from July through September, but peaks in late July. Postharvest field burning eliminates

all foliage and crop residue, probably making fields temporarily unsuitable for oviposition. I

suspect that females fly from burned fields to spring-seeded, green fields to oviposit. Bentgrass,

ryegrass and wild oats, as weeds in fine fescue fields, attract ovipositing females. High densities

of feeding larvae damage or kill the crop of fine fescue near weeds and become evident in late

autumn. Larvae 1.5-2 cm long are present in fields during winter and readily initiate feeding

after exposure to 21° C for 2 to 7 days in the laboratory. Nowickia latianulum (Tothill) and

Lissonota montana (Cresson) collectively parasitized 30% and 48% of larvae collected from two

field sites.

Key Words.— Insecta, Crymodes devastator, pest management, grass seed, insect parasites

The glassy cutworm, Crymodes devastator Brace, is a sporadic subterranean

pest that feeds primarily on the roots and crowns of grass but also attacks many

other hosts (Crumb 1929). Numerous reports document field infestations through¬

out the U.S. and Canada, but only scant biological information is available (Gil¬

lette 1891, Slingerland 1902, Rings & Arnold 1974). Various natural enemies

attack the larvae, but their impact on the larval population has never been quan¬

tified (Coquillett 1897, Gibson 1915, Gillette 1891, Strickland 1921). No infor¬

mation is available on the seasonal appearance, development, and activity of

larvae in fields of grass grown for seed in the Pacific Northwest.

This study determines: the seasonal development of larvae in relation to adult

flight, the feeding behavior of larvae, the degree of larval parasitization, and the

onset of diapause. Seasonal flight of adults was monitored, and oviposition by

females was related to cultural practices of grass seed production.

Materials and Methods

The relative seasonal abundance of adults was determined with a light trap

operated near Donald, Oregon. Eggs were obtained by placing adults in gallon

jars containing corrugated wax paper to provide cover for the moths and also an

oviposition substrate. Newly eclosed larvae were reared individually in stender

dishes (5 x 2.5 cm) whose bottoms were lined with damp blotting paper. Larvae

were fed freshly cut leaves of Chewings fescue (Festuca rubra var. commutata

Guad) every 1-3 days, depending on the size of the larvae. Pupae were kept in

stender dishes whose bottom had a 1 cm layer of moist peat. To estimate the

degree of field parasitism, larvae 1.5-2 cm long were collected in November from

commercial grass seed fields near either Peoria or Stayton, Oregon and reared as

previously described. All larvae were reared in controlled-environment chambers

1990

KAMM: THE GLASSY CUTWORM IN OREGON

JUNE JULY AUGUST SEPT OCT

DATE

Figure 1. Seasonal abundance of glassy cutworm adults attracted to a light trap operated near a

field of fine fescue.

at 20° C in a photoperiod of 16 h light, 8 h dark (LD 16:8) unless specified

otherwise.

The fine fescue fields and weeds within were sampled to determine larval density

in mid-summer and early autumn. Cores of crown and root zone (110 cm deep

by 10 cm diameter) of individual fescue or weed plants were removed and dissected

to determine larval numbers. Fine fescue plants were sampled when adjacent to,

or 1 m distant from, various weeds in the field. The number of larvae was de¬

termined in 20 plants of each species sampled. The weed species were bentgrass

(Agrostis tenus Sibth), ryegrass (Lolium perenne L. and L. multiflorum Lamarck),

and wild oats {Avena fatua L.). Because some of the latter were sprayed with the

Table 1. Mean time (days ± SD) that larvae and pupae of C. devastator required to develop at

20° C when exposed to different photoperiods.

Numher nf

Development time

Photoperiod

larvae

Larvae

Pupae

LD 16:8

102.9 ± 9.4 a

23.2 ± 2.5 a

LD 16:8 for 46 days, then LD 10:14

103.3 ± 17.6 a

23.6 ± 2.8 a

LD 10:14

126.8 ± 13.6

27.6 ± 2.9

a Larvae developed significantly faster compared with those exposed to the LD 10:14 regime (AN-

OVA, P < 0.001).

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Figure 2. Cumulative emergence of adults reared in various environmental regimens in controlled-

environment chambers.

herbicide glyphosate in mid-August, the numbers of larvae found on fine fescue

plants adjacent to sprayed and unsprayed plants were recorded in October.

Results and Discussion

Light trap data show that glassy cutworm adults were present in grass seed fields

from late June through September (Fig. 1). The seasonal abundance of adults

indicates that adults are most numerous during July and early August, when

spring-seeded fields of fine fescue are still green. At this time, older fields are

usually covered with postharvest straw or carbon residue from the cultural practice

of postharvest burning. Because the fire in burned fields moved rapidly and was

short-lived, cutworm pupae and larvae in plant crowns or soil escaped injury.

Upon emergence, adults are believed to fly out of burned fields into actively

growing, spring-seeded fields in search of oviposition sites.

Larval development was studied in the laboratory to determine the effect of

photoperiod on larval growth and diapause. Development proceeded at the same

rate when larvae several days old were exposed to either long days (LD 16:8) or

long days for 46 days followed by short days (LD 10:14) (Table 1). The latter

regime shows that early instar larvae developed more slowly in response to short

days; whereas, older larvae were not affected by the short-day regimen. Larvae

exposed to short days required 23.9 days longer to reach the pupal stage compared

with those exposed to long days. The cumulative emergence of adults is shown

in Fig. 2. Assuming oviposition temporarily parallels adult flight, the size variation

1990

KAMM: THE GLASSY CUTWORM IN OREGON

Table 2. Number and percentage of parasitized larvae of C. devastator collected from commercial

grass seed fields.

Date and location

Number of caterpillars

% parasitized

Parasites that pupated

Nowickia latianulum Lissonota montana

18 Jan 1974

Peoria, Oregon

159

10 Dec 1981

Stayton, Oregon

168

of larvae infesting fields in the autumn is probably due to egg deposition over

several months and to differential development of younger larvae exposed to the

shorter day lengths of late summer.

Some lepidopteran larvae that diapause in the Willamette Valley of Oregon

during winter may require a month or more to resume active development after

return of favorable conditions (Kamm 1973). The duration and stability of dia¬

pause vary among insects and in some cases diapause is little more than a transitory

delay in development (Beck 1980). When larvae of the glassy cutworm were taken

to the laboratory from the field during December and January, active feeding

began within 2 to 7 days. Larvae have also been observed to feed in the field

during periods of unseasonably warm winter temperatures in the Willamette Val¬

ley.

Many of the larvae collected from two field locations were parasitized by the

tachinid, Nowickia latianulum (Tothill), or the ichneumonid, Lissonota montana

(Cresson) (Table 2). Parasitized larvae were nearly mature before they ceased

feeding, and therefore it is doubtful that parasitization does much to reduce current

season feeding damage by larvae. However, subsequent adult populations will be

reduced according to the degree of parasitization in the larval population.

The distribution of larvae within a field of fine fescue indicated that significantly

more cutworms were found on fine fescue plants adjacent to weeds (nonfescue

grasses) than on fine fescue plants 1 m or more from the weeds (Table 3). When

the postharvest practice of burning temporarily eliminates all foliage in established

Table 3. Number of glassy cutworm larvae found on fine fescue near various weeds compared with

fine fescue 1 m from weeds (Stayton, Oregon, 13 Oct 1983).

Plant sampled

Weeds adjacent to fine fescue

Larvae/sample (X ± SD)

Fine fescue

None

0.70 ± 1.1

Fine fescue

Wild oats

4.00 ± 2.3 a

Fine fescue

Wild oats (sprayed)

1.60 ± 1.9 b

Bentgrass

—

3.85 ± 2.7 a

Fine fescue

Bentgrass

2.10 ± 1.6 a

Ryegrass

—

1.25 ± 1.5 b

Fine fescue

Ryegrass

5.00 ± 2.6 a

a Significantly more larvae present compared with fine fescue 1 m or more from weeds (ANOVA,

P < 0.001).

b Not significant.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

fields, females fly from burned fields to spring-seeded fields where green plants

are available for oviposition. The concentration of larvae on or near weeds suggests

that adults use these weeds for oviposition. Some larvae remained on perennial

ryegrass and bentgrass weeds but abandoned the annual wild oats upon senescence.

Also, fewer larvae were found on fine fescue plants adjacent to wild oats that were

killed with glyphosate compared with the number of larvae found on plants

adjacent to the unsprayed wild oats. Thus, early control of weeds reduced the

concentration of cutworms on fine fescue near nonfescue grassy weeds. Often the

fescue within a radius of 30-40 cm of living wild oat plants not treated with

glyphosate sustained considerable damage or was killed by feeding larvae in late

autumn. Other cutworm species are known to prefer weeds to crop plants for

oviposition (Busching & Turpin 1976).

Infestations may result in near elimination of crop plants within 20-30 cm of

weeds. Because the nonfescue grasses are often 30-80 cm taller than the fine fescue

in mid-summer, the taller plants may attract females. Regardless of the reason

for the concentration of larvae near weeds, control of weeds during early summer

reduced the concentration of larvae and feeding damage to the fine fescue.

Acknowledgment

I thank W. E. Gavin for assistance in conducting some of the tests and R. W.

Carlson and W. N. Mathis for identification of the parasites. This article is a

contribution of the ARS-USDA in cooperation with the Agric. Exp. Stn., Oregon

State Univ., Technical Paper No. 8822 of the Agric. Exp. Stn., Oregon State Univ.

Mention of a commercial or proprietary product does not constitute an endorse¬

ment of this product by USDA.

Literature Cited

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

Busching, M. K. & F. T. Turpin. 1976. Oviposition preferences of black cutworm moths among

various crop plants, weeds, and plant debris. J. Econ. Entomol. 69: 587-590.

Coquillett, D. W. 1897. Revision of the tachinidae of America north of Mexico. U.S. Dept. Agric.,

Div. Entomol., Tech. Bull., 7.

Crumb, S. E. 1929. Tobacco cutworms. U.S. Dept. Agric., Tech. Bull., 88.

Gibson, A. 1915. Cutworms and their control. Dom. Canada Dept. Agric., Entomol. Branch,

Bull., 10.

