Vol. 58

JANUARY 1982

No. 1

THE

Pan-Pacific Entomologist

In Memoriam

PHILIP HUNTER TIMBERLAKE

1883-1981

HURD, LINSLEY and HALL-Philip Hunter Timberlake, 1883-1981. 2

BENBOW and FOSTER—Biology of Eutreta diana Osten Sacken on Sand Sagebrush

Artemisia filifolia Torr. (Diptera: Tephritidae)... 19

WESTCOTT—Differentiating Adults of Apple Maggot, Rhagoletis pomonella (Walsh)

from Snowberry Maggot, R. zephyria Snow (Diptera: Tephritidae) in Oregon. 25

JOHNSON—A New Genus and Species of California Pill Beetle (Coleoptera: Byr-

rhidae). 31

CLEMENT and MILLER—Insect Seed Predation on Astragalus bisulcatus (Hook.) Gray

(Leguminosae). 38

YOUNG—Key to the Pupal Parasites of California Oakworm, Phryganidia californica

(Lepidoptera: Dioptidae), Based on Larval Exuviae. 42

PINTO, O ATM AN and PLATNER— Trichogramma australicum Girault (Hymenoptera:

Trichogrammatidae): Redescription and Lectotype Designation. 48

KULHAYY, SCHWANDT and HOBBS-Understory Plants as Indicators of Host Trees

of the Wounded Tree Beetle, Nosodendron californicum, in Northern Idaho (Co¬

leoptera: Nosodendridae). 53

OLKOWSKI, OLKOWSKI, van den BOSCH, HOM, ZUPARKO and KLITZ-The

Parasitoid Trioxys tenuicaudus Stary (Hymenoptera: Aphidiidae) Established on

the Elm Aphid Tinocallis platani Kaltenbach (Homoptera: Aphididae) in Berkeley,

California. 59

BARNARD—Flight Periodicity in Colorado Biting Midges (Diptera: Ceratopogonidae).. 64

KAMM — Protagrotis obscura Barnes and McDunnough (Lepidoptera: Noctuidae): A Pest

of Grasses Grown for Seed in the Pacific Northwest. 73

SMITH and OLSON —Confused Flour Beetle and Other Coleoptera in Stored Mari¬

juana. 79

SAN FRANCISCO, CALIFORNIA • 1982

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

rnnnorntinn with THF C AT TFORNTA A C' A T\I-'A TV 017 CrTT7\Tn?C

The Pan-Pacific Entomologist

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G. A. Marsh, Editor

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Pacific Coast Entomological Society

OFFICERS FOR 1982

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Title of Publication: The Pan-Pacific Entomologist.

Frequency of Issue: Quarterly (January, April, July, October).

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This issue mailed 27 May 1983

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The Pan-Pacific Entomologist (ISSN 0031-0603)

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Philip Hunter Timberlake, 1883-1981

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 2-18

Philip Hunter Timberlake

1883-1981

Philip Hunter Timberlake, Associate Entomologist Emeritus, University

of California, Riverside died April 17, 1981, at the age of 97. He was born

June 5, 1883 at Bethel, Maine to Isadore Margaret (nee Billings, 1850-1897)

and Davis True Timberlake (1844-1928) who were married on July 6, 1873.

Of the seven children, Philip H. Timberlake was the fifth born and the

second son of the family. His father was a school teacher who taught Latin

and Greek at the Lancaster Academy, Lancaster, New Hampshire.

Upon graduation from Bowdoin College, Brunswick, Maine, Mr. Tim¬

berlake was awarded an A.B. in 1908 having matriculated in the liberal arts.

Following graduation he entered Harvard University in 1909 and in 1910

was awarded an A.M. in biology. His first career employment was as an

Assistant Agent and Expert in the Gypsy Moth Laboratory, U.S.D.A., Mel¬

rose Highlands, Massachusetts between 1909-1910. During this time he de¬

veloped a sensitivity to moth scales and was transferred to a U.S.D.A.

laboratory, Bureau of Entomology at Whittier, California where he com¬

menced a study of beneficial insects as Agent and Expert in 1910-1912.

Later in 1912 he was transferred to a U.S.D.A. laboratory, Bureau of Ento¬

mology at Salt Lake City, Utah where he served until 1914 as Agent and

Expert investigating alfalfa weevil parasites. From 1914 until 1924 he was

stationed with the Hawaiian Sugar Planters Experiment Station in Honolulu,

Hawaii as an Associate Entomologist concerned with the biological control

of sugar cane insects. He also served as Curator of the collection and pub¬

lished a list of the types contained therein (1922, Proc. Hawaiian Entomol.

Soc., 5:174-177). It was during this period that Mr. Timberlake married

Edith Milhous in Otto Swezey’s home in Honolulu on November 26, 1917.

Mrs. Timberlake had been a secretary at the University of California, Berke¬

ley prior to moving to Whittier, California. Three children were born, all in

Hawaii, and now all reside in California: Elizabeth Paldanius in Riverside;

Philip Franklin Timberlake, M.D. in Newport Beach; and Priscilla MacLeod

in San Anselmo. Mrs. Timberlake passed away in July of 1972 in Riverside.

She was born in Jennings County, Indiana and her family had come to

California in 1896. Mrs. Timberlake was an aunt of former President Richard

Milhous Nixon.

By the early 1920’s Mr. Timberlake’s research and publications on ben¬

eficial insects, particularly on chalcidoid wasps and ladybird beetles, at¬

tracted international attention, and in 1924 he was offered and accepted a

position as Associate Entomologist in the biological control program of the

University of California at Riverside (then known as the Citrus Experiment

Station). As his appended bibliography reveals, Mr. Timberlake continued

research on the beneficially important parasitic Hymenoptera after assuming

his new duties, but within a few years he began to shift more and more of

PAN-PACIFIC ENTOMOLOGIST

his research effort to a taxonomic study of the bees, especially those of

southern California and the adjacent desert areas of the southwestern United

States. During this early period of transition in jobs, he began an intensive

program of collecting insects generally on the campus at Riverside. As time

and opportunity permitted he made numerous collecting trips especially in

southern California including the deserts. His collecting and research inter¬

ests were greatly influenced by T. D. A. Cockerell who identified much of

his early collection of bees and encouraged Mr. Timberlake to continue his

collecting of them. As time elapsed and much new material was collected

and identified, Cockerell deposited on loan with Mr. Timberlake a very

sizeable collection of his North American bee types. These types, eventually

to be deposited on the completion of Mr. Timberlake’s studies in the col¬

lections of the California Academy of Sciences, San Francisco, and Cock¬

erell’s continued interest and encouragement resulted in Mr. Timberlake’s

nearly complete concentration on the taxonomic study of bees, notably

those of western North America. This is clearly revealed in Mr. Timber¬

lake’s bibliography starting with item 33 and continuing with few exceptions

for the balance of his publications. The genus Perdita had fascinated Cock¬

erell and as Mr. Timberlake’s field efforts intensified and his collection grew

so did his own resolve to make known the taxonomy of this genus. Prior to

World War II Mr. Timberlake had completed a large manuscript on the

taxonomy of the genus Perdita, but he could not find a publisher. This

situation, however, changed after World War II and with his retirement in

1950 he was inundated with newly made collections of this genus from

virtually all parts of North America extending well south into Mexico. He

developed a plan to publish the work in a series of parts and the project

gained support for publication in the University of California Publications

in Entomology. After the published sections began to appear, his project

received additional support in the form of three grants from the National

Science Foundation. In all, seven parts of this revisional study of the genus

Perdita were published between 1954 and 1968 (see bibliography items 77,

82, 87, 88, 96, 98, and 101) plus two supplemental parts were subsequently

issued (bibliography items 105 and 114) along with several shorter papers

describing new species. At the beginning of his project on these bees about

180 species had been described from North America. When the Catalog of

Hymenoptera in America north of Mexico was published in 1979 more than

500 species were listed and this does not take into consideration the great

number of species described by Mr. Timberlake from Mexico. The study of

this genus of bees not only has contributed importantly to our understanding

of their taxonomy, but also has made known much about their floral rela¬

tionships and laid the foundation for a thoroughgoing analysis of their in¬

trafloral ecology and biology.

A glance through volumes 1 and 2 of the Catalog of Hymenoptera in

America north of Mexico reveals the breadth as well as an idea of the

VOLUME 58, NUMBER 1

number of the contributions that he made to the taxonomy of this order.

While much of his work on the North American bee fauna was concentrated

on the genus Perdita, it is abundantly clear from his bibliography that he

made major contributions to most of the recognized families of the Apoidea.

Although it is well known that Mr. Timberlake described a large number

of new taxa of Hymenoptera (genera, subgenera, species and subspecies)

from America north of Mexico, the exact number of these through 1976 was

unknown until recently because of the complexities in individually counting

them and then comparing these with those of other authors. On the basis

of the recently published Catalog of Hymenoptera in America north of Mex¬

ico (1979) this became possible owing to the computerized data base of that

catalog. Listed below according to decreasing total numbers of new taxa

proposed by authors, one finds that Mr. Timberlake ranks sixth among the

authors who have described the largest numbers of new taxa from America

north of Mexico.

New Taxa of Hymenoptera Described in America North of Mexico

Author

Volume 1

Volume 2

Total

1. Ashmead

2593

263

2856

2. Cresson

1293

951

2244

3. Cockerell

1882

1919

4. Viereck

919

436

1355

5. Provancher

948

159

1107

6. Timberlake

744

819

7. Girault

804

—

804

Additional new taxa of bees were subsequently proposed and described

by Mr. Timberlake after the cut-off dates of the above cited catalog so that

the total numbers of such taxa for him exceed that listed above. Thus, for

example, while the cut-off date in the catalog for the Apoidea is through

1976, Mr. Timberlake published six additional papers on bees in which many

new species were described (see items 109-114 in Timberlake bibliography).

These new taxa appreciably increase his total number described from Amer¬

ica north of Mexico. It also should be remembered that he described a large

number of new taxa from Mexico, especially pertaining to the genus Perdita.

It seems quite possible that the total number of new taxa described by him

during his lifetime exceeds 1000 for the Hymenoptera alone.

In the process of his own studies Mr. Timberlake accepted miscellaneous

collections of bees from most of North America for identification. As a

consequence of this practice, he made a major contribution to the curation

of North American collections by placing in order nearly all groups of bees.

The impact of this service to the scientific community extends into the

literature where Mr. Timberlake is acknowledged repeatedly for his help in

PAN-PACIFIC ENTOMOLOGIST

identification. Moreover, he freely loaned material that he had collected to

others for their studies of innumerable groups of bees and other insects. In

a very real sense he has had a lasting influence on the development of

knowledge pertaining to the North American Hymenoptera in general, and

to the Apoidea in particular.

Mr. Timberlake maintained membership in the following societies: Amer¬

ican Entomological Society, Cambridge Entomological Club, Entomological

Society of Washington, Hawaiian Entomological Society, Pacific Coast En¬

tomological Society.

Mr. Timberlake was an ardent and quietly enthusiastic field researcher.

He was thorough and meticulous in keeping and preserving his specimens

and the data associated with them. At almost every opportunity he was

ready to venture forth into the field, either by himself as he did for years

almost daily during the lunch hour on the Riverside Campus, or with others

on all day or on trips of longer duration. Aside from those trips made with

colleagues on the Riverside staff, he participated in a great number of field

trips to the deserts of California, as well as to other locations, in the com¬

pany of others like T. D. A. Cockerell, C. M. Dammers, Paul D. Hurd, Jr.,

Edmund C. Jaeger, E. Gorton Linsley, H. L. MacKenzie, Charles D. Mich-

ener, George A. Salt, and Pedro Wygodzinsky. He was an intellectually

stimulating companion and decidedly knowledgeable about the biota in which

collecting was being pursued. He reasoned that since insects were depen¬

dent directly or indirectly on the flora for their livelihood that anyone study¬

ing them should acquire as thorough a botanical knowledge as possible.

Observing him in the field revealed that this intimate knowledge of the biota

guided him in his collecting efforts. Thus, for example, Cockerell a few

months after Mr. Timberlake returned from one of his trips to the desert

wrote: “Last March [1927] Mr. Timberlake went out into the Colorado

Desert and brought back a series of remarkable new bees, including the

species now described, one a new genus (. Xeralictus ), the other a new sub¬

genus or genus ( Megandrena ) according to the point of view.” For each of

these new taxa the reader finds, as he nearly always will, that the specimens

are labeled as to the flowers from which they were collected. It is a hallmark

of the insect collection made by P. H. Timberlake that the material is not

only carefully prepared, but is labeled in a thoroughly informative manner.

These high data standards have provided the most comprehensive assem¬

blage of southwestern United States bees in existence and consequently the

collection is especially valuable not only because of the completeness of

data, but also because the material represents one of the most important

baseline collections of these and other insects from southern California and

adjacent areas. His field work and careful records of the species of bees

found at Riverside has contributed immensely to our knowledge of a local

fauna of bees, as well as to their relationships with the flora. Fieldwork

continued to be one of the most important aspects of Mr. Timberlake’s

VOLUME 58, NUMBER 1

research and only near the end of his life was he forced to abandon it,

primarily he repeatedly said, “because of my failing eye sight.”

In the mid-1930’s, Linsley was assigned temporarily to teach introductory

entomology at the University of California at Los Angeles and to build up

a collection of insects, primarily for use in teaching. Among the insects

assembled were a large number of endemic species of bees. His supervisor,

Professor A. M. Boyce, suggested that Timberlake might help with their

identification. They were taken to Riverside where Timberlake not only

named the bees but inspired Linsley’s interest in their taxonomy and biol¬

ogy, especially their flower relationships. Thus began a friendship, nurtured

by numerous trips to the desert and southern California mountains in search

of bees, which was sustained throughout Timberlake’s life and usually in¬

volved one or two visits to Riverside each year and much correspondence.

Previously he had given similar help and encouragement to Charles D. Mich-

ener when he was a student in high school and beginning to develop his

life-long interest in bees.

Timberlake enjoyed having students along with him in the field and be¬

cause of his considerable knowledge of the biota, the students also profited

by this association. Thus during the years of 1952-1959, 1961-1962, he par¬

ticipated in 10 of the annual spring collecting trips to the deserts and adjacent

areas of California. These field parties, led by Paul D. Hurd, Jr., were made

on behalf of the California Insect Survey, University of California, Berke¬

ley. They were camping trips of about two weeks duration and were de¬

signed to survey the insect fauna of the areas visited, as well as to train

students specializing in systematic entomology in the techniques of insect

collecting, preservation, and related operations under field conditions. Some

of the field sites have since become well known type localities and include

such place names in California as: Borego (currently Borrego), Box Canyon,

Chuchupate Ranger Station, Cronese, Descanso Ranger Station, Goldstone

Lake, Hopkins Well, Ivanpah, Kramer Hills, Plaster City, Pozo Ranger

Station, and Surprise Canyon. It was during these trips that Mr. Timberlake

shared his knowledge of insects and the California flora with students ma¬

joring in systematic entomology at Berkeley. Among these students were

Bernard J. Adelson, John K. Drew, Robert L. Langston, Evert E. Lind¬

quist, Gordon A. Marsh, Charles W. O’Brien, Jerry A. Powell, Jack R.

Powers, Jerome G. Rozen, Jr., Robert O. Schuster, Gerald I. Stage, Wallace

A. Steffan, Robbin W. Thorp, Catherine A. Toschi (now Tauber), and Mar¬

ius S. Wasbauer. During this decade of field activities on these spring trips

Mr. Timberlake went from 69 to 79 years of age and some of the students

commented that he was in better physical condition than they.

No matter how trying or difficult the conditions became in the field he

always maintained a good sense of humor. However, he had little patience.

If for example, someone got a vehicle stuck in the sand, he would simply

leave the scene, net in hand and collect until the vehicle was freed. Once

PAN-PACIFIC ENTOMOLOGIST

the center pole of a tent in which he and others were sleeping snapped due

to high winds and collapsed over everyone. Almost casually, he gently shook

the companion sleeping nearest him and suggested that it might be a good

idea if everyone abandon the tent before they suffocated. Only once was

he heard to utter an intemperate epithet. He observed that one of his favorite

collecting sites in Palm Springs was converted into a drive-in movie. One

of the students suggested that that was progress and to that Mr. Timberlake

quickly replied, “Progress HELL!”

In the fall of 1961 (September) a two week field trip was made by Mr.

Timberlake and Paul Hurd from Riverside, California to New Mexico with

the idea of collecting bees at as many localities as possible, but with special

emphasis on localities in southern New Mexico where T. D. A. Cockerell

had collected bees while on the staff of the University of New Mexico before

the turn of the century. The planned field work was particularly concerned

with sampling bees at fall flowering plants in order to obtain as many as

possible of the bee species described from the region by Cockerell. Inevit¬

ably the trip became known as: “Following in the Footsteps of T. D. A.

Cockerell.” During the trip a total of 45 collecting stops were made with

the majority of the sites being in New Mexico (28), fewer in Arizona (17),

and only one in southern California (Desert Center). Nearly 10,000 speci¬

mens were collected on that trip and the bulk of them from areas previously

collected by Cockerell. Mr. Timberlake, then 78, was especially pleased

with the results. In regard to Perdita he subsequently wrote (October 12,

1961) that on Baileya pleniradiata at 3 miles west of Bingham, Socorro

County, New Mexico: “I noted about 8 species of Perdita that evening

(September 12), but on mounting the refuse material you handed me from

that day or the next, I found about 15 species, and there may be more when

the whole material is studied.” The collecting results from this trip pertain¬

ing to Perdita are chronicled by Timberlake in his publications on that genus

(see especially items 97, 100, 104).

Among the many collecting trips Mr. Timberlake made after his retire¬

ment in 1950 with other members of the staff of his department, most were

made in the company of R. C. Dickson or Jack C. Hall, both exceptionally

outstanding collectors. It was also during this time that Jack Hall shared

Mr. Timberlake’s office and performed innumerable kindnesses by extend¬

ing much assistance both personal and professional that was so helpful to

Mr. Timberlake in his day to day research work. It was largely because of

this deeply appreciated support, as well as that from the Division of Bio¬

logical Control that Mr. Timberlake was able to maintain his “8:00 am to

5:00 pm” working day schedule almost to the end of his life.

Some Genera and Species Named in Honor of P. H. Timberlake

ORTHOPTERA

Mohavacris timberlakei Rehn (Tanaoceridae)

VOLUME 58, NUMBER 1

HEMIPTERA

Aleurotithius timberlakei Quaintance and Baker (Aleyrodidae)

Dysmicoccus timberlakei Cockerell (Pseudococcidae)

COLEOPTERA

Hippodamia tibialis timberlakei Capra (Coccinellidae)

LEPIDOPTERA

Ethmia timberlakei Powell (Ethmiidae)

DIPTERA

Aphoebantus timberlakei Melander (Bombyliidae)

Apolysis timberlakei Melander (Bombyliidae)

Brevitricliia timberlakei Kelsey (Scenopinidae)

Cophura timberlakei Wilcox (Asilidae)

Dionaea timberlakei Walton (Tachinidae)

Epidideicus timberlakei Hall (Bombyliidae)

Itolia timberlakei Wilcox (Asilidae)

Nannocyrtopogon timberlakei Wilcox and Martin (Asilidae)

Phytophaga timberlakei Felt (Cecidomyiidae)

Stenopogon timberlakei Bromley (Asilidae)

Tipula timberlakei Alexander (Tipulidae)

Urophora timberlakei Blanc and Foote (Tephritidae)

Volucella timberlakei Curran (Syrphidae)

HYMENOPTERA

Ammoplanops timberlakei Pate (Pemphredonidae)

Ancylandrena timberlakei Zavortink (Andrenidae)

Andrena timberlakei Cockerell (Andrenidae)

Ashmeadiella timberlakei Michener (Megachilidae)

Belomicrus timberlakei Pate (Crabronidae)

Brachycistis timberlakei Wasbauer (Tiphiidae)

Calliopsis timberlakei Shinn (Andrenidae)

Ceratina timberlakei Daly (Anthophoridae)

Coccophagus timberlakei Compere (Encyrtidae)

Colletes timberlakei Stephen (Colletidae)

Conostigmus timberlakei Kamal (Megaspilidae)

Dioxys pomonae timberlakei Hurd (Megachilidae)

Dufourea timberlakei G. E. Bohart (Halictidae)

Euparagia timberlakei R. M. Bohart (Masaridae)

Gnathopasites timberlakei Linsley (Anthophoridae)

Heriades timberlakei Michener (Megachilidae)

Heteranthidium subtimberlakei Schwarz (Megachilidae)

Heteranthidium timberlakei Schwarz (Megachilidae)

PAN-PACIFIC ENTOMOLOGIST

Hylaeus timberlakei Snelling (Colletidae)

Hypomiscophus timberlakei Bridwell (Larridae)

Melissodes timberlakei Cockerell (Anthophoridae)

Nomadopsis timberlakei Rozen (Andrenidae)

Nysson timberlakei R. M. Bohart (Nyssonidae)

Osmia timberlakei Cockerell (Megachilidae)

Oxybelus timberlakei R. M. Bohart and Schlinger (Crabronidae)

Peponapis timberlakei Hurd and Linsley (Anthophoridae)

Perdita timberlakei Cockerell (Andrenidae)

Plenoculus timberlakei Williams (Larridae)

Pseudopanurgus nebrascensis timberlakei Michener (Andrenidae)

Pseudopanurgus timberlakei Cockerell (Andrenidae)

Pterocheilus timberlakei R. M. Bohart (Eumenidae)

Solierella timberlakei Williams (Larridae)

Timberlakena Pate (Pemphredonidae)

Timberlakia Mercet (Encyrtidae)

Triepeolus timberlakei Cockerell (Anthophoridae)

Triopasites timberlakei Linsley (Anthophoridae)

Trypoxylon timberlakei Sandhouse (Larridae)

Xenosphex timberlakei Williams (Mellinidae)

Xeralictus timberlakei Cockerell (Halictidae)

BIBLIOGRAPHY OF PHILIP HUNTER TIMBERLAKE

1. 1910. Observations on the early stages of two aphidiine parasites of

aphids. Psyche, 17(4): 125-130, 2 figs. August 1910.

2. 1912. Technical results from the Gipsy Moth Parasite Laboratory.

V. Experimental parasitism: A study of the biology of Lim-

nerium validum (Cresson). U.S. Dep. Agric., Bur. Entomol.,

Tech. Ser., 19(5):71-92, figs. 32-41. May 29, 1912.

3. 1913. Preliminary report on the parasites of Coccus hesperidum in

California. J. Econ. Entomol., 6(3):293-303. June 1913.

4. 1916a. Note on an interesting case of two generations of a parasite

reared from the same individual host. Can. Entomol., 48(3):

89-91. March 14, 1916.

5. 1916b. Revision of the parasitic hymenopterous insects of the genus

Aphycus Mayr, with notice of some related genera. Proc. U.S.

Nat. Mus., 50(2136):561-640, pis. 26-31 (with 54 figs.). May 31,

1916.

6. 1918a. New genera and species of Encyrtinae from California para¬

sitic in mealybugs (Hymenoptera). Univ. Calif. Publ. Entomol.,

1(8):347—367, 7 figs. March 28, 1918.

7. 1918b. Notes on some of the immigrant parasitic Hymenoptera of the

Hawaiian Islands. Proc. Hawaii. Entomol. Soc., 3(5):399-404.

April 1918.

VOLUME 58, NUMBER 1

8. 1919a. Revision of the parasitic chalcidoid flies of the genera Homalo-

tylus Mayr and Isodromus Howard, with descriptions of two

closely related genera. Proc. U.S. Nat. Mus., 56(2293): 133-

194, pis. 38-41 (with 19 figs.). August 29, 1919.

9. 1919b. Notes on the North American species of Hippodamia (Coleop-

tera). J. N. Y. Entomol. Soc., 27(3/4): 162-174. June-Septem-

ber 1919.

10. 1919c. Observations on the sources of Hawaiian Encyrtidae (Hymen-

optera). Proc. Hawaii. Entomol. Soc., 4(1): 183-196. July 15,

1919.

11. 1919d. Descriptions of new genera and species of Hawaiian Encyrtidae

(Hymenoptera). Proc. Hawaii. Entomol. Soc., 4(1): 197-231.

July 15, 1919.

12. 1920a. Notes on the immigrant Hawaiian species of Ichneumonini or

Pimplini of authors (Hymenoptera). Proc. Hawaii. Entomol.

Soc., 4(2):266-275. June 1920.

13. 1920b. Descriptions of new genera and species of Hawaiian Encyrtidae

(Hymenoptera), II. Proc. Hawaii. Entomol. Soc., 4(2):409-437,

23 figs. June 1920.

14. 1921a. Notes on the Hawaiian bees of the genus Megachile (Hymen¬

optera). Proc. Hawaii. Entomol. Soc., 4(3):551-557. September

1921.

15. 1921b. Description of a new species Ootetrastichus from Formosa

(Hymen., Chalcid.). Proc. Hawaii. Entomol. Soc., 4(3):557-

564, 5 figs. September 1921.

16. 1922a. Identity of the Hawaiian carpenter bee of the genus Xylocopa

(Hymenoptera). Proc. Hawaii. Entomol. Soc., 5(1):51. October

1922.

17. 1922b. Observations on the phenomena of heredity in the ladybeetle,

Coelophora inaequalis (Fabricius). Proc. Hawaii. Entomol.

Soc., 5(1):121-133, 4 figs. October 1922.

18. 1922c. Descriptions of new genera and species of Hawaiian Encyrtidae

(Hymenoptera), III. Proc. Hawaii. Entomol. Soc., 5(1): 135—

167, 17 figs. October 1922.

19. 1922d. Notes on the identity and habits of Blepyrus insularis Cameron

(Hymenoptera, Chalcidoidea). Proc. Hawaii. Entomol. Soc.,

5(1):167-173, 2 figs. October 1922.

20. 1922e. A revision of the chalcid-flies of the encyrtid genus Chryso-

platycerus. Proc. U.S. Nat. Mus., 61(2): 1—10, pi. 1 (with 2

figs.). April 25, 1922.

21. 1923a. Review of Mercet’s work on the Encyrtidae (Hymenoptera)

of the Iberian Peninsula. Proc. Entomol. Soc. Wash., 25(3):

57-60. March 31, 1923.

22. 1923b. Descriptions of two new species of Encyrtidae from Mexico

PAN-PACIFIC ENTOMOLOGIST

23. 1924a.

24. 1924b.

25. 1924c.

26. 1924d.

27. 1925.

28. 1926a.

29. 1926b.

30. 1926c.

31. 1927a.

32. 1927b.

33. 1928a.

34. 1928b.

35. 1928c.

36. 1929a.

37. 1929b.

reared from mealy-bugs (Hym., Chalcidoidea). Proc. Hawaii.

Entomol. Soc., 5(2):323—333, pi. 11 (with 9 figs.), 1 fig. Sep¬

tember 1923.

Descriptions of new chalcid-flies from Hawaii and Mexico

(Hymenoptera). Proc. Hawaii. Entomol. Soc., 5(3):395—417,

8 figs. December 1924.

Records of the introduced and immigrant chalcid-flies of the

Hawaiian Islands (Hymenoptera). Proc. Hawaii. Entomol.

Soc., 5(3):418-449. December 1924.

Notes on Hawaiian Aphidae, with a list of food plants (Homop-

tera). Proc. Hawaii. Entomol. Soc., 5(3):450—460. December

1924.

The parasites of Pseudococcus maritimus (Ehrhorn) in Cali¬

fornia (Hymenoptera, Chalcidoidea). Part I. Taxonomic stud¬

ies. Univ. Calif. Publ. Entomol., (2):223-251, 8 figs. Septem¬

ber 16, 1924.

Description of new chalcid-flies from Panama and Hawaii

(Hymenoptera). Proc. Hawaii. Entomol. Soc., 6(1): 173-193,

pi. 9 (with 8 figs.). August 1925.

Miscellaneous new chalcid-flies of the hymenopterous family

Encyrtidae. Proc. U.S. Nat. Mus., 69(3): 1-34, pis. 1-2 (with

24 figs.). June 12, 1926.

New species of Hawaiian chalcid-flies (Hymenoptera). Proc.

Hawaii. Entomol. Soc., 6(2):305-321, pi. 10 (with 4 figs.).

July 1926.

Hymenoptera. [Insects of Hawaii, Johnston Island and Wake

Island]. Bull. Bernice P. Bishop Mus., 31:17—43, 6 figs. 1926.

New species of Hawaiian chalcid-flies (Hymenoptera)—II.

Proc. Hawaii. Entomol. Soc., 6(3):517—528, pi. 18 (with 6 figs.).

October 1927.

Biological control of insect pests in the Hawaiian Islands.

Proc. Hawaii. Entomol. Soc., 6(3):529-556, pis. 19-23, 5 figs.

October 1927.

Bees of the genus Perdita in the J. C. Bridwell collection.

Proc. Hawaii. Entomol. Soc., 7(1): 151-161. June 1928.

Bees of the genus Perdita Smith in the American Museum of

Natural History (Hymenoptera). Am. Mus. Novit., 321:1-13.

July 12, 1928.

Two new species of bees of the genus Perdita in the collec¬

tion of the California Academy of Sciences. Pan-Pac. Ento¬

mol., 5(l):25-33. July 1928.

Three new species of the hymenopterous family Encyrtidae

from New South Wales. Univ. Calif. Publ. Entomol., 5(2):

5-18, 5 figs. March 30, 1929.

New records and descriptions of bees of the genus Perdita

VOLUME 58, NUMBER 1

(Hymenoptera). J. N. Y. Entomol. Soc., 37(2): 111-125. Au¬

gust 2, 1929.

38. 1929c. A new species of the encyrtid genus Metaphycus from Wash¬

ington (Hymenoptera). Pan-Pac. Entomol., 6(l):43-45. Octo¬

ber 10, 1929.

39. 1929d. Records of western species of Perdita with descriptions of

two new species (Hymenoptera). Pan-Pac. Entomol., 6(2):

49-56. December 19, 1929.

40. 1932. Three new parasitic Hymenoptera from the Indo-Malayan re¬

gion. Proc. Hawaii. Entomol. Soc., 8(1): 153-162. November

1932.

41. 1933. A note on Andrena nudimediocornis Vier. Pan-Pac. Entomol.,

9(l):28-29. January 1933. (By Timberlake and T. D. A. Cock¬

erell).

42. 1937a. New species of Andrena from California (Hymenoptera). Pan-

Pac. Entomol., 13(l/2):69—74. January-April 1937.

43. 1937b. New anthophorid bees from California (Hymenoptera). Am.

Mus. Novit., 958:1-17. November 2, 1937.

44. 1938. New species of Andrena from California (Hymenoptera). Pan-

Pac. Entomol., 14( 1):24—29. March 1938.

45. 1939a. New species of bees of the genus Diadasia from California

(Hymenoptera, Apoidea). Bull. Brooklyn Entomol. Soc.,

34(1): 11-16. February 27, 1939.

46. 1939b. New species of bees of the genus Dufourea from California

(Hymenoptera, Apoidea). Ann. Entomol. Soc. Am., 32(2):

395_4i4. June 30, 1939.

47. 1939c. Two new species of Acmaeodera from California (Coleoptera,

Buprestidae). Pan-Pac. Entomol., 15(4): 179-182. October 28,

1939.

48. 1939d. A new species of Hippodamia from Mt. Rainier (Coleoptera:

Coccinellidae). Proc. Hawaii. Entomol. Soc., 10(2):265-266.

December 1939.

49. 1940a. New species of bees of the genus Diadasia from California

(Hymenoptera, Apoidea). Bull. Brooklyn Entomol. Soc., 35(1):

22-30. February 21, 1940.

50. 1940b. A new species of Encopognathus from California (Hymen¬

optera, Sphecoidea). Entomol. News, 51(6): 167-168. June 12,

1940.

51. 1940c. The genus Centris in California (Hymenoptera, Apoidea).

Pan-Pac. Entomol., 16(3): 138-141. July 29, 1940.

52. 1941a. A new species of Anthophora from California (Hymenoptera,

Apoidea). Pan-Pac. Entomol., 17(1):34-36. February 21, 1941.

53. 1941b. New or little known California bees. [Contributions from the

Los Angeles Museum-Channel Islands Biological Survey: 16].

Bull. South. Calif. Acad. Sci., 39(3): 190-196. March 31, 1941.

PAN-PACIFIC ENTOMOLOGIST

54. 1941c.

55. 1941d.

56. 1941 e.

57. 1941f.

58. 1942.

59. 1943a.

60. 1943b.

61. 1943c.

62. 1946.

63. 1947a.

64. 1947b.

65. 1947c.

66. 1948.

67. 1949.

68. 1950.

69. 1951a.

70. 1951b.

Synoptic table of North American species of Diadasia (Hy¬

menoptera, Apoidea). Bull. Brooklyn Entomol. Soc., 36(1):

2-11. April 1, 1941.

Encyrtidae of the Marquesas and Society Islands (Hymenop-

tera, Chalcidoidea). Occas. Pap. Bernice P. Bishop Mus.,

16(9):215—230, 1 fig. April 2, 1941.

Three new dufoureine bees from California (Hymenoptera,

Apoidea). Ann. Entomol. Soc. Am., 34( 1):38—42. April 7, 1941.

Ten new species of Stelis from California (Hymenoptera,

Apoidea). J. N. Y. Entomol. Soc., 49(2): 123-137. May 6, 1941.

A new species of Hippodamia from Mexico (Coleoptera,

Coccinellidae). Proc. Entomol. Soc. Wash., 44(3):39. March

31, 1942.

The Coccinellidae or ladybeetles of the Koebele collection—

Part I. Hawaii. Plant. Rec., 47(1): 1-67, pis. 1-2 (with 32 figs.),

1 fig. January 1943.

Racial differentiation in Nearctic species of Dianthidium

(Hymenoptera, Apoidea). J. N. Y. Entomol. Soc., 51(2):

71-109. May 6, 1943.

Bees of the genus Colletes chiefly from Colorado. Bull. Am.

Mus. Nat. Hist., 81 (5):385—410, 6 figs. November 12, 1943.

Two new species of Ptiloglossa from Arizona (Hymenoptera,

Apoidea). Pan-Pac. Entomol., 22(4): 156-158. December 3,

1946.

New species of Perdita from the southern states (Hymen¬

optera, Apoidea). Proc. Entomol. Soc. Wash., 49(3):81-84.

February 28, 1947.

Two new species of bees from Arizona (Hymenoptera, Apoi¬

dea). Pan-Pac. Entomol., 23( 1):26—30, 1 fig. April 9, 1947.

A revision of the species of Exomalopsis inhabiting the United

States (Hymenoptera, Apoidea). J. N. Y. Entomol. Soc.,

55(2):85-106. May 26, 1947.

Additions and corrections to the list of Nearctic species of

Dianthidium (Hymenoptera, Apoidea). J. N. Y. Entomol. Soc.,

56(3): 149-153. October 7, 1948.

The species of Dianthidium of Baja California (Hymenoptera,

Megachilidae). Pan-Pac. Entomol., 25(3): 129-132. July 29,

1949.

The bees of the genus Proteriades (Hymenoptera, Megachili¬

dae). Univ. Kans. Sci. Bull., 33, Part 2 (10):387-440, pi. 3

(with 16 figs.). March 20, 1950. (By Timberlake and C. D.

Michener).

New and little-known bees of the family Andrenidae from

California. Proc. U.S. Nat. Mus., 101(3281):373^H4. April

23, 1951.

New species of Anthophora from the western United States

VOLUME 58, NUMBER 1

(Hymenoptera, Apoidea). J. N. Y. Entomol. Soc., 59(1):

51-62. June 25, 1951.

71. 1951c. Western bees of the genus Colletes (Hymenoptera: Apoidea).

Wasmann J. Biol., 9(2): 181-238. September 18, 1951.

72. 1952a. Descriptions of new species of Nomadopsis from California

and Texas, and of a new allied genus from South America.

Ann. Entomol. Soc. Am., 45(1): 104-118. March 1952.

73. 1952b. New records of Perdita from the eastern United States (Hy¬

menoptera, Apoidea). Proc. Entomol. Soc. Wash., 54(4): 199-

204. August 26, 1952.

74. 1953a. A new name for the bee genus Ruiziella. Ann. Entomol. Soc.

Am. (1952), 45(4):528. January 30, 1953.

75. 1953b. Bees of the genus Perdita in the collection of the University

of Kansas (Hymenoptera, Apoidea). Univ. Kans. Sci. Bull.,

35, Part 2 (7):961-985. September 10, 1953.

76. 1954a. Two new species of Perdita from North Carolina (Hymenop¬

tera: Apoidea). Entomol. News, 65(1): 12-15. January 15, 1954.

77. 1954b. A revisional study of the bees of the genus Perdita F. Smith,

with special reference to the fauna of the Pacific Coast

(Hymenoptera, Apoidea). Part I. Univ. Calif. Publ. Entomol.,

9(6):345^132, pis. 13-26 (with figs. 1-172). January 15, 1954.

78. 1954c. Two new species of Nomada, subgenus Gnathias, from Cali¬

fornia (Hymenoptera: Apoidea). Pan-Pac. Entomol., 30(2):

133-136. June 24, 1954.

79. 1955a. A new genus for two new species of dufoureine bees from

California (Hymenoptera: Apoidea). Pan-Pac. Entomol., 31(3):

105-108. September 23, 1955.

80. 1955b. Notes on the species of Psaenythia of North America. Boll.

Lab. Zool. Gen. Agrar. Fac. Agrar. Portici, 33:398-409. No¬

vember 3, 1955.

81. 1956a. Description of two new species of Diadasia from North

America (Hymenoptera: Apoidea). Pan-Pac. Entomol., 32(2):

90-92. May 9, 1956.

82. 1956b. A revisional study of the bees of the genus Perdita F. Smith,

with special reference to the fauna of the Pacific Coast

(Hymenoptera, Apoidea). Part II. Univ. Calif. Publ. Ento¬

mol., ll(5):247-349, pis. 27-41 (with figs. 173-367). August

24, 1956.

83. 1957a. A new entedontine chalcid-fly from seed capsules of eucalyptus

in California (Hymenoptera: Eulophidae). Pan-Pac. Entomol.,

33(3): 109-110, 1 fig. September 26, 1957.

84. 1957b. Notes on the subgenus Chelostomoides of the genus Mega¬

chile Latreille (Hymenoptera: Apoidea). Pan-Pac. Entomol.,

33(3): 132. September 26, 1957.

85. 1958a. Temnosoma, a genus of bees new to the United States (Hymen-

PAN-PACIFIC ENTOMOLOGIST

optera: Halictidae). Pan-Pac. Entomol., 34(1):34. March 18,

1958.

86. 1958b. A new species of the genus Colletes from the Colorado Desert

of California (Hymenoptera: Apoidea). Pan-Pac. Entomol.,

34(3): 143-145. July 22, 1958.

87. 1958c. A revisional study of the bees of the genus Perdita F. Smith,

with special reference to the fauna of the Pacific Coast (Hy¬

menoptera, Apoidea). Part III. Univ. Calif. Publ. Entomol.,

14(5):303—410, pis. 4-15 (with figs. 368-521). October 10, 1958.

88. 1960a. A revisional study of the bees of the genus Perdita F. Smith,

with special reference to the fauna of the Pacific Coast (Hy¬

menoptera, Apoidea). Part IV. Univ. Calif. Publ. Entomol.,

17(1): 1—155, pis. 1-17 (with figs. 522-732). March 18, 1960.

89. 1960b. [Descriptions of Perdita drymariae and P. krombeini, new

species]. Pp. 324-325, 327-328. In Mitchell, T. B., Bees of the

eastern United States, Volume I. N. C. Agric. Exp. St. Tech.

Bull., 141:1-538, 134 figs., frontispiece.

90. 1960c. A peculiar new halictine bee from California (Hymenoptera:

Apoidea). Proc. Entomol. Soc. Wash., 62(2): 105-106, 1 fig.

July 5, 1960.

91. 1961a. [Book review]. Bees of the eastern United States, Volume I.

By Theodore B. Mitchell. Pan-Pac. Entomol., 37(1): 16. March

24, 1961. (By Timberlake and P. D. Hurd, Jr.).

92. 1961b. A review of the genus Conanthalictus (Apoidea: Halictidae).

Pan-Pac. Entomol., 37(3): 145-160. August 22, 1961.

93. 1961c. A new species of Tetralonia from the deserts of California

and Nevada (Hymenoptera: Apoidea). Pan-Pac. Entomol.,

37(4):209-212. November 7, 1961.

94. 1962a. An interesting new species of Emphoropsis from California

(Hymenoptera, Anthophoridae). Entomol. News, 73(2):36-38.

February 16, 1962.

95. 1962b. The conflict between the tribal and family names Mirini

(Hymenoptera) and Miridae (Hemiptera). Entomol. News,

73(3):66. March 8, 1962.

96. 1962c. A revisional study of the bees of the genus Perdita F. Smith,

with special reference to the fauna of the Pacific Coast (Hy¬

menoptera, Apoidea). Part V. Univ. Calif. Publ. Entomol.,

28(1): 1-123, pis. 1-13 (with figs. 733-904), figs. A-B. April

30, 1962.

97. 1964a. Some new species of Pseudopanurgus of the subgenus Hetero-

sarus Robertson (Hymenoptera, Apoidea). Am. Mus. Novit.,

2185:1-26. July 9, 1964.

98. 1964b. A revisional study of the bees of the genus Perdita F. Smith,

with special reference to the fauna of the Pacific Coast

(Hymenoptera, Apoidea). Part VI. Univ. Calif. Publ. Ento-

VOLUME 58, NUMBER 1

mol., 28(2): 125-387, pis. 1-22 (with figs. 905-1190), 77 figs.

November 2, 1964.

99. 1965. Notes on caupolicanine bees of Arizona (Hymenoptera: Apoi-

dea). J. N. Y. Entomol. Soc., 73(l):46-48. April 19, 1965.

100. 1967. New species of Pseudopanurgus from Arizona (Hymenoptera,

Apoidea). Am. Mus. Novit., 2298:1-23. August 18, 1967.

101. 1968. A revisional study of the bees of the genus Perdita F. Smith,

with special reference to the fauna of the Pacific Coast (Hy¬

menoptera, Apoidea). Part VII (including index to Parts I to

VII). Univ. Calif. Publ. Entomol., 49:1-196, pis. 1-14 (with

figs. 1191-1360), 31 figs. June 19, 1968.

102. 1969a. Metapsaenythia, a new panurgine bee genus (Hymenoptera,

Andrenidae). Entomol. News, 80(4):89-92. May 2, 1969.

103. 1969b. A contribution to the systematics of North American species of

Synhalonia (Hymenoptera, Apoidea). Univ. Calif. Publ. Ento¬

mol., 57:i-vi, 1-76, 80 figs. July 18, 1969.

104. 1971a. On the identity of Panurginus ineptus Cockerell (Hymenop¬

tera: Andrenidae). Pan-Pac. Entomol., 47(2): 148. June 8, 1971.

105. 1971b. Supplementary studies on the systematics of the genus Perdita

(Hymenoptera, Andrenidae). Univ. Calif. Publ. Entomol.,

66:i-vi, 1-63, figs. 1361-1417. November 5, 1971.

106. 1973a. Revision of the genus Pseudopanurgus of North America

(Hymenoptera, Apoidea). Univ. Calif. Publ. Entomol., 72:i-

vi, 1-58, pis. 1-4 (with 56 figs.), 4 figs. April 13, 1973.

107. 1973b. A new Synhalonia from New Mexico (Anthophoridae). Proc.

Entomol. Soc. Wash., 75(3):317—318. October 5, 1973.

108. 1975. The North American species of Heterosarus Robertson (Hy¬

menoptera, Apoidea). Univ. Calif. Publ. Entomol., 77:i—vi,

1-56, pis. 1-8 (with 87 figs.). June 6, 1975.

109. 1976. Revision of the North American bees of the genus Protandrena

Cockerell (Hymenoptera: Apoidea). Trans. Am. Entomol.

Soc., 102:133-227, 93 figs. July 30, 1976.

110. 1977a. Description of two new species of Heterosarus Robertson

(Hymenoptera, Apoidea). Pan-Pac. Entomol., 53( 1):56—59, 4

figs. June 23, 1977.

111. 1977b. Two new species of Perdita from Arizona and Utah (Hymen¬

optera, Andrenidae). J. N. Y. Entomol. Soc., 85(1): 18-20.

July 28, 1977.

112. 1977c. Descriptions of new species of Perdita (Hymenoptera, Apoi¬

dea) in the collection of the California Academy of Sciences.

Proc. Calif. Acad. Sci., (4)41( 10):281-295. October 20, 1977.

113. 1980a. Report on Perdita in the collection of the California Academy

of Sciences. Wasmann J. Biol. (1979), 37(l/2):30—39. January

30, 1980.

114. 1980b. Supplementary studies on the systematics of the genus Per-

PAN-PACIFIC ENTOMOLOGIST

dita (Hymenoptera, Andrenidae). Univ. Calif. Publ. Entomol.,

85:i—viii, 1-65, figs. 1418-1479. May 1980.

115. 1980c. Review of North American Exomalopsis (Hymenoptera, An-

thophoridae). Part I. The subgenus Anthophorula. Part II.

The subgenus Anthophorisca. Part III. The subgenera Phano-

malopsis and Megomalopsis, with a description of the new

subgenus Panomalopsis. Part IV. The subgenus Exomalopsis.

Univ. Calif. Publ. Entomol., 86:i-vi, 1—158, 258 figs. October

1980.

Additional information concerning the life and work of Mr. P. H. Tim-

berlake is contained in the below cited press releases and magazine article.

The articles accompanied by a photograph of him are so annotated.

Press Releases

—Retired scientist is simply wild over identity of wild bee types. Banning

Record, Banning, California, page reference missing, May 20, 1964

(photograph).

—Bee expert, 84, catches 5,000 on hunting trip. The Press, Riverside,

California, Section B, page 1, Monday, October 9, 1967.

—Nixon’s famous scientist kinfolk. Oakland Tribune, Oakland, Califor¬

nia, page 1-A, Sunday, December 12, 1971 (photograph).

—‘He has looked at more insects than any man alive or dead’ [still going

strong at 88]. Press-Enterprise, Riverside, California, page B-5, Sun¬

day, March 5, 1972 (photograph).

—Retirement full-time job for bee-expert—[Bee expert eschews retire¬

ment]. The Enterprise, Riverside, California, County page, concluded

on page 2, Tuesday, July 24, 1979 (photograph).

—University of California, Riverside [UCR’s] world famous entomologist

and bee specialist is dead at 97. The Enterprise, Riverside, California,

Section C, page 1, Tuesday, April 21, 1981 (photograph, reprinted from

March 5, 1972 article).

Magazine Article

Michael Rogers, Is this the face that named a thousand bugs? Esquire,

78(3): 108-109, 154, September 1972 (color photography).

This In Memoriam article was prepared by Paul D. Hurd, Jr., Senior

Scientist, Smithsonian Institution, Washington, D.C., and E. Gorton Lins-

ley, Professor of Entomology, Emeritus, University of California, Berkeley,

with assistance of Jack C. Hall, University of California, Riverside. The

photograph that accompanies this article was taken by the photographer of

the University of California, Riverside, on the occasion of Mr. Timberlake’s

87th birthday (June 5, 1970).

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 19-24

BIOLOGY OF EUTRETA DIANA OSTEN SACKEN ON SAND

SAGEBRUSH ARTEMISIA FILIFOLIA TORR.

(DIPTERA: TEPHRITIDAE)

Sandra M. Benbow

Dept, of Entomology, Texas Tech University, Lubbock 79409

AND

David E. Foster

Dept, of Entomology, Iowa State University, Ames 50011

Eutreta diana Osten Sacken is one of the most widely distributed and

frequently encountered gall-forming tephritids occurring in western North

America. Since its original description, E. diana has been reported from

most western states (Foote and Blanc, 1963). In his recent treatment of the

genus, Stoltzfus (1977) summarized what was known of E. diana biology:

its seasonal occurrence, distribution and host-plant associations. Two pub¬

lications have reported hymenopteran parasites reared from E. diana galls

(Fronk et al., 1964; Furniss and Barr, 1975).

The object of this paper is to make known for the first time the occurrence

of E. diana in Texas and its association with sand sagebrush, Artemisia

filifolia Torr., and to assess its impact on current year growth of that plant.

Methods and Materials

During the period of May 1976 through November 1977 studies were

conducted on the English Ranch, 4 miles east of Crosbyton, Crosby Co.,

Texas. During the seasonal occurrence of the larval, pupal and adult stages,

forty galls and associated current year growth were collected at two week

intervals and returned to the laboratory. Twenty galls from each collection

were measured to provide an index of gall development. The galls were then

dissected to monitor the occurrence and activity of various gall inhabitants.

The contents of each gall was recorded and the biological role of each

inhabitant assessed. The cephalopharyngeal skeletons of E. diana larvae

were measured and their length and shape recorded. First and second instar

skeletons were studied by preparing entire larvae in temporary glycerine

mounts; skeletons of third instar larvae were removed from the body prior

to mounting.

The remaining twenty galls from each collection were held for rearing.

After rearing activities were completed the galls were oven-dried and their

PAN-PACIFIC ENTOMOLOGIST

weights recorded. Concurrent with gathering galls for laboratory analysis 20

uninfested terminals were gathered from the same plants. These were re¬

turned to the laboratory where their lengths were measured. A paired-T-test

was used to compare dry weights of galled versus uninfested stems.

At the end of each growing season a straight line transect was laid out

across a portion of the study site not utilized for gathering samples. Galls

occurring on the first one hundred infested plants encountered along the

transect were counted and categorized according to their relative state of

decomposition: category 1, current year’s galls; category 2, last season’s

galls, appearing brown but with leafy covering still intact; category 3, galls

two years old, outer surface bare but not broken; category 4, galls three

years old and older, bare and broken, generally with only lower portions of

the gall remaining.

Results

Examination of the cephalopharyngeal skeleton provided the basis for

determining the instar of each E. diana larva examined. First instar skele¬

tons appear noticeably short with respect to their width when viewed lat¬

erally. Second and third instar skeletons are relatively long with respect to

their width and are easily separated from those of the first instar. Definitive

separation of all second and third instars is accomplished using the length

of the longest axis, which falls within a defined range for each instar (Fig.

1 ).

Seasonal occurrence .—First instar larvae begin hatching during the latter

half of March with hatching continuing until the middle of April. The length

of the first stadium is approximately 32 days. By the end of May all larvae

have attained the second instar. Third instar larvae first appear during the

third week of June. By mid-July all larvae have attained the third instar.

The lengths of the second and third stadia are approximately 63 and 64 days,

respectively. Pupae first appear during the third week of August. By the

second week of September all individuals have pupated. The length of pu¬

pation is approximately 30 days. Adult flies emerge between the third week

of September and the middle of October. They remain in the field until frost.

Oviposition begins in mid-October and continues as long as adults remain

active. Females deposit one egg per axillary bud. Eggs are placed lengthwise

between overlapping bud scales. A female frequently deposits several eggs

in the axillary buds of a single branch. As many as 17 galls of a single age

class have been found on a given branch. Undisturbed adults tend to remain

on the same plant. Females have been observed ovipositing on the same

plant for as many as four successive days.

Gall development and distribution .—At about the same time A. filifolia

begins to grow noticeably in March, newly hatched larvae bore through the

VOLUME 58, NUMBER 1

100

FW:

.• ;:

• •

Gy-vl

0 -v«

■ : : .•

;>7T

•,.v;

Vcf.'O

...

4 •»* •

r ..

• o •

V. ;‘ o

First instar Larvae

Second instar Larvae

Third instar Larvae

l.-JL'L

.. ■<

/“ : --

'r-v <:■> >.v. S'v.s

•

I"".

\ -V o V > y-'-CV • - •- :•••. '•

Fig.

5 10 15 20 25 30 35 40 45 50 55 60 65

Cephalopharyngeal Skeleton Length (mm)

Cephalopharyngeal skeleton lengths in Eutreta diana larvae.

base of the axillary bud and begin to feed at the base of the developing

stem. First instar feeding is restricted to the center of the developing stem.

Stem swelling is first noted after feeding has progressed for about two weeks.

Early second instar feeding occurs in both the developing gall and adjacent

stem but is eventually confined to the gall proper, a phenomenon apparently

corresponding to the hardening of stem tissue. As feeding continues the gall

chamber is sealed from the stem cavity by developing plant tissue and frass.

Late second and third instar feeding is confined to the surface of the gall

cavity. Mature third instar larvae feed in the distal portion of the gall to

produce an emergence portal covered by only a thin epidermal layer.

Gall development, as reflected by gall diameter (Fig. 2), corresponds di¬

rectly to larval feeding but is arrested about two weeks prior to pupation.

In instances where the larva succumbs prior to the third instar, gall for¬

mation ceases.

The direct effect of E. diana feeding on the host plant is the interruption

of internodal elongation. Both galled and normal stems increase in length

until mid-July, though normal stems increase at a much faster rate (Fig. 3).

Dry weights of galled and normal stems do not differ significantly at the five

percent level when compared using a paired-T-test.

PAN-PACIFIC ENTOMOLOGIST

Plants occurring along predetermined transects were examined at the end

of each growing season; galls were placed in one of four categories according

to their apparent age. Galls exhibited a clumped distribution, with an av¬

erage of 42 per plant (range 1-68). The average number of category 1 galls

Fig. 3. Average length of galled and normal Artemisia filifolia stems.

VOLUME 58, NUMBER 1

found on five plants not previously infested was 9 (range 4-17), whereas the

average number of category 1 galls on 81 plants exhibiting all four age

classes was 24 (range 1-43). When the data from all 100 infested plants were

pooled, the following averaged were obtained: category 1—21 (range 1-43);

category 2—9 (range 4-12); category 3—7 (range 3-10); and category A —5

(range 1-8) galls per plant.

Gall associates .—Six species of parasitic Hymenoptera have been asso¬

ciated with galls of E. diana. Of these, two occur commonly and are asso¬

ciated with larval or pupal stages. Four are uncommon and of undetermined

affinity. An undetermined Eurytoma sp. (Eurytomidae) infested 21 percent

of the E. diana galls reared during the study. It functioned as a solitary

ectoparasitoid of second and third instar larvae. Eurytoma larvae that have

devoured their host have been observed feeding on tissues of the gall wall.

Pupation occurs within the gall cavity. Adult Eurytoma emerge in Septem¬

ber.

A Tetrastichus sp. (Eulophidae) emerged from 24 percent of the galls held

for rearing. Larvae of this species are multiple endoparasitoids of E. diana

larvae and pupae. Their presence causes the host to succumb during pu¬

pation. Adult emergence occurs in September and October with as many as

27 individuals emanating from a single gall. Tetrastichus females gain access

to the host larva by chewing a hole through the gall, then ovipositing directly

in the host. Similar oviposital behavior has been recorded for Tetrastichus

cecidophagus Wangberg (Wangberg, 1977).

Two undetermined species of Eupelmus (Eupelmidae) were reared from

galls containing E. diana pupae. Their combined presence occurred in less

than one percent of the galls examined. One female Gastrancistrus sp.

(Pteromalidae) and one female representing an undetermined genus near

Comperilla (Encyrtidae) were reared from galls containing E. diana pupae

during September 1976.

Summary

Eutreta diana is reported from Artemisia filifolia for the first time, and

its known distribution expanded to include the Texas panhandle and High

Plains. The fly is univoltine and overwinters in the egg stage. Its three larval

instars are separated by the shape and length of the cephalopharyngeal

skeleton. Gall development is a response to feeding by first, second and

early third instar larvae. Galled stems grow at a reduced rate until mid-July

by which time they have reached their maximum length. Galls continue to

increase in diameter until the larva stops feeding. There is no significant

difference between two oven-dry weights of galled and uninfested stems.

The occurrence of relatively large numbers of current season galls on pre-

PAN-PACIFIC ENTOMOLOGIST

viously infested plants may be explained at least in part by the sedentary

behavior of undisturbed female E. diana.

Acknowledgments

The authors wish to thank Dr. W. Bryan Stoltzfus, Iowa State University

and Dr. Carl M. Yoshimoto, Canadian Biosystematic Research Institute for

the respective identifications of E. diana and its hymenopterous parasites.

Funding for this project was provided by the College of Agriculture, Texas

Tech University under whose auspices the results are published as contri¬

bution T-10-116.

Literature Cited

Foote, R. E., and F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif.

Insect Surv., 7:1-117.

Fronk, W. D., A. A. Beetle, and D. G. Fullerton. 1964. Dipterous galls on the Artemisia

tridentata complex and insects associated with them. Ann. Entomol. Soc. Amer.,

57:575-577.

Furniss, M. M., and W. F. Barr. 1975. Insects affecting important native shrubs of the north¬

western United States. USDA For. Ser. Gen. Tech. Rep., INT-19, 64 pp.

Stoltzfus, W. B. 1977. The taxonomy and biology of Eutreta (Diptera: Tephritidae). Iowa

State J. Res., 51:369-438.

Wangberg, J. K. 1977. A new Tetrastichus (Hymenoptera: Eulophidae) parasitizing tephritid

gall-formers on rabbitbrush in Idaho. Pan-Pac. Entomol., 53:237-241.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 25-30

DIFFERENTIATING ADULTS OF APPLE MAGGOT,

RHAGOLETIS POMONELLA (WALSH) FROM SNOWBERRY MAGGOT,

R. ZEPHYRIA SNOW (DIPTERA: TEPHRITIDAE) IN OREGON

Richard L. Westcott

Plant Division, Oregon Department of Agriculture, Salem 97310

Recent discovery of the apple maggot, Rhagoletis pomonella (Walsh), in

the Pacific Northwest (Anonymous, 1980) poses a serious threat to tree fruit

growers in this and surrounding areas. Therefore, it is imperative that ac¬

curate and timely identifications be made for ongoing survey and detection

activities, biological studies, and management decisions for regulatory ac¬

tivities and control. A discussion of the apple maggot and its distribution in

the Pacific Northwest was given by AliNiazee and Penrose (1981). Figure 1

updates this distribution, which now extends eastward in Washington to

Stevenson, Skamania Co.; southward in Oregon to Brookings, Curry Co.

The present paper attempts to expand and utilize earlier taxonomic studies

as they relate to R. pomonella in the western United States, thereby facil¬

itating identification in this area.

R. pomonella belongs to a group comprising four sibling species, and its

hosts are in the family Rosaceae, mostly the subfamily Pomoideae (Bush,

1966). In Oregon it has been reared from apple, crabapple and ornamental

hawthorn. Identification of R. pomonella in the eastern U.S. requires taxo¬

nomic discrimination from R. cornivora Bush and R. mendax Curran; while

in the West concern is, at least for the present, only with R. zephyria Snow,

a species restricted to snowberry, Symphoricarpos spp. This paper addresses

taxonomic differentiation of known Oregon populations of pomonella and

zephyria but its application should prove useful elsewhere, especially in the

western U.S.

Specimens utilized in this study were taken during the summer of 1980,

mostly from traps located in the Willamette Valley and immediate vicinity.

Most specimens of R. pomonella were from the greater Portland area, while

specimens of R. zephyria came from the northern and mid-Willamette Val¬

ley, and from the Columbia Gorge as far east as Hood River. Also utilized

was a series of R. zephyria reared from snowberries, Corvallis, 1933, S. C.

Jones (Oregon State Univ. Coll.). In the Pacific Northwest, actual and po¬

tential host plants for both species grow in association over a wide area.

Bush (1966), in his comprehensive work on the genus for North America,

presented an array of taxonomic characters, but those which proved most

useful were a wing band ratio, total wing length of female, length of the

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Known distribution of the apple maggot, Rhagoletis pomonella (Walsh), in the

Pacific Northwest: circles = 1980 survey data; triangles = 1981 survey data; stippled = gen¬

erally infested.

ovipositor and shape of the male claspers (surstyli). He studied various

populations of R. pomonella from different hosts. Wasbauer (1963) com¬

pared specimens of a New York population of that species with a California

population of R. zephyria, utilizing the above characters which were sug¬

gested to him by Bush (in litt.). Based on these studies and my work in

identifying hundreds of specimens of the two species, it is clear that the

most reliable and facile differentiating character is the configuration of the

surstyli, followed by the length of the ovipositor. Therefore, this paper

focuses on those characters in an attempt to simplify and clarify their use.

Males

In my opinion, males of R. pomonella and R. zephyria are readily sepa¬

rable if one utilizes the genital structures; in fact, this appears in many cases

to be the only way to positively identify them. It is not necessary to remove

and specially prepare these structures for study, thus saving time; and it is

best to view their posterior aspect. The detail and depth provided by SEM

VOLUME 58, NUMBER 1

20KV X150 100U 046 03021 flHS

Figs. 2-5. Fig. 2. Male genitalia showing surstyli of Rhagoletis pomonella (Walsh). Fig. 3.

Same, R. zephyria Snow. Fig. 4. Ovipositor, R. pomonella. Fig. 5. Same, R. zephyria (illus¬

tration shows variation common to both species).

photos (Figs. 2, 3) provides a more accurate comparison than available line

drawings, clearly showing the parallel surstyli of R. pomonella with their

broad surfaces facing directly laterad, versus the divergent surstyli of R.

zephyria with their broad surfaces arranged obliquely. This configuration is

PAN-PACIFIC ENTOMOLOGIST

best observed on a freshly killed or relaxed fly, so the legs may be moved

if they obscure one’s view; however, if a dry specimen is expendable, simply

break off the offending appendages! Occasionally, a specimen of R. po-

monella prepared from a sticky trap will have one surstylus (rarely both)

distorted from the natural position depicted in Fig. 2. However, this is a

minor problem and does not preclude positive identification. I have found

this distortion to be insignificant in R. zephyria.

An additional character of the surstylus which may be of value in sepa¬

rating the two species is the presence of much longer apical setae in R.

pomonella. However, these are sometimes difficult to see with a fight mi¬

croscope or may be broken off, especially on specimens prepared from

traps. I examined numerous males of R. zephyria and found no evidence of

these longer setae.

Females

When confronted with one or a very few flies for identification, most often

they were females. Traps captured many more females than males. Nor¬

mally this poses no problem, as identification can usually be made by mea¬

suring the length of the ovipositor, often from the combination of a wing

band ratio and wing length. Very large specimens can usually be determined

as R. pomonella on the the basis of size alone.

Some clarification is necessary with regards to measuring the ovipositor,

which exhibits similar variation in both species (Figs. 4, 5). Usually there

exists a variably-developed basal sclerotized dorsal extension or process

(Fig. 4). My measurements were made from the apex of the ovipositor to

the apex of this process; it can only be assumed that previous authors

mentioned herein did likewise. In some cases, especially when the ovipos¬

itor is darkly sclerotized, an accurate measurement can be made without

removing the ovipositor from the specimen. However, it usually is best to

place it on a slide in a mixture of mounting medium and glycerine sufficient

to restrict its movement. If the distal sheath is poorly translucent it may

have to be cleared. Problems may arise with lightly sclerotized or freshly

emerged specimens (see discussion below on R. zephyria).

Previous studies (Wasbauer, 1963; Bush, 1966) clearly showed that ovi¬

positor length was the most reliable character for differentiating females of

the two species. With the exception of a few anomalous specimens of R.

pomonella reared from plum and fire thorn in Florida and Texas, respec¬

tively, there was no overlap. In R. pomonella (N = 155) the length ranged

(R) from .90 mm (Florida specimens reared from hawthorn; otherwise the

smallest was .98 mm) to 1.49 mm, means (x) of the different populations

varied from 1.13-1.33 mm. In R. zephyria N - 47, R - .63-.88 mm, x

.75 and .78 mm. Data from an Oregon sample of specimens are as follows:

VOLUME 58, NUMBER 1

R. pomonella : N = 93, R = .92-1.38 mm, x - 1.14 mm. R. zephyria : N

121. R = .72-.88 mm, x = .81 mm. Although these figures are in close

agreement with those of prior workers, several specimens (for two of which

data were not included) indicate a very small character overlap between

species. Including both species, only five females had ovipositors measuring

in the “problem area” (.88-.98 mm). Three of these (.92, .93 and .97 mm)

were identified as R. pomonella based on supplementary objective criteria

(wing length; wing band ratio) and subjective characters. 1 Of those excluded

from the sample data, one (.95 mm) was questionably placed in R. zephyria

and another (.94 mm) combined characters of both species.

It should be noted that correlation in size of the fly (which is indicated

by wing length) with length of the ovipositor appeared insignificant. Al¬

though extensive comparisons were not made, some of the smallest speci¬

mens of R. pomonella possessed ovipositors of above average length, and

the opposite was true in R. zephyria.

In addition to the sample above I made hundreds of identifications, all

from trap catches. Eight females were found with ovipositor lengths ranging

from .90-.98 mm. All but one were determined as R. pomonella, including

one with the ovipositor measuring .90 mm. The eighth (.90 mm, too) was

identified as R. zephyria.

A sample taken from the extensive reared series of R. zephyria produced

the following data: N = 23, R = .69-.81 mm, x = .77 mm. The smaller

average ovipositor length could be an artifact of measurement, since scler-

otization was very light and a basal dorsal process was rarely evident. How¬

ever, it could also be a result of the picked host fruit deteriorating in quality

before larval maturity, as alluded to by Benjamin (1934:15). Investigations

to determine the latter effect are necessary, as the desirability of using

reared material in taxonomic studies is obvious. The need is specially evi¬

dent for further studies utilizing positively host-associated (preferably by

rearing) material from the western states, for biological investigations as

well.

In summary, it appears that a small percentage of female flies cannot be

identified with certainty; however, the chance of dependency on such spec¬

imens is low. If an identification is critical, such as in regulatory work, effort

must be made to secure additional material from which positive identifica¬

tion may be made.

Acknowledgments

I wish to thank R. L. Penrose, Oregon Department of Agriculture, for

providing pertinent data from his apple maggot program and for many help¬

ful suggestions. Gratitude is given for technical assistance from Diana Kim-

berling, of the same institution, who prepared Fig. 1. Appreciation goes to

PAN-PACIFIC ENTOMOLOGIST

A. H. Soeldner, Oregon State University, for SEM photography; and to R.

H. Foote, U.S. National Museum, for much encouragement and reviewing

the manuscript.

Literature Cited

AliNiazee, M. T., and R. L. Penrose. 1981. Apple maggot in Oregon: A possible new threat

to the Northwest apple industry. Bull. Entomol. Soc. Am., 27(4):245-246.

Anonymous. 1980. Coop. Plant Pest Rep. (U.S. Dep. Agric.), 5(25):472.

Benjamin, F. H. 1934. Descriptions of some native trypetid flies with notes on their habits.

U.S. Dep. Agric. Tech. Bull., 401:1-96.

Bush, G. L. 1966. The taxonomy, cytology, and evolution of the genus Rhagoletis in North

America (Diptera: Tephritidae). Bull. Mus. Comp. Zool., 134(11):431—562.

Wasbauer, M. S. 1963. Taxonomic discrimination of Rhagoletis pomonella from the east¬

ern United States and Rhagoletis zephyria from California. Calif. Dep. Agric., T-9:

1-6 (unpubl.).

Footnote

1 After examining many specimens, I came to recognize that most specimens of R. pomonella

possess a subtly lighter wing band color (perhaps perceptible only in fresh or recently collected

material) and, in the female, the wing is very slightly less broadly rounded apically.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 31-37

A NEW GENUS AND SPECIES OF CALIFORNIA PILL BEETLE

(COLEOPTERA: BYRRHIDAE)

Paul J. Johnson

1408 28th Street S.E., Auburn, Washington 98002

During routine examination of various collections of Byrrhidae, a new

species and genus was found in the miscellaneous material from the Uni¬

versity of California, Davis. My thanks to R. O. Schuster for the loan of

this material, and to Dr. L. K. Russell for checking the description.

This extremely interesting and unique byrrhid can be defined as follows:

Sierraclava, new genus

Type-species.—Sierraclava cooperi, new species.

Elongate-oval, convex; basal two-thirds of elytral margins parallel; head

entirely retractile, eyes and mouthparts not visible in repose; clypeus ob¬

solete; mandibles stout, tridentate; maxillary palps 4-segmented; labial palps

3-segmented; galea fleshy and heavily setaceous; antennae capitate-clavate,

scape and second article expanded, flagellum received into groove between

prosternum and pronotal epipleura; prosternum Y-shaped with short pro-

sternal process; proepisternum obsolete; mesosternum transversely narrow,

deeply excavated medially, obsolescent; metasternum broader than long;

abdomen longer than broad, five visible segments, apical segment not mod¬

ified; pro- and metatrochanters spurred basally; pronotum strongly con¬

stricted anteriorly, convex, laterally with tear-drop shaped excavation near

front angles, deeply concave epipleurae, basally subequal in width to elytra;

scutellum obsolecent, triangular; elytra twice as long as pronotum, convex,

deeply and serially punctate, bristle-like hairs arranged serially on interstrial

spaces, smaller appressed hairs giving a mottled appearance throughout

dorsum; entire venter modified for complete retraction of all appendages,

punctate and clothed with short bristle-like hairs; leg segments individually

modified for retraction of adjacent segments; tarsus essentially 4-4-4, first

article minute and completely fused to second article; aedeagus single-lobed,

spatulate, arcuate in lateral outline; female styli reduced, peg-like; length

2.0-2.6 mm.

Etymology. — Sierra- is in reference to the Sierra Nevada mountain range

in which the type locality is located; -clava refers to the distinctive capitate-

clavate antennae.

PAN-PACIFIC ENTOMOLOGIST

Sierraclava cooperi, new species

(Figs. 1-7)

Holotype male .—Form compact, elongate-oval, convex; basal two-thirds

of elytral margins parallel; length 2.0-2.6 mm; integument testaceous to

brown to piceous dorsally, testaceous to rufous ventrally; entire dorsum

densely clothed with variegated patterns of brown and white, decumbent,

scale-like setae; additionally sparsely to moderately clothed with straight,

stout, slightly clavate, truncated, dark colored hairs, these being randomly

distributed on head and pronotum, and serial on the interstrial spaces of

elytra; venter sparsely to moderately clothed with bristle-like hairs directed

posteriorly, arising from upon interpunctural spaces.