Gillette, C. P. 1891. Notes on habits and life histories of certain cutworms and cutworm moths.

Iowa Agric. Exp. Stn. Bull., 12.

Kamm, J. A. 1973. Role of environment during diapause on the phenology of Chrysoteuchia topiaria

(Lepidoptera: pyralidae). Entomol. Exp. Appl., 16: 407-413.

Rings, R. W. & F. J. Arnold. 1974. An annotated bibliography of the glassy cutworm. Ohio. Agric.

Res. Develop. Cntr. Res., Circ., 199.

Slingerland, M. V. 1902. Trap lanterns or “moth catchers.” N.Y. Cornell Agric. Exp. Stn., Bull.,

202.

Strickland, E. H. 1921. Parasites of the pale western cutworm in Alberta, Canada. Can. Entomol.,

53: 97-100.

Received 15 April 1989; accepted 9 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 71-78, (1990)

REPRODUCTION OF THE SAND WASPS

STICTIA SIGN AT A (L.) AND

BICYRTES VARIEGATA (OLIVIER)

(HYMENOPTERA: SPHECIDAE) ON THE

CARIBBEAN COAST OF QUINTANA ROO, MEXICO

Wayne F. Martin 1 and Robert F. Martin

Texas Memorial Museum and Department of Zoology,

The University of Texas at Austin, Austin, Texas 78705

Abstract.— Reproductive behavior of the sand wasps Stictia signata (L.) and Bicyrtes variegata

(Olivier) was studied from 1984 to 1988 on the Caribbean shoreline of the Yucatan Peninsula,

Mexico. Data on seasonal and diel activity patterns, on prey, and on excavation, orientation,

and dimensions of burrows are presented. A male flight polymorphism, previously undescribed

for the species, occurred in S. signata ; some males hovered in flight at various heights near

foliage, while others patrolled in sinuous flight near the surface of the beach. Burrows of B.

variegata, reported from other locations as polycellular, were with one exception unicellular.

Characteristics of B. variegata at our research site are discussed speculatively as possible con¬

sequences of drought.

Key Words. — Insecta, Hymenoptera, Sphecidae, Stictia signata, Bicyrtes variegata, reproduction,

male flight polymorphism, Mexico

The accumulation of observations on sand wasps has permitted syntheses of

hypotheses on their behavioral evolution (Evans 1966, Alcock et al. 1978, O’Neill

& Evans 1988). Comparisons between populations of single species from diverse

areas and climates have increased the knowledge of behavioral variability that is

necessary to understand differences between higher taxa, as well as behavioral

plasticity within single demes (Alcock 1979; O’Neill 1983; O’Neill & Evans 1983a,

b, 1988).

Stictia signata (L.) and Bicyrtes variegata (Olivier) have extensive Neotropical

distributions (Evans 1966); aspects of the reproduction of each have been reported

from widely separated locations (Janvier 1928, Callan 1954, Vesey-Fitzgerald

1956, Evans 1966, Post 1981, Philippi & Eberhard 1986). Here we describe

reproductive patterns of populations of these species that nest on the Caribbean

beaches of Quintana Roo, Mexico. We report several behavioral attributes that

differ markedly from those previously described for these species, including male

flight polymorphism in S. signata, and speculate on the role of unusual environ¬

mental conditions in influencing behavior.

Methods

Site Description. — We observed wasp behavior on beaches at Akumal (20°24'

N, 87° 18' W), Quintana Roo, Mexico, where sandy embayments alternate with

low rocky headlands. Variation in water level at the shore appeared to be more

a function of easterly trade wind velocity and direction than of minor semidiurnal

1 Present address: Department of Psychology, Virginia Commonwealth University, Richmond, Vir¬

ginia 23284.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

tidal changes (< 0.3 m, see Randall & Martin 1987) at the location. Thick vege¬

tation began 100+ m inland. Seasonally flooded dry areas alternated with nearly

permanent swampy areas along the coast. The general area was transitional vege-

tationally, and farther inland on slightly higher ground, elements of deciduous

dry forest and rainforest occurred. Development of rural communities with tourist

facilities (as at Akumal) presently is common in beachfront areas.

The year during which our primary data were taken (1985) was one of severe

local drought. Rains fell only occasionally and lightly at Akumal before and during

our work. This scarcity of rainfall interacted with a series of new and older

causeways (impassable to drainage from adjacent areas that received rain) to allow

a considerable portion of the normally swampy area inland to become dry and

partially devegetated.

Methods.— We did not attempt to investigate all aspects of reproductive be¬

havior equally, but arbitrarily concentrated our effort in areas we deemed most

significant and/or time effective. Thus, some behaviors are described in outline,

qualitatively, or only by reference to patterns described for the species in the

literature. When appropriate we present only ranges of behavior in such descrip¬

tions and for those based partly on observations begun or terminated in mid¬

behavior.

Our observations occurred from dawn to dusk until we were able to discern

the outlines of major activity patterns; we then concentrated our efforts over

shorter time periods. Observations occurred: 18 Dec 1984; 26-30 Apr, 5 Jun-10

Jul, and 6-15 Aug 1985; 22-27 Apr, 1-7 Jun, 26 Jun-5 Jul, and 20 Dec 1986;

7-18 Jun 1987; and 18 Mar 1988, for a total of 270 h. Data taken during June

and July 1985 (190 h) form the bases of most qualitative descriptions, and all

those presented in Tables 1 and 2.

Descriptions of male behavior are based on 74 observations of S. signata-, those

of female behavior, primarily on repeated observations at marked burrows of 26

S. signata and 30 B. variegata. We dug out burrows at which we had previously

observed repeated activity (26 of S. signata, 30 of B. variegata :), and we provide

statistical data (Table 1) for those at which we had previously observed provi¬

sioning and/or which contained provisions.

Wasps were disturbed only occasionally by us during observation and did not

appear to be confused (after brief reorientation flights) by the numbered beverage

bottle caps that we introduced as burrow markers while the wasps were under¬

ground.

Identifications were provided by Arnold Menke and James Gillaspy (sphecids),

T. J. Henry (hemipterans), and N. E. Woodley (dipterans).

Biology

At Akumal, S. signata and B. variegata aggregated most densely on moderately

compacted sand proximate to human habitation. Areas of the back beach near

the sea, with only scattered ground vegetation and partially shaded by coconut

palms, seemed most favored by both species. Female S. signata usually nested

in moderate density (— 1 per m 2 ) toward the periphery of male flight areas. Female

B. variegata nested in isolation or intermingled loosely with, and less densely

among, female S. signata. Dense aggregations (> 15 per m 2 ) of Microbembix

monodonta (Say) were also common.

Table 1. Descriptive statistics for burrows of Stictia signata and Bicyrtes variegata at Akumal, Quintana Roo, Mexico. ab

Species

Entry direction (degrees)

Distance from shore (m)

Slope down (degrees)'

Burrow length (cm)

Burrow depth (cm)

± SE

S. signata

157.6

18.1

41.1

2.5

32.5

2.4

32.9

2.9

21.2

1.2

36.5

2.0

B. variegata

149.2

8.5

30.8

2.8

36.3

1.8

13.1

0.7

9.0

0.5

38.5

2.1

a All measurements were not possible at all burrows; thus, sample sizes vary.

b Only included are burrows observed being provisioned or those that contained prey.

c Upper figures represent initial slope; lower, final slope.

1990 MARTIN & MARTIN: REPRODUCTION OF SAND WASPS

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 66(1)

Table 2. Numbers and taxa of nymphal hemipteran prey of B. variegata at Akumal, Quintana

Roo, Mexico.

Cell

Family

Alydidae

Coreidae

Pentatomidae

Scutelleridae

1 (Hyalemenus sp.)

5 {Arvelius sp.)

1 (Hyalemenus sp.)

2 ( Arvelius sp.)

1 (Hyalemenus sp.)

1 (Arvelius sp.)

3 ( Arvelius sp.)

1 (Leptoglossus sp.)

3 ( Hyalemenus sp.)

2 ( Hyalemenus sp.)

2 ( Arvelius sp.)

6 (Prob. Homaemus sp.)

Stictia signata Female Activity.— Females were observed digging during April,

June-August, and December. December sightings were rare, but a large larva was

recovered from an exhumed cell on 18 Dec 1984. During June, females dug or

provisioned burrows as early as 05:20 h. Activity continued through early after¬

noon, slackened during mid-afternoon, then increased again, and tapered off after

17:00 h. Sand temperatures (taken on 14-15 Jun 1987 in sunny areas just below

the surface of sand recently shaded by us) in wasp aggregations approximated 27°

C at 06:00 h and 37° C from 10:00-14:00 h. Air temperature 7.6-15.2 cm above

sand surfaces ranged from 28° C at 06:00 h to approximately 32° C from 10:15-

14:00 h, and descended subsequently.

Burrow initiation usually began early in the day. Digging, closing, and leveling

usually followed the descriptions of Evans (1966) for Stictia Carolina (Fabr.). With

few exceptions, possibly due to our disturbance, an outer closure was maintained

even during regular provisioning of large larvae. Females arriving at nests fre¬

quently attracted small, possibly kleptoparasitic or scavenger dipterans that often

succeeded in entering burrows with little reaction from the female wasps.

Entries to burrows usually were situated on the back beach in loose aggregations

in compacted sand and averaged 41.1 m from the edge of the sea (Table 1).

Burrows led toward the SSE (Table 1), in the general direction of the sea and into

the prevailing direction of the wind. Burrows were longer than deep, and if mul-

1990

MARTIN & MARTIN: REPRODUCTION OF SAND WASPS

tiply-pitched, they were slightly but not significantly (P > 0.1, Mann-Whitney

U- test) steeper farther from the entrance (Table 1). Of 18 provisioned burrows

that we excavated, all were unicellular with loose inner closures.

Provisioning involved dipteran prey, carried in the usual (see Evans 1966)

position. Members of three families of Diptera were identified from exhumed

cells (other taxa may have been represented in the fragmented material): Calli-

phoridae— Chrysomya megacephala (Fabr.) and Cochliomyia macellaria (Fabr.);

Sarcophagidae— Helicobia sp. and three unidentified spp. of Sarcophaginae; Ta-

chinidae— Muscopteryx sp. In addition to the wasp larva, small, probably scav¬

enging or kleptoparasitic larvae occurred in most cells, and were usually buried

in frass and loose sand.

Stictia signata Male Activity. —Sinuous, patrolling flight usually occurred ap¬

proximately 0.5 m above the surface of extensive non-vegetated sandy areas. The

flight was similar to that described for S. Carolina by Evans (1966) and was

conspicuous during most sunny mornings in April and from June through August.

This activity usually began after 06:15 h, increased until 08:00-09:00 h, then

slackened, and ceased between 10:30 and 12:00 h. Males frequently patrolled

semi-exclusive, irregularly shaped areas of over 20 m 2 . The sinuous flight patterns

of males from adjoining patrol areas were occasionally interrupted by face-offs,

head butting, and then upward flight by both males.

Stationary hovering also occurred in this population. At 07:00 h or usually

later, males frequently appeared in hovering flight at relatively fixed positions.

Hovering was not observed after 12:15 h. Hovering males were less common than

those in patrolling flight. Males hovered in isolation or in groups; those in groups

usually were dispersed horizontally and separated by at least 4 m. Individual

hovering positions were held for as long as 1 h. Brief rests on nearby palm fronds

and quick forays to interact with neighboring males occurred during hovering

periods. Vertical stratification of hovering males also existed. In one situation (28

Jun 1986), five individuals hovered at various levels near shrubs—two spaced

vertically between 1-1.6 m, and three spaced vertically between 2-4 m, while

circuitous patrolling flight of other males occurred within 5 m. We observed

hovering flight on 23 Apr 1986 and during June of 1985-1987, but made no

attempt to define its seasonal limits beyond these dates.

Bicyrtes variegata Female Behavior. — Females were observed digging burrows

during April, and from June through August, but not during December. Females

became active later in the day than did S. signata females; their earliest digging

was observed at 07:30 h (June) and earliest provisioning at 07:44 h (June). Digging

involved accentuated body tilting. Stones up to 0.4 cm diameter occasionally were

carried with the mandibles to the burrow entrance from inside and were tossed

sideways away from it. Burrow initiation usually took place in the forenoon;

completion took 45 min to 2.5+ h. Arriving females frequently attracted small,

possibly kleptoparasitic or scavenging dipterans that frequently succeeded in en¬

tering the burrow. Female B. variegata exhibited only cursory or moderate neg¬

ative reaction toward these flies.

After digging or provisioning, the wasps’ exits from the burrow were usually

head first. They maintained an outer closure of loose sand that was casually kicked

inward or was more carefully constructed by entering the burrow backwards while

kicking. Final abdominal tamping of the closure occurred in some instances. The

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

wasps exited the site, after one to several orientation flights, in an upward departing

flight.

Entries to B. variegata burrows were most frequently situated on the back beach

in compacted sand, loosely scattered among S. signata burrows. Burrows led

predominately toward the SSE, averaged 30.8 m from the sea, were longer than

deep, and were not significantly (P > 0.4, Mann-Whitney U- test) steeper toward

the rear (Table 1). Of 24 burrows exhumed, 23 had one cell and one had two

cells. No inner closures were noted.

Provisioning with nymphal hemipterans occurred from 07:44 h to 17:04 h.

Provisioning usually began on the same day as burrow initiation, and either was

completed during that day, or extended to a second day. When wasps returned

to burrows with prey, they descended by a vertical hovering flight. Wasps spent

from 1-5 min inside the burrow during each provisioning visit; the shortest

interprovisioning interval was 3 min, but provisioning intervals usually lasted

10-30+ min.

Four families of nymphal hemipterans were collected from 25 cells of 24 bur¬

rows between 8 Jun-5 Jul 1985; most were completely paralyzed. The wasp laid

its egg in a semi-erect position on the mid-ventral line of the first hemipteran

brought to the burrow, usually between the meso- and metacoxal bases. Penta-

tomids were used most frequently, followed by coreids, scutellerids and alydids

(Table 2). Individuals of three families occurred in one cell, two families in six

cells, and one in 18 cells (Table 2). In one cell, we found a quite large wasp larva

24 h after an adult had provisioned the burrow; in another, a large larva was

found within one hour of a provisioning visit. This suggests that some progressive

provisioning occurs in this species. A maximum of seven prey individuals was

found in a cell (Table 2); this total, however, is biased by an early exhumation

protocol but may be due in part to a scarcity of prey during drought conditions.

In addition to the wasp larva, small, probably scavenging or kleptoparasitic larvae

occurred in most cells.

The sizes of female B. variegata captured while digging varied considerably and

may reflect resource shortage and foraging success. Dimensions of four specimens

collected at Akumal ranged from 3.7-5.2 (x ± SD: 4.3 ± 0.7) mm for head width,

12.3-19.2 (15.1 ± 3.2) mm for total length, and 9.5-14.0 (11.2 ± 2.1) mm for

wing length.

Discussion

The digging behavior of female S. signata at Akumal was typical of that reported

for the genus (Evans 1966). Burrow length was similar to that recorded for this

species in Brazil (Post 1981), but shorter than that of other populations of S.

signata (Evans 1966) or for other species of Stictia (Evans 1966, Evans & Matthews

1974, Matthews et al. 1981, Sheehan 1984).

Prey items of S. signata were from families of Diptera previously reported as

utilized by this wasp (Evans 1966, Post 1981); Cochliomyia macellaria (Fabr.),

taken for provisioning at Akumal, is also captured by S. signata on Dominica

and Puerto Rico (Evans 1966) and in Brazil (Post 1981).

Females of B. variegata at Akumal are typical of those of other Bicyrtes (Evans

1966) in that they are only loosely (if at all) gregarious, and nested in isolation

or among aggregations of another sand wasp. In contrast to the situation reported

1990

MARTIN & MARTIN: REPRODUCTION OF SAND WASPS

as typical by Evans (1966), burrows at Akumal were (one exception) unicellular;

this characteristic and the considerable range of size of adults may reflect a patchy

scarcity of prey occasioned by drought conditions in the area.

At Akumal, B. variegata utilized only immature hemipterans; Evans (1966)

noted that adults were less common than nymphal stages among Bicyrtes prey.

Also, as Evans (1966) summarized, pentatomids and coreids were the most com¬

mon prey; however, nymphal alydids, not mentioned as prey of this genus by

Evans (1966), nor by Evans & Matthews (1974), were also present.

Our observations of several concurrent intrademe flight patterns by male S.

signata augment the alternative mating strategies reported for other sphecids (i.e.,

Alcock 1975, 1979; Evans & O’Neill 1978; O’Neill & Evans 1983a, b, 1988).

Low, moving flight has been described for male S. signata in Quintana Roo by

Evans (1966). Low hovering flight was reported by Post (1981) in Brazil. William

Sheehan (personal communication) noted interpopulational differences in male

S. signata flight at separate Costa Rican sites during different years: in one pop¬

ulation, a low rapid directional patrol along a path in which females nested was

seen; in another, males hovered within the spreading branches of a shrubby tree

from near ground level to 4-5 m. Sheehan (personal communication) also ob¬

served alternative male flight patterns within populations of Stictia heros (Fabr.)

in Costa Rica, where some males patrolled an open beach with fast low (10-20

cm) sinuous flight, while others held territories at the edge of the beach; individuals

which were marked performed both behaviors.

Our data supplement those of studies that suggest that broad behavioral plas¬

ticity exists within sphecid species as an adaptive response to a variable environ¬

ment (Alcock 1979, Alcock et al. 1978, O’Neill & Evans 1983b). We speculate

that an unusual suite of attributes of B. variegata reported herein (reduced prey

number, nymphal prey only, unicellular burrows, and small adult size) may reflect

this adaptation and may have been mediated by a drought-influenced depauperate

food chain.

Acknowledgment

We thank K. Tedin, A. Hook, D. Watson, L. Wheeler, J. Edwards, L. Adkins,

D. Gottschall, L. Hamilton, M. Martin, and W. Sheehan for field and reference

assistance. Arnold Menke, T. Henry, and N. Woodley of the Taxonomic Services

Unit, Systematic Entomology Laboratory, U.S. Department of Agriculture, Belts-

ville, Maryland, and J. Gillaspy identified specimens. Two anonymous referees

provided editorial and content assistance. The project was supported in part by

the Texas Memorial Museum, The University of Texas at Austin.

Literature Cited

Alcock, J. 1975. Male mating strategies of some philanthine wasps (Hymenoptera: Sphecidae). J.

Kans. Entomol. Soc., 48: 532-545.

Alcock, J. 1979. The evolution of intraspecific diversity in male reproductive strategies, pp. 381-

402. In Blum, M. S. & M. A. Blum (eds.). Sexual selection and reproductive competition in

insects. Academic Press, New York.

Alcock, J., E. M. Barrows, G. Gordh, L. J. Hubbard, L. Kirkendall, D. W. Pyle, T. L. Ponder & F.

G. Zalom. 1978. The ecology and evolution of reproductive behavior in the bees and wasps.

Zool. J. Linn. Soc., 64: 293-326.

Callan, E. McC. 1954. Observations on Vespoidea and Sphecoidea from the Paria Peninsula and

Patos Island, Venezuela. Bol. Entomol. Venez., 9: 13-27.

THE PAN-PACIFIC ENTOMOLOGIST

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Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps. Harvard University

Press, Cambridge, Massachusetts.

Evans, H. E. & R. W. Matthews. 1974. Observations of the nesting behavior of South American

sand wasps (Hymenoptera). Biotropica, 6: 130-134.

Evans, H. E. & K. M. O’Neill. 1978. Alternative mating strategies in the digger wasp Philanthus

zebratus. Proc. Nat. Acad. Sci. (Washington, D.C.), 75: 1901-1903.

Janvier, H. 1928. Recherches biologiques sur les predateurs du Chile. Ann. Sci. Nat. Zool., 10(11):

67-207.

Matthews, R. W., R. A. Saunders & J. R. Matthews. 1981. Nesting behavior of the sand wasp Stictia

maculata (Hymenoptera: Sphecidae) in Costa Rica. J. Kans. Entomol. Soc., 54: 249-254.