Head rounded, convex, completely retractile into pronotum, eyes not

visible from front and mouthparts not visible when head retracted; frons

densely, deeply punctate, punctures becoming smaller, less dense towards

vertex, pubescence as above; frons becoming depressed anteriorly with

broad, shallow impressions laterally above mandibular insertions; circum-

frontal carina strong from vertex to eyes, becoming finer anteriorly where

it may represent fronto-clypeal suture; space between anterior “clypeal”

margin and “clypeal suture” densely, deeply punctate, this space and an¬

terior clypeal margin deflexed inward from clypeal suture; entire frons mod¬

erately microreticulate, shining between reticulations. Eyes hemispherical,

flattened, coarsely faceted, gray in color. Antennae as in Fig. 4, 11-seg-

mented, capitate-clavate, not reaching base of pronotum, entire flagellum

sparsely pubescent with short pale hairs, apical segments moderately pu¬

bescent; antennae received into grooves between prosternum and prono¬

tum. Labrum small, twice as wide as long, one-eighth as wide as widest

portion of frons; shallowly, sparsely punctate; shining; sparsely pubescent

on disc, denser marginally; broadly, shallowly emarginate apically; anterior

angles broadly rounded. Mandibles stout, 1.5 times longer than broad; tri-

dentate; membranous and ciliated medially; notched medially with notch

extending towards base; molar area chitinized, bluntly toothed; ventral con¬

dyle small. Ventral mouthparts as in Fig. 3; maxillary palps 4-segmented,

ultimate segment narrowly pear-shaped, cylindrical, apex truncate and

membranous; penultimate one-half length of ultimate; segment 2 subequal

to penultimate; basal one-half length of segment 2. Galea fleshy, lobe-like,

heavily setaceous apically. Lacinia similar to galea but shorter. Labial palps

3-segmented; utltimate segment broadly pear-shaped, cylindrical, apex trun¬

cate and membranous; penultimate subequal in length to ultimate; basal

segment one-half length of penultimate. Ligula cordate, shallowly emargin¬

ate apically; membranous. Mentum subtrapezoidal, short, 5 times wider

than long, broadly and shallowly emarginate anteriorly. Sub mentum quad¬

rate, 4.5 times wider than long. Gula trapezoidal, twice as wide as long,

sides sinuate anteriorly.

VOLUME 58, NUMBER 1

t i

1mm.

Figs. 1-7. Sierraclava cooperi Johnson, new genus, new species. Fig. 1, Adult, lateral view,

elytral pubescence removed. Fig. 2. Adult, dorsal view, left side pubescence removed. Fig. 3.

Ventral mouthparts, left labial palp and right maxillary palp removed. Fig. 4. Antenna. Fig. 5.

Female genitalia. Fig. 6. Male genitalia, lateral view. Fig. 7. Male genitalia, ventral view.

PAN-PACIFIC ENTOMOLOGIST

Pronotum convex, trapezoidal, strongly constricted anteriorly, appearing

broadly rounded in dorsal view; anterior angles obtuse, slightly deflexed

and depressed inside of angle; lateral margin double in anterior half, en¬

closing a depression with beaded margins and widest at anterior angle; lat¬

eral margin beaded basally; hind angles subacute, depressed, right-angled;

posterior margin finely beaded, depressed, slightly sinuate; integument shin¬

ing, strongly microreticulate, impunctate; epipleura deeply excavated for

reception of proleg, shining, finely microreticulate. Scutellum very small,

smooth, triangular.

Elytra 2.5 times longer than and subequal in width to pronotum, slightly

sinuate and depressed basally; integument strongly microreticulate, shining;

11-striate; striae composed of deep, serial punctures, often coalescing; sub-

sulcate apically; striae 2, 3, 4, 6, 7, and 9 ending just after declivity; sutural,

5, 8, and 10 extending subapically; stria 11 merging with stria 10 at meta-

femoral emargination; humeral angles broadly obtuse; epipleura deeply ex¬

cavated at base for reception of mesoleg, form continuous with pronotal

epipleura; epipleura narrowing and ending at distal end of excavation; elytra

with indexed lobe immediately distad from excavation, elytron excavated

shallowly for reception of metaleg; excavation gently curving, margin then

straight to apex; entire margin finely beaded; apex extended, slightly re¬

flexed, angles narrowly obtuse, adjacent; sutural margin finely beaded.

Prosternum Y-shaped; densely, deeply punctate; margins beaded; front

margin broadly and evenly emarginate; lateral margins broadly and sinuately

emarginate; prostemal process obtusely rounded apically; moderately clothed

with short pale bristles.

Proepisternum obsolete. Proepimeron obsolete.

Protrochantin small, triangular, tectiform.

Procoxae transverse, excavated for reception of femora; posterior surface

of coxa with a punctate plate-like process covering retracted trochanter.

Trochanter one-quarter length of femora, triangular, with basal spur.

Femora stout, extending to pronotal margin, flattened, modified for com¬

plete retraction, punctate, with pale bristles.

Tibia subequal in length to femora, flattened, only slightly narrower than

femora, moderately pubescent with short scale-like setae, more sparsely

with bristles; apical spurs paired, subequal, minute.

Tarsus one-half length of tibia; four segmented; retractile into tibial grooves;

claws simple.

Mesosternum deeply excavated medially for reception of prosternal pro¬

cess, visible portion very narrow, subcarinate laterally, 3 times wider than

long, front portion deeply depressed, excavated for reception of procoxae,

exposed portion densely and deeply punctate.

Mesoepisternum elongate, triangular. Mesoepimeron elongate, pentago¬

nal.

VOLUME 58, NUMBER 1

Mesotrochantin quadrate. Mesocoxae subglobose; anterior surface of coxa

forming a small, punctate plate. Mesotrochanter short, triangular, not

spurred. Mesofemora similar to profemora. Mesotibia similar to protibia.

Mesotarsus similar to pro tarsus.

Metasternum broad, 3 times wider than long, subequal in length to ab¬

domen; anterior intercoxal process one-third as wide as full sternum, broad¬

ly, shallowly, evenly emarginate; anterior quadrants deeply excavated for

reception of mesolegs; margins of excavations carinate; lateral margins lin¬

ear, subparallel; posterior margin straight except for a small intercoxal pro¬

cess; median longitudinal dark line extending from tip of posterior intercoxal

process to about two-thirds length of sternum, shallowly and narrowly im¬

pressed; sternum medially, latitudinally depressed; moderately to densely,

deeply punctate; clothed sparsely with pale bristles.

Metepisternum elongate, deflexed; visible portion anterior, triangular,

slightly excavated for mesolegs; covered portion triangular, elongate, nar¬

row.

Metepimeron membranous, narrow, elongate, quadrate.

Metatrochantin obsolete externally.

Metacoxae approximate, extending to elytral margin; plate-like; strongly

sinuate posteriorly, widest mesally; moderately, deeply punctate; deeply

internally excavated for reception of femora.

Metafemora extending to elytral margin, fitting into shallow emargination;

otherwise similar to preceding femora.

Metatibia subequal in length to femora; similar to preceding tibiae.

Metatarsus similar to preceding tarsi.

Abdomen five-sevenths as wide as long; moderately to densely punctate;

sparsely bristled; sternite 1 deeply excavated laterally for reception of meta¬

legs; intercoxal process rounded, convex, margins beaded; sternite 2 con¬

vex; sternite 3 slightly convex; sternite 4 flattened; sternite 5 convex me¬

dially, broadly rounded apically; sternite length ratio 3:2:1:1:3. Entire venter

finely microreticulate.

Genitalia as in Figs. 6 and 7; pars basalis convex dorsally, concave ven-

trally; parameres obsolete; median lobe slightly flattened, spatulate apically.

Female .—Similar to male; abdomen very slightly broader; antennal scape

and segment 2 slightly broader, more flattened; genitalia as in Fig. 5; coxites

lightly sclerotized, large, stout, sparsely setaceous; stylus small, peg-like,

apically setaceous.

Fittle overall variation, other than size, is apparent in the type series.

Etymology .—I take great pleasure in dedicating this unique, enigmatic

moss beetle to the honor of Kenneth W. Cooper; for many years of devotion

to studying bryophagous insects.

Holotype male, allotype female, and one-half of the paratype series will

be returned to the University of California, Davis. Fabelling for the holo-

PAN-PACIFIC ENTOMOLOGIST

type, allotype and most paratypes (58) reads: Sequoia National Park, Cali¬

fornia; altitude 2000-3000 ft.; V-12-1937; A. T. McClay. Additional paratype

data as follows: same data, VI-13-1929 (2), VI-2-1929 (1); Sequoia National

Park, California, Ash, Mt. Forebay, IV-25-1949, R. C. Bechtel (10); same

data, Ash Mt. R., IV-21-1950 (2) and IV-28-1950 (1). Paratypes will be de¬

posited in the collections of the author; K. W. Cooper, Riverside; L. K.

Russell, Corvallis; Oregon State University, Corvallis; U.S. National Mu¬

seum, Washington, D.C.; California Academy of Sciences, San Francisco;

Canadian National Collection of Arthropods, Ottawa; British Museum (Nat¬

ural History), London; Australian National Insect Collection, Canberra.

At this time, little is known of the exact habitat for this species. R. C.

Bechtel (pers. comm.) mentioned that the beetles were collected by sheet¬

drying debris which had been washed down a flume at the Kaweah Power

House no. 3 Reservoir. This flume initiated some distance upstream from

the reservoir. Sierraclava cooperi is probably a moss feeder as are its near¬

est relatives, Curimopsis spp., and I would suggest that this beetle may be

found in mosses growing on boulders, talus, gravel, or mineral soil but

receiving enough moisture to keep the microhabitat moist for at least part

of the year.

Sierraclava cooperi is taxonomically associated with the byrrhid subfam¬

ily Syncalyptinae as arranged by El-Moursy (1969) and Arnett (1973); that

is, the genera Curimopsis Ganglbauer, Chaetophora Kirby et Spence ( Syn-

calypta Dillwyn, auct. (Johnson, 1978)), and Microchaetes Hope. Within

this subfamily, Sierraclava is closest to Curimopsis, and will key out to this

genus in Arnett (1973). The following key modification will allow for quick

and easy separation from Curimopsis and all other Nearctic byrrhids:

13 (1). Frons with two oblique grooves; integument without scales, but

with simple clavate hairs; length 1.0-2.0 mm .

. Chaetophora Kirby et Spence

Frons without two oblique grooves; integument with decum¬

bent scales or hairs of various forms, and bristle-like hairs of

various shapes; length 2.0-3.0 mm . 14

14(13). Elytral striae fine, not deeply impressed; punctures fine to mod¬

erate, diffuse; bristle-like hairs randomly distributed .

. Curimopsis Ganglbauer

Elytral striae serially, deeply punctate; bristle-like hairs serial

on interstrial spaces . Sierraclava, n. g.

The above character separation of Sierraclava from Curimopsis is rather

obvious and can be easily distinguished even with a low-power handlens in

the field. Additionally, differentiating characters separating these two genera

include the shorter and more compressed pars basalis, the more spatulate

VOLUME 58, NUMBER 1

median lobe, the more reduced styli of the female, and the overall convex

and elongated form of Sierraclava. Opposingly, Curimopsis has a longer,

subcylindrical, arcuate pars basalis; a narrower median lobe; less reduced

female styli; and the overall form is more ovate and dorsally depressed.

Literature Cited

Arnett, R. H. 1973. The Beetles of the United States (a Manual for Identification), 4th ed.

Am. Entomol. Inst., Ann Arbor, Michigan, 1112 pp.

El-Moursy, A. A. 1969. The North American genera on the family Syncalyptidae El-Moursy.

Bull. Soc. Entomol. Egypte, 53:131-135.

Johnson, C. 1978. Notes on Byrrhidae (Col.): With special reference to, and a species new

to, the British fauna. Entomol. Rec., 90:141-147.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 38-41

INSECT SEED PREDATION ON ASTRAGALUS BISULCATUS

(HOOK.) GRAY (LEGUMINOSAE ) 1

Stephen L. Clement 2

Ohio Agricultural Research and Development Center, Wooster 44691

AND

Donald H. Miller

Lyndon State College, Lyndonville, Vermont 05851

A few Astragalus (Leguminosae) species assume special significance in

the intermountain region of western United States and Canada because they

are highly poisonous to livestock. The poisonous nature of these plants is

associated with their ability to accumulate large (>1000 ppm) quantities of

selenium (Se) (Trelease and Martin, 1936; Rosenfeld and Beath, 1964). One

such poisonous plant, A. bisulcatus (Hook.) Gray, is a widely distributed

perennial species in the intermountain region (Barneby, 1964).

Although the phytochemistry of A. bisulcatus has been the subject of

several studies (see Nigram and McConnell, 1969; Chow et al., 1971), there

is only one published account of insect seed predation on this species. In

this publication, Trelease and Trelease (1937) found that the seeds from an

A. bisulcatus population located near Laramie, Wyoming were heavily in¬

fested with a bruchid, Acanthoscelides fraterculus (Horn), and a seed-chal-

cid identified as a Bruchophagus species (probably mexicanus (Ashmead)).

These investigators were surpsied to find insects completing their devel¬

opment on A. bisulcatus seeds containing about 1475 ppm of Se, especially

in view of the known toxicity of seleniferous vegetation to various mammals

and arthropods such as red spiders (Gnadinger, 1933) and aphids (Hurd-

Karrer and Poos, 1936).

The present study was designed to assess the relative intensity of insect

seed predation in 4 Wyoming populations of A. bisulcatus. We document

the potential role of insect seed predators on the population dynamics of A.

bisulcatus because this poisonous weed seems to be spreading along sele¬

niferous horizons in and east of Grand Teton National Park (GTNP). More¬

over, A. bisulcatus has become well established on newly reclaimed areas

of a uranium mine in the Powder River Basin of Wyoming (J. D. Love,

pers. comm.).

VOLUME 58, NUMBER 1

Methods and Study Areas

A collection consisting of one randomly selected raceme from 7 plants

was made in the Gros Ventre Canyon (lower site), 2.5 km east of the GTNP

boundary on July 26, 1978. In 1979, collections were taken at this site on

July 19, 27 and August 19. Collections were taken at 3 additional sites in

1979: Gros Ventre Canyon (upper site), 6.1 km east of GTNP boundary

(sampled July 27); 2.4 km northeast of Wolff Ranch, GTNP (sampled July

20); and 24.2 km NW Dubois (sampled July 27 along Hwy. 26). Each 1979

collection consisted of one raceme from 10 plants although in 2 instances

(Wolff Ranch collection and July 27 collection at the lower site in Gros

Ventre Canyon) it was difficult finding one intact raceme per plant because

of high seed predation by rodents. It appeared that these rodents were

indiscriminate in their selection of seed pods.

The lower and upper Gros Ventre Canyon and Dubois study sites were

small (<0.2 ha) roadside areas, each supporting about 20-50 scattered plants.

At the Wolff Ranch site an undetermined number of plants (<100) were

distributed for a few hundred meters along a small ridge that overlooked a

large meadow to the north. A. bisulcatus plants at each site were found on

dry gravelly, seleniferous soils.

Samples were returned to a laboratory at the Ohio Agricultural Research

and Development Center (OARDC) where fully developed seed pods from

each raceme were individually dissected to determine the extent of insect

seed predation. Mean percentage seed predation values (±SD) for each

collection were calculated from 7 or 10 replicates of one raceme each.

Results and Discussion

An average of 68.5 ± 18.5% of the seed pods collected in Gros Ventre

Canyon (lower site) on July 26, 1978 were infested with larvae of the an-

thomyiid fly, Hylemya anane (Walker). In 1979, seed pod infestation rates

of this fly were uniformly high in all collections: 66.3 ± 23.1% and 89.5 ±

7.1% in the July 19 and 27 collections, respectively, at the lower site in

Gros Ventre Canyon; 72.2 ± 28.3% in the collection from the upper site in

Gros Ventre Canyon; 61.7 ± 22.1% in the Wolff Ranch collection; and 79.5

± 23.0% in the Dubois collection. Each of these infested pods contained

one developing H. anane larva and this larva consumed, or at least de¬

stroyed, all the seeds within the pod (Fig. 1). An additional collection was

taken on August 19 at the lower site in Gros Ventre Canyon, but by this

date most of the insect seed predators had vacated their seed pods; however,

dissections of all the pods revealed a seed pod infestation rate of 82.4 ±

14.3%. The mean number of seed pods per raceme ranged from a low of

16.1 ± 8.6 (August 19 collection, lower site in Gros Ventre Canyon) to a

high of 29.1 ± 14.6 (Dubois collection).

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Larva of Hylemya anane within a seed pod of Astragalus bisulcatus.

An unidentified seed chalcid wasp was responsible for additional seed

destruction. However, our only estimate of the amount of damage caused

by this wasp was obtained by dissecting the seeds in 45 randomly selected

seed pods from the July 27, 1979 Gros Ventre Canyon (lower site) collection.

In this sample, a developing chalcid larva was found in 8 of 74 (10.8%)

mature seeds that were dissected. All attempts to rear larvae to adults failed.

Heretofore, nothing has appeared in the literature concerning the biology

of H. anane or closely related species (G. Steyskal, pers. comm.). Thus,

biological aspects of H. anane gleaned from our study are noteworthy. For

instance, observations on the development of this species in an outdoor

screenhouse at OARDC between early August 1978 and late June 1979 sug¬

gest that H. anane is univoltine and has an obligatory diapause. In this

context, larvae vacated their seed pods in late August to pupate on the soil

surface (3 cm deep) in rearing containers (1 qt ice cream cartons). Adults

emerged from overwintering pupae during May and June.

Hymenopterous parasitoids attacked H. anane in Wyoming. These para-

sitoids were recovered from screenhouse rearing containers in late August

1978 and early May 1979, and sent to the USDA Systematic Entomology

Laboratory in Washington, D.C., where they were identified as Chlorocytus

sp. and Halticoptera sp. (Pteromalidae).

The selenium in A. bisculcatus seeds does not prevent substantial pre-

VOLUME 58, NUMBER 1

dispersal seed destruction by at least 3 insect species in several Wyoming

populations. In addition, insect seed predators readily attack at least 2 other

Astragalus species whose foliage is toxic to livestock: A. cibarius Sheld.

by a bruchid beetle ( Ac antho sc elides frater cuius Horn), a seed weevil

(Tychius soltaui Casey), a seed chalcid ( Bruchophagus mexicanus Ash-

mead), stink bugs {Chlorochroa uhleri Stal, C. ligata Say), and lepidop-

terous larvae ( Glaucopsyche lygdamus Dbldy., Strymon melinus Hiibner)

(Green and Palmbald, 1975); and A. pectinatus Dough ex Hook, by a fly

(probably Pseudotephritis sp.) (see Moxon, 1939).

Acknowledgments

We are indebted to Dr. J. D. Love for his support and encouragement

with this study. We thank Drs. B. Bowman, S. Duffey, A. Moxon, and R.

Rust for general assistance, and Mr. G. Steyskal and Dr. E. Grissell, USDA

Systematic Entomology Laboratory, for taxonomic assistance.

Literature Cited

Bameby, R. C. 1964. Atlas of North American Astragalus. Mem. N.Y. Bot. Gard., 13:1-

1188.

Chow, C. M., S. N. Nigam, and W. B. McConnell. 1971. Biosysthesis of Se-methylseleno-

cysteine and S-methylcystome in Astragalus bisulcatus: Effect of selenium and sulphur

concentrations in the growth medium. Phytochemistry, 10:2693-2698.

Gnadinger, C. B. 1933. Selenium insecticide material for controlling red spider. Ind. Eng.

Chem., 25:633-637.

Green, T. W., and I. G. Palmbald. 1975. Effects of insect seed predators on Astragalus

cibarius and Astragalus utahensis (Leguminosae). Ecology, 56:1435-1440.

Hurd-Karrer, A. M., and F. W. Poos. 1936. Toxicity of selenium containing plants to aphids.

Science, 84:252.

Moxon, A. L. 1939. The selenium content of grasshoppers found feeding on seleniferous

vegetation. Proc. South Dakota Acad. Sci., 19:69-70.

Nigam, S. N., and W. B. McConnell. 1969. Seleno amino compounds from Astragalus

bisulcatus isolation and identification of r-L-glutamyl-Se-methylseleno-L-cysteme. Bio-

chem. Biophys. Acta, 192:185-190.

Rosenfeld, I., and O. A. Beath. 1964. Selenium. Academic Press, New York, 411 pp.

Trelease, S. F., and A. L. Martin. 1936. Plants made poisonous by selenium absorbed from

the soil. Bot. Rev., 2:373-396.

Trelease, S. F., and H. M. Trelease. 1937. Toxicity to insects and mammals of foods con¬

taining selenium. Am. J. Bot., 24:448-451.

Footnotes

1 Approved for publication as Journal Article No. 119-80 of the Ohio Agricultural Research

and Development Center, Wooster 44691.

2 Present address: USDA Biological Control of Weeds Laboratory, 1050 San Pablo Ave.,

Albany, California 94706.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 42^17

KEY TO THE PUPAL PARASITES OF CALIFORNIA OAKWORM,

PHRYGANIDIA CALIFORNIA (LEPIDOPTERA: DIOPTIDAE),

BASED ON LARVAL EXUVIAE

Lester C. Young

Department Plant and Soil Science, California State

Polytechnic University, Pomona 91768

The California oakworm, Phryganidia californica Packard, is a common

defoliator of coast live oak, Quercus agrifolia Nee, particularly in the San

Francisco Bay area. The moth is bivoltine with a 9 month spring generation

followed by a 3 month fall generation. Cyclic outbreaks occur for 2-3 years

followed by relative scarcity for 4-7 years (Horn, 1974). A detailed account

of the life history of P. californica is given by Harville (1955) who noted

regular fluctuations in its population dynamics since 1851 with no apparent

harm to trees. However, since Q. agrifolia is highly valued as a shade and

ornamental tree in public parks and residential areas, highly defoliated trees

are considered to sustain aesthetic and monetary losses (Harville, 1955;

Wickman, 1971).

The majority of the parasite complex of P. californica is associated with

the pupal stage (Harville, 1955; Horn, 1974; Young, 1977). However an egg

parasite, Tetrastichus sp., has been found (Young, 1977) and two tachinid

flies, Actia sp. and Zenillia virilis Aldrich and Webber, were reared from

larvae (Harville, 1955; Young, 1977).

Primary pupal parasites include ichneumonids, Itoplectis behrensii (Cres-

son) and Ephialtes Ontario (Cresson), and a chalcidid, Brachymeria ovata

(Say). I. behrensii was originally described in 1896 with a more complete

description given by Townes and Townes (1960) in their revision of the

subfamily Pimplinae. This parasite is specific to P. californica with no rec¬

ord of an alternate host. B. ovata is polyphagous with over 100 recorded

hosts (Burks, 1960) and was first recorded on P. californica by Burke and

Herbert (1920). E. Ontario was first reported on P. californica by Hagen

(1949) and is a facultative parasite known mostly to attack lepidopterous

pupae found on conifers (Townes and Townes, 1960).

Hyperparasites, in the pupal parasite complex, have been reported to

attack only I. behrensii (Harville, 1955; Horn, 1974). These include ichneu¬

monids, Gelis tenellus (Say), Mastrus aciculatus (Provancher) and Bathy-

thrix sp., and a pteromalid, Dibrachys cavus (Walker). However, both D.

cavus and G. tenellus were found hyperparasitizing B. ovata and E. Ontario

(Young, 1977).

VOLUME 58, NUMBER 1

0.1 mm

Fig. 1. Ephialtes Ontario, last stage larva. Buccal armature showing stipital sclerite (SS).

During a previous study it became necessary to identify parasites from

post emerged P. californica pupae (Young, 1977). The major advantage of

this method was that a more accurate assessment of parasite abundance and

effectiveness could be made.

Parasite Determinations

Identification of parasite species was based on remnants left in pupae by

emerged adults. During fall 1974, several hundred suspected parasitized

pupae were collected from oak trees on the campus of the University of

California, Berkeley and were individually reared for adult parasites. The

adults were identified and P. californica pupae from which they emerged

were examined. Meconium, cast larval skin, head capsule, emergence hole

size and other remnants were used to construct the following key to the

pupal parasites of P. californica. The hyperparasite Bathythrix sp. was ex¬

cluded from this key because of insufficient recoveries.

PAN-PACIFIC ENTOMOLOGIST

0.1 mm

Fig. 2. Itoplectis behrensii, last stage larva. Buccal armature.

Key to the Pupal Parasites of Phryganidia californica Packard

1. Pupal wall thick, opaque, ranging from cream color to black; emer¬

gence hole(s) present. 2

Pupal wall thin, translucent, split laterally along antennae, termi¬

nating to open flap just below head ... Host Phryganidia californica

2. Meconium present posteriorly or occasionally anteriorly at opposite

end of emergence hole . Primary Parasite 3

Meconium absent at either end of pupa; shriveled remains of primary

parasite larva present .Secondary Parasite 5

3. Meconium appears as mass of grayish ovoid pellets, each pellet ap¬

proximately 0.5 mm long; pupa often showing dark spot between

wing pads; cast skin of last larval molt including head capsule of

primary parasite present.. 4

Meconium packed into smooth mass often filling Vs of host pupa;

VOLUME 58, NUMBER 1

0.1 mm

Fig. 3. Mastrus aciculatus, last stage larva. Buccal armature showing epistomal arch (EA),

lacinial sclerite (LS), prelabial sclerite (PS), hypostoma (H).

parasite remains consisting of pupal envelope rather than cast skin

of last larval molt of primary parasite . Brachymeria ovata

4. Emergence hole often exceeding 2.2 mm in diameter; buccal arma¬

ture of head capsule of last larval molt with stipital sclerite present

(Fig. 1), not common . Ephialtes Ontario

Emergence hole seldom exceeding 2.2 mm in diameter; buccal ar¬

mature of head capsule of last larval molt lacking stipital sclerite

(Fig. 2), very common . Itoplectis behrensii

5. Emergence hole less than 1.0 mm in diameter, pupa containing sev¬

eral parasite pupal envelopes . Dibrachys cavus

Emergence hole greater than 1.0 mm in diameter, pupa containing

remains of solitary parasite . 6

6. Pupa containing flimsy, loosely woven cocoon; head capsule with

complete, lightly sclerotized epistomal arch, lacinial sclerites pres¬

ent, hypostomal arm short barely extending beyond stipital scler¬

ite, prelabial sclerite present (Fig. 3) . Mastrus aciculatus

Pupa containing thickened cocoon approximately 7.0 mm long and

2.0 mm wide; head capsule with incomplete epistomal arch, la-

PAN-PACIFIC ENTOMOLOGIST

!_ Q. I mm _ J

Fig. 4. Gelis tenellus, last stage larva. Buccal armature showing hypostoma (H).

cinial sclerite absent, hypostomal arm long extending Vs beyond

stipital sclerite, prelabial sclerite absent (Fig. 4). Gelis tenellus

Acknowledgments

I wish to thank Drs. B. D. Burks and G. Gordh for help on the identifi¬

cations of some of the parasites.

Literature Cited

Burke, H. E., and F. B. Herbert. 1920. California oakworm. U.S. Dep. Agric. Farmers’ Bull.,

1076:1-14.

Burks, B. D. 1960. A revision of the genus Brachymeria Westwood in America north of

Mexico (Hymenoptera: Chalcididae). Trans. Amer. Entomol. Soc., 86:225-273.

Hagen, K. S. 1949. Two new ichneumonid host records. Pan-Pac. Entomol., 25:3.

Harville, J. P. 1955. Ecology and population dynamics of the California oak moth, Phrygan-

idia californica (Lepidoptera: Dioptidae). Microentomology, 20:83-166.

Horn, D. J. 1974. Observations on primary and secondary parasitoids of California oakworm,

Phryganidia californica, pupae (Lepidoptera: Dioptidae). Pan-Pac. Entomol., 50:53-59.

Townes, H. K., and M. Townes. 1960. Ichneumon-flies of America north of Mexico. U.S.

Nat. Mus. Bull., 216(2):i—vii, 1-676.

VOLUME 58, NUMBER 1

Wickman, B. E. 1971. California oakworm. U.S. Dep. Agric., For. Serv., For. Pest Leaf!.,

72:1-4.

Young, L. C. 1977. Pupal parasites of the California oak moth, Phryganidia californica

Packard: A biological and ecological study. Ph.D. Dissertation, University of California,

Berkeley, 126 pp.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 48-52

TRICHOGRAMMA AUSTRALICUM GIRAULT (HYMENOPTERA:

TRICHOGRAMMATIDAE): REDESCRIPTION AND

LECTOTYPE DESIGNATION

J. D. Pinto, E. R. Oatman, and G. R. Platner

Department of Entomology, University of California, Riverside 92521

Nagarkatti and Nagaraja (1979) summarized the history of the name T.

australicum. The name was improperly applied to one of the most common

Australasian species of Trichogramma by Nagarkatti and Nagaraja (1968).

This error was first noted by Viggiani (1976) upon examination of a portion

of Girault’s material of T. australicum. It now appears that most previous

references to T. australicum are instead assignable to T. chilonis Ishii.

We recently examined most of the series of T. australicum which Girault

had before him at the time of his description, including the male syntype

which Viggiani (1976) assumed to be lost. A designation of this male as

lectotype and a redescription of T. australicum is presented below. All

material examined is deposited in the Queensland Museum, Queensland,

Australia. Girault’s specimens are on glass slides and mounted in Canada

Balsam. We have remounted the lectotype in Hoyer’s medium.

Trichogramma australicum Girault

Trichogramma australicum Girault, 1912:109. Viggiani, 1976:182. Nagar¬

katti and Nagaraja, 1979:115.

Type information .—The original description of T. australicum was based

on two females and one male. The male and only one female were listed as

types by Girault. The male “captured by sweeping grass along the left bank

of the Pioneer River, Mackay, Queensland,” 15 October 1911, is herein

designated as lectotype (Type No. Hy/801, Queensland Museum, exam¬

ined). It is not clear which of the two females Girault considered a type.

Contrary to his published statement, both are on separate slides bearing the

Hy/801 type label. We have labelled the female from the “Mulgrave River,

near Pyramid Mountain, Nelson (Cairns),” 25 November 1911, as paralec-

totype. Locale for the other female is not specified on its slide nor in Gi¬

rault’s original description. Other material, identified by Girault as T. aus¬

tralicum, also was examined by him at the time of his description. These

are listed below.

Lectotype male .—Dusky black with scutellum, metanotum, and vertex

bright orange-yellow (fide Girault, 1912). Legs distinctly lighter except hind

coxae dark.

VOLUME 58, NUMBER 1

Antennae (Fig. If) with flagellum slightly curved, relatively short, 0.95 as

long as hind tibia, 0.24 as wide as long, flagellar setae short, stout, relatively

blunt apically, 50-55 in number, length of longest seta 1.44 as long as max¬

imum width of flagellum.

Fore wing (Fig. la) with vein tracts distinct, setae between tracts moderate

in number, area between 4th and 5th tracts (i.e., the 2 tracts posterior to

the RS 2 ) with 18 setae; longest seta on postapical margin 0.18 as long as

maximum width of wing.

Hindwing (Fig. lb) with only middle vein tract complete; anterior tract

apparently absent; posterior tract composed of 5 widely spaced, short setae

extending only to apical of wing.

Mesoscutellum with anterior pair of setae damaged, broken at extreme

base but obviously much finer and presumably much shorter than posterior

pair.

Genital capsule (Fig. Id) relatively narrow, 0.36 as wide as long; dorsal

expansion of gonobase (DEG) moderately narrowed apically, narrowing

gradually, sides subsinuate, shallowly constricted at base, apex distinctly

sclerotized; DEG and chelate structures (CS) both attaining 0.89 the length

of genital capsule; median ventral projection (MVP) long, robust, distinctly

pointed, almost at level of apex of DEG, attaining 0.84 the length of genital

capsule. Aedeagus (Fig. le) 0.81 as long as hind tibia, apodemes relatively

long, comprising 0.54 the length of entire structure (Fig. lg).

Paralectotype female .—Badly damaged. Color as in male (fide Girault,

1912). Antenna (Fig. lc) with funicular segments wider than long; 1st funic¬

ular segment 0.67 as long as wide; 2nd 0.80 as long as wide, their combined

length distinctly less than that of pedicel. Ovipositor 1.22 the length of hind

tibia (Fig. lg).

Other material .—Girault (1912) identified 19 additional specimens as T.

australicum, five of which were males. These were taken from the following

Queensland locales: Nelson (5 females, 3 males); Mareeba (1 female); Her-

berton (2 females); Cairns (1 male); Innisfail (1 female); and Cooktown (5

females, 1 male). Eleven of these specimens, including two males, were

examined in our study. Other material from Girault’s collection were col¬

lected either after his description of T. australicum or were ambiguously

labelled.

Substantial variation exists within the original series. Girault stated that

one of the females from Nelson was distinctly lighter (“nearly uniformly

yellow’’) than others. We also note that the length of the MVP varies. In

the lectotype, its apex is clearly below that of the DEG and CS. In the male,

probably collected from Cairns (labelled as from Innisfail in Girault’s ma¬

terial), it attains the same level as both of these structures. Also, the entire

genital capsule is somewhat narrower in this male (0.29 as wide as long).

The genitalia figured by Viggiani (1976) shows a considerably shorter MVP

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Trichogramma australicum. a, b, d-g, lectotype 8. c, g, paralectotype ?. a, fore-

wing. b, hind wing, c, $ antenna, d, genital capsule, e, aedeagus. f, 8 antenna, g, ratio of

8 + 9 genitalia to hind tibia.

VOLUME 58, NUMBER 1

than occurs in the males examined by us. We have determined that the male

examined by Viggiani was collected from Indooroopilly, 29 September 1920,

and clearly was not a part of Girault’s original series. Unfortunately, the

condition of this specimen is not conducive to a comparison of other struc¬

tures.

The flagellar setae of the lectotype are stout and short (longest seta 1.4 as

long as maximum width of flagellum). Although the antennae of the other

two males examined are shriveled, their setae are substantially finer and

longer, with a corresponding ratio of ca. 1.9 in both.

In both females used by Girault in his original description, the ovipositor

is ca. 22% longer than the hind tibia. It is only ca. 4% longer in all other

females in his series.

Distribution .—Known only from Girault’s original material collected at

various locales in Queensland, Australia.

Remarks. —Males of T. australicum are most similar to those of T. cali-

fornicum described by Nagaraja and Nagarkatti (1973) from northeastern

California. The only differences we can detect in a comparison of the types

of these species are the structure of the MVP and flagellar setae. The dif¬

ference in the MVPs is minor. In T. californicum, the MVP is slightly more

narrow than that in T. australicum (cf. Fig. Id below and Fig. 10 in Nagaraja

and Nagarkatti, 1973). The flagellar setae are longer in T. californicum. In

T. australicum, the longest flagellar setae are 1.4 as long as the maximum

flagellar width. Nagaraja and Nagarkatti (1973) state that in T. californicum

they are “nearly twice” the maximum width. Our measurements on the

holotype of T. californicum indicate that the flagellar setae are 1.7 as long

as the maximum flagellar width, but the flagellum appears to have been

abnormally inflated in preparation. Thus, the true value may be closer to

2.0. Certain other important characters, however, viz. hind wings, length

of the aedeagus, and the nature of the mesoscutellar setae, cannot be com¬

pared because of the condition of material available for study.