O’Neill, K. M. 1983. The significance of body size in territorial interactions of male beewolves

(Hymenoptera: Sphecidae, Philanthus). Anim. Behav., 32: 404-411.

O’Neill, K. M. & H. E. Evans. 1983a. Alternative male mating tactics in Bembecinus quinquespinosus

(Hymenoptera: Sphecidae): correlation with size and color variation. Behav. Ecol. Sociobiol.,

14: 39-46.

O’Neill, K. M. & H. E. Evans. 1983b. Body size and alternative mating tactics in the beewolf

Philanthus zebratus (Hymenoptera: Sphecidae). Biol. J. Linn. Soc., 20: 175-184.

O’Neill, K. M. & H. E. Evans. 1988. The natural history and behavior of North American beewolves.

Comstock Publishing Associates, Cornell University Press, Ithaca, New York.

Phillipi, T. & W. G. Eberhard. 1986. Foraging behavior of Stictia signata (Hymenoptera: Sphecidae).

J. Kans. Entomol. Soc., 59: 604-608.

Post, D. C. 1981. Observations on female nesting and male behavior of Stictia signata (Hymenoptera:

Sphecidae) in Brazil. Rev. Biol. Trop., 29: 105-113.

Randall, C. W. & R. F. Martin. 1987. Distribution, abundance, and movement patterns of shoreline

chitons of the Caribbean coast of Mexico. Nautilus, 101: 75-79.

Sheehan, W. 1984. Nesting biology of the sand wasp Stictia heros (Hymenoptera: Sphecidae: Nys-

soninae) in Costa Rica. J. Kans. Entomol. Soc., 57: 377-386.

Vesey-Fitzgerald, D. 1956. Notes on Sphecidae (Hym.) and their prey from Trinidad and British

Guiana. Entomol. Month. Mag., 92: 286-287.

Received 12 March 1989; accepted 14 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 79-88, (1990)

FURTHER OBSERVATIONS ON THE BIOLOGY AND

HOST SPECIFICITY OF PROCHOERODES TRUXALIATA

(GUENEE) (LEPIDOPTERA: GEOMETRIDAE),

A BIOLOGICAL-CONTROL AGENT FOR

BACCHARIS HALIMIFOLIA L. IN AUSTRALIA

L. E. Ehler , 1 M. G. Kinsey 1 and W. A. Palmer 2

department of Entomology, University of California, Davis, California 95616

2 North American Field Station, Queensland Department of Lands,

2801 Arrowhead Circle, Temple, Texas 76502

Abstract.— Studies on the developmental and reproductive biology of Prochoerodes truxaliata

(Guenee) were conducted on Baccharis pilularis de Candolle at Davis, California. Under labo¬

ratory conditions, mean developmental time from egg to adult was slightly shorter for males (49

days) than for females (50.6 days). Developmental times for fourth and fifth instar larvae and

for pupae were significantly different between sexes. Mean generation time for females was

approximately 53 days. There were five larval instars and measurement of head capsules revealed

a relatively constant growth ratio (0.64-0.68). Longevity of adults was approximately 10 days.

Following a short preoviposition period (approximately two days), females which laid fertile

eggs produced an average of 489 eggs per female; however, not all the eggs were always deposited

prior to the death of a given female. Other females, despite their exposure to males, laid only

infertile eggs; however, total number of eggs produced per female was lower than that observed

for fertile females (x = 339) and only one third of these were actually deposited. Field observations

revealed that moths which were active early in the evening laid predominantly fertile eggs,

whereas those active later in the evening laid predominantly infertile ones. In quarantine facilities

in Australia, host specificity of larvae was tested on a wide range of economic and native plant

species unavailable in the U.S. Larvae fed on a number of asteraceous plants but were unable

to complete development on any plant other than Baccharis halimifolia L. Consequently, P.

truxaliata has been cleared for release in Australia for control of B. halimifolia.

Key Words.— Insecta, Lepidoptera, Geometridae, Prochoerodes truxaliata, biological control/

weeds, Baccharis

There are over 200 species of insects associated with Baccharis pilularis de

Candolle in California (Tilden 1951). This insect fauna has been of considerable

ecological interest over the years, and more recently certain species in the com¬

munity attained practical importance in biological control. A related host plant,

Baccharis halimifolia L., which is native to the southeastern U.S., is a serious,

introduced rangeland weed in Australia. During foreign exploration for natural

enemies of B. halimifolia in the U.S., it was discovered that a number of species

previously recorded only from B. pilularis would also feed and complete devel¬

opment on B. halimifolia (WAP, unpublished data). At least two were sufficiently

stenophagous to permit their importation into Australia. The cecidomyiid midge

Rhopalomyia californica Felt, which develops in conspicuous terminal galls on

B. pilularis, was introduced into Queensland in 1982; it established, and by 1986

had spread throughout the range of the target weed (McFadyen 1985; WAP,

unpublished data). The second species, Prochoerodes truxaliata (Guenee), is one

of the least conspicuous insects associated with B. pilularis in California. The

cryptic larvae of this geometrid can cause severe defoliation and thus this species

shows considerable promise as a biological-control agent.

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

The pre-introductory investigations required for clearing P. truxaliata for im¬

portation into Australia resulted in a considerable amount of new information

on this rather poorly known species. Palmer & Tilden (1987) reported on its

developmental biology and host specificity; the latter studies involved plants

available in the U.S. (primarily Texas), including native species. This paper reports

additional data on developmental and reproductive biology of P. truxaliata, and

the results of host-specificity tests conducted in Australia.

Materials and Methods

Laboratory Studies.—The insects used in the laboratory studies were the progeny

of 20 female moths collected from B. pilularis pilularis on the Davis campus of

the University of California (Solano County). The moths were held in a sleeve

cage in the laboratory and observed daily for oviposition. Eggs were collected

each day and placed in ventilated microcentrifuge tubes. One cohort of eggs was

stored at 5° C for 18 days before being placed in the rearing room, whereas all

other eggs were immediately placed in the rearing room and held at 25° C. The

eggs held in cold storage were a subset of those eggs allowed to accumulate in

storage prior to shipment to Australia for use in host-specificity tests. During

incubation, all eggs were observed several times daily and the date and time of

eclosion were recorded for each larva.

Newly hatched larvae were transferred to rearing chambers constructed from

30 ml plastic containers and held at 25° C (one larva per chamber). A circular

hole (one cm diameter) was cut in the bottom of the container and covered with

90-mesh nylon organdy. The severed end of a cutting taken from a growing tip

of B. p. pilularis from the Davis campus was inserted through a small hole in the

lid of the plastic cup so that about one cm of stem protruded. The lid was placed

on the cup, the entire unit was inverted, and then allowed to float on styrofoam

in a tray of distilled water allowing the stem of the cutting to remain submerged.

As larvae grew larger, these containers were replaced with 60 ml plastic containers

of similar construction. Rearing chambers were cleaned and fresh foliage was

added daily. Larvae were observed several times a day and behavioral activities

were noted.

A minute droplet of silver conducting paint (Ladd Research Industries, Inc.,

stock number 60805) was placed on the dorsal surface of the head of the newly

hatched larva. This was repeated after each molt and the head capsules were later

recovered for each successive instar. The sequence of head capsules for each larva

was then glued to a microscope slide; each capsule was measured at 50 x using a

Nikon stage micrometer electronically wired to an Autometronics dual-axis digital

readout. Measurements were expressed in millimeters.

Pupae were sexed and held in the rearing room until emergence. As the moths

emerged, individual male/female pairs were confined in 450 ml plastic cylindrical

containers. The bottom of the container was removed and replaced with a double¬

weave brass screen. One mm-square openings in the screen permitted the 0.8 mm

eggs to drop through, and into a collecting tray beneath the unit. A circular hole

was cut in the lid of the container and replaced with 90-mesh nylon organdy. A

cotton wick saturated with 20% sucrose solution was inserted through a hole in

the side of the cage. Eggs were collected daily and held in ventilated microcen¬

trifuge tubes at 25° C for determination of fertility. Moths were observed daily,

1990

EHLER ET AL.: BIOLOGY OF PROCHOERODES

and dead individuals were removed. Shortly after death, females were dissected

in physiological saline solution and the condition of the reproductive tract and

the number of eggs remaining was recorded.

Field Observations.— Moth behavior was observed in the campus arboretum

during the evenings of 11-12 Jul 1988. On each night, moths were observed and

captured during two intervals: 20:30-22:10 h and 23:00-00:10 h. Observations

were terminated shortly after midnight due to the general absence of active female

moths. Behavior of moths was observed with the aid of a flashlight. All females

were returned to the laboratory, held individually in 60 ml plastic containers, and

observed for oviposition. Egg fertility was recorded for each female.

Host Specificity. — In the quarantine laboratory at the Alan Fletcher Research

Station (Sherwood, Qld., Australia), all plants listed in Table 1 were tested against

neonate larvae. A leaf of each plant was selected and examined to ensure that

there was no insect damage. It was then placed in a plastic petri dish with five

newly hatched, unfed larvae. The larvae used in this and subsequent experiments

were obtained from eggs shipped directly to Australia from Davis. The source

population was the same as that used in the laboratory studies. Every two days

the contents of the dishes were examined and the foliage replaced. Live larvae,

frass, head capsules or feeding marks on the leaf were noted. Three replications

of each plant species were made. The tests were conducted in randomized batches

of 20 plants which always included B. halimifolia as a control.

Eighteen species were selected from those in Table 1 to ascertain whether larvae

behaved similarly on potted plants and on cut foliage. The selection included all

the Astereae and a random selection of other species. Five neonate larvae were

placed on foliage of each plant; a fine-mesh bag was placed over them to confine

them to a portion of the plant. The plants were placed in a greenhouse, examined

at regular intervals, and the surviving larvae noted. The experiment was replicated

twice.

A test was also conducted to determine whether late-instar larvae had a wider

host range than neonate larvae. In this test, which was replicated twice, all the

species in the Asteraceae in Table 1 were used. Larvae were reared from eclosion

on B. halimifolia foliage until they were estimated to be third instar. Five larvae

were then transferred to cuttings of a test plant held in souffle cups filled with

water. These cuttings were then placed individually in 500 ml plastic containers.