The two females used in Girault’s description of T. australicum are dis¬

tinct from those of T. californicum. The funicular segments of the former

are wider than long. They are typically quadrate in Trichogramma as they

are in T. californicum. The only other females known to us which have

similar funicular segments are those of T. retorridum (Pinto et al., 1978).

The hosts of T. australicum are unknown. Reports of T. australicum in

Java on various Lepidoptera eggs (e.g., Girault, 1914, 1915) are apparently

based on a different species (Girault, 1922).

Since none of Girault’s T. australicum specimens were reared, the con-

specificity of variants and the sexes requires confirmation. Considering the

magnitude of variation described above we feel that his original series prob¬

ably represents more than one species.

PAN-PACIFIC ENTOMOLOGIST

Acknowledgments

We are thankful to E. C. Dahms, Senior Curator, Entomology, Queens¬

land Museum, Queensland, Australia, for providing Girault’s type speci¬

mens of Trichogramma australicum, and to Nancy A. Browning for pre¬

paring the illustrations.

Literature Cited

Girault, A. A. 1912. Australian Hymenoptera Chalcidoidea—1. The family Trichogramma-

tidae with descriptions of new genera and species. Mem. Queensland Mus., 1:66-116.

Girault, A. A. 1914. Notes on the Hymenoptera Trichogrammatidae and Mymaridae. Can.

Entomol., 46:327-330.

Girault, A. A. 1915. Australian Hymenoptera Chalcidoidea—I. Second Supplement. Mem.

Queensland Mus., 3:142-153.

Girault, A. A. 1922. New chalcid flies from Australia. The Entomologist, 55:205-208.

Nagaraja, H., and S. Nagarkatti. 1973. A key to some New World species of Trichogramma

(Hymenoptera: Trichogrammatidae), with descriptions of four new species. Proc. Ento¬

mol. Soc. Wash., 75:288-297.

Nagarkatti, S., and H. Nagaraja. 1968. Biosystematic studies on Trichogramma species. I.

Experimental hybridization between Trichogramma australicum Girault, T. evanescens

Westwood and T. minutum Riley. Commonw. Inst. Biol. Control Tech. Bull., 10:

81-96.

Nagarkatti, S., and H. Nagaraja. 1979. The status of Trichogramma chilonis Ishii (Hym.:

Trichogrammatidae). Orient. Insects, 13(1-2): 115-118.

Pinto, J. D., G. R. Platner, and E. R. Oatman. 1978. Clarification of the identity of several

common species of North American Trichogramma (Hymenoptera: Trichogrammati¬

dae). Ann. Entomol. Soc. Amer., 71:169-180.

Viggiani, G. 1976. Ricerche sugli Hymenoptera Chalcidoidea XLIX. Trichogramma confu-

sum n. sp. per T. australicum Nagarkatti & Nagaraja (1968) nec Girault (1912), con note

su Trichogrammatoidea Girault e descrizione di Paratrichogramma heliothidis n. sp.

Boll. Lab. Entomol. Agr. “Filippo Silvestri” di Portici, 33:182-187.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 53-58

UNDERSTORY PLANTS AS INDICATORS OF HOST TREES OF THE

WOUNDED TREE BEETLE, NOSODENDRON CAL1FORNICUM,

IN NORTHERN IDAHO (COLEOPTERA: NOSODENDRIDAE) 1

D. L. Kulhavy, 2 J. W. Schwandt, 3 and S. D. Hobbs 4

College of Forestry, Wildlife and Range Sciences,

University of Idaho, Moscow 83843

The wounded tree beetle, Nosodendron californicum Horn, occurs in the

northwestern United States on slime fluxes of white fir, Abies concolor

(Gord. and Glend.) Lindl., grand fir, A. grandis (Douglas) Lindl., Douglas-

fir, Pseudotsuga menziesii (Mirb.) Franco, and in California on A. concolor

and California black oak, Quercus kelloggii Newb. (Sokoloff, 1959, 1964;

Hatch, 1961; Arnett, 1968; Zack et al., 1979). The habitat of the wounded

tree beetle in northern Idaho is slime fluxes of tree wounds of grand fir

infected with the Indian paint fungus, Echinodontium tinctorium (Ell. and

Ev.) Ell. and Ev. The wounded tree beetles are apparently attracted to the

slime flux odor and are a component of the slime flux ecosystem (Sokoloff,

1959, 1964). In Idaho, wounded tree beetles are found in A. grandis IPachis-

tima myrsinites and Thuja plicata/P. myrsinites habitat types (h.t.) (Osborne

and Kulhavy, 1975) as described by Daubenmire and Daubenmire (1968).

Adults and larvae are found singly or in groups on frost cracks, basal

stem wounds, or in ooze puddles at the base of tree wounds. The adults

and larvae overwinter under loose bark or in duff at the base of host trees.

Colonization of host trees is thought to occur from July through August and

the beetles are sedentary during the months of September through March

(Osborne and Kulhavy, 1975).

In this paper, plant species groups were evaluated during the 1974 and

1975 field seasons as possible indicators of suitable host trees of N. cali¬

fornicum in the T. plicata/P. myrsinites h.t. of northern Idaho.

Study Areas

Two 50-hectare areas were established in the T. plicata/P. myrsinites h.t.

southwest of Elk River, Clearwater County, Idaho. Study area A, located

9 km SW of Elk River on the Tired Wolf and the Butterfield drainages, is

in the xeric portion of the T. plicata/P. myrsinites h.t. There have been

repeated entries into the stand to remove high value western white pine,

Pinus monticola Douglas. The residual stand consists of over-mature grand

fir with pockets of T. plicata Donn understory. Many of the grand fir contain

E. tinctorium decay and have basal logging wound scars and frost cracks.

PAN-PACIFIC ENTOMOLOGIST

Table 1. Structure of a 2 x 2 chi-square contingency table used in Table 2.

(a) Trees having both beetles and

associated plant groups

(b) Trees having no beetles, but having

indicator plant groups

plant groups

(d) Trees having neither indicator plants

nor beetles

ab interaction is calculated as [a/(a + b)] x 100; the ac interaction is calculated as [a/

(a + c)] x 100.

Slime fluxes and ooze puddles from the slime flux runoff have formed at

the base of the wounded grand fir and in logging wounds. The crown canopy

is not continuous, resulting in a mosaic of understory vegetation. The av¬

erage elevation is 920 m with a south to southeast aspect.

Area B, located 7 km SSE of Elk River in the mesic portion of the T.

plicata/P. myrsinites h.t., borders a fork of Oviatt Creek. The area is a cool

air drainage with T. plicata as the major understory tree species. The over¬

story is dominated by over-mature grand fir, western white pine, and rem¬

nant western larch (. Larix occidentalis Nutt.), with Douglas-fir, Pseudotsuga

menziesii (Mirb.) Franco, on the drier slopes. The area has no recent indi¬

cations of logging or fire, and the tree crowns form a canopy-like cover of

the forest floor. The majority of the dominant and codominant grand fir have

E. tinctorium decay. The average elevation is 895 m with a west to north¬

west slope.

Methods

In Areas A and B, all dominant and codominant grand fir with oozing

wounds (n = 60 and n = 40, respectively) were selected for study. A vege¬

tation species list, including all perennial shrubs, forbs, and graminoides,

was compiled on a 7 50 circular hectare (V 20 acre) plot centered at each tree.

On each sample tree, the numbers and location of any life stages of N.

californicum were recorded. To determine if any groups of plants were

indicators of the presence of N. californicum, the data were analyzed using

a 2 x 2 contingency table and chi-square computer program developed by

Miller (1974). The 2x2 contingency table segregates the beetle-plant group

associations into four categories (Table 1). The ab interaction is the condi¬

tional probability of finding a particular plant group given the presence of

the beetle. This gives the percentage of trees having beetles and also having

indicator plant groups. The ac interaction is the conditional probability of

finding the beetles given the presence of the indicator plant groups.

Results

Utilizing the 2x2 contingency tables and chi-square tests, two 3-plant

groups and two 4-plant groups in Area A; and three 3-plant groups, one 4-

VOLUME 58, NUMBER 1

Table 2. Indicator plant groups for Nosodendron californicum host trees in the Thuja

plicata/Pachistima myrsinites habitat type, northern Idaho, 1974—1975.

Contingency table cell 1 Probability

Indicator plant group

Chi-square

Area A

Hieracium albijiorum Hook.,

Vaccinium membranaceum

Douglas, Tiarella trifoliata L.

13 76*** 1 2

46%

92%

H. albijiorum, V.

membranaceum, Pachistima

myrsinites (Pursh) Raf.

11 92 ***

42%

92%

H. albijiorum, P. myrsinites, T.

trifoliata, Thermopsis

montana Nutt.

11 92 ***

42%

92%

H. albiflorum, T. trifoliata, T.

montana, Galium triflorum

Michx.

11 92 ***

42%

92%

Area B

Polystichum munitum (Kaulf.)

Presl, V. membranaceum,

Lonicera ciliosa (Pursh) DC.

11.88***

68%

94%

P. munitum, V.

membranaceum, Disporum

trachycarpum (Wats.) Benth.

and Hook.

11.95***

60%

100%

P. munitum, V.

membranaceum, L. ciliosa,

Rubus parviflorus Nutt.

11.95***

60%

100%

P. munitum, V.

membranaceum, Goodyera

oblongifolia Raf.

9.00**

60%

94%

P. munitum, V.

18.03***

76%

95%

membranaceum, and either

D. trachycarpum or L. ciliosa

(or both D. trachycarpum and

L. ciliosa )

1 a, both N. californicum and indicator plants present; b, indicator plants alone; c, N. cal¬

ifornicum alone; d, neither N. californicum nor indicator plant groups present.

2 ** P 0.01; *** P 0.001.

plant group, and the combination of two of the 3-plant groups in Area B

indicated the presence of N. californicum at least 92% of the time (Table

2). The best indicator plant group in Area A, with an ab conditional prob¬

ability of 46% and an ac conditional probability of 92% consisted of Hier-

PAN-PACIFIC ENTOMOLOGIST

Table 3. Common names and usual habitat of indicator plants for host Abies grandis of

Nosodendron californicum, northern Idaho, 1974—1975.

Indicator plant species

Common name

Usual habitat 1

Hieracium albiflorum Hook.

Hawkweed

Fairly moist slopes and open

woods

Vaccinium membranaceum

Douglas

Huckleberry

Mountain slopes

Tiarella trifoliata L.

False mitrewort

Damp woods

Pachistima myrsinites (Pursh) Raf.

Mountain lover

Midmontane

Thermopsis montana Nutt.

Buckbean

Widespread in Pacific Northwest

Galium triflorum Michx.

Bedstraw

Widespread in Pacific Northwest

Goodyera oblongifolia Raf.

Rattlesnake-

plain tain

Dry to moist forests

Lonicera ciliosa (Pursh) DC.

Orange

honeysuckle

Widespread native twining vine

Disporium trachycarpum (Wats.)

Benth. and Hook.

Fairy-bell

Wooded slopes, often near streams

Rubus parviflorus Nutt.

Thimbleberry

Moist to dry, wooded to open

areas

Polystichum munitum (Kaulf.) Presl

Christmas fern

Moist conifer forests, in open, or in

deep shade

1 Scientific names, common names and usual habitat from Hitchcock and Cronquist (1973).

acium albiflorum Hook, (hawkweed), Vaccinium membranaceum Douglas

(huckleberry) and Tiarella trifoliata L. (false mitrewort). The best indicator

plant group in Area B comprised the combination of two 3-plant groups with

an ab conditional probability of 76% and an ac conditional probability of

95%. These plant groups were Polystichum munitum (Kaulf.) Presl (Christ¬

mas fern), V. membranaceum, with either Disporium trachycarpum (Wats.)

Benth. and Hook., or Lonicera ciliosa (Pursh) DC. (or bothD. trachycarpum

and L. ciliosa ) present. The common names and usual habitats of the indi¬

cator plant groups are given in Table 3.

Discussion

Our analysis indicates that we can predict the occurrence of N. califor¬

nicum with a greater than 90 percent probability of being correct in areas

where indicator plant groups are present (ac conditional probabilities, Table

2) in the T. plicata/P. myrsinites h.t. near Elk River, Idaho. However, as

indicated by the ab conditional probabilities (Table 2), we cannot say that

wherever the beetles occur, the indicator plants will occur. This means that

VOLUME 58, NUMBER 1

the range of N. californicum within the sample area exceeds that of the

indicator plant groups.

The higher ab probabilities in Area B indicate considerable overlap be¬

tween the ecological amplitude of N. californicum and the indicator plants.

The canopy-like coverage of the forest floor, coupled with the high occur¬

rence of old-growth grand fir (mean ± SD X =117 years ± 17) with E. tinc-

torium decay combine to provide macro- and microenvironmental factors

necessary for the occurrence and propagation of N. californicum. In con¬

trast, many of the host grand fir in Area A are colonized fortuitously in that

N. californicum invades logging wounds. Many of these wounds will not

provide suitable overwintering sites, as the preferred overwintering site is

the root collar below the duff (Osborne and Kulhavy, 1975). As Area A

recovers from the effects of logging and stand disturbance, the numbers of

grand fir colonized by N. californicum should diminish.

As N. californicum is often cryptically concealed on its host or is present

only below the duff (Osborne and Kulhavy, 1975), finding the indicator plant

groups in areas of grand fir with E. tinctorium decay will aid in locating N.

californicum. The plant groups from Area B indicate the preferred habitat

of N. californicum are cool air drainages in the T. plicata/P. myrsinites h.t.,

while those of Area A are indicative of marginal sites for N. californicum.

Literature Cited

Arnett, R. H., Jr. 1968. The Beetles of the United States (a Manual for Identification). American

Entomological Institute, Ann Arbor, Michigan, xii + 1112 pp.

Daubenmire, R., and J. B. Daubenmire. 1969. Forest vegetation of eastern Washington and

northern Idaho. Wash. State Univ. Agric. Exp. Stn., Tech. Bull., 60:1-104.

Hatch, M. H. 1961. The beetles of the Pacific Northwest. Part III: Pselaphidae and Di-

versicornia I. University of Washington Press, Seattle, 503 pp.

Hitchcock, C. L., and A. Cronquist. 1973. Flora of the Pacific Northwest. University of

Washington Press, Seattle, 730 pp.

Miller, D. L. 1974. Root decays and their interaction in Idaho grand fir. Ph.D. Dissertation,

University of Idaho, Moscow, Idaho, 58 pp.

Osborne, H. L., and D. L. Kulhavy. 1975. Notes on Nosodendron californicum Horn on

slime fluxes of grand fir, Abies grandis (Douglas) Lindley, in northern Idaho (Coleoptera:

Nosodendridae). Coleopt. Bull., 29:71-73.

Sokoloff, A. 1959. The habitat-niche of American Nosodendridae. Coleopt. Bull., 13:97-98.

Sokoloff, A. 1964. Studies on the ecology of Drosophila in the Yosemite region of California.

V. A preliminary survey of species associated with D. pseudoobscura and D. persimilis

at slime fluxes and banana traps. Pan-Pac. Entomol., 40:203-218.

Zack, R. S., E. J. Davis, and K. Raffa. 1979. A new host record and notes on Nosodendron

californicum Horn (Coleoptera: Nosodendridae). Coleopt. Bull., 33:74.

Footnotes

1 Published with approval of the Director, Forest, Wildlife and Range Experiment Station,

University of Idaho, Moscow, as contribution No. 0237. Supported in part by Stillinger Me¬

morial funds.

PAN-PACIFIC ENTOMOLOGIST

2 School of Forestry, Stephen F. Austin State University, Nacogdoches, Texas.

3 Current address: Department of Lands, Box 670, Coeur d’Alene, Idaho.

4 Current address: School of Forestry, Oregon State University (stationed at Medford, Or¬

egon), Medford.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 59-63

THE PARASITOID TRIOXYS TENUICAUDUS STARY

(HYMENOPTERA: APHIDIIDAE) ESTABLISHED ON THE

ELM APHID TINOCALLIS PLATANI KALTENBACH

(HOMOPTERA: APHIDIDAE) IN

BERKELEY, CALIFORNIA

William Olkowski and Helga Olkowski

The John Muir Institute, 1010 Grayson Street,

Berkeley, California 94710

Robert van den Bosch 1 and Richard Hom

Division of Biological Control,

University of California, Albany 94706

Robert Zuparko and William Klitz

The John Muir Institute, 1010 Grayson Street,

Berkeley, California 94710

Tinocallis platani Kaltenbach is an arboricolous aphid in the Drepanosi-

phinae, tribe Phyllaphidini. Members of this tribe have alate fundatrigeniae

and viviparae (Eastop, 1977). This species is monophagous on elms {Ulmus

spp.) and is known widely from Europe and Russia (Mackauer and Stary,

1967). Stary (1966) lists it attacking Ulmus effusa Willd. in Czechoslovakia.

It has been introduced into western North America and is known there from

British Columbia, Utah and California (Richards, 1967). High summer pop¬

ulations of the aphid excrete sizable amounts of honeydew, which causes a

nuisance and irritation for people beneath elms planted as shade trees.

In Berkeley, California insecticides were used against this aphid from

about 1945 to 1971. Two treatments were applied per season, diazinon being

the material reportedly used in recent years. Berkeley maintenance person¬

nel report that these applications had decreasing effectiveness. In 1971,

water-washing of the trees to reduce aphid populations was substituted for

insecticide use as an interim measure while parasitoid introduction pro¬

ceeded.

In 1972, R. van den Bosch collected two parasitoid species from elm

aphids in the area of Prague and South Moravia, Czechoslovakia. These

parasitoids were identified as Trioxys hortorum Stary and a new species,

Trioxys tenuicaudus Stary (Carver and Stary, 1974; Stary, 1978). Both were

released on Los Angeles Street in Berkeley. Contrary to earlier reports

(Olkowski et al., 1976), T. hortorum has not become established. This paper

PAN-PACIFIC ENTOMOLOGIST

documents field observations prior and subsequent to parasitoid releases,

showing the establishment and spread of T. tenuicaudus.

Materials and Methods

Three areas were sampled in this study. Los Angeles Street with 35 elms

was the release site. A half-mile away is Hopkins Street with a group of

approximately 50 elms, and located 3 miles across town on Ashby Avenue

are 5 elms. By sampling the sites at Hopkins Street and Ashby Avenue, we

were able to record the dispersal of the aphid from its point of release.

Aphid population size was assessed by randomly sampling leaves from

the canopy of selected trees at each of the study sites and counting the

number of aphids on each leaf. For the years 1971, 1975, 1976 and 1979,

mean values are presented based on sample sizes of 80, 40, 40 and 80 leaves,

respectively, for each sample date. Ladders or mechanized lifts were used

to reach tree canopies.

After collection in Czechoslovakia, parasitoids were shipped to the Uni¬

versity of California Division of Biological Control quarantine laboratory at

Albany, where they were reared on field-collected twigs infested with aphids.

Fj adults were released directly onto field populations in 1972. Aphids from

the field were dissected to assess parasitism. Leaves with aphids were col¬

lected, placed in plastic bags, and transported to the laboratory for dissec¬

tion. In 1974 a mean of 39.2 aphids were dissected on each of five dates,

with total dissections ranging from 22 to 75. In 1975, 50 aphids were dis¬

sected on each of 3 dates. In 1976 a mean of 28.3 aphids were dissected on

each of 10 dates, with 3 to 50 dissections per date. Those dates for which

sample sizes were from 3 to 11 (or less than 50) are indicated in Fig. 1. In

1975, 25 aphids were sampled and dissected on each of 19 dates.

Results

In May 1971, when aphid monitoring of elms was initiated for the first

time in many years, no insecticide treatments had been applied. The un¬

treated aphid populations produced copious honey dew excretions, resulting

in numerous citizen complaints to the city about trees. Sidewalks beneath

such trees became exceedingly sticky and darkened by honey dew rain.

Leaves became sticky to touch. Starting on June 4 and June 14, 1971, more

detailed population counts were made on four trees with particularly heavy

aphid populations along Los Angeles Street. An average for the combined

samples for June 4 was 14.8 aphids/leaf and for June 14 was 42.1 aphids/leaf,

with 100% of the leaves infested on both dates. These pre-importation num¬

bers, shown in the box on Fig. 1, serve as a rough scale of comparison to

population levels after the parasitoid introductions which occurred the fol¬

lowing year. The vertical axes in the graphs of mean number of aphids per

7c Parasitism by Trioxys tenuicaudus and Mesidiopsis sp.* Mean Number of Aphids per Leaf.

Los Angeles Street

40 -

10 -

4 -

/ Preimportation

/ numbers

June 1971

1 -

No 1974

aphid counts

0 -

100 - 1974

80 -

60 -

- Trioxys tenuicaudus

Mesidiopsis sp.

40 -

No dissections

in early 1975

May June July Aug Nos

Los Angeles Street

Ashby Avenue

. 1 .

1976

J_l-1-1-1

1979

April May June July Aug Sept

Fig. 1. Population density of the Elm Aphid Tinocallis platani and degree of parasitism by Trioxys tenuicaudus and Mesidiopsis sp. in

Berkeley, California. * Sample sizes from 22 to 75 except those 1976 samples indicated by t where sample sizes were from 3 to 11.

VOLUME 58, NUMBER 1

PAN-PACIFIC ENTOMOLOGIST

leaf (Fig. 1) are log scales. The points in the graphs were determined by

calculating log(N + 1), where N = aphids/leaf.

In June and July 1972, 60 T. tenuicaudus adults were released on elms

along Los Angeles Street. In 1973, monitoring was started to determine

establishment of the released parasitoids. The only parasitoid detected in

these samples was a Mesidiopsis sp. (at the time believed to be M. subfla-

vescens Westwood, but we regard it as an undescribed species). This aphe-

linid appears to be common in and probably native to California, where it

is present on native oak aphids such as Tuberculatus sp. At the end of the

1973 season, a detailed examination of release trees on Los Angeles Street

was conducted. No Trioxys tenuicaudus parasitized aphids were found;

however, Mesidiopsis mummies were again common.

The first recoveries of T. tenuicaudus occurred in the spring of 1974 at

the release site. This was the first indication of a successful colonization.

Dissections later that season indicated parasitism as high as 28% (Fig. 1).

Aphid populations were low throughout the spring of 1975 except for the

week of June 14, when populations reached 6 aphids per leaf. T. tenuicaudus

parasitism in 1975 (ca. 20%) became evident in the autumn before leaf fall,

when aphid numbers were extremely high, between 35 and 40 aphids/leaf.

In 1976, T. tenuicaudus showed a high of 68% parasitism in the spring,

while Mesidiopsis sp., based on a few dissections, appeared in the fall.

Except for the fall peak in 1975, aphid numbers have remained low on Los

Angeles Street since, with no aphid complaints received.

In order to follow the spread of the parasitoid, aphids were sampled and

dissected from sites one-half mile and three miles from the release site on

Hopkins Street and Ashby Avenue respectively, where no parasitoids had

been released. Samples taken in June 1974 after the first recoveries at the

release site at Los Angeles Street revealed no parasitism at either Hopkins

or Ashby, based on 35 and 15 dissections respectively. In 1977, T. tenui¬

caudus was found for the first time on Hopkins Street, based on emergence

of the wasp from collected aphid mummies. In 1978, both T. tenuicaudus

and Mesidiopsis sp. were found on Hopkins Street, and T. tenuicaudus

appeared for the first time on Ashby Avenue. Thorough sampling on Ashby

Avenue in 1979, when the aphid population peaked at 19 aphids/leaf (Fig.

1) again revealed parasitism of both T. tenuicaudus and Mesidiopsis sp.,

with the imported parasitoid especially active throughout the first half of

the season.

Discussion

Trioxys tenuicaudus, an aphidiid parasitoid of the elm aphid Tinocallis

platani, was introduced in Berkeley in 1972. This parasitoid was first re¬

covered two years later at the release site. Resident complaints diminished

as aphid populations no longer reached numbers causing excessive honey-

VOLUME 58, NUMBER 1

dew drip. This constitutes the second instance of classical biological control

applied to a shade tree aphid pest.

The aphelinid Mesidiopsis sp., also found to parasitize the elm aphid,

showed a distribution often occurring separately from the introduced T.

tenuicaudus. Mesidiopsis generally occurred later in the season and over¬

lapped minimally with T. tenuicaudus. Further work is required to reveal

whether this pattern reflects an ecological preference for a given portion of

the season, a direct interaction between the species or a disequilibrium in

parasitoid-prey relations.

Spread of aphidiid parasitoids from the initial site of colonization may be

related to the stage of host attacked. Presumably, winged adults carry the

parasitoids to new locations. T. tenuicaudus prefers to attack early instars,

particularly I and II; we rarely found it in adult hosts. It spread very slowly,

taking six years to move a few blocks from the site of initial colonization.

In contrast, another aphidiid, T. curvicaudus Mackauer, which parasitizes

the linden aphid, Eucallipterus tiliae (Linnaeus), oviposits in adults and later

instars (Olkowski, in prep.). This parasitoid spread three miles across Berke¬

ley in a single season.

Literature Cited

Carver, M., and P. Stary. 1974. A preliminary review of the Aphidiidae (Hymenoptera:

Ichneumonoidea) of Australia and New Zealand. J. Aust. Entomol. Soc., 13:235-240.

Eastop, V. F. 1977. Worldwide importance of aphids as virus vectors. In K. F. Harris and

K. Maramorosch (eds.), Aphids as Virus Vectors. Academic Press, San Francisco, 559

pp.

Mackauer, M., and P. Stary. 1967. Hym. Ichneumonoidea. World Aphidiidae. In V. Delucchi

and G. Remaudiere (eds.), Index of Entomophagous Insects. Paris, 195 pp.

Olkowski, W., H. Olkowski, R. van den Bosch, and R. Horn. 1976. Ecosystem management:

A framework for urban pest control. BioScience, 26:384-389.

Richards, W. R. 1967. A review of the Tinocallis of the world (Homoptera: Aphidiidae).

Can. Entomol., 99:536-553.

Stary, P. 1966. Aphid Parasites of Czechoslovakia, A Review of the Czechoslovak Aphidiidae

(Hymenoptera). Academia, Publishing House of the Czechoslovak Academy of Sciences,

Prague, 242 pp.

Stary, P. 1978. Parasitoid spectrum of the arboricolous callaphidid aphids in Europe (Hy¬

menoptera: Aphidiidae; Aphidoidea: Callaphididae). Acta Entomol. Bohemoslov. 75:

164-177.

Footnote

1 Deceased.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 64-72

FLIGHT PERIODICITY IN COLORADO BITING MIDGES

(DIPTERA: CERATOPOGONIDAE) 1 2

Donald R. Barnard

Lone Star Tick Research Laboratory, Agricultural Research,

SEA, USDA, P.O. Box 588, Poteau, Oklahoma 74953

With the possible exception of some species of Forcipomyia, which act

as pollinators of cacao (Billies, 1941), biting midges (Diptera: Ceratopogon-

idae) in the genera Atrichopogon, Bezzia, Dasyhelea, Forcipomyia, and

Palpomyia have little if any striking economic importance compared with

pestiferous bloodsucking species (Wirth, 1956; Chan and LeRoux, 1967).

As a consequence we know much less of the basic biology and ecology of

species in these genera than we do for Culicoides and Leptoconops. Eco¬

nomic importance, however, is not an absolute prerequisite for study. The

diversity of adult feeding behavior within the Ceratopogonidae, for example,

poses an array of behavioral and biological problems which continue to

challenge contemporary observers of insect natural history (Wirth, 1956;

Downes, 1978).

In northeastern Colorado, virtually nothing is known about the flight hab¬

its of adult Ceratopogonidae. Consequently, I undertook this study to make

observations of seasonal abundance and diel patterns of flight activity for

Atrichopogon fusculus (Coq.), Bezzia pulverea (Coq.), B. setulosa (Loew),

Dasyhelea grisea (Coq.), D. mutabilis (Coq.), Forcipomyia bipunctata (L.),

F. brevipennis (Macquart), and Palpomyia tibialis (Meigen). In this paper

are reported the results of this study.

Materials and Methods

The flight activity of airborne Ceratopogonidae was assessed between

January and December 1978 by identifying and counting those collected with

a vehicle-mounted interception trap (Barnard, 1979). Throughout this period

collections were made once every 14 days and each time specimens were

collected over a 24-hr period. Diel flight activity was related to important

temporal events, using times of sunrise and sunset as reference points. Each

day of collection was divided into 20 periods: period one began at morning

nautical twilight and ended at sunrise; periods 2-11 were derived by dividing

the time between sunrise and sunset into 10 equal parts; period 12 began at

sunset and ended at the end of evening nautical twilight; the time from the

end of evening nautical twilight to the beginning of morning nautical twilight

was divided into 8 equal parts and formed periods 13-20. During the year,

VOLUME 58, NUMBER 1

periods one and 12 ranged from 59 to 74 min. Four collection runs were

made during each of these 2 periods: one at the beginning of the period, one

at 22 ± 5 min, one at 45 ± 5 min, and one at the end of the period. I made

3 collection runs, equally spaced in time during each of periods 2-11 and

13-20. A mean value for flight activity was calculated for each period by

summing the number of individuals collected in each run in a given period

and dividing by the number of runs for that period.

Throughout this study, each collection run was made over the same course

on a lightly travelled, graded-surface road 0.25 km west of Wattenburg,

Colorado (104°50'W, 40°01'N) in the South Platte River drainage system.

The course passed through partially flooded, low-lying areas and higher

irrigated pastures on which horses and cattle grazed. Two drylot dairies and

1 drylot sheep operation were located, respectively, 500 and 1000 m south,

and 1500 m west of the course. On each collection run I travelled the course

in one direction (2.0 km) and then returned to the starting point, a total

distance of 4.0 km. Runs were made at ca. 40 km/hr and lasted ca. 7 min.

During all dusk to dawn periods I made collection runs with only parking

lights on. Specimens collected during each run were stored separately and

temporarily (in the collection bag used for that run) in an ice chest over dry

ice.

Results

Data from observations of seasonal activity and diel patterns of flight

for each species of biting midge collected during this study are given in Ta¬

ble 1.

Atrichopogon

Seasonal flight activity of male and female A. fusculus commenced in

May and airborne females were most abundant in July. Population levels of

males diminished in June, but increased and remained high thereafter until

October. A. fusculus exhibited peak diel flight activity during periods 11

and 12, before or after sunset, and during periods 1 and 2 before and after

sunrise.

Forcipomyia

Forcipomyia bipunctata was active from April to October, population

levels of both sexes were highest in July. F. brevipennis commenced flight

activity in June and terminated activity in mid-November. Population levels

for females were high from July to September, whereas males were most

abundant in September. Few males or females of F. bipunctata or F. brev¬

ipennis were collected after September.

The diel flight period in F. bipunctata was bimodal from July to Septem¬

ber with the main peak at sunset and a smaller peak near sunrise. Adults

PAN-PACIFIC ENTOMOLOGIST

Table 1. Mean number of airborne adult Ceratopogonidae collected in each period in each

month, and the percentage of total numbers of adults collected in periods 1,2, 11, and 12, and

the mean percentage of total adults collected in periods 3 thru 10 and 13 thru 20. Wattenburg,

Colorado, 1978.

(SR = sunrise, SS

= sunset, PH

= photophase, SC

= scotophase)

Period

Month

Sex

1 -

-SR—2

3-10

11 -

-SS—12

13-20

Atrichopogon fusculus

May

1.4

3.2

4.5

0.2

1.2

1.3

4.0

June

0.2

1.3

1.1

3.5

3.2

1.4

July

2.9

1.4

5.6

0.2

4.4

1.8

14.1

19.9

1.1

August

2.2

3.3

3.1

0.2

7.1

4.0

1.1

September

1.3

12.9

1.6

7.1

1.3

1.1

October

1.1

1.3

1.1

0.2

1.3

0.2

% of total

6.2

10.2

5.2

55.1

19.7

3.6

5.1

7.2

3.2

39.9

39.0

5.6

Forcipomyia bipunctata

April

0.3

1.4

May

1.4

1.5

1.4

1.3

2.5

4.5

1.2

June

2.8

1.2

5.6

2.2

1.1

July

1.3

1.4

0.1

3.5

4.0

1.7

3.5

2.5

0.2

25.1

50.1

4.3

August

1.6

0.1

3.2

4.0

1.3

2.5

0.2

5.0

6.3

1.4

September

1.3

3.3

0.3

2.5

1.2

1.8

5.6

1.3

October

0.2

1.2

1.3

1.8

% of total

7.5

8.4

9.7

26.5

38.7

9.2

2.5

5.4

4.8

29.8

50.8

6.7

Forcipomyia brevipennis

June

1.6

4.5

1.8

1.2

1.3

VOLUME 58, NUMBER 1

Table 1. Continued

Period

Month

Sex

-SR—2

3-10

11—SS—

-12

13-20

July

6.3

4.9

1.3

0.3

4.5

2.5

1.3

August

0.2

3.2

2.2

0.5

0.1

2.2

4.9

1.1

September

35.5

3.5

0.1

14.1

9.8

October

0.1

2.8

0.2

1.8

November

2.3

% of total

2.7

77.2

17.5

2.6

4.4

45.8

41.9

5.3

Dasyhelea grisea

April

0.2

May

0.2

1.2

1.8

1.6

0.5

June

1.2

3.1

1.8

July

10.2

3.1

6.3

1.8

2.2

31.6

6.2

56.2

39.8

August

2.8

3.2

4.3

19.9

2.5

7.9

4.7

63.1

12.6

September

1.8

1.4

89.1

31.6

5.9

3.1

501.2

158.5

October

1.1

3.5

31.6

% of total

1.5

9.2

6.3

64.4

18.6

0.4

5.3

2.3

69.6

22.4

Dasyhelea mutabilis

April

2.9

1.9

May

12.2

4.0

1.3

3.2

36.5

31.6

June

1.3

81.7

1.6

3.1

365.2

15.8

1.3

July

5.0

31.6

54.6

79.4

9.9

3.9

125.8

158.4

147.3

141.3

August

3.9

7.9

29.8

75.8

38.0

1.4

10.4

74.9

141.2

103.1

PAN-PACIFIC ENTOMOLOGIST

Table 1. Continued

Period

PH SC

Month

Sex

1—SR-

-2

3-10

11 -

-SS—12

13-20

September

2.9

35.5

1258.9

2.8

7.6

122.3

757.6

288.3

October

8.7

3.2

105.9

125.9

3.2

November

0.5

1.1

% of total

0.5

2.5

12.9

81.1

3.0

5.4

31.1

43.8

19.6

Palpomyia tibialis

May

1.4

10.0

June

1.6

2.5

34.6

1.3

July

9.7

0.9

% of total

. M

100

6.4

89.7

3.6

Bezzia pulverea

May

1.3

2.2

9.7

June

3.0

7.0

8.2

2.5

35.4

6.4

37.9

53.7

5.5

July

3.3

1.6

7.1

August

3.6

2.1

September

1.6

% of total

12.1

41.5

44.8

1.6

1.5

23.1

3.8

28.9

38.9

3.8

Bezzia setulosa

May

1.6

2.0

1.8

12.7

June

7.1

1.8

. 2.5

15.8

17.8

2.2

July

13.9

1.3

0.2

1.8

28.8

24.5

1.4

August

31.6

1.0

281.8

354.8

5.0

97.8

1.2

56.2

292.9

19.9

September

1.4

12.6

1.6

9.8

2.5

October

1.4

% of total

4.6

0.2

44.1

50.3

0.8

17.2

19.2

59.0

4.0

VOLUME 58, NUMBER 1

were active throughout the scotophase in July, and during the early parts

of the scotophase in August and September. F. brevipennis exhibited a

unimodal flight period with peak activity in period 11 (period 12 for females

in August). High levels of flight activity took place during the photophase

in June and flight during the scotophase was noted in July and August.