The contents of the containers were then examined regularly and live larvae, frass,

exuviae and feeding damage noted. The foliage was replaced every three days.

Results and Discussion

Laboratory Studies.—Our observations of the eggs of P. truxaliata were gen¬

erally similar to those of Palmer & Tilden (1987). The egg is nearly spherical

(slightly ovoid), and approximately 0.8 mm in diameter. The chorion is glossy

with a textured surface, and has no known adhesive substances. The newly de¬

posited egg is turquoise green. The fertile egg turns brown after about 48 h (some

may require up to 72 h), whereas an unfertile egg retains its original color. As it

hatches, the first instar larva chews a circular opening in the chorion. The chorion

is seldom if ever eaten by the larva. Mean developmental time for eggs was

approximately 10 days at 25° C, with no significant difference between males and

females (Table 2). In contrast, chilled eggs (at 5° C for 18 days prior to incubation

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Table 1. The plants against which neonate larvae were tested in the cut foliage, no-choice exper¬

iment.

Anacardiaceae: Mangifera indica L.

Apiaceae: Apium graveolens L.

Asteraceae: Tribe Astereae: Aster novi-belgii L., Brachycome multifida de Candolle, Callistephus chi-

nensis (L.) Nees Von Esenbeck, Calotis cuneata (F. Mueller ex. Bentham) G. L. Davis, Conyza

sumatrensis (Retzius) E. H. Walker, Olearia nernstii F. Mueller, Bentham, Vittadinia sulcata N. T.

Burbidge. Tribe Heliantheae: Cosmos bipinnatus Cavanilles, Dahlia variabilis (Willdenow) Desfon-

taines, Ecliptaprostrata (L.) L., Gaillardia aristata Pursh, Glossogyne tenuifolium Cassini, Helianthus

annuus L., Wedelia biflora de Candolle, Zinnia linearis Bentham. Tribe Inuleae: Cassinia laevis R.

Brown, Gnaphalium sphaericum Willdenow, Helichrysum bracteatum (Ventenat) Andrews. Tribe

Senecioneae: Emilia sonchifolia (L.) de Candolle, Flaveria australasica Hooker, Senecio lautus

Solander, ex. Willdenow. Tribe Anthemideae: Chrysanthemum carinatum Schousboe, Cotula aus¬

tralis (Sieber) Hooker f. Tribe Eupatorieae: Adenostemma lavenia (L.) Kuntze. Tribe Vemonieae:

Vernonia cinerea (L.) Lessing. Tribe Lactuceae: Cichorium intybus L. Tribe Cynareae: Carthamus

tinctorius L. Tribe Calenduleae: Calendula officinalis L.

Brassicaceae: Brassica oleracea L.

Caricaceae: Carica papaya L.

Chenopodiaceae: Beta vulgaris L.

Cucurbitaceae: Cucurbita maxima Duchesne

Fabiaceae: Arachis hypogaea L., Medicago sativa L., Phaseolus vulgaris L.

Liliaceae: Allium cepa L.

Malvaceae: Gossypium hirsutum L.

Myrtaceae: Eucalyptus sp.

Passifloraceae: Passiflora edulis J. Sims

Poaceae: Triticum aestivum L., Saccharum officinarum L., Sorghum vulgare Persoon, Zea mays L.

Proteaceae: Macadamia integrifolia J. Maiden & E. Betche

Rosaceae: Fragaria vesca L.

Rubiaceae: Coffiea arabica L.

Rutaceae: Citrus sinensis (L.) P. Osbeck

Solanaceae: Solanum tuberosum L., Lycopersicon lycopersicum L.

Vitaceae: Vitis vinifera L.

at 25° C) hatched at least three days sooner. Such an effect has been documented

in other insects (Lin et al. 1954, Kimberling & Miller 1988). In such cases, although

the insect’s embryonic development evidently continues at temperatures below

the thermal threshold for eclosion, the entire developmental sequence leading to

larval eclosion cannot be completed. This finding may be helpful in the mass

production and colonization of P. truxaliata, particularly when cold storage of

eggs is necessary. However, the eclosion rate among chilled eggs was considerably

less (54%, n = 69) than for non-chilled eggs (78.1%, n = 170).

Larval eclosion occurred between dusk and dawn. The neonate larva actively

searched the host plant and generally displayed a negative geotropism (also noted

by Palmer & Tilden 1987). When given a choice, the larva fed on young leaves

in the meristematic region. In the absence of such leaves, the larva would feed

on well developed leaves and showed a marked preference for younger foliage.

As noted by Palmer & Tilden (1987), first instar larvae (in contrast to later instar

larvae) commonly fed during the day in addition to nocturnal feeding. When

resting, the smaller larva assumed an erect position, mimicking defoliated petioles;

larger larvae mimicked small twigs. The larva held the substrate with its prolegs

Table 2. Developmental times and head-capsule widths for P. truxaliata in the laboratory.

Developmental time (days ) 3

Mean ± SEM C

Range

Mean (± SEM) b head-capsule width (mm) c

Male

Female

Male Female

Egg

9.63 ± 0.09

9.60 ± 0.09

9-10

1 st in star

4.81 ± 0.18

4.63 ± 0.21

4-7

0.430 ± 0.003

0.429 ± 0.003

2nd instar

4.66 ± 0.2

4.43 ±0.11

3-8

3-6

0.656 ± 0.005

0.659 ± 0.006

3rd instar

4.41 ± 0.19

4.33 ± 0.23

3-7

3-8

1.025 ± 0.014

1.039 ± 0.01

4th instar

5 th instar

4.47 ± 0.14

5.23 ± 0.14

3-6

4-7

1.541 ± 0.01

* 1.611 ±0.011

Active

6.66 ± 0.17

8.67 ± 0.32

5-9

6-14

2.294 ± 0.033

2.363 ± 0.043

Prepupa

2.25 ± 0.13

2.43 ± 0.11

1-4

1-3

—

Pupa

12.16 ± 0.2

11.33 ± 0.22

10-14

8-13

—

Egg to adult

48.97 ± 0.63

50.6 ± 0.75

42-58

44-60

—

a Based on 32 male and 30 female individuals which completed development to adult.

b Based on 17 male and 19 female individuals which completed development to adult.

c Asterisk indicates significant difference (Mest, P < 0.05).

1990 EHLER ET AL.: BIOLOGY OF PROCHOERODES

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

2.5 -

2.0 -

1.5 -

1.0 -

.5 -

1st

2nd

3rd

Instar

4th

5th

Figure 1. Mean and range of head-capsule widths for the five larval instars of P. truxaliata (data

from both sexes).

and assumed a 45° angle to the stem. This position was maintained with the aid

of a silken strand, from the spinnerettes, also attached to the stem. The larva

would drop when disturbed, and remain suspended from the foliage by the silken

strand. Mean developmental time for first-instar larvae was slightly less than five

days and did not vary with sex (Table 2).

Subsequent instars were nocturnal feeders. Second and third instar larvae showed

a similar preference for young leaves in the meristematic region and when dis¬

rupted, dropped from the plant as did the first instar. Mean developmental times

for these instars were approximately the same and did not vary with sex (Table

2). For fourth instar larvae, however, mean developmental time for males was

significantly shorter than for females; that for females was almost one day longer

than for females of the third instar (Table 2). Both fourth and fifth instar larvae

behaved differently than previous instars by showing less preference for younger

leaves, and not readily dropping from the plant when disturbed.

The fifth instar consisted of an active feeding phase followed by a non-feeding

prepupal period. Mean developmental time for the active phase was the longest

for any instar, and that for the males was significantly shorter than for females

(Table 2). The prepupal period lasted slightly more than two days and did not

vary with sex (Table 2). During this period, a silken structure was spun which

eventually housed the pupa. The pupal stage was significantly shorter for females

1990

EHLER ET AL.: BIOLOGY OF PROCHOERODES

2 4 6 8 10 12 14 16 18 20 22

DAYS FOLLOWING ADULT ECLOSION

Figure 2. Mean number of eggs laid per female per day for P. truxaliata under laboratory conditions.

than for males. Sexual dimorphism, expressed in the terminal abdominal seg¬

ments, was sufficient to enable sorting of pupae according to sex.

The average head-capsule widths for the five instars were consistent with the

“Brooks-Dyar Rule” (cf. Daly 1985). With the exception of fourth instar larvae,

there was no significant difference between sexes (Table 2). Average values in¬

corporating data from both sexes are plotted in Fig. 1. There is no overlap among

the instars and thus head-capsule width would appear to be a suitable indicator

of instar in field collections. The range for fifth instar larvae was relatively large.

We attribute this to the fact that dorsal sutures of these head capsules seldom

remained intact; this caused difficulty in obtaining accurate measurements. The

growth ratio was relatively constant (0.64-0.68) and is generally consistent with

previous studies oflepidopterous larvae (Drooz 1965, Dupree 1965, Enders 1976).

In two cases, we observed six instars, but neither survived to adulthood. Palmer

& Tilden (1987) reported up to seven and sometimes eight instars in this species.

This is not particularly surprising, as number of instars in Lepidoptera can vary

according to environmental conditions (Long 1953).

Mean developmental time from egg to adult was slightly longer for females

(50.6 days) than males (49 days) (Table 2). Adults eclosed and were active only

at night. Adult longevity for males (10.7 days) was higher (not significant) than

for females (9.6 days). Following a 2-3 day preoviposition period in the laboratory,

two groups of females were evident: fertile versus unfertile. Fertile females laid

approximately 50% of their eggs within the first four days (Fig. 2), averaging 445.7

(n = 13) eggs per female. However, dissections revealed that an average of 42.9

eggs per female remained in the reproductive system at death. Thus, the total

potential fecundity per fertile female was considered to be 488.6. Infertile females

laid a relatively constant number of eggs per female per day (Fig. 2), but the total

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Table 3. Percent survival of neonate larvae at 7 and 21 days on those plants on which any survival

was observed at 7 days.

Plant species

% survival

7 days

21 days

Baccharis halimifolia

Callistephus chinensis

Calotis cuneata

Cotula australis

Eclipta prostrata

Vittadinia sulcata

eggs deposited per female was approximately 25% (average 110.2, n = 12) of that

for fertile females. Furthermore, nearly twice as many eggs remained in the re¬

productive system at the time of death (average 228.5) as were deposited and

many unlaid eggs were undeveloped. Total potential fecundity for these females

averaged 338.7. We cannot explain why infertile females, despite being given the

same opportunity to mate, failed to deposit fertile eggs. Average longevity of

fertile and infertile females did not differ significantly.

Field Observations. — Moths which were active (readily visible in the upper strata

of the foliage) early in the evening laid predominantly fertile eggs whereas those

active later in the evening laid predominantly infertile eggs. Thirty-seven of 41

females (90.2%) collected between 20:30 and 22:00 h laid eggs prior to the fourth

day after capture. Virtually all of the eggs for each female were fertile. Thirty-

three of the 37 moths (89.2%) laid eggs prior to dawn of the following day,

indicating the moths had mated at least one day prior to capture. The remaining

Table 4. Percent survival of neonate larvae on foliage of potted plants.

Plant species

% survival

Week 1

Week 4

Week 8

Allium cepa

Arachis hypogaea

—

Aster novi-belgii

—

Baccharis halimifolia

Brachycome multifida

—

Callistephus chinensis

Calotis cuneata

—

Cassinia laevis

—

Conyza sumatrensis

—

Cosmos bipinnatus

—

Cotula australis

—

Cucurbita maxima

—

Eclipta prostrata

—

Gossypium hirsutum

—

Helichrysum bracteatum

—

Olearia nernstii

—

Vernonia cinerea

—

Vittadinia sulcata

Zinnia linearis

—

1990

EHLER ET AL.: BIOLOGY OF PROCHOERODES

Table 5. Percent survival when 3rd instar larvae were placed on bouquets of foliage.

Plant species

% survival

Week 1

Week 3

Week 6

Adenostemma lavenia

Baccharis halimifolia

100

Brachycome multifida

—

Calendula officinalis

—

Callistephus chinensis

—

Calotis cuneata

—

Carthamus tinctorius

—

Cassinia laevis

Chrysanthemum carinatum

—

Cosmos bipinnatus

—

Cotula australis

—

Dahlia variabilis

—

Eclipta prostrata

—

Emilia sonchifolia

—

Gaillardia aristata

—

Glossogyne tenuifolium

Gnaphalium sphaericum

—

Helianthus annuus

—

Helichrysum bracteatum

—

Olearia nernstii

—

Senecio lautus

—

Vernonia cinerea

—

Vittadinia sulcata

—

Wedelia biflora

—

Zinnia linearis

—

four females oviposited but none of their eggs hatched. All of the 28 females

collected between 23:00-00:10 h laid eggs, although only 12 (42.9%) laid pre¬

dominantly fertile eggs. None of the eggs of the remaining 16 females were fertile.

Males and females were caught in a one-to-one sex ratio. Apparently virgin females

are active later in the evening, presumably seeking mates; once mated, they become

active earlier in following evenings. This hypothesis, however, should be tested

further.

Host Specificity. — After seven days, neonate larvae placed on cut foliage of most

of the plants listed in Table 1 had died without any appreciable feeding. However,

surviving larvae and evidence of feeding were found on five species (Table 3). By

the end of three weeks, all larvae except one on Eclipta prostrata (L.) L. had died.

This individual remained alive for 12 more days, although it developed at a much

slower rate than control larvae on Baccharis. No larvae (other than the controls)

survived longer than 33 days, nor did any complete development.

Neonate larvae placed on potted plants survived for less than a week on all but

five species. Three of these species [Callistephus chinensis (L.) Nees Von Esenbeck,

Cotula australis (Sieber) Hooker f., and Vittadinia sulcata N. T. Burbidge] were

common to those supporting larvae for a week in the cut-foliage experiment (Table

4). On these five, there was evidence of feeding and development to at least the

second instar. On only Callistephus chinensis was the growth rate similar to that

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 66(1)

observed on B. halimifolia. The larvae on C. chinensis grew rapidly through the

first three instars but then ceased development and died three weeks later. The

increased mortality and the failure of the surviving larvae to develop indicate

that this plant is not a suitable host.

Late instar larvae could not complete their life cycle on cut foliage of any of

the test plants (Table 5). In many cases the larvae took longer than a week to die,

even if they did not feed. On only two plants, Cassinia laevis R. Brown and

Glossogyne tenuifolium Cassini, was there any protracted feeding. On these, how¬

ever, there was negligible growth of the larvae and they eventually died.

This insect exhibited an ability for early instar larvae to feed on a number of

Asteraceae, almost all in the tribe Astereae and thus has been comprehensively

tested. In the United States and Australia, it has been tested against plants in (1)

12 genera of the Astereae (approximately 10% of the genera of this tribe throughout

the world) and (2) 31 genera in other tribes of the Asteraceae (see also Palmer &

Tilden 1987). This testing against closely related plants is one of the most com¬

prehensive of any insect candidate for biological-control. All the evidence gathered

indicates that P. truxaliata is host-specific to certain Baccharis spp. In addition,

there is no record in the United States of it ever being found on any plant other

than a Baccharis spp. Because host-specificity experiments indicated that P. trux¬

aliata was able to utilize only Baccharis spp., it was considered safe to release in

Australia. Permission for release from quarantine was granted in February, 1989.

Acknowledgment

Gordon Paris undertook all the laboratory work at the Alan Fletcher Research

Station and J. Chippendale and T. Ellis assisted in the supply of test plants. We

thank J. A. DeBenedictis, A. H. Porter, and A. M. Shapiro for critical review of

the manuscript, and J. McCaskill for taxonomic assistance.

Literature Cited

Daly, H. V. 1985. Insect morphometries. Annual Rev. Entomol., 30: 415-438.

Drooz, A. T. 1965. Elm spanworm head capsule widths and instars. J. Econ. Entomol., 58: 629-

631.

Dupree, M. 1965. Observations on the life history of the lesser cornstalk borer. J. Econ. Entomol.,

58: 1156-1157.

Enders, F. 1976. Size, food-finding, and Dyar’s constant. Environ. Entomol., 5: 1-10.

Kimberling, D. N. & J. C. Miller. 1988. Effects of temperature on larval eclosion of the winter moth,

Operophtera brumata. Entomol. Exp. Appl., 47: 249-254.

Lin, S., A. C. Hodson & A. G. Richards. 1954. An analysis of threshold temperatures for the

development of Oncopeltus and Tribolium eggs. Physiol. Zool., 27: 287-311.

Long, D. B. 1953. Effects of population density on larvae of Lepidoptera. Trans. Roy. Entomol.

Soc., Lond., 104: 543-585.

McFadyen, P. J. 1985. Introduction of the gall fly Rhopalomyia californica from the USA into

Australia for the control of the weed Baccharis halimifolia. pp. 779-787. In Delfosse, E. S.

(ed.). Proceedings of the VI International Symposium on Biological Control of Weeds. Agric.

Canada, Ottawa.

Palmer, W. A. & J. W. Tilden. 1987. Host specificity and biology of Prochoerodes truxaliata (Guenee)

(Geometridae), a potential biocontrol agent for the rangeland weed Baccharis halimifolia L. in

Australia. J. Lepid. Soc., 41: 199-208.

Tilden, J. W. 1951. The insect associates of Baccharis pilularis De Candolle. Microentomology, 16:

149-188.

Received 6 July 1989; accepted 17 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 89-92, (1990)

DESCRIPTION OF THE IMMATURES OF

STYRINGOMYIA NEOCALEDONIAE ALEXANDER

(DIPTERA: TIPULIDAE) AND NOTES ON ITS BIOLOGY

C. Dennis Hynes

Department of Biological Sciences, California Polytechnic State University,

San Luis Obispo, California 94307

Abstract. — The immatures of Styringomyia neocaledoniae Alexander are described. Information

on the habitat of the last larval instar and pupa, and on the resting behavior of the adult is given.

Comments on the taxonomic placement of the genus are also given.

Key Words. — Insecta, Diptera, Tupulidae, Styringomyia neocaledoniae, habitat, behavior, clas¬

sification

Most of the information about the more than 100 species of Styringomyia

consists of the original description, location of the holotype and paratypes, and

some comparisons to other species. Edwards (1914) considered the group isolated,

possibly primitive on the basis of the strange construction of the adult form, and

suggested that it should be removed from its original placement in the Antochini

but did not indicate its placement upon removal. For this purpose, Alexander

(1920) erected a special tribe, the Styringomyini, but later placed it in the Eriop-

terini as a subtribe, Styringomyaria. I suggested recently that the genus would be

better situated in the Eriopteraria (Hynes 1987), based on the presence of the so-

called L-shaped cervical plate (Crampton 1925).

Both Edwards (1914, 1924) and Alexander (1920, 1947) reported observations

made by others as to rearing and the habitats of the larvae, as well as peculiar

behaviors of the adults. No larval specimens or descriptions accompanied these

rearings. Within the Limoniinae, the larval instars give better evidence of rela¬

tionships than do the adult. The larvae were needed to answer the question of

relationships among these genera.

Recently, I was able to collect and rear immature Styringomyia neocaledoniae

Alexander in New Caledonia, and to gather some preliminary information on

their habitats. The following description is based on observations from four larvae

and three pupae.

Taxonomy and Biology

Description.—Larva. Length 7.2 mm; dorsoventral and dextrosinistral measurements at third ab¬

dominal segment 0.5 mm. Body white-yellow, cylindrical, becoming slightly smaller at eighth abdom¬

inal segment, then abruptly larger at disk. Spiracular disk (Fig. 1) with seven fleshy lobes (three smaller

lobes on the dorsal rim, two larger lateral lobes, and two ventrolateral lobes), structured as a rectangular

ventral margin of the plate; face slightly convex; white with light brown markings at center of ven¬

trolateral arms. Spiracles with very light rims, darker centers; the medial outer margin with two small

invaginations. Head capsule length (tip of labrum to posterior end) 0.48 mm (Fig. 2); width 0.14 mm

at mandibular articulation. Labrum with basal plate projecting forward, projections heavily sclerotized,

with two ovate rows of setae extending from ventral surface (messores?). Mandible strongly rotated,

nearly vertical; six “bars” with membranes between them forming three plates; dorsal and lateral

plates membranous, sclerotized areas at junction of dorsal and dorsolateral bars. Maxillary plate not

toothed, expanded slightly at anterior end near junction with heavily sclerotized maxillary cardo.