Dasyhelea

Dasyhelea grisea was active from April to October, while D. mutabilis

commenced flight activity in April and ceased activity in November. D.

mutabilis population levels were highest between June and October, and

levels for each sex fluctuated only slightly during this time. D. grisea pop¬

ulation levels were highest from mid-July to mid-September.

D. grisea showed a bimodal diel flight period from July to September,

with the main peak of activity observed before sunset and the second peak

after sunrise. In October and April, flight occurred principally in the late

afternoon, whereas activity in May and June was greatest during the morn¬

ing hours. From May to October, flight activity by D. mutabilis took place

between periods 1 and 12; adults were never collected at night. In April,

flight occurred during midday only, and in November, only females were

collected, these in periods 10 and 11.

Bezzia

Both sexes of B. setulosa were active between May and September; only

females were collected in November. Overall B. setulosa population levels

were highest in August, and except for this month and September, when

males were most abundant, females dominated collections. B. pulverea was

active between May and September. Males and females were most abundant

in June, after which population levels decreased; males were not collected

after August.

Between May and November, both sexes of B. setulosa were active in

periods 11, 12, or both (before and after sunset) but a preference was shown

for neither period. Activity diminished in period 13, and periods 14-20 and

period 1 (before sunrise) were devoid of flight activity in all months except

July. In July, B. setulosa females were active sporadically throughout the

scotophase. In August and September, males and females were active in

the morning during period 2 (following sunrise) and in June flight activity

by females continued into period 4 (ending at 1000). Bezzia setulosa was

generally inactive during the daytime hours.

In June, B. pulverea was active throughout the photophase but activity

by both sexes peaked in period 12 (after sunset) and by females again in

period 2 (following sunrise). Also in June, flight continued into period 15

(ending at 2405) and commenced again during the morning crepuscule (pe¬

riod 1). The principal flight time between July and September was period

11 (before sunset) as was period 12 in May, although activity by females

PAN-PACIFIC ENTOMOLOGIST

extended into period 16 in July. Male B. pulverea were not collected after

July.

Palpomyia

P. tibialis was active between May and July; female population levels

peaked in June, the only month in which males were collected.

P. tibialis exhibited a unimodal diel flight period. Females were active

principally during the evening crepuscule between May and July, but in

June and July flight continued for several hours into the scotophase. Fe¬

males were also active during period 11 (before sunset) in May and June

and during period 9 in June. Male P. tibialis were collected only during

period 12 in June.

Discussion

The diel flight period in biting midges collected during this study was

seasonally variable. Flight periods observed in April, October, and Novem¬

ber, for example, appeared to be temperature-mediated; temperatures in

these months regularly fall to below 5°C during the mid-late scotophase and

early photophase (Barnard, unpubl. data). In contrast, flight periods ob¬

served in July and August when air temperatures were ^10°C, presumably

are those expressed in the absence of flight-inhibiting low temperatures.

Such seasonally-influenced flight periods are best illustrated by flight activity

data for A. fusculus, F. bipunctata, and D. grisea (and to a lesser extent,

D. mutabilis ). In these species, a secondary peak of flight activity at sunrise

was not observed until July. Moreover, in these species diurnal flight activ¬

ity during the summer is displaced toward either end of the photophase, in

an apparent response to increasing daytime temperatures.

The influence of season on flight period is less apparent for other species.

In F. brevipennis, for example, flight does not extend into the scotophase

until July and August. Bezzia setulosa exhibits a unimodal flight period in

May and October, a bimodal flight period in August and September, and a

unimodal period for males and bimodal period for females in June and July.

Flight activity in P. tibialis and B. pulverea was of too little duration and

concentrated in one month to be affected by season (64 and 83%, respec¬

tively, of all P. tibialis and B. pulverea were collected in June).

I characterized diel flight during periods of biting midges collected during

this study by calculating the percentage of total males and females in flight

in each of periods 1, 2, 11, and 12; and for periods 3 thru 10 and 13 thru 20,

by calculating the mean percentage of total males and females in flight per

period. The percentages thus calculated are weighted in favor of months in

which flight activity was most concentrated; however, it is in these months

that conditions for flight are most favorable and the flight activity observed

free from the inhibiting effect of low temperature. Each of the biting midge

VOLUME 58, NUMBER 1

species collected was active principally during the late afternoon and eve¬

ning crepuscule. Atrichopogon fuse ulus males, females of F. bipunctata and

B. pulverea, and males and females of D. grisea and B. setulosa, in addition,

showed a morning-crepuscular flight peak of lower amplitude than the eve¬

ning peak. Males and females of each species except male P. tibialis (which

were collected only during period 12) were active during the daytime and

all species except male P. tibialis, D. grisea, and D. mutabilis were active

at night. Consistent with their flower-seeking food habits, Dasyhelea species,

particularly female D. mutabilis were active primarily during daylight hours.

Kaufmann (1974) described the diel flight period of Forcipomyia inorna-

tipennis (Austen), a cacao pollinator in Ghana, as bimodal with peak activity

at dawn and dusk. And because of their biological diversity and in some

cases economic importance (Bystrak and Wirth, 1978), other Forcipomyia

species have been studied in detail. Hematophagous species suck blood

from various insects and from phalangids (Wirth, 1956; Wirth and Stone,

1973), whereas species in the subgenus Euprojoannisia are important pol¬

linators of cacao (Billies, 1941; Saunders, 1924; Bystrak and Wirth, 1978)

and Para rubber (Wirth, 1956).

Little is known about the feeding habits of species in the remaining gen¬

era. Some species of Atrichopogon (e.g., A. epicautae Wirth, A. farri Wirth,

A. meloesugans Kieffer, and A. oedemararum Stora) suck the blood of

oedemerid and meloid beetles (Downes, 1955; Wirth and Stone, 1973; Wirth,

1956). Other species, such as A. pollinivorus Downes and A. pavidus (Win-

nertz), fly to flowers and feed solely upon nectar and pollen (Downes, 1955).

In Bezzia and Palpomyia, flight at dawn and dusk apparently serves at least

2 functions: meeting of the sexes and procurement of food (Downes, 1978).

Downes (1978) observed flight activity by B. setulosa in late afternoon and

noted several instances of this species preying upon male chironomid midges

and one instance of B. setulosa feeding upon B. setulosa. He also noted

that Palpomyia and Bezzia spp. feed upon chaoborids, chironomids (Dip-

tera), and upon baetids (Ephemeroptera), each of which as a taxon was

most active in this study in periods 10-12, the same time P. tibialis, B.

setulosa, and B. pulverea adults were most active.

Acknowledgment

Dr. Willis W. Wirth, Systematic Entomology Laboratory, AR, SEA,

USDA, Washington, D.C. provided the species determination for Atricho¬

pogon, Bezzia, Dasyhelea, Forcipomyia, and Palpomyia.

Literature Cited

Barnard, D. R. 1979. A vehicle-mounted insect trap. Can. Entomol., 111:851-854.

Billies, D. J. 1941. Pollination of Theobroma cacao in Trinidad, B. W. I. Trop. Agric.

(Trinidad), 18:151-156.

PAN-PACIFIC ENTOMOLOGIST

Bystrak, P. G., and W. W. Wirth. 1978. The North American species of Forcipomyia,

subgenus Euprojoannisia (Diptera: Ceratopogonidae). U.S. Dep. Agric., Tech. Bull.,

1591 : 1 - 51 .

Chan, K. L., and E. J. LeRoux. 1967. Ecological studies on three pond midges (Diptera:

Ceratopogonidae) in Quebec. Ann. Entomol. Soc. Que., 12:14-68.

Downes, J. A. 1955. The food habits and description of Atrichopogon pollinivorus sp. n.

(Diptera: Ceratopogonidae). Trans. R. Entomol. Soc. Lond., 106:439^148.

Downes, J. A. 1978. Feeding and mating in the insectivorous Ceratopogonidae (Diptera).

Mem. Entomol. Soc. Can., No. 104, 62 pp.

Kaufmann, T. 1974. Behavioral biology of a cocoa pollinator, Forcipomyia inornatipennis

(Diptera: Ceratopogonidae) in Ghana. J. Kan. Entomol. Soc., 47:541-548.

Saunders, L. G. 1924. On the life history and the anatomy of the early stages of Forcipomyia

(Diptera: Ceratopogonidae). Parasitology, 16:164-213.

Wirth, W. W. 1956. New species and records of biting midges ectoparasitic on insects (Dip¬

tera: Heleidae). Ann. Entomol. Soc. Am., 49:356-364.

Wirth, W. W., and A. Stone. 1973. Aquatic Diptera. Pp. 293-372 in R. L. Usinger, Aquatic

Insects of California. University of California Press, Berkeley, 508 pp.

Footnotes

1 This research was undertaken while the author was postdoctoral fellow, Department Zo¬

ology-Entomology, Colorado State University, Ft. Collins, and was supported in part by Co¬

operative Agreement No. 12-14-5001-257 with AR, SEA, USDA.

2 This paper reports the results of research only. Mention of a proprietary product does not

constitute a recommendation or an endorsement by the USDA.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 73-78

PROTAGROTIS OBSCURA BARNES AND McDUNNOUGH

(LEPIDOPTERA: NOCTUIDAE): A PEST OF GRASSES

GROWN FOR SEED IN THE PACIFIC NORTHWEST 1

J. A. Kamm 2

Agricultural Research, Science and Education Administration,

USD A, Corvallis, Oregon 97331

Several species of cutworms and other Lepidoptera are sporadic pests of

bluegrass and fine fescue grown for seed in eastern Oregon and Washington

(Crawford and Harwood, 1964; Oetting, 1977). For years, Oregon grass seed

producers in Union County were thought to have a serious pest problem

with Cry modes devastator (Brace) (Noctuidae), based on larval determi¬

nations. In April 1976, many fields were badly damaged by larval feeding,

and over 100 larvae were collected that appeared to be C. devastator. These

larvae were reared to adults in the laboratory; 2% percent were C. devas¬

tator , and the remainder were Protagrotis obscura Barnes and Mc-

Dunnough. Adult and larval voucher specimens of these P. obscura have

been deposited in the collection of the U.S. National Museum, Washington,

D.C.

A study of P. obscura was undertaken to determine the seasonal cycle,

feeding damage, and certain aspects of its biology as a destructive pest of

grasses grown for seed.

Materials and Methods

The phenology of adults was determined with two battery-powered black-

light traps operated in commercial fields of bluegrass grown for seed near

La Grande, Oregon. Traps were operated 4 nights weekly during the moths’

flight season. Specimens for taxonomic study were obtained from these

light-trap collections or from larval specimens collected in the fields and

reared in the laboratory. The seasonal occurrence of the larval population

was determined by removal of cores of sod (20 cm diameter) at irregular

intervals during the year. Larvae were extracted from the sod with Berlese

funnels or removed by hand dissection of the sod. Some larvae removed

from the field were reared to adults on fine fescue using methods described

elsewhere (Kamm, 1970). The degree of parasitism and the sizes of head

capsules were determined from these field collections. The sizes of the head

capsules of larvae collected at different times during the season were de¬

termined by measuring across the widest portion of the head using a bin¬

ocular microscope fitted with an ocular micrometer.

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Adult (top) and larva (bottom) of P. obscura.

Observations

Description of life states .—Adults are about 2 cm long (body length) and

vary in color from reddish-brown to tan with tan markings on the wings

(Fig. 1). Darker colored moths have conspicuous light-tan markings on each

VOLUME 58, NUMBER 1

120

JUNE JULY AUGUST

Fig. 2. Black light captures of adult P. obscura in commercial seed fields of bluegrass, La

Grande, Oregon, 1979.

wing, which are difficult to discern on lighter-colored moths (Barnes and

McDunnough, 1911). Eggs are cream colored, sculptured, and shaped like

a pumpkin. They are 0.57 to 0.61 mm in horizontal diameter. Larvae are

grayish-white and have a brown head capsule. Mature larvae are about 2.5

cm long (Fig. 1). Larvae of P. obscura and C. devastator are so similar in

appearance that existing larval keys are not adequate to separate these

species.

Distribution. — P. obscura was found in seed production areas surround¬

ing La Grande in eastern Oregon and previously was reported in the Spo¬

kane area of eastern Washington (Crawford and Harwood, 1964). In 1978,

I collected larvae from a commercial seed field of bluegrass near Rockford,

Washington, and reared adults of P. obscura, thus, confirming the presence

of the species in eastern Washington. To my knowledge, this insect has

never been collected in western Oregon or Washington.

Seasonal history .—Adults began to emerge in June, and maximum num¬

bers occurred in early July, based on light-trap captures (Fig. 2). Adult

emergence in La Grande occurs close to the time of emergence in the Spo¬

kane area (Crawford and Harwood, 1964). The flight season of adults indi¬

cates that the species is univoltine. In general, the moths are nocturnal but

PAN-PACIFIC ENTOMOLOGIST

APRIL

MAY

JUNE

JULY

AUGUST

SEPT.

OCT.

NOV.

APRIL

MAY

JUNE

JULY

AUGUST

SEPT.

OCT.

NOV.

Fig. 3. Seasonal ^sequence of the life stages of P. obscura in commercial seed fields of

bluegrass, La Grande, Oregon, 1979.

readily fly when disturbed during the day. They are capable of strong sus¬

tained flight. Females reared from larvae in the laboratory had immature

ovaries the day of emergence and had a 6- to 7-day preoviposition period.

The females were first to emerge in the laboratory and preceded the emer¬

gence of the first males by about the length of the preoviposition period.

Eggs were deposited on leaves or within the leaf sheath of grass culms,

usually in clusters. The eggs hatched in 9 to 10 days in the laboratory (21°C).

In the field, the small larvae burrow into the plant crown where they actively

feed until the onset of cold weather in October. Larvae overwinter in the

plant crown but do not construct a hibernaculum. Feeding resumes in early

April, and mature larvae begin to pupate in the crowns of grass during May.

One lot of 205 larvae collected from the field on April 13 and reared in the

laboratory required an average of 26 days to pupate (range 22-36) and re¬

mained in the pupal stage an average of 28 days (range 24-34). The seasonal

occurrence of life stages is shown in Fig. 3.

Larvae collected from the field in October and reared in the laboratory

were not in an intense diapause. Most larvae molted within 3 to 4 days and

then resumed feeding. One lot of 28 larvae collected from the field on Oc¬

tober 11 had a mean head capsule width of 1.7 mm (range 1.2 to 2.3).

Another lot of 147 larvae collected from the field on April 15 had a mean

head capsule width of 3.0 mm (range 2.2 to 3.9) and about one-half were

actively feeding. The head capsule width of several full grown larvae av¬

eraged 4.3 mm. These observations and measurements demonstrate that

larvae clearly resume feeding the following spring after wintering as partially

grown larvae.

Host plants and feeding damage .—The presence of larvae in infested

VOLUME 58, NUMBER 1

fields is difficult to detect during July and August. Larval feeding damage

becomes evident in September and October, when partially or entirely dead

crowns appear in the field. Larvae are often found in the remaining green

part of partially damaged crowns. In effect, the larvae sever the roots from

the shoots in the crown, this being accompanied by an accumulation of frass.

Larvae do not move about on the foliage or burrow into the soil, but remain

in the crown as long as food is available there. When larval feeding results

in the death of the plant, the larvae move to adjacent plants.

In general, feeding damage is most severe during October and April be¬

cause the larvae are larger at these times. Limited larval feeding probably

occurs from November to April during periods of warmer temperatures.

The grass tillers that grow in late summer and early fall must be vernalized

by winter temperatures to produce seed the following year. Destruction of

these tillers, either in the fall or spring, will reduce seed yields. Dense

infestations of larvae may also damage the stand to the point that the field

must be reseeded. All varieties of bluegrass, fine fescue, and ryegrass grown

in Union County proved susceptible to infestation of P. obscura. Heavily

damaged grass fields are usually fall-plowed and reseeded to wheat. Most

larvae survive the tillage, and the wheat seedlings are destroyed by larval

feeding the following spring. Larvae have not been observed to complete

their life cycle on wheat or damage established wheat fields.

Parasites. —The ichneumonid parasite Lissonota clypeator montana

(Cresson) emerged from 6.3 percent of larvae of P. obscura collected from

the field and reared in the laboratory. No other species of parasite was

observed. In the Lissonotini (Banchinae), 8-12 species of Lissonota occur

in the Pacific Northwest and parasitize a wide range of caterpillars (Krom-

bein and Hurd, 1979). Among these, Lissonota montana (Cresson) is a

known parasite of C. devastator and Protagrotis obscura.

Discussion

The biological observations presented here are the first reported for this

univoltine cutworm P. obscura, a serious pest of grasses grown for seed.

Larval infestations must be controlled to maintain stands of grass and keep

fields productive. The lush regrowth that occurs when irrigation is resumed

after harvest of seed probably favors survival of the pest species; otherwise

the grasses remain dormant without irrigation until autumn rains.

There is the potential for rapid infestation of new seedings because adults

are strong fliers and since females have a 6-day preoviposition period, they

may go through an active dispersal phase before their ovaries mature and

oviposition can begin (Johnson, 1969). The mobility of females is probably

reduced once they become heavily laden with eggs and oviposition begins.

Infestations of P. obscura, Chrysoteuchia topiaria (Zeller) (Pyralidae:

Crambinae) and Chionodes psiloptera (Barnes and Busck) (Gelechiidae) were

PAN-PACIFIC ENTOMOLOGIST

common in the same field. In fact, large numbers of C. psiloptera were

encountered in fine fescue, a new host for this gelechiid. The larvae of this

latter species are easily distinguished from the other two (Oetting, 1977).

The feeding damage of C. psiloptera differs from that of P. obscura and C.

topiaria in that larvae of the former sever individual tillers at the base of

the plant but consume little of them. After these severed tillers desiccate

and turn brown, the plant appears to be predominantly green with a few

brown tillers scattered throughout the crown. Dense infestations eventually

kill the plant, and the entire crown turns brown. Unlike C. psiloptera, dam¬

age by larvae of P. obscura and C. topiaria first appears as large dead spots

in the crown, which gradually enlarges, killing the entire crown. Large dead

areas appear throughout the field when larval populations are dense. Dam¬

age usually appears first on the high points or other well-drained areas in

the field.

Acknowledgments

I thank R. W. Carlson for identification of Lissonota clypeator montana

(Cresson) and D. C. Ferguson for identification of P. obscura.

Literature Cited

Barnes, W., and J. McDunnough. 1911. New species and genera of North American Lepi-

doptera. J. N.Y. Entomol. Soc., 19:153-154.

Crawford, C. S., and R. F. Harwood. 1964. Bionomics and control of insects affecting

Washington grass seed fields. Wash. State Univ. Agric. Exp. Stn. Tech. Bull. 44, 25

pp.

Johnson, C. G. 1969. Migration and Dispersal of Insects by Flight. Methuen and Co. Ltd.,

London.

Kamm, J. A. 1970. Effects of photoperiod and temperature on Crambus trisectus and Cram-

bus leachellus cypridalis. Ann. Entomol. Soc. Am., 63:412-416.

Krombein, K. V., and P. D. Hurd, Jr. 1979. Catalog of Hymenoptera in America north of

Mexico. Smithsonian Institution Press, Washington, D.C., 1:1-1198.

Oetting, R. D. 1977. Immature stages and biology of Chionodes psiloptera a pest in bluegrass

fields (Lepidoptera: Gelechiidae). Pan-Pac. Entomol., 53:258-263.

Footnotes

1 Contribution of Agricultural Research, SEA, USD A, in Cooperation with the Agricultural

Experiment Station, Oregon State University. Technical Paper No. 5536 of the latter.

2 Mailing address: Legume and Grass Seed Production Laboratory, Department of Ento¬

mology, Oregon State University, Corvallis, Oregon 97331.

PAN-PACIFIC ENTOMOLOGIST

January 1982, Vol. 58, No. 1, pp. 79-80

CONFUSED FLOUR BEETLE AND OTHER COLEOPTERA

IN STORED MARIJUANA

Robert L. Smith and Carl A. Olson

Department of Entomology, University of Arizona, Tucson 85721

On 17 January 1979, we received a request for assistance from Mr. Harry

Merritt, a Douglas, Arizona, pest control operator, who services the Federal

Drug Enforcement Administration Building in that city. He explained that

clerical workers and administrative officers were complaining about large

numbers of small beetles which had become a nuisance in the offices. The

infestation had not declined in spite of scheduled pest control service and

Mr. Merritt suspected that the “evidence room” with its several tons of

confiscated marijuana ( Cannabis sativa Linnaeus) was the source of beetles.

Specimens provided by Mr. Merritt were Tribolium confusum Jacquelin du

Val. On 26 January 1979, we visited the facility to examine the infestation.

We found that virtually all of the large “evidence boxes” contained at least

some infested 1 kg bricks, and that all of the bricks in some boxes were

infested. Several infested bricks were broken and their contents carefully

examined. We found approximately 20% of the contained Cannabis seed

had been hollowed, presumably by Tribolium larvae. To test this, we set

aside several g of whole seeds in each of two petri dishes sealed with mask¬

ing tape. Upon examining this material several weeks later, we discovered

that one dish contained one adult confused flour beetle and a damaged seed.

In addition to Tribolium, we discovered numerous specimens of Trogo-

derma variabile Ballion and Attagenus megatoma (Fabricius) (Dermesti-

dae), and Adistemia watsoni (Wollaston) and Microgramme arga (Reitter)

(Lathridiidae). Representatives of these families were not uniformly distrib¬

uted as were the Tribolium. The dermestids were generally found on the

floor of the evidence room and the lathridiids on the tops of evidence boxes

and entrapped in the adhesive tape used to bind the bricks of Cannabis.

Since lathridiids are fungus feeders, we inquired as to the usual condition

of the illicit product when it was confiscated. Agents informed us that the

material was frequently of very poor quality and often moldy because it was

stored in the open air on the Mexican side of the border while awaiting

transportation to the U.S.

Arnaud (1974) surveyed the arthropods associated with Cannabis stored

in the Hall of Justice in San Francisco. He found one species of mite, four

species of flies, one species of Lepidoptera, and six species of beetles. Of

the beetles, only Microgramme arga was common to both surveys.

PAN-PACIFIC ENTOMOLOGIST

Acknowledgments

We wish to thank Harry Merritt for calling our attention to the problem;

J. M. Kingsolver, Systematic Entomology Laboratory, U.S.D.A., for iden¬

tifying the dermestids; and F. G. Andrews, California Division of Plant

Industry, for determining the lathridiids. Arizona Agricultural Experiment

Station ms. No. 3354.

Literature Cited

Arnaud, P. H. 1974. Insects and a mite associated with stored Cannabis sativa Linnaeus.

Pan-Pac. Entomol., 50:91-92.

A MESSAGE TO SUBSCRIBERS AND MEMBERS

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California Academy of Sciences, Golden Gate Park, San Francisco,

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Howell V. Daly

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Vol. 58

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THE

Pan-Pacific Entomologist

DOYEN—New Species of Tenebrionidae from Western North America (Coleoptera). 81

BYERS—A New Neopanorpa (Mecoptera: Panorpidae) from Laos .. 92

LANE, MILLER, and COLLINS—Ticks (Acari: Argasidae and Ixodidae) from the Cal¬

ifornia Channel Islands. 96

REIMER and GOEDEN—Life History of the Delphacid Planthopper Stobaera tricarinata

(Say) on Western Ragweed, Ambrosia psilostachya Decandolle, in Southern Cali¬

fornia (Hemiptera-Homoptera: Delphacidae). 105

KIMSEY—The Female of Pompilus ( Ammosphex) wasbaueri Evans (Hymenoptera: Pom-

pilidae)... 109

PARKER and BOHART—Notes on the Biology of Andrena ( Callandrena) helianthi

Robertson (Hymenoptera: Andrenidae). 111

KIMSEY and BACKUS—Observations on the Biology and Life History of the Net-winged

Midge Dioptopsis sequoiarum (Alexander) (Diptera: Blephariceridae). 117

DORR and NEFF —Pseudomasaris marginalis Nesting in Logs in Colorado (Hymenop¬

tera: Masaridae). 124

STEINER—Use of the Proboscis for Prey-piercing and Sucking by Sphecid Wasps of the

Genus Prionyx (Hymenoptera: Sphecidae), a Case of Convergent Evolution. 129

GARRISON —Paltothemis cyanosoma, a New Species of Dragonfly from Mexico (Odo-

nata: Libellulidae). 135

O’BRIEN and ATSATT—Life History and General Bionomics of Trirhabda sericotra-

chyla Blake (Coleoptera: Chrysomelidae) in Southern California. 139

HARTMAN and HYNES—The Immature Stages of Tipula simplex Doane and T. acuta

Doane (Diptera: Tipulidae). 153

STONE—The Peach Beetle, Cotinis mutabilis (Gory and Percheron), in California (Co¬

leoptera: Scarabaeidae). 159

SCIENTIFIC NOTES

SHAPIRO—A New Elevational Record for Pieris protodice in California (Lepidoptera:

Pieridae). 162

GIULIANI—Notes on a Collection of Intertidal Beetles from the Farallon Islands, Cal¬

ifornia . 163

CLARK, FROHLICH, and COMANOR—Shrike Predation on the Scorpion Anuroctonus

phaiodactylus (Wood) and on a Solpugid (Scorpionida: Vaejovidae; Solpugida). 164

SAN FRANCISCO, CALIFORNIA • 1982

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

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OFFICERS FOR 1982

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Frequency of Issue: Quarterly (January, April, July, October).

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

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PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 81-91

NEW SPECIES OF TENEBRIONIDAE FROM WESTERN

NORTH AMERICA (COLEOPTERA)

John T. Doyen

Department of Entomological Sciences,

University of California, Berkeley 94720

The species dealt with here belong to obscure genera which are rarely

encountered except by specialists, and are seldom mentioned in the litera¬

ture. The new species of Adelonia and Triphalopsis represent significant

range extensions for the genera. The Chilometopon exhibits characters un¬

usual for the genus, suggesting possible relationships with the tribe Thino-

batini of southern South America.

Observations were made with Leitz and Wild stereomicroscopes and an

American Optical compound scope. Genitalia were mounted on glycerine

jelly slides and drawn with an optical grid. The excellent illustrations of

beetles were executed by Carolyn Mullinex. The California Academy of

Sciences (D. Kavanaugh), San Francisco, and the American Museum of

Natural History (L. H. Herman), New York, and K. W. Brown, Stockton,

California, kindly loaned some of the specimens.

Chilometopon brachystomum, new species

(Fig. 1)

Form ovate, elytra slightly inflated, brachypterous; uniformly castaneous,

feebly shining, in life faintly obscured by thin deposit of powdery white

wax. Cuticle thin, fragile.

Head about as long as wide, broadest across eyes, slightly narrower across

epistoma just above antennae; epistoma impressed laterally between epi-

stomal sutures and supraorbital carinae; slightly emarginate on each side

just mesad of epistomal suture, then abruptly projecting anterad about Vio

head width, truncate anteriorly; epistomal suture distinct laterally, obsolete

medially; supraorbital carina distinct above eye, fading anteriorly and not

reaching margin of head; dorsal cranial surface set with shallow punctures,

separated by about one puncture diameter on vertex, becoming denser and

anastomosing into fine, irregular, longitudinal ridges between eyes, and be¬

coming contiguous along epistomal margin. Eyes slightly ovoid, bulging,

barely indented by epistomal canthus. Mentum hexagonal, twice as wide as

long, shallowly emarginate anteriorly. Antennae slender, long, reaching well

beyond base of pronotum; ratio of segment lengths (base to apex) 11:9:16:

9:8:7:7:6:6:5:8; basal 7 segments slightly serrate, becoming gradually broad-

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Chilometopon brachystomum female.

er; apical 4 segments sub-moniliform; 9th and 10th segments about as wide

as long; 11th 1.5 times longer than wide.

Pronotum subrectangular, broadest near middle; anterior and posterior

widths subequal, slightly broader than head, anterior margin slightly raised

except medially; anterior angles nearly 90°, slightly projecting with tuft of

about 6-8 yellow setae reaching to middle of eye; lateral margins sharp,

slightly raised, evenly arcuate almost to base, becoming straight just before

VOLUME 58, NUMBER 2

sharp, 90° posterior angles; posterior margin weakly bisinuate, slightly raised,

especially medially. Pronotal disk shallowly punctate; punctures about as

large as eye facet, separated by about one puncture diameter medially,

becoming contiguous and coarser on lateral thirds. Hypomera and proster¬

num set with shallow, coarse, nearly contiguous punctures. Prosternal pro¬

cess about Vs coxa width, abruptly declivous posteriorly.

Elytral width across humeri about 1.25 times width of pronotal base,

widest slightly behind middle, apices pointed; disk estriate, irregularly set

with shallow punctures about as large as eye facet and separated by about

one puncture diameter centrally, becoming smaller and sparser posteriorly.

Epipleura nearly glabrous, complete, with sharply raised margins; widest at

humerus, gradually narrowing to apex. Metathoracic wings reduced, about

2 /s length of elytra.

Mesosternum planar, not excavated between coxae, set with coarse, con¬

tiguous setigerous punctures; mesepisterna set with very coarse, contiguous

punctures; mesepimera glabrous or with a few, nearly obsolete punctures.

Metasternum and metepimera set with setigerous punctures separated by

about one puncture diameter. Abdominal sternites glabrous except for few,

obsolescent punctures near anterior margin of 1st sternite and fine sparse,

yellow, appressed setae, these becoming denser on 5th sternite. Abdominal

sternite length ratios (base to apex) about 33:30:25:15:18; intercoxal process

triangular, apex rounded.

Legs slender; femora slightly clavate; sparsely clothed with appressed

setae; tibia gradually enlarged apically, densely clothed with short, spinose

setae; tibial spurs about as long as penultimate tarsomere; tarsomere length

ratios as follows: fore tarsus 10:8:8:7:12; middle tarsus 12:7:7:5:13; hind tar¬

sus 20:9:7:16. Tarsal claws slender, about as long as 2nd hind tarsomere.

Aedeagus (Fig. 2) with tegmen inverted; median lobe sclerotized ventral-

ly, membranous dorsally, simply upcurved with sparse, apical setae. Median

lobe free, with linear, parallel baculi.

Body dimensions: elytral length 3.9-4.6 mm (6 6), 4.6-5.2 mm (9 9);

pronotal length 1.0-1.1 mm (dd), 1.0-1.3 mm (9 9); greatest elytral width

2.5-2.9 mm (d d), 2.8-3.6 mm (9 9); greatest pronotal width 1.3-1.6 mm

(dd), 1.5-1.7 mm (9 9).

Holotype male and 24 male, 24 female paratypes from Mexico, Baja Cal¬

ifornia del Norte, Bahia San Luis Gonzaga, April 3, 1973. J. Doyen, J.

Powell and S. L. Szerlip. The holotype is deposited in the California Acad¬

emy of Sciences, San Francisco. Paratypes are in the Essig Museum of

Entomology, University of California, Berkeley.

The beetles were collected at night on the surface of low, coastal dunes

and sand hummocks no more than a few meters above sea level. Other

species of Chilometopon are active during the hottest months of the year,

when they commonly aggregate at night on flowers of Petalonyx and other

PAN-PACIFIC ENTOMOLOGIST

Figs. 2—4. Male genitalia. Fig. 2. Chilometopon brachystomum, dorsal aspect of aedeagus

(a), lateral aspect (b), dorsal aspect of median lobe (c). Fig. 3. Adelonia insularis, lateral (a)

and dorsal (b) aspects of aedeagus. Fig. 4. Triphalopsis californicus, lateral (a, b) and ventral

(c, d) aspects of aedeagus and median lobe.

summer-blooming plants. The name brachystomum refers to the episto-

mum, which is shorter than in other species of Chilometopon.

Discussion.—Chilometopon brachystomum is distinguished from other

North American Trimytini by the very short medial epistomal lobe. In other

Trimytini the medial lobe is at least as long as the labrum. According to the

characters used by Casey (1907) to characterize the genera of Trimytini, C.

brachystomum would establish a distinct genus. However, in all features

except the size of the epistoma, it is exceedingly similar to other species of

Chilometopon. Moreover, the size and shape of the epistomal lobe is vari¬

able. In C. abnorme Horn, C. castaneum Casey and C. ensifer Casey the

medial lobe is about as long as the labrum and apically rounded. In C.

pallidum the medial lobe is shorter than the labrum and apically truncate,

as in Prometopion. The apical antennal segment of C. brachystomum is

unmodified in males. In this feature brachystomum is most similar to C.

pallidum Casey, in which the apical segment is only slightly elongate in

males. No other described species of Chilometopon are brachypterous.

Clearly, the short winged condition in C. brachystomum is of relatively

recent origin, since the wings are only slightly reduced in size and still show

VOLUME 58, NUMBER 2

distinct anterior venation. The metanotum is essentially unmodified in struc¬

ture.

Blaisdell (1943) recorded three similar species of Chilometopon from Baja

California, namely C. castaneum Casey, C. rugiceps Blaisdell and C. crib-

ricolle Blaisdell. In all these species the medial epistomal lobe is at least as

long as the labium, and arcuate or arcuato-truncate anteriorly. In males of

these three the apical antennal segment is at least as long as the preceding

3 segments combined. The tribe Trimytini is presently under study by Mr.

William MacLachlan, University of Arizona. Consequently, no key to species

is provided here.

In general appearance C. brachystomum resembles closely Thinobatis

Eschscholtz, with which it shares the abbreviated epistoma, very similar

aedeagus, metendosternite, tentorium and mouthparts. Thinobatis is apter¬

ous, differing from Chilometopon in associated thoracic features, has the

eyes entire anteriorly (very weakly emarginate in Chilometopon ) and has a

distinct tooth on the middle of the dorso-lateral ridge of the mandible. The

middle tooth is not developed in Chilometopon, but similar teeth are present

on both mandibles of Trimytis. The dorsal surface of the mandibles of Pro-

metopion are thickened and coarsely sculptured, as in Trimytis, but lack

distinct teeth. Lacordaire (1859) distinguished his Thinobatides from Tri-

bolocarides (=Trimytini) primarily by the structure of the epistoma—rela¬

tively short and broad in Thinobatini; trilobed and exposing the bases of the

mandibles in the Trimytini. It is premature to suggest that Thinobatini and

Trimytini are synonymous, but the character distributions described above

indicate that the higher classification of these and related tribes needs to be

critically reexamined.