THE PAN-PACIFIC ENTOMOLOGIST

Yol. 66(1)

2.0 mm

Figures 1-6. Styringomyia neocaledoniae. Figure 1. Spiracular disk of larva. Figure 2. Head capsule

of larva (D = dorsal, Y = ventral). Figure 3. Metathoracic breathing horn. Figure 4. Terminal segments

of pupa of male. Figure 5. Adult female at rest. Figure 6. Portion of abdomen of adult male that

extends beyond wings while at rest.

1990

HYNES: IMMATURES OF STYRINGOMYIA

Pupa.— Length 4.2 mm; dextrosinistral width at base of wing pad 1.1 mm; dorsoventral width at

base of wing pad 1.25 mm. Body yellow-white. Two conspicuous tubercles behind base of antennae

each having a long stout seta. Mesonotal breathing horns flattened dorsoventrally, short (0.14 mm),

extended laterally and slightly cephalad (Fig. 3). Mesonotum without crest of spines. Wing pads dark

in older specimens, end at posterior margin of second abdominal segment. Leg sheaths short, ending

at posterior edge of third abdominal segment; sheaths nearly same length but metathoracic slightly

shorter, prothoracic pair slightly longer. The posterior edges of abdominal segments sparsely armed

with very small spines. Male terminalia (Fig. 4) with eighth stemite possessing a double toothed

tubercle at each extreme lateral margin, each with a short, thick seta at tip. Dorsal portion of eighth

tergum with a lateral tubercle on each side, armed with one or two short spines. Dorsum of ninth

tergum with two pointed, flattened tubercles directed laterad, fleshy serrations on posterior edge.

Sheaths of dististyles smooth, conical, directed caudad at base, curving gently laterad to a point, a

short thick spine at tip.

Taxonomic Placement. — Data from the larval instars show that both Alexander

and Edwards were correct in moving Styringomyia from the Antochini to the

Eriopterini in that the genera in the former tribe have “massive” head capsules.

The larval head capsule of Styringomyia is that of typical eriopterine larvae having

the six so-called “bars” and slanted mandible. However, the degree of difference

between the larvae of Styringomyia neocaledoniae and the larvae of other Eriop-

teraria is no greater than that between the larvae of Erioptera and Molophilus.

There is no evidence that a special tribe or subtribe is necessary to indicate generic

relationships. Indeed, the “strange” differences of structuring in the adult stage

could just as easily be interpreted as indicating a more specialized condition,

rather than primitive as suggested by Edwards (1914). On the basis of the larval

data, along with the presence of the L-shaped cervical plate in the adult, Styrin¬

gomyia should be placed in the tribe Eriopterini, subtribe Eriopteraria.

Biology. — The larvae of Styringomyia neocaledoniae were found in three dif¬

ferent habitats. The first habitat is best described as a gathering of small branches

into which leaves from various trees have accumulated, forming a mat of decaying

organic material. These accumulations, located on the forest floor, are rather large,

attaining lengths of two to three meters and widths of one to two meters. They

are very probably formed when heavy rains flood an area causing runoff. Many

remain wet or damp for long periods of time. The second habitat is the organic

remains old tree trunks, especially the interior sections where large amounts of

dark brown organic material form. The material remains damp throughout ex¬

tended dry periods. The third habitat is at the base or petiole of rotting palm

fronds. This habitat resembles the bases of the North American Yucca whipplei

Torrey, the habitat for Gnophomyia comstocki Alexander, in that the rotted ma¬

terial is mainly between the epidermal layers which sandwich the lignified rods

between them. The material between the rods is brown and sticky, and remains

damp through extended dry periods. This habitat is very protective and serves

as a media for many species of eriopterine larvae. Associated with Styringomyia

neocaledoniae are Erioptera ( Meterioptera) rhaphidostyla Alexander, Elephanto-

myia ( Elephantomyia ) garrigouana Alexander, and Toxorhina (C.) caledonica

Alexander. Larval specimens of as yet unidentified species of Helius, Molophilus,

and Amphineurus were also found. The larvae were not easily collected and had

to be washed and literally scrubbed out of the sticky material. The pupae are even

more difficult to obtain. They are found in a strong case that is closely appressed

to the drier rotting areas. Within rearing cages containing humic materials, the

larvae of S. neocaledoniae form fine earthen tubes through which they traveled;

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

this behavior is similar to that of Limonia. The tubes are thin, but extend over

edges and through the spaces to other large particles of habitat.

The adults walk with short, rapid, and quite erratic movements. Both Edwards

(1924) and Alexander (1947) noted the peculiar resting position of adults which

reputedly resembles a spider web (Fig. 5). This would infer that the animal is

mimicking spiders as a matter of protection. If this infers spider mimicry, however,

I found no spiders with even a remote resemblance. If correct, however, the

terminal portion of the male sticks out from under the wings looking like a head

region from above (Fig. 6)? From this angle the dark dististyles appear as eyes,

presenting two possible “heads.” Alternatively, however, the dorsal and lateral

views give light and dark disruptive patterning. The resting position brings the

legs approximately parallel, causing the dark and light bands. The lighter wing

areas and the dorsomedial on the thorax and head enhances the disruptive col¬

oration. All other Styringomyia possess this coloration. While it is unlikely that

all Styringomyia mimic the spiders, or have a spider to mimic, all have disruptive

coloration.

Alexander (1920) also noted life history information on Styringomyia didyma

that leads one to the conclusion that it may have the shortest life cycle of any

crane-fly species; there may, however, have been no assurance that the media

contained no other larvae. His noted hatching period is given as 10-15 days,

approximating that of eriopterines generally. My data on neocaledoniae indicate

that the pupal stage takes 7 days, again normal for eriopterines. There was no

indication that S. neocaledoniae is multivoltine. I suspect Alexander’s media was

contaminated with other larvae mature enough to emerge and confuse the noted

development time; thus S. didyma should be reared again to confirm the infor¬

mation given. If Alexander’s information is true, S. didyma would be useful as a

laboratory species with which to gain information on crane-fly biology.

Acknowledgment

I thank the French Institute of Research for Development (ORSTOM), Noumea,

for help during my study in New Caledonia, and especially Jean Chazeau for his

support at the Riviere Bleue Project in New Caledonia. I also thank Gaylene

Thomas (Cal Poly, San Luis Obispo) for help in the illustrations.

Literature Cited

Alexander, C. P. 1920. The crane-flies of New York, Part II. Biology and phylogeny. Cornell Univ.

Mem., 38.

Alexander, C. 1947. Notes on the Tropical American species of Tipulidae (Diptera) III. The spe¬

cialized Eriopterini: Rhabdomastix, Cryptolabis, Erioptera, Molophilus, Styringomyia, Tox-

orhina and allies. Rev. de Entomol., 18: 317-360.

Crampton, G. C. 1925. Evidences of relationship indicated by the thoracic sclerites of certain

eriopterine tipulid Diptera. Insect. Inscit. Menst., 13: 197-213.

Edwards, F. W. 1914. A revision of the tipulid genus Styringomyia. Trans. Royal Entomol. Soc.,

London, 1914: 206-227.

Edwards, F. W. 1924. New data concerning Styringomyia (Diptera, Tipulidae). Ann. Mag. Nat.

Hist., Series 9, 13: 265-274.

Hynes, C. D. 1987. New species of of the genus Styringomyia from the South Pacific and Southeast

Asia (Tipulidae, Diptera). Pan-Pacif. Entomol., 63: 92-97.

Received 4 April 1989; accepted 29 November 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 93-95, (1990)

Scientific Note

THE SELLING OF GAMPSOCLEIS GRATIOSA BRUNNER

(ORTHOPTERA: TETTIGONIIDAE) AS SINGING PETS

IN CHINA

Insects have been more prominent in Chinese culture than in cultures of most

other countries, particularly those of the west. Insects are important as drug sources

in traditional Chinese medicine (Read, B. E. 1941. Chinese materia medica, insect

drugs, Peking Nat. History Bull., Reprinted 1982, Southern Materials Center,

Taipei, Taiwan), and have been often used for human food (Mitsuhashi, J. 1984.

Edible insects of the world. Kokinshoin, Tokyo). They have been frequent subjects

of Chinese art such as the jade crickets and other insects (Shang and Western

Chow dynasties, approximately 1600-771 BC), the “Flowers and Insects” scroll

paintings (Shou Ping, Yun Hsi school 1633-1690 AD), and the paintings of Chi

Bai Shi (20th century). Insects have long been a source of amusement including

the releasing of large quantities of fireflies for imperial entertainment (Liu, G.

1939. Psyche, 46: 23-28), the gambling sport of fighting crickets, and the keeping

of singing insects as pets (Laufer, B. 1927. Field Mus. Nat. Hist., Chicago, An-

thropol. Leaf., 22).

The more vociferous members of the famly Gryllidae and Tettigoniidae, often

known collectively as “singing crickets,” have been popular pets in China for

more than 1000 years. During the Tang dynasty, the ladies of the palace caught

singing crickets, put them in golden cages and placed them near their pillows.

This custom (minus the gold) was imitated by the common people. A special

literature on keeping crickets was created in the Sung dynasty (960-1279 AD)

and cage making developed into a palace art during the early Ching dynasty (1644—

1911 AD) (Laufer 1927; Soloman, B. J. 1984. Arts of Asia, 4: 76-87). The court

practice of keeping singing crickets, often in beautifully molded gourd cages made

from Lagenaria vulgaris Seringe, continued into this century as depicted in the

1987 American film “The Last Emperor.” Singing crickets woven into bamboo

cages were sold on the streets of China in the 1920s (Laufer 1927).

After the 1949 Revolution, the keeping of pets was considered to be a bourgeois

pastime, reminiscent of the decadent court and idle rich of the past, who were

often cricket fanciers (Wang Ren, personal communication).

In August 1987, while visiting Beijing, I encountered a seller of singing crickets

(Fig. 1 ) with several hundred woven, split bamboo cages (7 cm diameter) tied to

the back of his bicycle. Each cage contained a large (4-5 cm) green or tan male

Gampsocleis gratiosa Brunner which produced loud ringing “cries” with their

brachypterous wings. The calls and the sight of the seller attracted many people

willing to pay the 30 fen (US $0.10, Aug 1987 exchange rates) asking price. A

few weeks later in Pao-tou, Inner Mongolia Autonomous Region, I saw another

seller, with the same cricket species in identical bamboo cages, at the outdoor

market. From this man and another cricket seller in Beijing, I learned about the

marketing of G. gratiosa. The Pao-tou seller had come from Bao-ding, Hebei

Province (about two days travel by train) to sell his singing insects.

Bao-ding has a wholesale market where G. gratiosa are sold to sellers. The

collectors of G. gratiosa are from four Hebei villages (Shuang He Zhuang—Twin

THE PAN-PACIFIC ENTOMOLOGIST

Vol. 66(1)

Figure 1. Gampsocleis gratiosa, woven into bamboo cages, being sold as singing pets on a Beijing

street in August, 1987.

River, Da Long Hua—Great Dragon River, Quing Xi Ling—Western Ching Tomb,

and Qiao Jai He—Cho’s River). The collectors go out to the countryside and to

other villages to collect and buy G. gratiosa during June and July. The Pao-tou

seller, who is also a collector, said that 20-30 of the 1000 people in his village

are involved in the trade. These insects are reportedly collected mainly in hilly

areas on wild jujubes (Zizyphus sp.) and tamarisk (Tamarix sp.), with the green

ones taken on north facing slopes and the tan on south facing slopes.

The sellers pay the collectors 25 fens (US $0.08) per insect plus 6 fens (US

$0.02) for a cage, early in the season when there are few adults. Later in the season

when adults are more common, sellers pay only 7-8 fens (US $0.03) per insect.

Sellers then travel to larger, often quite distant cities, such as Beijing, Pao-tou,

Shanghai, Canton, and Lan-chou in the western part of China. They can make

the most money in Shanghai and distant cities like Lan-chou early in the season,

when G. gratiosa sell for 1-3 yuan each. Three yuan is approximately US $1.00,

and equal to half a day’s pay for many Chinese workers (Wang Ren, personal

communication). The retail price drops to approximately 30 fens (US $0.10), later

in the season as G. gratiosa becomes more common. Even this amount presumably

permits the seller to travel to a distant city, pay for lodging and food, and still

have enough money left to return home with a good profit.

Gampsocleis gratiosa is occasionally sold at the farmer’s market in Guan-yuan

Park, Beijing, during the winter (Ye Zheng Chu, personal communication). These

1990

SCIENTIFIC NOTE

insects are produced in small quantities by artificial rearing and are consequently

very expensive. In winter 1988, a male G. gratiosa sold for 50 yuan (US $ 16.70)—

more than one-third of an ordinary worker’s monthly salary. The singing crickets

are frequently kept in gourd cages fitted with a wire ring that prevents their escape

during feeding and vibrates to enhance the insect’s song. Owners often place these

cages in their undershirt pockets to keep the insects warm. Under such conditions

the G. gratiosa sing “happy songs” bringing joy and mental contentment to their

owners.

The popularity of G. gratiosa may be due to its large size and lack of shyness

in addition to its ringing song. They readily protrude their impressively large

mandibles through the weave or wire rings of their cages to take slices of apple,

cabbage, cucumber, green onions or other vegetables offered. They can put their

legs through the spherical bamboo cages and walk and roll the cages along; a

behavior which causes the cages to be hung. The katydids will live from several

weeks to months in captivity, depending on their age and the quality of care.

Gampsocleis gratiosa may also be a symbol of prosperity, as is a very similar

looking (perhaps the same) tettigoniid in southern China. In Hong Kong, one

sees carved ivory (or imitations) of elongate Chinese cabbages with the “ Gamp¬

socleis ” perched on top, sold for a good luck or prosperity symbol.

The marketing of G. gratiosa is an indication that China’s ancient interests in

singing insects still persist in this modem era. It is also an unusual, but culturally

relevant product, of China’s newly permitted free market economics.

Acknowledgment. — I thank Wang Ren (Biological Control Laboratory, Chinese

Academy of Agricultural Sciences, Beijing) for helping me interview and under¬

stand the cricket sellers; Ye Zheng Chu (same institute) for providing the infor¬

mation on the selling and keeping of G. gratiosa in the winter; and the anonymous

cricket sellers willing to share the details of their fascinating trade. I also appreciate

the manuscript reviews by Gene Defoliart (University of Wisconsin, Madison),

Wang Ren, Ye Zheng Chu and Natalia Vandenberg.

Robert W. Pemberton, Asian Parasite Laboratory, U.S. Department of Agri¬

culture, Agricultural Research Service, Seoul, Korea (% American Embassy, APO

San Francisco, California 96301.

Received 19 July 1989; accepted 10 October 1989.

PAN-PACIFIC ENTOMOLOGIST

66(1): 96, (1990)

EDITORIAL NOTICE:

CHANGES IN JOURNAL FORMAT AND

EDITORIAL PROTOCOL

Beginning with volume 66(1), The Pan-Pacific Entomologist has changed format

to provide a more attractive publication. With this format change, the journal is

also modifying the requirements for submitted manuscripts to enhance consis¬

tency. A detailed Information for Contributors is provided (66 [1]: 1-8) for the

convenience of prospective authors. Among these new requirements, note that

taxonomic manuscripts that describe new taxa must have separate paragraphs

accompanying each description detailing (a) information on the types, and (b) a

diagnosis; furthermore, a listing of the material examined, where appropriate,

must be presented in a separate paragraph using a new standard format for data.

Taxonomic keys are optional, but if present are subject to guidelines including

reversibility (“back-tracking”) provisions.

Also beginning with volume 66 (1), the editorial handling of manuscripts will

follow a newly defined protocol to upgrade the quality of the journal. Upon receipt,

manuscripts will be reviewed by two peer reviewers and accepted or rejected on

the basis of their recommendations. Review of manuscripts will be carried out

by the journal, regardless of whether the manuscript has been previously reviewed

before submission. When the assigned reviewers disagree on the acceptability of

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of suitability. The editor, however, reserves the right to initially reject those

manuscripts that are poorly written or deviate strongly from the scope or format

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may also be required to incorporate certain changes suggested by the reviewers,

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authors should maintain a file copy of the manuscript on a computer or word-

processor until formal acceptance is granted.

Alterations to, or disagreements with, either the edited copy or the required

addressing of reviewer’s concerns as noted by the editor must have a statement

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author’s preference is deemed to be less suitable than the edited version, despite

the stated rationales, the case will be subject to independent arbitration by a third

party with taxonomic, biological and editing experience; these arbitrators will be

chosen by the editor and their ruling will be binding before the final acceptance

of the manuscript for publication.

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¬

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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

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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

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).

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FOR DIAGNOSES, TYPES AND MATERIAL EXAMINED (INCLUDING A SPECIFIC FORMAT). List the unabbreviated

taxonomic author of each species after its first mention.

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.

Illustrations. — Illustrations must be of high quality and large enough to ultimately reduce to 117 x 181 mm while maintaining label

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Mount all illustrations. Label illustrations on the back noting: (1) figure number, (2) direction of top, (3) author’s name, (4) title of

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Volume 66

THE PAN-PACIFIC ENTOMOLOGIST

January 1990

Number 1

Contents

The Pan-Pacific Entomologist: format information for contributors. 1

PARKER, F. D. & V. J. TEPEDINO—Bee pollination of Cuphea (Lythraceae) species in

greenhouse and field... 9

BOUSQUET, Y. & H. GOULET—Description of a new species of Metrius (Coleoptera: Ca-

rabidae: Paussini) from Idaho with comments on the taxonomic status of the other taxa

of the genus.... 13

O’NEILL, K. M. —Female nesting behavior and male territoriality in Aphilanthops subfrigidus

Dunning (Hymenoptera: Sphecidae). 19

GOEDEN, R. D.—Life history of Eutreta diana (Osten Sacken) on Artemisia tridentata Nuttall

in southern California (Diptera: Tephritidae). 24

GOEDEN, R. D.—Life history of Eutreta simplex Thomas on Artemisia ludoviciana Nuttall

in southern California (Diptera: Tephritidae)___ 33

GUENTHER, S. J. & W. CHAPCO—A morphometric analysis of Melanoplus females (Or-

thoptera: Acrididae). 39

SORENSEN, J. T., R. J. GILL, R. V. DOWELL & R. W. GARRISON-The introduction of

Siphoninus phillyreae (Haliday) (Homoptera: Aleyrodidae) into North America: niche

competition, evolution of host plant acceptance, and a prediction of its potential range

in the Nearctic_ 43

HARRIS, A. C.—Podagritus cora (Cameron) and P. albipes (F. Smith) (Hymenoptera: Spheci¬

dae: Crabroninae) preying on Ephemeroptera and Trichoptera_ 55

DOWELL, R. V. & B. STEINBERG—Influence of host plant on fecundity of Aleurocanthus

woglumi Ashby (Homoptera: Aleyrodidae). 62

KAMM, J. A. —Biological observations of glassy cutworm (Lepidoptera: Noctuidae) in western

Oregon. 66

MARTIN, W. F. & R. F. MARTIN—Reproduction of the sand wasps Stictia signata (L.) and

Bicyrtes variegata (Olivier) (Hymenoptera: Sphecidae) on the Caribbean coast of Quin¬

tana Roo, Mexico. 71

EHLER, L. E., M. G. KINSEY & W. A. PALMER—Further observations on the biology and

host specificity of Prochoerodes truxaliata (Guenee) (Lepidoptera: Geometridae), a

biological-control agent for Baccharis halimifolia L. in Australia. 79

HYNES, C. D. —Description of the immatures of Styringomyia neocaledoniae Alexander (Dip¬

tera: Tipulidae) and notes on its biology_ 89

SCIENTIFIC NOTE

PEMBERTON, R. W.—The selling of Gampsocleis gratiosa Brunner (Orthoptera: Tettigoniidae)

as singing pets in China. 93

Editorial notice: changes in journal format and editorial protocol. 96

The

PAN-PACIFIC

ENTOMOLOGIST

Volume 66 April 1990 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

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