Adelonia insularis, new species

Form elongate, flattened, winged; body uniformly brownish black, shin¬

ing, with castaneous appendages.

Head subhexagonal in dorsal view, broadest across posterior margin of

eyes, anterior epistomal margin evenly, shallowly emarginate, lateral epi¬

stomal margins nearly straight, then converging just before eyes; epistomal

sutures distinct laterally, obscured medially; dorsal cranial surface uniform¬

ly set with shallow punctures slightly smaller than eye facet and separated

by about one puncture diameter, becoming finer, more closely set along

anterior epistomal margin. Eyes reniform, ventral lobe about twice as large

as dorsal. Mentum subquadrate, about 1.5 times wider than long, lateral

margins arcuate, slightly recurved, almost parallel just before base. Anten¬

nae clavate, reaching about distance to pronotal base; ratio of segment

lengths (base to apex) 10:5:8:6:6:5:7:7:7:7:10; segments 2-6 submoniliform,

segments 7-11 about 1.4-1.5 times broader than long, asymmetrical and

larger medially; segment 11 ovoid, 1.1 times longer than broad.

PAN-PACIFIC ENTOMOLOGIST

Prothorax subquadrate, 1.1 times broader than long, about 1.1 times wider

across base than apex; anterior border shallowly, evenly emarginate with

raised margin laterally; anterior angles nearly 90°, rounded; lateral margins

slightly, evenly arcuate, with narrowly upturned, rounded margin; posterior

angles slightly obtuse, slightly rounded at apex; posterior margin faintly

bisinuate with narrow raised margin. Pronotal disk shallowly punctate;

punctures about as large as eye facet, separated by 1-2 puncture diameters

medially, becoming gradually contiguous and reticulate near lateral margins;

shallow round foveae located near posterior margin about halfway from

angles to midline, and 2nd pair usually near lateral margins about 5 /9 distance

to posterior angle; shallow transverse depression located in medial Vx near

posterior margin. Hypomera scabrous except for smooth coxal cowling;

prosternum finely punctatorugose, becoming finely, sparsely punctate be¬

tween coxae; prostemal process declivous behind coxa, truncate, finely,

sparsely punctate.

Elytral width subequal to pronotal width at base, nearly parallel sided to

third abdominal segment, 10 striate; striae set with rounded, posteriorly

open, slightly transverse punctures separated by 1.5-2 times puncture di¬

ameter; puncture diameter on disk about 4 times that of pronotal punctures,

decreasing to half this size on declivity; interstriae finely, sparsely punctate,

obtusely rounded; 7th-9th interstriae weakly inflated in humeral region; epi-

pleura strongly elevated, forming deep gutters with narrowly rounded mar¬

gins; broadest basally, gradually narrowing to elytral apex; finely, sparsely

and obscurely punctate.

Mesosternum shallowly concave before coxae, set with shallow punctures

separated by about one puncture diameter and bearing short, declined setae;

mesopleura coarsely punctatorugose; metasternum, metepistema, and ab¬

domen set with fine punctures separated by about 1 puncture diameter on

metepisterna and sternite 5, by 2-3 diameters on metastemum and sternites

1-4; intercoxal process triangular, apically rounded.

Legs robust, femora inflated, tibiae slender, gradually enlarged to apex;

mesofemoral tooth short, blunt, obtusely triangular; metafemoral tooth about

as long as tibial width, right angled or slightly acute.

Aedeagus (Fig. 3) with tegmen dorsal, mostly membranous ventrally; me¬

dian lobe adnate, not protrusible.

Body dimensions: elytral length 5.2-6.2 mm ($9), 5.1-6.0 mm (6 6);

pronotal length 1.7-2.0 mm (9 9 and 6 6); greatest elytral width 2.7-3.0

mm (9 9), 2.6-3.1 mm (dd); greatest pronotal width 2.4-2.9 mm (9 9);

2.3-2.8 mm (dd).

Holotype male and 20 male, 32 female, 14 unsexed paratypes from Mex¬

ico, Tres Marias Islands, Magdalena Island, May 20, 1925. H. H. Keifer.

Holotype and 50 paratypes, California Academy of Sciences; 15 paratypes,

Essig Museum of Entomology, University of California, Berkeley.

VOLUME 58, NUMBER 2

Discussion.—Adelonia insularis is similar to A. sulcatula (Champion),

differing in the following features: in insularis the epistomal margin is shal¬

lowly, evenly emarginate; the pronotal disk is punctate medially, reticu-

lately punctate laterally; the meso- and metapleura and sterna and abdomen

are punctate; the mesofemoral tooth is very blunt and obtuse, the meta-

femoral tooth right angled. In sulcatula the epistomal margin is almost

straight; the pronotal disk is reticulately punctate medially, rugulose later¬

ally; the pleura, sterna and abdomen are set with squat, flattened tubercles,

at least laterally; the femoral teeth are larger and usually more acute. In

Adelonia costipennis (Blair) from the Galapagos Islands the mesofemora

lack teeth and the metafemoral teeth are very obtusely dentate. Adelonia

filiformis Laporte from southern Baja California, is only 5-6 mm long, has

the pronotum evenly, sparsely punctate and has much smaller elytral punc¬

tures.

The collection of Adelonia made by Keifer contained 9 individuals of

sulcatula, as well as the series of insularis. The specimens of sulcatula are

not significantly different from populations of the same species from Mexico

and Central America, suggesting a recent, probably historical, introduction

onto the Tres Marias Islands.

Key to the Species of Adelonia Excluding South America

1. Mesofemora armed with distinct teeth on mesal surface. 2

Mesofemora without teeth . costipennis (Blair)

2. Pronotum, mesosternum, metastemum and abdomen punctate .... 3

Pronotum, mesosternum and metasternum punctate medially, tu-

berculate laterally. sulcatula (Champion)

3. Elytral interstriae convexly rounded . 4

Elytral interstriae flat . quadricollis (Champion)

4. Pronotal disk with punctures separated by about 1 puncture diam¬

eter, slightly closer laterally, but never reticulate; lateral margins

without foveae . filiformis Laporte

Pronotal disk with punctures separated by less than 1 puncture di¬

ameter medially, becoming reticulately punctate laterally; lateral

margins usually with distinct foveae near middle ... insularis, n. sp.

Triphalopsis calif or nicus, new species

(Fig. 5)

Form ovate, elytra moderately inflated, apterous; color uniformly casta-

neous, feebly shining; dorsum sparsely clothed with slender, pale, erect

hairs about as long as combined length of antennal segments 2 and 3.

Head slightly deflexed, about Vs wider than long, broadest across eyes;

epistoma trilobed, with lateral lobes broadly, evenly rounded, separated

from medial lobe by deep emarginations; medial epistomal lobe about 1.3

PAN-PACIFIC ENTOMOLOGIST

Fig. 5. Triphalopsis californicus, female.

times broader, about 4 times longer than lateral lobes, arcuately triangular

with serrate margin; epistomal suture absent; low supraorbital carinae ex¬

tending from middle of eyes almost to epistomal emarginations; dorsal cra¬

nial surface sparsely, finely punctate; vertex and frons carinulate, with ridges

occasionally anastamosing, becoming obsolete medially just behind strigate

VOLUME 58, NUMBER 2

epistomal lobe; eyes slightly ovoid, indented about l A by epistomal canthus.

Mentum hexagonal, about twice as wide as long, punctatorugose; submen-

tum invaginated anterodorsad as a shallow pocket above mentum; postgenal

processes (Brown, 1971) prominent, acutely angulate. Antennae filiform,

last 3 segments slightly enlarged; ratio of segment lengths (base to apex) 14:

9:12:9:8:7:7:6:7:7:10. ’

Pronotum about 1.4 times wider than long, widest just behind middle;

anterior border straight, unmargined, angles barely exserted; anterior angles

sharp, almost 90°; lateral borders narrowly, weakly margined, nearly straight

anteriorly, arcuate behind middle; posterior angles obtuse, not rounded;

posterior border convexly arcuate with broad, flat, finely punctate margin;

base about 1.1 times wider than apex. Pronotal disk coarsely punctatoru-

gulose, rugae predominantly longitudinal; punctures each set with single

long seta. Hypomera and prosternum coarsely punctatorugulose; hypomeral

punctures bearing short, procumbent setae; sternum with a few long, straight

setae; prosternal process about l A coxa width, declivous just behind coxae,

then subhorizontal, apically truncate.

Elytra subequal in width at base to pronotal base; basal margin slightly

raised; lateral margins evenly arcuate, widest at about middle; disk regularly

set with deep, coarse punctures, each bearing single, long seta and separated

by 1-2 puncture diameters. Epipleura faintly rugulose, set with a few short

setae, widest at humerus, abruptly narrowed just behind acutely rounded

humeral angles then gradually narrowing to elytral apices.

Thoracic pleura and sterna set with very large, deep, setigerous punc¬

tures; mesosternum barely excavated between coxae; metasternum about

as long as mesocoxa; mesocoxae almost contiguous; metacoxae separated

by about half mesocoxal diameter. Abdominal sternites set with deep setig¬

erous punctures about half as large as those on metastemum, separated by

1-2 puncture diameters; setae reclined (about 60°), about half as long as

elytral setae. Abdominal sternite length ratios (base to apex) about 35:27:

20:12:20; intercoxal process about half as wide as mesocoxa, apex broadly

rounded.

Legs finely punctate, setose; femora slightly clavate, short, barely ex¬

ceeding body margins; tibiae nearly cylindrical, gradually enlarged apically

and bearing 2 rows of about 8-10 spines on outer surface, these shortest on

fore tibiae; tibial spurs about as long as basal protarsomere; basal 4 tarso-

meres with ventral tufts of stiff, yellow setae about as long as 4th protar¬

somere; tarsomere length ratios as follows: fore tarsus 5:4:3:3:13; middle

tarsus 7:5:4:4:14; hind tarsus 12:7:5:15.

Aedeagus (Fig. 4) with tegmen inverted; apical piece curved dorsad; with

auriculate sclerotized processes mid-dorsally; basal piece sclerotized later¬

ally; median lobe free, sinuately curved; baculi nearly meeting along dorsal

midline, fused apically.

PAN-PACIFIC ENTOMOLOGIST

Body dimensions: elytral length 3.8-4.9 mm; pronotal length 1.4-1.7 mm;

greatest elytral width 2.9-3.4 mm; greatest pronotal width 2.1-2.6 mm.

Holotype male and 1 female paratype from California, Imperial County,

9 mi. W Coyote Wells, March 26, 1961, W. A. Steffan. Paratypes: Califor¬

nia, Imperial County, Mountain Springs, el. 3400', March 27, 1979 (1 9,1

<3); San Diego County, .5 mi. W Yaqui Well, Cholla-Ocotillo, el. 1500',

February 19-23, 1978, ethylene glycol can trap (4 3 3); San Diego County,

Mountain Springs, nr. Desert View Tower, 1960 (1 3). Mexico, Baja Cali¬

fornia Norte, 6.2 mi. W Bahia de Los Angeles, July 11, 1979, pit trap (1 3,

2 9 9); 2 mi. N Arroyo Catavina, December 27, 1979 (1 9). Holotype,

California Academy of Sciences, San Francisco. Paratype distribution: Es-

sig Museum of Entomology, University of California, Berkeley (5); Amer¬

ican Museum of Natural History, New York (1); California Department of

Food and Agriculture Collection, Sacramento (2); collection of K. W. Brown

(5); collection of R. Aalbu (2); University of California, Irvine (1).

Discussion.—Triphalopsis californicus is similar in size, body configura¬

tion and cuticular sculpturing to T. partida Blaisdell and T. minor Blaisdell.

In partida and minor the long setae on the dorsum are deflexed at the tip;

the setae of californicus are straight. In partida and minor the raised pos¬

terior border of the pronotum usually becomes much broader near the lateral

angles, which are distinctly exserted and nearly 90°. In californicus the

posterior pronotal border is slightly broadened at most and the angles are

obtuse and slightly or not exserted. Triphalopsis impressicollis Blaisdell has

straight, erect setae, as in californicus, but has the anterior prothoracic

angles exserted and acute (nearly right angled, slightly or not at all promi¬

nent in californicus ). In impressicollis the pronotal disk is distinctly im¬

pressed near the lateral margins and before the hind angles. In californicus

the disk is evenly convex.

All four species of Triphalopsis have an obvious secondary sexual char¬

acter which is widespread in Triorophini. On the first abdominal sternite of

males is a small, dense, oval patch of pale recumbent setae. These setae

are absent from females. Configuration of the patch and length of setae vary

among other genera and species. In Triorophus the short, very dense, erect

setae occupy a nearly circular area. In Micromes and Eschatomoxys the

patch is oval or teardrop shaped and occupies a depression in the cuticle.

In Stibia and Triphalus a small circular depression bears the setae.

Triphalopsis was omitted from Arnett (1960). The following changes in

Arnett’s key (p. 648) will separate Triphalopsis from similar genera:

4(3). Tarsi spinose or with sparse spiniform vestiture ventrally, not

pubescent . 5 ( Oxygonodera, Micromes, Trichiotes )

Tarsi with coarse, dense, pale setae at least as long as 4th pro-

tarsomere on ventral surfaces

VOLUME 58, NUMBER 2

7(4). Dorsum sparsely covered with slender, erect setae more than half

as long as anterior tarsus . Triphalopsis Blaisdell

Dorsum nearly glabrous or with much shorter, recumbent setae

. Triphalus LeConte

Key to the Species of Triphalopsis

1. Setae on dorsum with decurved tips . 2

Setae on dorsum straight to apices . 3

2. Pronotal disk coarsely rugosopunctate . partida Blaisdell

Pronotal disk coarsely punctate with intervals flat .... minor Blaisdell

3. Pronotal disk evenly convex; anterior angles right angled, slightly

prominent at most . californicus, n. sp.

Pronotal disk impressed near lateral margins and before posterior

angles; anterior angles acute and prominently exserted .

. impressicollis Blaisdell

Literature Cited

Amett, R. H., Jr. 1960. The Beetles of the United States. Catholic University Press, Wash¬

ington, D.C.

Blaisdell, F. E. 1943. Contributions toward a knowledge of the insect fauna of Lower Cali¬

fornia. No. 7. Coleoptera: Tenebrionidae. Proc. Calif. Acad. Sci., 24:171-288.

Brown, K. W. 1971. Redefinition of the genera Pelecyphorus and Philolithus with a key to

the genera of the tribe Asidini (Coleoptera: Tenebrionidae). Coleopt. Bull., 25:17-30.

Casey, T. L. 1907. A revision of the American components of the tenebrionid subfamily

Tentyriinae. Proc. Wash. Acad. Sci., 9:275-522.

Lacordaire, T. 1859. Histoire Naturelle des Insectes. Genera de Coleopteres .... Tome 5.

Roret, Paris.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 92-95

A NEW NEOPANORPA (MECOPTERA: PANORPIDAE) FROM LAOS 1

George W. Byers

Department of Entomology, University of Kansas, Lawrence 66045

Although Mecoptera, chiefly of the genus Neopanorpa, have been re¬

corded from numerous localities in Vietnam to the east and Thailand to the

west, there have been no species reported from either Laos or Cambodia,

in central Indo-China (see distribution map, Byers, 1965:743, Fig. 113). I

have long been aware of certain undescribed species from Laos (Byers,

1965:705), but the species described below is different from those seen ear¬

lier, of which only females had been collected. It is certain that at present

our knowledge of the Mecoptera of the mountainous interior of Indo-China

is only fragmentary.

Following is a description of the first species of Neopanorpa and of the

order Mecoptera recorded from Laos. For the opportunity to study these

specimens, I am indebted to the collector, Mr. Gary L. Peters of Oregon

State University.

Neopanorpa globulifera, new species

Description based on 3 <5, 1 $ pinned, 1 8, 2 9 preserved in alcohol.

For some common characters of the genus, see Byers, 1965:706.

Head .—Dorsum of head shiny black from upper edges of antennal sockets

to neck, continued as narrow black fine along lower edge of each eye.

Rostrum dark yellowish brown to brown, slightly darkened at tip of labrum,

paler around anterior tentorial pits; palps brown. Antennal scape pale yel¬

lowish brown, pedicel brown, flagellum blackish brown, with 43 flagello-

meres (<3), each slightly shorter than those before it, apical segments about

half as long as those near base.

Thorax .—Pronotum blackish brown, slightly paler at sides and along pos¬

terior margin; 2-3 marginal setae at each side of broad, shallow median

emargination in two specimens (absent in others but may be broken off).

Mesonotum black on anterior half, with broad median black stripe extending

backward over scutellum and metanotum; posterolateral comers of meso¬

notum and sides of metanotum sordid yellowish brown. Pleural surfaces,

coxae, legs and basitarsi sordid yellowish brown, distal tarsomeres grading

through brown to dark brown.

Wings faintly tinged with brown, iridescent; markings smoky brown. Api¬

cal band entire but ending posteriorly at vein M x or near it, with small

proximal spur into cell 2nd R 5 in holotype and one female paratype. Ptero-

VOLUME 58, NUMBER 2

stigmal band entire, forked posteriorly in front wing, distal branch of fork

faint or absent in hind wing. Basal band represented by transverse spot over

base of Rs to M and larger, more distal spot from M to hind margin of wing.

Marginal spot small, ending anteriorly at vein R x .

Abdomen of male .—Terga 2-5 blackish brown; corresponding sterna pale

yellowish brown; segment 6 mostly blackish brown, grading into pale brown

near posterior margin; segments 7-9 dark yellowish brown except hypo-

valves of ninth sternum light brown, dististyles brown with darker bases.

Posterior process of third tergum (Figs. 1, 2) roughly triangular, extending

about half way across tergum 4, apex slightly prolonged, with downcurved

terminal and lateral setae. Beneath process, on tergum 4, a smooth pale,

shallow depression surrounded by short, black setae. Hypovalves (Figs. 3,

4) expanded near mid-length, with somewhat narrowed tips and each with

mesal lobe near base. Ninth tergum slightly widened before shallowly emar-

ginate, nearly truncate apex; a smooth, strongly sclerotized (not darkened),

subapical process curved inward (ventrad) from each side around proctiger

(Fig. 7). Outer surface of dististyle shallowly indented (Figs. 3,5), covered

with short setae; longer, more sparse setae on mesal surfaces; basal lobe

concave, cup-like, with pendent process subspherical in shape, smooth,

densely sclerotized. Dorsal surface of basal lobe of dististyle bearing vertical

row of six black spines (Fig. 6). Aedeagus with pale ventral valves longer

than darkly sclerotized dorsal valves and thus concealing them in ventral

aspect; ventral parameres slender, pale, weakly sclerotized, arising and di¬

verging from lower corners of ventral valves; dorsal parameres slender,

acutely tipped; lateral processes conspicuous, rounded.

Abdomen of female .—Terga 2-6 dark brown to blackish brown, except

posterior margin of tergum 6 grading into light brown; sterna dull light brown.

Apical segments dark yellowish brown, cerci black. Subgenital plate of ster¬

num 8 deeply notched posteriorly; apical setae not conspicuous (Fig. 9).

Axial portion of genital plates approximately 3.5 times as long as wide, with

projecting but not divergent anterior apodemes (Fig. 8); arms of distal plate

spatulate, twisted at base; entire structure flattened, thus thin in lateral

aspect.

Body length. —Male, about 11 to 12 mm (holotype 11 mm); female about

10 to 11 mm (allotype 11 mm). Length of fore wing, male, 11.2 to 12.3 mm

(holotype 12.2 mm); female, 11.9 to 12.6 mm (allotype 12.6 mm).

Types. —Holotype, male, Sam Thong, Xieng Khouang Province, 90 miles

(145 km) northeast of Vientiane, Laos, 12 May 1968, collected by Gary L.

Peters. Allotype and paratypes, same locality but 9 May 1968 (1 3), 25 June

1968 (1 3, 1 9), 28 July 1968 (allotype ?), 31 July 1968 (1 9, damaged by

psocids), 1 May 1969 (1 <3, callow). Holotype, allotype and most paratypes

are in the collection of Oregon State University, Corvallis, Oregon. One

male, one female paratypes in the Snow Entomological Museum, University

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-9. Neopanorpa globulifera, new species. Figs. 1-7. Male paratype. Figs. 8, 9.

Female allotype. Fig. 1. Notal organ, abdominal terga 3 and 4, dorsal aspect. Fig. 2. Same,

left lateral aspect. Fig. 3. Genital bulb, ventral (posterior) aspect, hairs mostly omitted except

on left hypovalve and left side of ninth sternum. Fig. 4. Right hypovalve, right lateral aspect.

Fig. 5. Base of left dististyle, apex of basistyle, and most of aedeagus, ventral aspect; lp—

lateral process, vp—ventral paramere, vv—ventral valve. Fig. 6. Same, dorsal (anterior) as¬

pect; dp—dorsal paramere, dv—dorsal valve, vv—ventral valve. Fig. 7. Ninth abdominal ter-

gum and proctiger, right lateral aspect; cr—cercus, pr—proctiger (tenth segment). Fig. 8. Gen¬

ital plates, ventral aspect. Fig. 9. Subgenital plate of eighth sternum, ventral aspect. Scale a.

Figs. 1-4; scale b, Figs. 5-9.

of Kansas, Lawrence, Kansas. The type locality is at approximately 19°12'N,

102°54'E, at an elevation of 1170 m (3800 ft). In 1968, it was a village and

military airbase. Note: this is not the town of Sam Thong, or Ban Sam

Thong, which is more likely to appear on maps of Laos; that town is at

19°51'N, 103°51'E. The scorpion-flies were found on upper surfaces of leaves

of shrubs and herbaceous undergrowth 3-5 feet high at the edge of a forest.

Neopanorpa globulifera most closely resembles N. parvula Willmann

(1976) of northern Vietnam but does not much resemble any other species.

It differs from parvula in wing pattern and several details of the male genital

bulb; the female of parvula is not known. N. parvula has only one small

VOLUME 58, NUMBER 2

spot in the fore wing proximal to the Ptb, a remnant of the basal band

between Cu t and Cu 2 . Willmann illustrated the basal lobe of the dististyle

in parvula with a very irregular pendent process, clearly not with a sub-

spherical termination as in globulifera, and he neither showed nor described

thick, black setae on the dorsal surface of the basal lobe (cf. his Fig. 10).

He found no ventral parameres in parvula, and the dorsal parameres while

acutely tipped are broadly triangular, not slender as in globulifera. There

are minor differences in the hypovalves, but these are difficult to describe

and can best be appreciated by comparison of Willmann’s Figs. 9 and 10

with Figs. 3 and 4 of globulifera.

In my key to Neopanorpa of Indo-China (Byers, 1965), males of globu¬

lifera will be identified as N. burmana. The two species are similar in wing

pattern, structure of the notal organ, and in the thick, black setae on the

basal lobe of the dististyle, of which there are only two in burmana. In the

structure of the aedeagus and hypovalves, however, they are not much

alike. The female of burmana is unknown.

The genital plates and subgenital plate of the female of globulifera are

similar to those of N. byersi Webb and Penny (1979), a species from north¬

ern Thailand that is otherwise decidedly different from globulifera. Most

regional species have the axial portion of the genital plates either very short

and lacking anterior apodemes, or much elongated and longer than the arms

of the distal plate.

The species takes its name from the shape of the pendent process of the

basal lobe of the dististyle (Latin globulus, a little ball + fero, to bear or

carry).

Literature Cited

Byers, G. W. 1965. The Mecoptera of Indo-China. Pacific Insects, 7:705-748.

Webb, D. W., and N. D. Penny. 1979. Neopanorpa byersi (Mecoptera: Panorpidae), a new

species from Thailand. Pacific Insects, 20:63-66.

Willmann, R. 1976. Zwei neue Neopanorpa -Arten aus Vietnam (Mecoptera). Ann. Historico-

Naturales Musei Nationalis Hungarici, 68:65-68.

Footnote

1 Contribution no. 1772 from the Department of Entomology, The University of Kansas,

Lawrence 66045.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 96-104

TICKS (ACARI: ARGASIDAE AND IXODIDAE) FROM

THE CALIFORNIA CHANNEL ISLANDS

Robert S. Lane

Department of Entomological Sciences,

University of California, Berkeley 94720

AND

Scott E. Miller and Paul W. Collins

Santa Barbara Museum of Natural History,

Santa Barbara, California 93105

The tick fauna of the California Channel Islands, a group of 8 islands lying

20 to 98 km off the coast of southern California (Philbrick, 1967), has been

little studied. Of the 20 species of argasid and 31 species of ixodid ticks

known from California (D. P. Furman, pers. comm.), only 1 argasid, Or-

nithodoros talaje (Guerin-Meneville), and 3 ixodids, Ixodes peromysci Au-

gustson, I. pacificus Cooley and Kohls, and I. signatus Birula reportedly

occur there (Banks, 1908; Augustson, 1939; Cooley and Kohls, 1945; Miller

and Menke, 1981; Schwan and Kelly, 1981). The record of O. talaje from

San Clemente Island (Banks, 1908) is of doubtful validity, however.

Heretofore the only Channel Islands survey of significant numbers of

vertebrates for ectoparasites was that of the Los Angeles Museum-Channel

Islands Biological Survey from 1939 to 1941. Ticks collected during that

survey were partially reported on by Augustson (1939) and Cooley and

Kohls (1945), and recently studied by D. P. Furman and E. C. Loomis,

University of California, who are preparing a bulletin on ticks of California.

From 1975 to 1979, personnel (P. W. Collins, S. E. Miller, J. Storrer, K.

W. Rindlaub, J. M. Greaves, L. Laughrin, and F. G. Hochberg) of the Santa

Barbara Museum of Natural History (SBMNH) made a survey of certain

mammals and their ectoparasites on the following islands: East, Middle, and

West Anacapa (Ventura County), San Miguel, Santa Barbara, Santa Cruz,

and Santa Rosa (all Santa Barbara County). Coincidentally, lizards were

captured and their ectoparasites also removed. Two of the several species

of ticks found during this study were previously unrecorded from any of the

islands. The purpose of this report is to document the geographical, host,

and temporal distributions of these ticks.

VOLUME 58, NUMBER 2

Materials and Methods

Study area. —The California Channel Islands comprise 2 groups of 4 major

islands each as follows: the Northern Channel Islands made up of San Miguel,

Santa Rosa, Santa Cruz, and Anacapa; and the Southern Channel Islands

composed of Santa Barbara, San Nicolas, Santa Catalina, and San Clemen¬

te. The islands range in total land area from 2.6 km 2 for Santa Barbara to

249 km 2 for Santa Cruz (Philbrick, 1967). Coastal sage scrub is the predom¬

inant vegetational type; chaparral and oak woodland also occur on the larg¬

est islands (Savage, 1967). Temperatures are milder and climate moister than

the adjacent mainland areas.

Thirty-four species of modern land mammals have been recorded on the

islands, 14 of which are native to California, including a single species,

Urocyon littoralis (Baird), endemic to the islands (von Bloeker, 1967; John¬

son, 1978). Man and 19 species deliberately or accidentally introduced by

him constitute the remainder. The herpetofauna consists of 3 species of

salamanders, 1 species of frog, 5 species of lizards, and 7 species of snakes

(Savage, 1967).

Mammalian and reptilian collections .—Approximately 585 deer mice,

Peromyscus maniculatus (Wagner) sspp. were captured with Museum Spe¬

cial snap traps (baited with rolled oats and peanut butter) on Anacapa, San

Miguel, and Santa Barbara Islands, and 1013 P. maniculatus elusus Nelson

and Goldman were taken with Sherman live traps (baited with rolled oats)

on Santa Barbara Island, examined, and then released. Sherman traps were

set overnight in lines of 60 each. Black rats, Rattus rattus (Linnaeus), were

collected with Victor rat traps (baited with rolled oats and peanut butter)

on Anacapa and San Miguel Islands. One hundred island gray foxes, U. 1.

littoralis, were live-trapped on San Miguel Island, and 15 U. littoralis sspp.

found dead on various islands also were inspected. European rabbits, Oryc-

tolagus cuniculus (Linnaeus), and Canadian elk, Cervus canadensis Erx-

leben, were taken by shooting on Santa Barbara and Santa Rosa Islands,

respectively. Southern alligator lizards, Gerrhonotus multicarinatus (Blain-

ville), were collected on Anacapa, San Miguel, and Santa Rosa Islands. In

1980, after the present survey had been completed, a spotted skunk, Spilog-

ale gracilis amphialus Dickey, found dead on Santa Cruz Island, was also

examined for ectoparasites.

Ticks were removed with forceps from live-trapped animals in the field

or from frozen specimens in the laboratory. Occasionally, collections from

2 or more individuals of the same mammalian species captured within the

same trapline were pooled. All ticks were preserved in 70% ethanol prior

to identification. Larval ticks were usually mounted in Berlese or Hoyer’s

media on micro slides and examined with a compound microscope, whereas

PAN-PACIFIC ENTOMOLOGIST

unmounted nymphal and adult ticks were inspected with a dissecting mi¬

croscope. All material has been deposited in the SBMNH except for 2 larvae

and 4 nymphs of I. peromysci that were placed in the tick collection of the

Rocky Mountain Laboratories, Hamilton, Montana.

Results

As summarized in Table 1, a total of 1934 mammals representing 7 species,

and 32 alligator lizards, yielded 28 collections of ticks of 5 species (1 argasid,

4 ixodids). In addition, 117 P. maniculatus streatori Nelson and Goldman

from San Miguel and 9 U. littoralis sspp. from various islands were exam¬

ined for ticks with negative results. Only 11 (0.7%) of 1599 deer mice, P.

maniculatus sspp., 3 (2.6%) of 115 island foxes, U. littoralis sspp., and 5

(1.6%) of 322 black rats, R. rattus, had ticks. However, field examination

of live mammals may have missed some small ticks. Ixodes pacificus and

I. peromysci exhibited the broadest host ranges by occurring on 3 species;

the remaining ticks were each collected from 1 or 2 host species.

The records of Otobius megnini (Duges) from Santa Rosa, Haemaphy-

salis leporispalustris (Packard) from Santa Barbara, I. pacificus from San

Miguel and Santa Rosa, I. peromysci from West Anacapa, and /. rugosus

Bishopp from Santa Cruz constitute new distributional records for these

ticks. New host records include I. pacificus from U. /. littoralis, I. pero¬

mysci from G. multicarinatus, P. maniculatus anacapae von Bloeker, and

R. rattus, and I. rugosus from U. littoralis santacruzae Merriam and S.

gracilis amphialus. The collection of I. peromysci from G. multicarinatus

also represents the first record of this tick from a reptile.

Immatures of I. peromysci were found on animals predominantly in Oc¬

tober with 2 records in March, whereas most adults of this tick were collected

during March with isolated records in June and October (Table 1). Collection

records for the other ticks are too scanty to warrant discussion of their

seasonal distributions.

Discussion

The ear tick, O. megnini, is widely distributed in warmer regions of the

United States and appears to be common in parts of southern California

(Cooley and Kohls, 1944). Ten of 11 Californian collection records published

by Cooley and Kohls (1944) were based upon material collected in southern

California, and all collections were taken from cattle. Although O. megnini

has been found on elk before (e.g., Bishopp and Trembley, 1945; Rich,

1957), this is apparently the first published report of its occurrence on elk

in this state. Elk were unsuccessfully introduced onto Santa Rosa Island in

1905. A second introduction in 1930 involving approximately 15 animals

from Yellowstone National Park, Wyoming, resulted in the present popu¬

lation. O. megnini may have been brought to Santa Rosa Island directly on

Table 1. Records of argasid and ixodid ticks collected from the California Channel Islands, 1975-1980.

Number

TTnst tnn infp«tprl/

no. examined)

Locality 1

Collector

Date

Argasidae

Otobius megnini

Cervus canadensis

(1/2)

Upper Windmill Canyon,

Santa Rosa I.

S. Miller

21.IV. 1976

Ixodidae

Haemaphy salis

leporispalustris

Oryctolagus

cuniculus (1/20)

NE Signal Peak, Santa

Barbara I.

P. Collins

6.III. 1979

Ixodes pacific us

Gerrhonotus

multicarinatus (4/

25) 2

SBMNH Nos. 301-302,

Johnson’s Lee, Santa

Rosa I.

P. Collins

13. V. 1975

Gerrhonotus

multicarinatus (4/

25) 2

SBMNH No. 360,

Windmill Canyon,

Santa Rosa I.

P. Collins

27. V. 1976

Gerrhonotus

multicarinatus (4/

25) 2

SBMNH No. 426, mouth

of Old Ranch Canyon,

Santa Rosa I.

P. Collins

11.V.1978

Gerrhonotus

multicarinatus (4/

25) 2

Canada do Mar, San

Miguel I.

P. Collins

27.IV. 1979

Urocyon l. littoralis

(2/100)

San Miguel I.

L. Laughrin

10.11.1979

U. 1. littoralis

(2/100)

San Miguel I.

L. Laughrin

10.11.1979

Homo sapiens (1/1)

Nidever Canyon, San

Miguel I.

P. Collins

25.IV. 1979

VOLUME 58, NUMBER 2

Table 1. Continued

Number

Host (no. infested/

no. examined)

Locality 1

Collector

Date

Ixodes peromysci 0

G. multicarinatus (1/

7) 2

SBMNH No. 488, Oak

Canyon, West Anacapa

D. Johnson, S.

Miller

I.X. 1978

Peromyscus

maniculatus

anacapae (4/196) 3

SBMNH No. 104, West

Anacapa I.

P. Collins, S.

Miller

1.X.1978

Peromyscus

maniculatus

anacapae (4/196) 3

SBMNH No. 110, West

Anacapa I.

P. Collins, S.

Miller

2.X.1978

Peromyscus

maniculatus

anacapae (4/196) 3

SBMNH No. 110, West

Anacapa I.

P. Collins, S.

Miller

3.X.1978

Peromyscus

maniculatus

anacapae (4/196) 3

SBMNH No. 110, West

Anacapa I.

J. Greaves

3.X.1978

P. maniculatus

elusus (7/1286)

Santa Barbara I.

J. Storrer, J.

Trager

12. VI. 1978

P. maniculatus

elusus (7/1286)

SBMNH Nos. 74-75,

Santa Barbara I.

J. Storrer

6.III. 1979

P. maniculatus

elusus (7/1286)

SBMNH Nos. 74-75,

Santa Barbara I.

P. Collins

7.III. 1979

P. maniculatus

elusus (7/1286)

SBMNH No. 76, Santa

Barbara I.

P. Collins

8.III. 1979

P. maniculatus

elusus (7/1286)

SBMNH No. 75, Santa

Barbara I.

J. Storrer

8.III. 1979

PAN-PACIFIC ENTOMOLOGIST

Table 1. Continued

Number

Host (no. infested/

no. examined)

Locality 1 2 3

Collector

Date

P. maniculatus

elusus (7/1286)

SBMNH No. 75, Santa

Barbara I.

J. Storrer

8.III. 1979

P. maniculatus

elusus (7/1286)

SBMNH No. 76, Santa

Barbara I.

P. Collins, S.

Miller

9.III. 1979

Rattus rattus (5/322) 4

SBMNH No. 105, West

Anacapa I.

P. Collins,

S.Miller

1.X.1978

Rattus rattus (5/322) 4

West Anacapa I.

P. Collins

l.X.1978

Rattus rattus (5/322) 4

SBMNH No. 107, West

Anacapa I.

P. Collins, S.

Miller

2.X. 1978

Rattus rattus (5/322) 4

SBMNH No. 105, West

Anacapa I.

P. Collins, S.

Miller

3.X.1978

Rattus rattus (5/322) 4

SBMNH No. 106, West

Anacapa I.

P. Collins, S.

Miller

3.X.1978

Ixodes rugosus 0

U. littoralis

santacruzae (1/6)

Main ranch (Stanton

property), Santa Cruz

L. Laughrin

3.IV. 1978

Spilogale gracilis

amphialus (1/1)

Nr. Bosque Cabrillo,

Santa Cruz I.

P. Schuyler

22.XI.1980

1 SBMNH numbers refer either to herpetology specimen numbers or mammalian census station (=trapline) numbers.

2 Fourteen alligator lizards were collected from Santa Rosa, 11 from San Miguel, and 7 from Anacapa Islands.

3 Of this total, 96 deer mice were examined from West Anacapa, 99 from Middle Anacapa, and 1 from East Anacapa Islands.

4 In total, 283 specimens were examined from East, Middle, and West Anacapa Islands, and 39 from San Miguel Island.

VOLUME 58, NUMBER 2

102

PAN-PACIFIC ENTOMOLOGIST

elk or it may have been transported there on cattle from the mainland and

subsequently transferred to elk by mutual grazing of both hosts on the same

rangelands. Cattle have been present on Santa Rosa Island since about 1902

and, recently, the owners of Santa Rosa Island have had trouble with ticks

on cattle.

Ixodes peromysci is 1 of only 2 Ixodes species (out of a total of 36 species)

from the U.S.A. that has a strictly insular distribution (Keirans and Clifford,

1978). Previously it had been found solely on deer mice, P. maniculatus

elusus, from Santa Barbara Island, Santa Barbara County (not Los Angeles

County), the type locality, and on P. maniculatus dementis Meams from

San Clemente Island, Los Angeles County, California. Thus, the host rec¬

ords from P. maniculatus anacapae, R. rattus, and G. multicarinatus are

new for this tick. None of the other 19 species of Ixodes recorded from

California, except for I. pacificus, is known to use reptiles as hosts for its

immatures. The collections from West Anacapa Island extend the known

distribution of I. peromysci approximately 69 km NNW of its former north¬

ernmost distribution.

The low infestation levels recorded for each of the 4 host species and

subspecies of I. peromysci, especially the rodents, probably are underesti¬

mates because most of the black rats and some of the deer mice were snap-

trapped, frozen, and subsequently examined, and some live-trapped deer

mice were examined cursorily and then released. As noted by Westrom and

Yescott (1975), the vagility of ectoparasites interferes with attempts to ac¬

curately estimate their abundance, particularly when an exodus from dead

hosts occurs prior to collection attempts. These investigators showed that

fleas, chiggers, and mesostigmatid mites began leaving California ground

squirrels, Spermophilus beecheyi (Richardson), within 45 minutes post mor¬

tem in contradistinction to lice, which remained on the host for approxi¬

mately 5 hours before leaving. Unfortunately, ticks were not found on any

of the animals they inspected.

The rabbit tick, Haemaphysalis leporispalustris, has been recorded from

Alaska, Canada, all the U.S.A., and southward to Argentina. A single spec¬

imen was collected from the ear of a “rabbit” on East Anacapa Island, 26

August 1940, by G. P. Kanakoff during the 1939-1941 Channel Islands Bi¬

ological Survey (unpublished determination by G. F. Augustson recently

confirmed by D. P. Furman and associates). The European rabbit, Oryc-

tolagus cuniculus, the only lagomorph documented from East Anacapa Is¬

land (not Lepus europaeus Pallas as stated by von Bloeker, 1967), has been

extirpated from there since the mid-1960’s (Banks, 1966). On Santa Barbara

Island, 1 of 20 O. cuniculus inspected in this study was parasitized by H.

leporispalustris, which is considerably lower than the frequency per host

reported for this tick on native lagomorphs (i.e., Lepus californicus Gray

and Sylvilagus bachmani (Waterhouse)) from the Californian mainland

VOLUME 58, NUMBER 2

103

(Coultrip et al., 1973; Lane et al., 1981). In the U.S.A., important hosts of

the rabbit tick include a number of species and subspecies of Lepus (hares)

and Sylvilagus (rabbits) as well as a variety of birds, small mammals, and

other hosts (Cooley, 1946).

In western U.S.A., /. pacificus, a vicious man-biter, reportedly occurs

from California to Washington and in Nevada, Utah, and Idaho (Keirans

and Clifford, 1978). The only published record of it from the Channel Islands

was that of a nymph taken from a California Quail, Lophortyx californica

catalinensis Grinnell, on Santa Catalina Island, 23 January 1941 (Cooley

and Kohls, 1945). Although /. pacificus has not been recorded hitherto from

island foxes, it has been found before on Urocyon sp. in California (Arthur

and Snow, 1968). At least 53 additional species of vertebrates including man

and many other mammals, 5 species of birds, and possibly 9 species of

lizards have been reported as hosts for I. pacificus on the mainland. In

California, this tick commonly attacks man and has been incriminated in

several mild cases of paralysis in children (Herms, 1950) and in a single

human case of erythema chronicum migrans (Naversen and Gardner, 1978).

Finally, in the U.S.A., I. rug os us parasitizes skunks, foxes, weasels, and

domestic dogs from California to Washington (Keirans and Clifford, 1978).

The only spotted skunk examined from Santa Cruz Island yielded a female

I. rugosus, and island foxes found dead there had an infestation rate of 0.17

/. rugosus ticks per host (n = 6). Although none of 100 island foxes from

San Miguel yielded /. rugosus, it cannot be assumed that this tick does not

parasitize foxes there because tick infestation rates found in this study were

generally quite low.

Acknowledgments

We are indebted to J. E. Keirans and C. M. Clifford, Rocky Mountain

Laboratories, for determining the specific identities of some ticks and other

favors; D. P. Furman, University of California, Berkeley, for confirming

the specific identities of most ticks and for review of an earlier version of

the manuscript; J. T. Doyen, UCB, for critique of the manuscript; W. Z.

Lidicker, UCB, for taxonomic advice on mammals; and to F. G. Hochberg,

SBMNH, for support of fieldwork and reviewing the manuscript. Collection

of specimens from San Miguel, Anacapa, and Santa Barbara Islands was

funded by National Park Service Contract No. CX-2000-8-0040. Collections

were made by permission of the National Park Service, Al Vail of the Vail

and Vickers Company, and the Nature Conservancy.

Literature Cited

Arthur, D. R., and K. R. Snow. 1968. Ixodes pacificus Cooley and Kohls, 1943: Its life-

history and occurrence. Parasitology, 58:893-906.

Augustson, G. F. 1939. Contributions from the Los Angeles Museum-Channel Islands Bi-

104

PAN-PACIFIC ENTOMOLOGIST

ological Survey. 4. A new Ixodes (Acarina: Ixodidae). Bull. S. Calif. Acad. Sci., 38:

141-147.

Banks, N. 1908. A revision of the Ixodoidea, or ticks, of the United States. U.S. Dep. Agr.

Entomol. Tech. Ser., 15:1-61.

Banks, R. C. 1966. Terrestrial vertebrates of Anacapa Island, California. Trans. San Diego

Soc. Natur. Hist., 14:173-188.

Bishopp, F. C., and H. L. Trembley. 1945. Distribution and hosts of certain North American

ticks. J. Parasitol., 31:1-54.

Cooley, R. A. 1946. The genera Boophilus, Rhipicephalus, and Haemaphysalis (Ixodidae)

of the New World. Nat. Inst. Health Bull. No. 187, 54 pp.

Cooley, R. A., and G. M. Kohls. 1944. The Argasidae of North America, Central America

and Cuba. Amer. Midland Natur. Monogr. No. 1, Univ. Press, Notre Dame, Indiana,

152 pp.

Cooley, R. A., and G. M. Kohls. 1945. The genus Ixodes in North America. Nat. Inst.

Health Bull. No. 184, 246 pp.

Coultrip, R. L., R. W. Emmons, L. J. Legters, J. D. Marshall, Jr., and K. F. Murray. 1973.

Survey for the arthropod vectors and mammalian hosts of Rocky Mountain spotted

fever and plague at Fort Ord, California. J. Med. Entomol., 10:303-309.

Herms, W. B. 1950. Medical Entomology with Special Reference to the Health and Well¬

being of Man and Animals, 4th ed. The MacMillan Co., New York, 643 pp.

Johnson, D. L. 1978. The origin of island mammoths and the Quaternary land bridge history

of the Northern Channel Islands, California. Quat. Res., 10:204-255.

Keirans, J. E., and C. M. Clifford. 1978. The genus Ixodes in the United States: A scanning

electron microscope study and key to the adults. J. Med. Entomol. Suppl. No. 2,

149 pp.

Lane, R. S., R. W. Emmons, D. V. Dondero, and B. C. Nelson. 1981. Ecology of tick-borne

agents in California. I. Spotted fever group rickettsiae. Amer. J. Trop. Med. Hyg., 30:

239-252.

Miller, S. E., and A. S. Menke. 1981. Entomological bibliography of the California Islands.

Occas. Pap. Santa Barbara Mus. Natur. Hist., No. 11, 78 pp.

Naversen, D. N., and L. W. Gardner. 1978. Erythema chronicum migrans in America. Arch.

Dermatol., 114:253-254.

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

Islands. Santa Barbara Bot. Gard., Santa Barbara, Calif., 363 pp.

Rich, G. B. 1957. The ear tick, Otobius megnini (Duges) (Acarina: Argasidae), and its record

in British Columbia. Can. J. Comp. Med., 21:415-418.

Savage, J. M. 1967. Evolution of the insular herpetofaunas. Pp. 219-227 in R. N. Philbrick

(ed.). Proceedings of the Symposium on the Biology of the California Islands. Santa

Barbara Bot. Gard., Santa Barbara, California, 363 pp.

Schwan, T. G., and P. R. Kelly. 1981. Ixodes signatus (Ixodoidea: Ixodidae) parasitizing

Pigeon Guillemots on the Channel Islands, California. J. Med. Entomol., 18:171-172.

von Bloeker, J. C., Jr. 1967. Land mammals of the Southern California Islands. Pp. 245-263

in R. N. Philbrick (ed.). Proceedings of the Symposium on the Biology of the California

Islands. Santa Barbara Bot. Gard., Santa Barbara, California, 363 pp.

Westrom, D., and R. Yescott. 1975. Emigration of ectoparasites from dead California ground

squirrels Spermophilus beecheyi (Richardson). Calif. Vector Views, 22:97-103.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 105-108

LIFE HISTORY OF THE DELPHACID PLANTHOPPER

STOBAERA TRICARINATA (SAY) ON WESTERN RAGWEED,

AMBROSIA PSILOSTACHYA DECANDOLLE, IN SOUTHERN

CALIFORNIA (HEMIPTERA-HOMOPTERA: DELPHACIDAE)

N. J. Reimer and R. D. Goeden

Department of Entomology, University of California,

Riverside 92521

Stobaera tricarinata (Say) is a common insect associate of western rag¬

weed, Ambrosia psilostachya Decandolle, a native, perennial herb (Com-

positae) in southern California (Goeden and Ricker, 1976). The life history

of this heretofore little-known delphacid planthopper is reported herein.

Taxonomy.—Stobaera tricarinata adults were redescribed by Kramer

(1973) in his revision of the genus. The egg and nymphs were described by

Reimer and Goeden (1981).

Distribution and host plants.—Stobaera tricarinata was collected from

A. psilostachya by the authors and D. W. Ricker at the following locations

in southern California: Los Angeles Co.—El Toro, San Juan Capistrano,

Yorba Linda; Riverside Co.—Beaumont, Rancho California, Temecula,

Wildomar; San Bernardino Co.—Chino, East Highland, Mill Creek; San

Diego Co.—Alpine, Encinitas, Jamul, La Mesa, Pala, Poway, Rainbow,

Valley Center; Santa Barbara Co.—Carpenteria, Goleta, Santa Barbara;

Ventura Co.—Fillmore, Piru, and Ventura.

Kramer (1973) described tricarinata as the most commonly encountered

species of Stobaera in the United States. He recorded this species from 27

states as well as Baja California, Mexico. Other plant records in Kramer

(1973) included A. confertiflora Decandolle, A. chamissonis (Lessing)

Greene, and Helianthus argophyllus Torrey and Gray. The first 2 records

involved only 2 males and 1 female, respectively, probably transients, that

Goeden and Ricker (1974, 1975) who supplied these specimens, interpreted

as unconfirmed plant associations in their faunistic surveys of these 2 rag¬

weeds. Nymphs and adults also have been collected from H. annuus L.

ssp. lenticularis (Douglas) Cockerell and Xanthium strumarium L. through¬

out southern California by J. H. Hilgendorf, Department of Entomology,

University of California, Riverside (pers. comm.). Thus, S. tricarinata

apparently is selectively, not broadly, oligophagous on host plants belong¬

ing to the tribe Heliantheae of the Compositae (Munz and Keck, 1959).

Biology. —Field populations were studied at a 1-ha site located 2 km east

106

PAN-PACIFIC ENTOMOLOGIST

and above Mountain Home Village in Mill Canyon in the San Bernardino

Mountains. Laboratory and insectary studies were conducted at the Uni¬

versity of California, Riverside, where insectary cultures were maintained

on potted A. psilostachya at 27 ± 1°C, 45-70% relative humidity, and a 14/

10-hr (light/dark) photoperiod.

Egg .—The fusiform-ellipsoidal eggs were inserted singly, blunt end first,

into the pith of stems of A. psilostachya. The tapered end pointed outward

and upward toward the stem apex, penetrated the vascular tissues, and

occasionally protruded from an ovipositional slit in the epidermis. Each egg

occupied a separate ovipositional cavity. All of 22 eggs observed in the

insectary hatched in 15 days.

Nymph .—There are 5 instars. The duration of 27 each, first-fifth instars

averaged 3.0 ± 0.02 days, 5.5 ± 0.46 days, 6.5 ± 0.38 days, 8.4 ± 0.8 days,

and 11.3 ± 1.4 days, respectively.

After eclosion, the first instars crawled to the undersides of the leaves

and fed in the intervenal spaces and smaller veins. The second instars fed

in the same leaf areas. However, the third instars fed on the upper surfaces

of the leaves as well as on the stems. Most fourth and fifth instars fed on

the stems. Moulting occurred in the same areas as feeding.

The apical tibial spur or calcar appears in the second instar (Reimer and

Goeden, 1981). This instar and the remaining 3 instars jumped readily by

aid of this movable spine.

Adult .—Adults fed in the midribs on the upper surfaces of the leaves and

on the more succulent, distal parts of the stems and branches. Both nymphs

and adults excreted honey dew. Seventeen individual feedings by 15 adults

observed in the insectary lasted an average of 42 ± 3 (range: 15-63) min.

No courtship behavior was observed. Mating was observed only twice.

Both instances involved virgin females and occurred late in the afternoon

in the insectary. Copulation by 1 pair occurred on a stem; by the other pair,

on a cage floor. The pair on the stem fined up vertically with the female

facing upward and the male facing downward. The female curved her ab¬

domen dorsally and the male twisted his abdomen laterally until his termi-

nalis contacted the genital opening of the female. This pair remained in

copula for 8 min; the other pair, for 11 min. While united, the female twisted

her abdomen sporadically, while the male remained stationary.

Nine mature females collected in Mill Canyon when dissected had a mean

number of 9.9 ± 0.20 (range: 9-11) ovarioles on their right sides and 10.0 ±

0.23 (range: 9-11) ovarioles on their left sides. Twenty ovarioles from these

females averaged 3.34 ± 0.43 mm in length. The ovarioles fie dorsal to the

digestive tract in the fourth, fifth, and sixth abdominal segments.

The female oviposited in the distal third of a vertical stem while facing

upward toward the stem apex. She raised her abdomen and extended her

ovipositor so that it was perpendicular to her abdomen and to the plant

VOLUME 58, NUMBER 2

107

surface. She then inserted her ovipositor into the plant tissues until it was

fully buried. The ovipositor initially was inserted perpendicularly, but once

it entered the plant, it formed a more or less pyriform cavity expanded

downward and away from the stem apex. This mirrored the curved shape

of the ovipositor. The egg was extruded as the female slowly withdrew her

ovipositor from the stem.

Under insectary conditions, 14 females laid an average of 6.8 ± 0.45

(range: 1-15) eggs per day. Oviposition occurred a mean number of 1.3 ±

0.12 (range: 0-4) times per week. These females laid a mean total of 32.4 ±

4.1 (range: 8-57) eggs during their lifetimes.

The mean longevity of 12 males was 38.7 ± 4.3 (range: 21-55) days com¬

pared to 50.5 ± 2.5 (range: 34-60) days for 12 females. In the field over¬

wintered adults live an additional 3 to 4 months.

Seasonal history.—Stobaera tricarinata was bivoltine in Mill Canyon dur¬

ing 1977 and 1978. The generation occurred during June-August, followed

by a second generation in October and November. The sex ratio of both

generations approximated 1:1. The F 2 generation overwinters as adults with

undeveloped gonads. No females with fully developed ovaries were found

at the study site after October 31 during 1978. The fall (F 2 ) generations of

nymphs and adults are more darkly pigmented than the summer (F x ) gen¬

erations.

The overwintered females oviposited during active vegetative growth by

western ragweed in April and May. A week interval occurred between the

2 generations in mid-September during 1977 and 1978, when 50 weekly sweeps

of ragweed shoots with a standard sweep net yielded neither nymphs nor

adults. The F 2 generation presumably was present as eggs at this time.

Nymphs of the F 2 generation were collected from late September until mid-

November during both years.

Parasitization. —The nymphs and adults of S. tricarinata in Mill Canyon

were parasitized by a dryinid wasp identified as Pseudogonatopus arizoni-

cus Perkins (Hymenoptera: Dryinidae). This dryinid has been reported from

Stobaera sp. elsewhere; however, this was the first record for this parasite

from southern California and from an identified host species (P. H. Freytag,

pers. comm.). This parasite was most abundant in Mill Canyon during Au¬

gust.

Acknowledgments

The technical assistance of D. W. Ricker, Department of Entomology,

University of California, Riverside, is gratefully acknowledged. Dr. Paul H.

Freytag, Department of Entomology, University of Kentucky, Lexington,

identified the dryinid. From a thesis submitted by the senior author in partial

satisfaction of the M.S. degree in Entomology, University of California,

Riverside.

108

PAN-PACIFIC ENTOMOLOGIST

Literature Cited

Goeden, R. D., and D. W. Ricker. 1974. The phytophagous insect fauna of the ragweed,

Ambrosia chamissonis, in southern California. Environ. Entomol., 3:835-839.

Goeden, R. D., and D. W. Ricker. 1975. The phytophagous insect fauna of the ragweed,

Ambrosia confertiflora, in southern California. Environ. Entomol., 4:301-306.

Goeden, R. D., and D. W. Ricker. 1976. The phytophagous insect fauna of the ragweed.

Ambrosia psilostachya, in southern California. Environ. Entomol., 5:1169-1177.

Kramer, J. P. 1973. Revision of the American planthoppers of the genus Stobaera (Homop-

tera: Delphacidae) with new distributional data and host plant records. Proc. Entomol.

Soc. Washington, 75:379-402.

Munz, D. A., and D. D. Keck. 1959. A California Flora. Univ. California Press, Berkeley

and Los Angeles, 1681 pp.

Reimer, N. J., and R. D. Goeden. 1981. Descriptions of the immature stages of Stobaera

tricarinata (Say) (Hemiptera-Homoptera: Delphacidae). Pan-Pacific Entomol., 57:429-

433.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 109-110

THE FEMALE OF POMP1LUS (AMMOSPHEX) WASBAUERI EVANS

(HYMENOPTERA: POMPILIDAE)

Lynn Siri Kimsey

Department of Entomology, University of California,

Davis 95616

When Evans (1966:396) first described Pompilus wasbaueri, the female

was unknown to him. Recently, while determining specimens with M. S.

Wasbauer in preparation of a study of the Pompilinae of California, we

located three females of this species. For convenience, I chose one speci¬

men as a pleisiallotype.

The other two females, from Marin and Alameda Counties, California

vary somewhat from the specimen described below in the wing venation

and clypeal length. In all three specimens, the clypeus is at least three times

as wide as long; in one it is 3.7 times. The second submarginal cell in all

three is narrower than the third, varying from two-thirds as wide to half as

wide. There is variation in the distance from the apex of the marginal cell

to the wing tip, from 1.3 to 2.0 times the marginal cell length. All three

specimens are in the collection of the California Academy of Sciences in

San Francisco.

The following abbreviations are used: middle interocular distance (MID),

transfacial distance (TFD), lower interocular distance (LID), upper inter¬

ocular distance (UID), postocellar length (POL), ocellocular length (OOL).

Pompilus (Ammosphex) wasbaueri Evans

Pleisiallotype

Female. —Length 6.5 mm; forewing 5.0 mm. Body black overlaid with

fine black pubescence which reflects various shades of brown and somewhat

silvery on lower front; fore- and hind wings evenly infuscate. Head with a

few weak erect setae, thorax (including propodeum) without erect setae.

Head 1.18 times as wide as high; labrum not exserted; clypeus 3.2 times as

wide as high, apical margin very slightly concave. Front wide, MID 0.62

TFD; MID 1.09 LID; UID 0.85 LID. Temples about two-thirds as wide as

eye; vertex strongly arched above eyes. Ocelli in a right triangle; POL:

OOL = 7.2. Scape, pedicel and first two flagellomeres in a ratio of 11:6:21:

12, first flagellomere length 0.75 UID. Pronotum broadly angulate poste¬

riorly. Postnotum medially half as long as metanotum; propodeum weakly

sloping anteriorly, with an oblique, slightly concave declivity on posterior

third. Fore-basitarsus with three evenly spaced stout combspines; second

110

PAN-PACIFIC ENTOMOLOGIST

tarsomere with a stout medial combspine; combspines about as long as

maximum width of tarsomeres; apical tarsomeres with spines beneath ba-

sally; longer hind tibial spur 0.61 times basitarsus length. Forewing marginal

cell 1.3 times its length from wing tip; SMC2 two-thirds as wide as SMC3,

half as wide anteriorly as posteriorly, SMC3 about as wide as long.

Pleisiallotype female. —California, Marin Co., Mill Valley, July 23-25,

1965 (P. H. Arnaud).

Literature Cited

Evans, H. E. 1966. A revision of the Mexican and Central American spider wasps of the

subfamily Pompilinae. Mem. Amer. Ent. Soc., 20:1^142.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 111-116

NOTES ON THE BIOLOGY OF

ANDRENA (CALLANDRENA) HELIANTHI ROBERTSON

(HYMENOPTERA: ANDRENIDAE)

F. D. Parker and G. E. Bohart

Bee Biology and Systematics Laboratory, Agricultural Research,

USDA, Utah State University, Logan 84322

The nesting habits of andrenid bees, one of the largest groups of North

American aculeates, are poorly known (Hurd, 1979). Perhaps their exclusive

habit of nesting in the ground is partly responsible for this paucity of knowl¬

edge. The 52 U.S. species of Callandrena, a Nearctic subgenus of Andrena,

exemplify this dearth of biological information; the nest of only one, An¬

drena accepta Viereck, has been described (Rozen, 1973). A brief descrip¬

tion of nest digging, however, has been reported by Hicks (1926) for An¬

drena helianthi Robertson, the subject of this study. For the past four years,

our laboratory has conducted studies of sunflower pollination, and during

this time a sizeable adult population of A. helianthi has been observed. In

the summer of 1979, many nests of this sunflower bee were found in our

plots, and we were able to observe various aspects of its biology, including

pollen collection, nest architecture, larvae, nest associates, and seasonal

occurrence.

Nesting Site

Adult females were found nesting between irrigation furrows in plots

planted to sunflower near Logan, Utah, and observations were made during

July and August 1979. The one-acre plot where the bees nested was nearly

flat, and the soil type was Millville Silt Loam. When Andrena nests were

discovered during the course of the pollination studies, they were marked

by a stake, and in September four of them were excavated. The number of

nests found was not recorded, but there were at least ten. They were scat¬

tered among the rows of sunflower, and the nest entrances were usually

near sunflower stalks. Although nest entrances were usually a meter or more

apart, some were within a few centimeters of one another.

Nest Architecture

Entrance hole. —We were unable to observe bees starting to dig their

nests, but, instead, we located bees entering established nests or found

newly made tumuli. The oval tumulus surmounting a typical nest excavation

measured 7 cm across and 1.5 cm high. Tumuli did not develop around the

112

PAN-PACIFIC ENTOMOLOGIST

entrances of a few nests that were made under soil clods (Fig. 1) on the

sloping side of irrigation furrows. Most nest entrances opened to the south

and averaged 8 mm wide. A turret constructed in the side of the tumulus

acted as a nearly horizontal passageway leading from the actual tunnel en¬

trance to the edge of the tumulus (Fig. 2).

Burrows .—When the nests were excavated, the main burrow was plugged

at intervals, but the lateral burrows were so completely packed with soil

that they could not be traced. The main burrow of a typical nest was 8 mm

wide, horizontal and plugged for 3 cm before it became vertical (for 10 cm)

and again horizontal and plugged (for 2-3 cm). Beyond the second plug, the

main burrow descended nearly vertically (for 5 cm), and then it was plugged

again (for 1 cm). Finally, it sloped downward for another 24 cm, from which

point the lateral branches were presumed to diverge (Fig. 3A). The main

burrow ranged in depth from 30 to 37 cm. It was unlined, but had smooth

walls. The distance from the main burrow to its associated cells ranged from

5 to 19 cm. In most nests, these cells were clustered, an indication that only

1 or 2 lateral burrows had been constructed (Fig. 3B).

Cells .—Cells were found at depths ranging from 37 to 45.5 cm. In most

nests, they were in short series (1-3) and were separated by about one cm

of tightly packed soil. Some cell series within a few centimeters of each

other were probably connected to the same lateral burrow. The number of

cells/nest ranged from 1 to 8 and averaged 4.2. The oval cells were horizontal

and ranged in inner width from 7 to 9 mm and length from 15 to 20 mm.

The bees constructed the cell walls by compacting about 2 mm of soil against

the roughed-out excavations and then coating the basal 2 A of the smooth

cell walls with a thin, brownish layer of wax (Fig. 4). The cell was narrower

at the neck region (7 mm) than at the base (9 mm). It was capped by a

clockwise spiral of 1.5 mm wide rings of soil. From the margin to the center

were 5 rings, each slightly farther than its predecessor from the base of the

cell, thus producing a domed cap (Fig. 5). The position and shape of the

pollen-nectar provisions were not observable since in all cells the provisions

had been entirely consumed.

The yellowish fecal material was deposited at the base of the cell in a pile

of 1-mm-wide flattened pellets (Fig. 6).

Larvae .—The naked larvae rested on their dorsal surface, supported by

their large, dorsolateral lobes (Figs. 7, 8). Often, fecal material was stuck

to the larvae between these lobes. The larvae were creamy-white, rigid, and

averaged 12 mm in length. Overwintering took place in the prepupal stage.

Nest Associates

Two nests with a total of 5 cells were parasitized by an undescribed

species of Nomada, subgenus Pachynomada. This species was described

recently in an as yet unpublished revision of Pachynomada by A. Moalif,

VOLUME 58, NUMBER 2

113

Figs. 1, 2, 4-11. Fig. 1. Entrance to nest of Andrena. Fig. 2. Nest entrance of Andrena

showing plug of soil used to close the nest during the afternoon and night. Fig. 4. Cell of

Andrena illustrating wax lining at the base. Fig. 5. Cell cap of Andrena. Note the spinal pattern.

Fig. 6. Mat of fecal material deposited at base of Andrena cell. Fig. 7. Cell of Andrena that

contains the overwintering prepupal larva. Fig. 8. Lateral view of prepupal Andrena larva.

Note the large projection on larvae. Fig. 9. Prepupal larva of Nomada n. sp. in Andrena cell.

Fig. 10. Ventral view of prepupae of Nomada n. sp. Fig. 11. Fecal pellets of Nomada n. sp.

in Andrena cell.

PAN-PACIFIC ENTOMOLOGIST

114

A B

Fig. 3. Diagram of Andrena nest—(A) cross section illustrating slope of burrow and cells,

(B) arrangement of lateral burrows and cells.

Dept, of Biology, Utah State University. The parasitic larvae were easily

distinguished by their yellow color and rigid form (Figs. 9, 10). Their fecal

pellets were short (2 mm wide and 0.7 mm long) and scattered about the

cell walls (Fig. 11). Egg placement was not determined, but some cells had

a shallow puncture at the base or on the side of the cell. Linsley and

MacSwain (1955) described similar shallow holes as being made for egg

deposition by another species, Nomada ( Gnathias ) opacella Timberlake.

Two other cells contained hyphal mats of an unknown fungus in the As-

cophaeraceae (det. N. Youssef, Utah State University). Most of the fecal

pellets were broken down and encased by these hyphae. This fungus was

also seen growing in a cell occupied by a healthy larva of Nomada.

Adults

Seasonal occurrence .—Since sunflowers were planted three times during

the spring, bloom was available from July to September. Counts of all bees

on flower heads were made by 0900, 1100, and 1300 hours every Monday,

Wednesday, and Friday. Sunflowers began to bloom on July 25, and the first

Andrena were observed on July 27. Both sexes were abundant but their

ratio was not recorded. The seasonal appearance of Andrena helianthi in¬

dicates that it has but one generation/year. The graph of their seasonal

No. sunflower heads inbloom/day

VOLUME 58, NUMBER 2

115

Fig. 12. Seasonal pattern of sunflower bloom and seasonal abundance of Andrena helianthi

and Nomada n. sp.

appearance appears bimodal, but these peaks are influenced by the blooming

pattern of the three plantings (Fig. 12).

Daily activity. — Andrena. During the first part of the season more adults

were observed at the 0900 hour count than during the 1100 and 1300 hour

periods combined. This pattern changed in late August, when the 0900 and

1100 hour counts were nearly the same. Only 9.8% of all Andrena counted

were recorded at 1300 hours. Their absence from the flowers and the closure

of the nests during the afternoon indicates that females remain in the bur¬

rows at this time of day. Males were not seen clustering on the flower heads,

and it is not known where they spend the night.

Nomada .—These cuckoo bees did not appear until late in the season

(August 10), but they were observed on all subsequent days during which

counts were made (Fig. 12). More Nomada were observed on the flower

heads at 1300 hours (55) than at 0900 (38) or 1100 hours (23). Apparently

the cuckoo bees enter the host nests in the morning when the Andrena are

116

PAN-PACIFIC ENTOMOLOGIST

out foraging and forage on the sunflowers in the afternoon when the host

bees are working in their nests.

Provisioning.—Andrena helianthi foraged on and provisioned its nests

exclusively with sunflower pollen. The maximum pollen load carried by

females was estimated (by washing the pollen grains from the body hairs

and counting them with a hemocytometer) at 250,000 grains.

Discussion

Andrena accepta, the only other species of Callandrena whose biology

is known, has communal nests (Rozen, 1973). Although burrows of A. he¬

lianthi were not monitored during the nesting season, the number of cells/

nest indicates that the females were solitary in their nesting behavior.

Acknowledgments

Appreciation is extended to our technicians D. Veirs and C. Hatley for

their assistance in conducting the field work. Thanks are also due those

persons who reviewed the manuscript, N. Youssef, Department of Biology,

Utah State University, and R. W. Thorp, Department of Entomology, Uni¬

versity of California, Davis.

Literature Cited

Hicks, C. H. 1926. Nesting habits and parasites of certain bees of Boulder County, Colorado.

Univ. Studies, Series A, 15:217-252.

Hurd, P. D„ Jr. 1979. Apoidea. Pp. 1741-2209 in K. V. Krombein, P. D. Hurd, Jr., D. R.

Smith, and B. D. Burks, Catalog of Hymenoptera in America North of Mexico, Vol. 2.

Smithsonian Institution Press, Washington, D.C.

Linsley, E. G., and J. W. MacSwain. 1955. The habits of Nomada opacella Timberlake with

notes on other species. Wasmann J. Biol., 13:253-276.

Rozen, J. G. 1973. Biological notes on the bee Andrena accepta Viereck. J. N.Y. Entomol.

Soc., 81:54-61.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 117-123

OBSERVATIONS ON THE BIOLOGY AND LIFE HISTORY OF THE

NET-WINGED MIDGE DIOPTOPSIS SEQUOIARUM (ALEXANDER)

(DIPTERA: BLEPHARICERIDAE)

R. B. Kimsey and E. A. Backus

Department of Entomology, University of California, Davis 95616

The species sequoiarum Alexander was originally described in the genus

Philorus Kellogg and subsequently moved to Dioptopsis by Alexander

(1958). Dioptopsis includes species from Japan, Kashmir, southeast Eu¬

rope, central Asia, and western North America (Alexander, 1958). Diop¬

topsis sequoiarum is one of five species that occur in the western United

States (Hogue, 1970, 1973). 1

Few biological and behavioral data are available on blepharicerids occur¬

ring in the United States. Observations have been made on an eastern

species, Blepharicera capitata Loew (Kellogg, 1900) (cited by Gibo, 1964

as B. tenuipes Walker). Also available are short notes on several species

(Kellogg, 1903), an account of collection habitats for Dioptopsis dismalea

Hogue (Hogue, 1970), and extensive studies on Blepharicera micheneri

Alexander and Philorus yosemite (Osten Sacken) in California (Gibo, 1964).

In the present paper we provide new observations and data on the biology

and behavior of a little-known species, Dioptopsis sequoiarum. An attempt

is made to correlate morphological features, illustrated with scanning elec¬

tron micrographs, and hydrophobic properties of the teneral adult cuticle.

We will relate sex ratios in various life stages and habitats to sexually di¬

morphic behavior patterns. Salient features of the biology of D. sequoiarum

are compared with those of the better known California genera.

Methods and Materials

Study site .—Our study was conducted along an 11.5 km section of Sage-

hen Creek, Nevada County, California. Beginning at the intersection of

Sagehen Creek and Rt. 89, the section extended west and parallel with

Sagehen Creek Road.

The area we studied most extensively was 10 km west of the Rt. 89

junction, a point where Sagehen Creek passes under Sagehen Road through

a large culvert. We designated this area the culvert study site. It includes a

rectangular area 10 m either side of the creek, 25 m either side of the culvert.

The creek flows from the northwest through the culvert and then bends to

the east. The study site is bounded at either end by stands of aspen, lodge-

pole pine, and fir. Above and below these boundaries the creek is narrow,

118

PAN-PACIFIC ENTOMOLOGIST

deep and winding with numerous waterfalls and logjams. The canopy is

dense, and most of the creek is in deep shade.

Within the culvert study site the creek is straight, wide (4.5 m) and uni¬

formly shallow (2-10 cm). The creekbed passes down a uniform 7% grade

and is composed of fine gravel densely overlaid with rocks ranging in size

from 4-80 cm. Vegetation along the margin of the culvert study site was

largely cleared away during construction of the culvert, so, with the excep¬

tion of a few clumps of willow, the creek is open to the sky.

The rest of Sagehen Creek is typical of high, cold eastern Sierra streams

passing through meadows or over beaver dams. The elevation varies from

7000' at the western end of the study area to 5200' at the intersection of the

creek and Rt. 89.

Methods .—Standardized sweepnet surveys (Bowen et al., 1980) were

conducted at various distances away from the creek at several localities

including the culvert study site. Undisturbed adults were also counted in

the culvert and other resting sites. In both of these surveys, sex ratio data

were also collected. Emergence behavior was observed by removing rocks

with pupae to more calm margins of the creek. These rocks were positioned

so that the pupae were constantly washed with a few millimeters of water.

Various substrates were characterized and surveyed for pupae and larvae.

Samples of all life stages were collected and preserved in 4% isotonic (pH

7.4) glutaraldehyde or 70% ethanol. The glutaraldehyde specimens were

then dehydrated and critical-point dried for examination with a scanning

electron microscope.

Results

All life stages, except eggs, were found in a few high density patches

along or in the 11.5 km portion of Sagehen Creek. During both seasons, the

densest population discovered was at the culvert study site. D. sequoiarum

did not occur below 6000', although occasional adult females were taken at

a lower elevation.

Larvae .—The larvae were found exclusively on submerged, smooth rock

substrates. Exposed roots, submerged logs, organic debris, spray-moistened

rocks and the like, though available, were not used. Larvae were found

most commonly in shallow, 1-4 cm, fast-moving white water, on the top,

lateral and downstream aspects of rocks. They could also be found at greater

depths but only at the base of small waterfalls or where water passed over

the top of large rocks creating a deep, turbulent white water basin. In these

circumstances, larvae were found as deep as 15 cm. The specific locality in

the stream seemed to be independent of the distance from the shore, but

seemed to be correlated with areas of higher light intensities.

Two forms of larval locomotion were observed. The first, a slow, forward

progression was achieved by successively detaching, advancing, and reat-

VOLUME 58, NUMBER 2

119

Table 1. Sex ratio data from surveys conducted at Sagehen Creek.

Date

Study site

Distance

from creek

Life stage

Female

Male

Ratio

23 July 1980

Culvert

Pharate adult

2.7:1.0

—

Adult

1.0:9.5

1978, 1980*

All

Less than 10 m

Adult

166

1.0:8.4

Greater than 10 m

Adult

4.5:1.0

* The 1978, 1980 data are summed over the entire study period, at all sites along the 11.5

km Sagehen Creek section.

taching each ventral sucker individually. Dorsally, this appeared similar to

rhythmic caterpillar motion and constituted normal locomotion. The second,

a quick, lateral movement was observed only after the application of intense

stimuli. The anterior or posterior suckers were rapidly detached, and the

free end of the larva was adducted laterally so that the larva assumed a

slight crescent shape. The free suckers were then reattached and, on the

opposite end, were detached. Three or four complete replications of this

sequence, the average number observed in a single event, displaced the

larvae roughly 1 cm from the point of stimulation. This is apparently an

evasive response.

The larval density on individual rocks was fairly low, 1 per 4 cm 2 . This

distribution seemed highly consistent and regular as if each larva was ex¬

erting some form of territoriality.

Pupae .—Pupae were restricted to the same microhabitats and substrates

as the larvae. However, pupae were nearly always found in very dense

clusters of 3-180, rarely deeper than 3 cm.

Pupal orientation was governed by the direction of current flow. The

median, longitudinal axis of the pupal case was parallel to the microcurrent,

with the head oriented downstream. Because the turbulence around the

irregularly shaped and placed rocks was so complex, the orientation of any

two pupae in the same cluster was usually somewhat different, although in

some clusters, even subtle orientation differences were indistinguishable.

Of 54 pupae collected on 23 June 1980 (see Table 1) at the culvert site,

40 were females and 14 were males, a ratio of 2.9:1.0.

Eclosion .—On several occasions teneral adults were observed emerging

directly from torrential water, short distances downstream from pupal clus¬

ters. The following account is a synopsis of the behavior of three eclosing

adult female flies observed on 13 and 22 July 1980 at the culvert study site.

The pupae were on rocks which were moved to slow, clear water at the

creek margin.

The pupal skin split down a median, dorsolongitudinal line through which

120

PAN-PACIFIC ENTOMOLOGIST

the head emerged first. Slowly, over the next 5 min, the thorax appeared,

and the halteres were erected. In the next 3 min, the wings longitudinally

unfolded and began flapping. The wings were very soft and grey and did

not unfold to full width until after flapping had commenced. Next, the very

narrow abdomen was pulled from the pupal skin, followed by the legs. The

insects remained attached to the pupa for 1-2 min, and then detached. They

were carried downstream a short distance and, reaching the surface, im¬

mediately took flight. The wings never ceased beating during the entire

process. The teneral adult, including wings, was completely grey, except

for the eyes. The upper half of the eye was red and the lower was black.

The whole insect was very soft and easily damaged. Eclosion, under these

unnatural circumstances, required 8 min.

During the entire eclosion process, both under water and once they reached

the surface, the insects remained completely dry. When submerged, the

entire fly was covered with a silvery sheen presumably due to a thin film of

air. When attempts were made to wet freshly killed specimens with glutar-

aldehyde fixative, an acutely negative meniscus formed at points of contact.

Subsequent ultrastructural investigations revealed the entire cuticular sur¬

face of the adult to be covered with a dense mat of microtrichia (Figs. 1-

7). This mat varied in structure and density on different areas of the body

and was present even on the wings, eyes, and halteres.

Adults .—Adults were found in groups most commonly underneath logs

overhanging the creek in shady situations. Large numbers were observed

on the ceiling of the culvert. Individuals were also collected with sweepnets

at various distances from the stream (Table 1).

Adult D. sequoiarum were normally abundant on the culvert ceiling at

midday and rare or absent after 1800 hr. On four nonconsecutive days,

adults were counted between 1100 and 1400 hr and again between 1800 and

1900 hr. The mean frequency at midday was 14 whereas, in the evening, it

had dropped to 0.25.

Discussion

Dioptopsis sequoiarum closely resembles D. dismalea in general habitat

choice (Hogue, 1970). Common characteristics include occurrence in open

sections of stream, proximity to meadows, aspens, small willow thickets,

Figs. 1-7. Fig. 1. Head and mesoscutum of pharate male, x8. Fig. 2. Tarsal claw of teneral

female, x300. Fig. 3. Wing dissected from pharate adult female, x40. Fig. 4. Haltere of teneral

female, xl48. Fig. 5. Terminalia of pharate male, x79. Fig. 6. Magnification of wing micro¬

trichia, in Fig. 3, x700. Fig. 7. Magnification of dorsal microtrichia on abdomen in Fig. 5,

x800.

VOLUME 58, NUMBER 2

121

122

PAN-PACIFIC ENTOMOLOGIST

and for the most part, the most torrential water. Descriptions of larval and

pupal habitat correspond almost exactly.

Of the two species, Blepharicera micheneri and Philorus y os emit e, stud¬

ied by Gibo (1964), B. micheneri was closest to D. sequoiarum. The larvae

of B. micheneri were found on submerged rocks or rocks at the base of

waterfalls, at a depth (5.08 cm) comparable to that of D. sequoiarum (1-4

cm). The distribution of larvae was similar; 1 cm apart in B. micheneri and

1 per 4.0 cm 2 for D. sequoiarum. Pupal orientation was somewhat similar;

Gibo described pupae of B. micheneri aligned with the current in fast-mov¬

ing water, but not strictly so in the slow parts of the stream. All pupae we

found were in fast-flowing streams, where they were aligned with the cur¬

rent. The distance from the surface at which pupation occurred was very

similar; 1.27 cm for B. micheneri and not deeper than 3.0 cm for D. se¬

quoiarum.

Aggregations of adult Dioptopsis sequoiarum were found in resting sites

similar to those occupied by Blepharicera micheneri. These sites were char¬

acterized by moist cool shade and close proximity to the stream. D. se¬

quoiarum were never found in aggregations at distances greater than a few

meters from the stream or in direct sunlight, as were Philorus yosemite. D.

sequoiarum prefer to hang upside down from surfaces overhead, rather than

on vertical or horizontal surfaces. These aggregations were composed prin¬

cipally of males.

The sex ratio data, reported above for the various times and localities,

provided some insight into the various aspects of adult behavior. On 23

July, the ratio of soon-to-emerge pupae were skewed to females, 2.7:1.0,

whereas, in the culvert resting site, males were the most common, at 1.0:9.5.

The latter figure is closely correlated with the ratio observed 10 m or

less from the stream, during the entire study. However, the ratio reverses

in favor of females at distances greater than 10 m from the creek. We hy¬

pothesize this shift in ratios is probably caused by a dimorphism between

male mating behavior and female oviposition site selection. Males may tend

to stay near the stream where the likelihood of mating with newly-emerged,

virgin females is fairly good. Females, on the other hand, could be the

invasive, colonizing form, flying considerable distances after copulation, in

search of new oviposition sites. Variation in this behavior could account for

the few females that stay close to the eclosion site assuring the population

for the following year, and for the occasional stray male found great dis¬

tances from the stream. This hypothesis was supported by the observation

of two females apparently ovipositing in very small rivulets, completely

unsuitable for larval development, 3.2 km from any known larval habitat.

Males may also emerge earlier in the season than females, similar to what

one observes in a generation of mosquitoes. This phenology might explain

the dominance of mature female pupae on 23 July, late in the season.

VOLUME 58, NUMBER 2

123

There are at least two specializations of teneral midges that adapt them

to torrential habitats. The first is the wing-folding pattern; visible as the net-

like pattern between veins after which the group is named. The wings are

developed to full length and width in the pupae and at eclosion are imme¬

diately available for flight. Actually, they are already beating before emer¬

gence from the pupae is complete. The second adaptation is the mat of

hydrophobic microtrichia which, on D. sequoiarum, cover the entire body.

These structures probably retain the plastron, keeping the insect dry, and

break the surface tension as the midge takes flight from the torrential stream.

Additionally, the soft, unsclerotized cuticle common to all teneral insects

may be especially important to blepharicerids, preventing permanent dam¬

age by the action of white water during eclosion and before taking flight. In

support of this idea, teneral insects collected as they emerged had legs bent

and curled almost everywhere but at the real joints. Teneral flight is com¬

monly observed in other Nematocera, as well as Ephemeroptera, and Odo-

nata. But, blepharicerids seem to have developed this habit to an extreme.

It is interesting to note that the unique qualities of the culvert study site

(i.e., its straightness, overhanging ceiling, width and shallow depth) are

completely artificial. Most of these attributes were created when the bridge

and culvert were constructed in the early 1960’s. The culvert study site has,

by far, the most dense population of D. sequoiarum, an example of envi¬

ronmental perturbation by man that may have increased the frequency of

an otherwise rare insect.

Literature Cited

Alexander, C. P. 1958. Geographical distribution of the net-winged midges (Blepharoceridae,

Diptera). Proc. Tenth Intemat. Cong. Entomol., 1:813.

Bowen, W. R., V. E. Burton, K. S. Hagen, V. M. Stern, C. G. Summers, and N. C. Toseana.

1980. Insect control guide for alfalfa hay. Ext. Lflt. Univ. Calif. Coop. No. 2763.

Gibo, D. L. 1964. Notes on the biology of Blepharocera micheneri and Philorus yosemite

(Diptera: Blepharoceridae) in southern California. Bull. So. Calif. Acad. Sci., 63:44-53.

Hogue, C. L. 1970. Description of a new species of net-winged midge from the Great Basin,

with a key to the North American species of the genus Dioptopsis (Diptera: Blephari-

ceridae). Cont. Sci. L.A. County Mus., 178:1-10.

Hogue, C. L. 1973. The net-winged midges or Blephariceridae of California. Bull. Calif.

Insect Surv., 15. 83 pp.

Kellogg, V. L. 1900. Notes on the life-history and structure of Blepharocera capitata Loew.

Entomol. News, 11:305-318.

Kellogg, V. L. 1903. The net-winged midges (Blepharoceridae) of North America. Proc.

Calif. Acad. Sci., (3)3:187-232.

Footnote

1 “Blepharicerid taxonomists advise that this and other North American species in the genus

‘Dioptopsis ’ are not congeneric with the type from southeast Europe. The generic placement

used here is provisional only until the relationships of the species are known.” (C. L. Hogue,

pers. comm.)

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 124-128

PSEUDOMASARIS MARGINALIS NESTING IN LOGS IN

COLORADO (HYMENOPTERA: MASARIDAE)

Laurence J. Dorr and John L. Neff

Department of Botany, University of Texas, Austin 78712 and

7307 Running Rope, Austin 78731

Pseudomasaris wasps have attracted attention because of their pollen

collecting activities. Nevertheless, information concerning the nesting bi¬

ology of most of the 15 species of Pseudomasaris recognized by Richards

(1963) is sparse. Mud nests attached to rocks have been described for P.

coquilletti Rohwer (Richards, 1963), P. occidentalis (Cresson) (Hungerford,

1937), P. phaceliae Rohwer, P. zonalis (Cresson) and P. maculifrons (Fox)

(Parker, 1967). Similar mud nests attached to plant stems are known for P.

texanus (Cresson) (Bequaert, 1940). Two species, P. edwardsii (Cresson)

and P. vespoides (Cresson), also construct mud nests, but attach them to

either plant stems or rocks (Davidson, 1913; Cockerell, 1913; Hicks, 1927,

1929, 1931; Torchio, 1970). We here report the discovery of yet a third type

of nest substrate: abandoned beetle tunnels in logs which are used by P.

marginalis (Cresson). This appears to be the first report of a nest substrate

of this type in the Masaridae.

Nest Site

On 11 July 1980 a female of P. marginalis was observed entering and

exiting an abandoned beetle tunnel in a fallen bristlecone pine {Pinus aris-

tata Engelmann) log. The log was located at timberline (3550 m elevation)

on the southeastern side of Pennsylvania Mountain, west of Fairplay, Park

County, Colorado. It was oriented so that the nest entrance was on the

upper surface of the log approximately 5 dm above the ground. When the

nest was reexamined on 14 July 1980 the entrance had been sealed with

mud.

Nest Architecture

Upon opening the nest in mid August 1980, it was found to consist of a

linear series of four cells (Fig. 1). The nest occupied the initial curved 60

mm of the beetle tunnel; the distal portions of the tunnel being filled with

sawdust. The two innermost cells contained prepupae with their heads ori¬

ented toward the nest entrance, while the two outer cells were empty. The

innermost cell (Fig. 1) was lined at the bottom with a thin layer of mud

which abutted the sawdust left in the tunnel by the beetle (Fig. 1, a). The

VOLUME 58, NUMBER 2

125

Figs. 1-3. Nest of Pseudomasaris marginalis. Fig. 1. Lateral section through nest showing

the orientation of the four cells. The cells are numbered 1 through 4. The partitions are labelled

a, b, c, d and e. Fig. 2. Outline of cross section through cell number one. Fig. 3. Spiral closing,

lower surface of partition e which separates cells three and four.

top of this cell was closed with a thin, 0.9 mm thick, mud plug (Fig. 1, b).

This cell was 11.2 mm long and 3.9 mm wide and as with all the other cells

it was somewhat flattened in cross section (Fig. 2) because the beetle tunnel

itself was flattened in cross section.

There was a small space between the closing of the lowermost cell and

the base of the next cell. The mud base of this second cell (Fig. 1, c) was

2.1 mm thick and slightly depressed in the center. This second cell was 10.1

mm long and 4.3 mm wide. The mud partition separating the second and

third cells (Fig. 1, d) was 0.4 mm thick and the mud extended along the cell

walls several mm below the partition.

The third cell was empty, containing no signs of provisions, feces or any

other indication of occupation or use. This was the shortest cell, 8.2 mm

long and 4.0 mm wide. It was separated from the fourth cell by a 1.5 mm

thick partition (Fig. 1, e) which had an obscure spiral pattern on its inner

surface (Fig. 3). This closure might alternatively be considered to be formed

of three uneven concentric circles.

The fourth cell was also empty. There was a suggestion of an incomplete

partition halfway up the cell. This cell was 13.0 mm long and 4.4 mm wide.

It was capped by the nest closing, which had a nipple at its base. Cells three

and four appeared to have a thin lining of mud while cells one and two

lacked this lining.

The nest closing was not flush with the wood surface, but was slightly

126

PAN-PACIFIC ENTOMOLOGIST

depressed in the center. It was made of the same fine grained, very hard

clay as were the other mud partitions and was 9.9 mm thick. At the surface

the nest closing was 11.4 mm long and 10.5 mm wide, but it tapered toward

the top of cell four.

Provisions

Grains of Phacelia sericea A. Gray pollen were found adjacent to the

cocoon in cell two. Adult females of Pseudomasaris marginalis were col¬

lected on Pennsylvania Mountain in July and August foraging on Phacelia

sericea at 3550 and 3660 m elevations. A single male was also taken on

Pennsylvania Mountain in late June on Polemonium delicatum Rydberg at

approximately 3500 m elevation.

Cocoons

Cells one and two were lined with transparent cocoons. These cocoons

consisted of fine threads and a matrix of very thin sheet-like material, the

thickness of the cocoon lining varying considerably. For the most part, it

was very thin, strictly adhering to the cell walls. Where the cocoons abutted

the wood substrate, they frequently did not form a complete layer. Elsewhere

cocoon finings were thicker, forming complete sheets and they were readily

extractible from the cell walls. This latter pattern was most evident on the

lateral sections of cocoons in the posterior portion of the first cell and the

anterior portion of the second cell.

Feces

The bottom of the innermost cell was covered with a grey excretory mass

and that of the second cell with a greyish-green excretory mass that was

similar to the first in texture. Small, hard, shiny black fecal pellets were in

the outer fining of the cocoon and adjacent to the cocoon in the basal part

of cell one and the anterior portion of cell two. These pellets were from 0.9

to 1.0 mm in length and 0.5 to 0.6 mm in width.

Parasites

No parasites were present in the nest.

Discussion

The most detailed report on the nesting biology of a Pseudomasaris wasp

is that of Torchio (1970) for P. edwardsii. While the principal difference

between our report and that of Torchio (1970) concerns the nest substrate

chosen by the wasps, the nest of P. marginalis also differs from the nests

of P. edwardsii in a few other aspects.

The cells of the P. marginalis nest are in a linear series. This is undoubt¬

edly due to the constraints imposed by the form of the beetle burrow. Tor-

VOLUME 58, NUMBER 2

127

chio (1970) noted that generally cells of P. edwardsii nests are joined along

their lateral margins although he did observe one nest placed in a long

narrow groove in which the cells were arranged in a linear series.

The outermost two cells in the P. marginalis nest are empty. Torchio

does not report empty cells for P. edwardsii nests, but empty cells are

apparently common in nests of P. vespoides (Tepedino et al., 1979). The

latter suggest that these empty cells may represent a defense against para¬

sitism, but such explanations are questioned by Krombein (1967).

The cocoons of the P. marginalis larvae appear to be essentially similar

to those described for P. edwardsii. The nature of the feces differ, however.

Whereas the feces of P. marginalis are individual pellets, those of P. ed¬

wardsii are aggregated into fecal cakes (Torchio, 1970).

Finally, although the cells in the P. marginalis nest we examined are

represented by soil partitions and wooden walls, other Pseudomasaris con¬

struct complete soil cells in which both the cell cap and walls are made of

mud (Torchio, 1970). Torchio (1970) reports that the cell closures of P.

edwardsii cells are formed by smooth concentric circles of mud. The caps

of the two empty cells of our P. marginalis nest are definitely nippled on

their inner surfaces and appear to have a spiral structure. The inner surfaces

of the cell caps of the provisioned cells are relatively smooth with the fine

structure obscured by the closely adhering cocoons. It is not clear if this is

a real difference between the taxa or an artifact due to the absence of empty

cells in P. edwardsii nests.

While our report is the first concerning the use of a beetle burrow in wood

as a nest by any species of Pseudomasaris, the nest we describe does not,

as we have indicated, represent a radical departure from the type of nests

previously described. It does suggest that there is more variation within the

genus than previous reports have suggested. It would be of interest to know

if this use of beetle tunnels is typical of all P. marginalis and if similar nests

are constructed by P. macneilli R. M. Bohart, an apparently closely related

species of high altitudes in California and Utah.

Acknowledgments

Field work was supported by NSF grant DEB-79-10786 to P. G. Kevan.

Vouchers have been deposited in the United States National Museum.

Literature Cited

Bequaert, J. 1940. Notes on the distribution of Pseudomasaris and on the foodplants of the

Masaridinae and Gayellinae. Bull. Brooklyn Entomol. Soc., 35:37-45.

Cockerell, T. D. A. 1913. Pseudomasaris bred in California. Proc. Entomol. Soc. Wash.,

15:107.

Davidson, A. 1913. Masaria vespoides. Bull. Southern Calif. Acad. Sci., 12:17-18.

128

PAN-PACIFIC ENTOMOLOGIST

Hicks, C. H. 1927. Pseudomasaris vespoides (Cresson), a pollen provisioning wasp. Can.

Entomol., 59:75-79.

Hicks, C. H. 1929. Pseudomasaris edwardsii Cresson, another pollen provisioning wasp,

with further notes on P. vespoides (Cresson). Can. Entomol., 61:122-125.

Hicks, C. H. 1931. Notes on pollen-user wasp, Pseudomasaris edwardsii Cresson. Bull.

Southern Calif. Acad. Sci., 30:23-29.

Hungerford, H. B. 1937. Pseudomasaris occidentalis (Cresson) in Kansas (Hymenoptera—

Vespidae). J. Kansas Entomol. Soc., 10:133-134.

Krombein, K. V. 1967. Trap-nesting wasps and bees: Life histories, nests, and associates.

Smithsonian Press, Washington, D.C., 570 pp.

Parker, F. D. 1967. Notes on the nests of three species of Pseudomasaris Ashmead (Hy¬

menoptera: Masaridae). Pan-Pac. Entomol., 43:213-216.

Richards, O. W. 1963. The species of Pseudomasaris Ashmead. Univ. Calif. Publ. Entomol.,

27:283-310.

Tepedino, V. J., L. L. McDonald, and R. Rothwell. 1979. Defense against parasitization in

mud-nesting Hymenoptera: Can empty cells increase net reproductive output? Behav.

Ecol. Sociobiol., 6:99-104.

Torchio, P. F. 1970. The ethology of the wasp, Pseudomasaris edwardsii (Cresson) and a

description of the immature forms (Hymenoptera: Yespoidea, Masaridae). Los Angeles

Co. Mus. Nat. Hist. Contrib. Sci. No. 202, 32 pp.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 129-134

USE OF THE PROBOSCIS FOR PREY-PIERCING AND

SUCKING BY SPHECID WASPS OF THE GENUS PRIONYX

(HYMENOPTERA: SPHECIDAE), A CASE OF

CONVERGENT EVOLUTION

Andre L. Steiner

Department of Zoology, University of Alberta,

Edmonton, Alberta T6G 2E9, Canada

Most sphecid wasps feed mainly on nectar of flowers, sap, exudations or

aphid honeydew, which they lick. Some are also known to use body fluids

diverted from the prey they catch for their larvae. This supplements their

usual diet or perhaps in some cases represents their exclusive diet. Ampulex

wasps, for instance, have been seen amputating part of both antennae of

their paralyzed roach-prey and feeding on the exuding body fluids (Williams

and others, in Bohart and Menke, 1976, p. 75). Other wasps, such as Xy-

locelia (Powell, 1963, p. 162) or Pemphredon (in Bohart and Menke, 1976,

p. 180) suck dry some of their aphid prey and do not use them for nesting

(also pers. obs.). Still other wasps catch even a different prey for their own

consumption. Thus the nyssonine wasp Stictia signata (Linnaeus) was ob¬

served on the Amazon River feeding on Aedes aegypti (Linnaeus) mosqui¬

toes whereas it fed horseflies to its larvae (Howard et al., 1912). Oxybelus

emarginatus Say captured Diptera much smaller than the usual prey, crushed

them and consumed the body fluids (Snoddy, 1968, p. 1030). Many more

instances can be found in the literature.

Most sphecid wasps have rather short, unspecialized mouthparts, which

can also be used for chewing or crushing prey or opening a hole to extract

body fluids. In contrast, many bees have often developed highly specialized

and elongate mouthparts, adapted to the exploitation of very specialized

flowers, with deep corollas. A few sphecid wasps, notably some bembicine

wasps, have evolved similar specializations. Bembix and even more so

Steniolia wasps have a long or very long proboscis they use to exploit

flowers with deep or very deep corollas, as hummingbirds and hawkmoths

do. In contrast to bees, however, this long, pointed and stiff proboscis can

also be used to puncture the Diptera they take as prey and to suck them

dry, as robberflies (Asilidae) do. Ferton (1897, 1899, 1902) was one of the

first to report such behavior from Bembix wasps (B. oculata Panzer, B.

rostrata (Linnaeus)). These prey were never used for nesting. Microbembex

monodonta (Say) also sucks Diptera (Hartman, 1905, p. 24). Janvier (1928)

observed in South America Bembix brullei Guerin-Meneville that stabbed

130

PAN-PACIFIC ENTOMOLOGIST

Diptera with the proboscis, on the ventral side of the thorax, in front of the

forelegs, sucked them dry and then discarded the remains. Nielsen (1945,

p. 32) thinks that such behavior is motivated by thirst, not hunger, as Ferton

originally suggested.

Prionyx wasps have a much less elongate proboscis but they can also use

it for stabbing, piercing their grasshopper-prey (pers. obs.) in addition to

licking nectar in moderately deep corollas. The latter is probably the primary

function and the former a derived function. Such stabbing, followed by

licking of body fluids, was consistently observed as part of a detailed be¬

havioral study of Prionyx parked Bohart and Menke, conducted in S.E.

Arizona (1972-1973), then in west-central Oregon (1977) and summarized

by Steiner (1976). The prey were Acrididae, mainly Oedipodinae, but also

some Cyrtacanthacridinae (Steiner, in prep.) taken as adults or last nymphal

instars. This behavior usually followed prey stinging and was preceded by

a thorough investigation of the latero-dorsal surface of the neck membrane

(Fig. 1).

First the wasp was usually seen compressing rhythmically this neck area

with the jaws (“kneading,” also observed by Peckham and Kurczewski,

1978, on crickets paralyzed by Chlorion aerarium Patton). Then the wasp

appeared to stab the neck membrane with the proboscis and started to lap

some body fluids deep in the wound, perhaps as deep as the crop. Subse¬

quent examination of such grasshoppers invariably revealed the presence

of one latero-dorsal “stabbing wound” on only one side of the neck (Fig.

2). This single wound did not appear to result from the action of the jaws

otherwise two more or less symmetrical wounds would be observed, one

on each side of the neck. Such symmetrical wounds were found regularly

on each side of the foreleg bases of crickets and grasshoppers used as prey

by Liris and Tachysphex wasps, respectively (Steiner, 1962, 1976). These

wasps, and also apparently Larra analis Fabricius, that preys on Gryllotalpa

hexadactyla Perty mole crickets (Smith, 1935), performed vigorous chewing

motions at the base of these forelegs and then lapped the fluids which oozed

from the wounds. For Liris wasps, this behavior often occurred right after

prey stinging (feeding behavior?) and was consistently repeated inside the

burrow (“malaxation”), just before egg-laying. Now, however, the forelegs

of the prey were vigorously compressed and at the same time pushed for¬

wards, immediately followed by egg-laying, right behind the forelegs. This

second behavior, although superficially very similar to the first one, except

for pushing, is therefore no longer feeding but pre egg-laying behavior in¬

stead, in other words part of nesting behavior (Steiner, 1962, 1971). In con¬

trast, neck puncturing of the grasshopper by Prionyx wasps is not repeated

before egg-laying and the oviposition site is no longer located in the same

area but at the base of one hind leg. In Prionyx it is therefore not pre egg-

laying or a “preparation” of the oviposition site. The primary or sole func¬

tion of this behavior appears to be feeding, absorption of fluids or at least

VOLUME 58, NUMBER 2

131

Fig. 1. Prionyx parked about to perform “malaxation” behavior on the dorsal side of the

neck of an oedipodine grasshopper, shortly after prey-stinging.

sampling of the latter. A somewhat comparable behavior was also observed

in other Prionyx species, for instance P. albisectus Lepeletier and Serville

by Roth (1925) and P. subfuscatus Dahlbom by Ferton (1902). In the latter

case, however, wasp and prey were mouth to mouth, therefore this behavior

is perhaps different from the one described in P. parked. Ferton described

this as feeding behavior and thought it was related with a great scarcity of

flowers in the arid region considered. Such grasshoppers were discarded

and not subsequently used. In the Arizona study of P. parked 59 instances

of neck puncturing were recorded on a total of 44 different grasshoppers

(some were punctured several times); the status of 21 other grasshoppers

in this respect was unknown and only 5 remaining grasshoppers were known

with certainty not to have been punctured. Therefore, the majority of grass¬

hoppers (at least 44 out of 70 and probably many more) were treated in this

way. Position of the wound varied relatively little except for an occasional

wound located near the median dorsal line rather than latero-dorsally, as

shown in Fig. 2.

As an alternative or complementary explanation, this behavior might rep¬

resent licking up of the defensive fluid often used by the attacked grasshop¬

per and regurgitated through the mouth (Steiner, 1976, also detailed study

in prep.). Prionyx wasps have been seen lapping up fluids exuding from the

132

PAN-PACIFIC ENTOMOLOGIST

Fig. 2. Paralyzed adult grasshopper (Psoloessa delicatula Scudder) showing “stabbing

wound” in latero-dorsal neck region, resulting from “malaxation” by a Prionyx parked wasp.

mouth of the prey (pers. obs.; see also for instance Ferton, 1902; Piel, 1935

for P. subfuscatus and Evans, 1958 for P. atratus Lepeletier). This fluid

has a clear repelling, unpleasant or even deleterious effect on the wasps,

when it comes in contact with their body, abdomen tip, during stinging. It

triggers vigorous body rubbing and can stop the attack (Steiner, 1976). It

might also be noxious for the egg and larva. Conceivably, the wasp might

try to eliminate this defense by removing the fluid and lapping it up, from

the mouth, or even through the neck wound, internally. The first paralyzing

sting in the throat also prevents or stops regurgitation, among other effects,

if delivered quickly enough (Steiner, 1976). Besides or instead of feeding,

neck puncturing could then have a protective function for the wasp, egg, or

both. More research is clearly needed.

Podalonia wasps and their agrotid caterpillar-prey (cutworms) exhibit a

very comparable or identical behavior (pers. obs.), namely head-neck

“kneading” and/or puncturing with the rather pointed proboscis for the

wasp, mouth regurgitation of a defensive fluid for the prey. Lapping of this

fluid was also observed, dorsally or ventrally, and also the vigorous body

rubbing, even contortions (Fabre’s “victory dance”?), following contact of

the fluid with the body of the wasp. The compression motions of the jaws

have also been interpreted by some as an aid to paralysis, due to better

venom diffusion and/or direct mechanical action on the brain. Mole crickets

attacked by Larra wasps release an even more potent, very viscous, defen¬

sive fluid in which the wasps can become entangled (Williams, 1928).

Various “preparations” of prey by sphecid wasps might therefore have

a different functional significance and evolutionary origin. In some cases no

link with feeding or absorption of fluid is apparent and only a nesting func¬

tion can be detected. Thus Oxybelus and some other crabronine wasps

prevent the egg-bearing fly from falling on the side by extending laterally

one wing of the prey. This is clearly pre egg-laying behavior but contrary

to Liris wasps it is apparently completely divorced from feeding or lapping

of body fluids.

VOLUME 58, NUMBER 2

133

Like Liris and Tachysphex, Prionyx parkeri wasps do regularly use the

prey for nesting, after the neck-puncturing has been performed. Such prey,

which are never sucked dry, are not necessarily discarded as Ferton stated

for P. subfuscatus. Therefore, this treatment must be compatible with sat¬

isfactory preservation of the prey and successful nesting, otherwise wasps

using such prey would probably have been selected against. Of the 44 punc¬

tured grasshoppers, 23 were subsequently used for nesting (these wasps

store only one prey per nest). The remaining 21 were not used, but not

necessarily because they had been punctured, since some non-punctured

ones were also discarded. A meaningful comparison of nesting success of

punctured vs. non-punctured grasshoppers is difficult in this study, how¬

ever, first because no special effort was made to raise the larvae in optimal

conditions, second too many grasshoppers were of unknown status because

they were stored before their neck area could be checked. At any rate here

are the inconclusive results: a total of 21 nests with punctured grasshoppers

and 14 of unknown status or non-punctured grasshoppers were dug up. Two

nests of each category were destroyed accidentally, leaving 19 and 12, re¬

spectively. Nesting success was as follows; larvae that reached maturity

and spun a cocoon: 5 (=26.3%) vs. 4 (=33.3%), respectively; adults pro¬

duced the next year: 3 (=15.8%) vs. 1 (=8.33%). Clearly these numbers are

too small to warrant reliable conclusions. It is clear, however, that neck-

puncturing is compatible with nesting success and did not reduce the latter

drastically, when compared with the other category.

In conclusion, apparent derivation of prey-piercing behavior from nectar¬

licking behavior by evolution of a stronger, more specialized, piercing pro¬

boscis, as seen in some sphecid wasps (particularly Bembix ), occurred at

least once more, in another group of unrelated insects, namely a few fruit¬

piercing and skin-piercing (blood-sucking) noctuid moths (Banziger, 1971,

1975). They must also have evolved from the more common nectar-feeding

forms, with a non-piercing proboscis. This is apparently a case of conver¬

gent evolution.

Furthermore, at least one sphecid wasp, Oxybelus emarginatus, is known

to lick blood droplets from the cattle exposed to their blood-sucking (sim-

uliid) fly prey (Snoddy, 1968). They do not, however, pierce the skin of the

cattle like the blood-sucking moths do. The latter might well have gone

through a similar stage of dependence from blood excreted by mosquitoes

before becoming able to pierce the skin themselves (Alcock, 1975, p. 401).

Acknowledgments

The above observations are part of a sabbatical project conducted at the

Southwestern Research Station, at Portal, Arizona, of the American Mu¬

seum of Natural History, New York. Research was supported in part by a

grant from NRCC (A3499). Help from the staff and guests of these institu-

134

PAN-PACIFIC ENTOMOLOGIST

tions is gratefully acknowledged. A. S. Menke, U.S. National Museum

(Entomology), Washington, kindly identified the wasp specimens and D. C.

Rentz, The Academy of Natural Sciences, Philadelphia, the grasshoppers.

Literature Cited

Alcock, J. 1975. Animal Behavior, an Evolutionary Approach. Sinauer, Sunderland, Mas¬

sachusetts, 547 pp.

Banziger, H. 1971. Blood sucking moths of Malaya. Fauna, 1:4-6.

Banziger, H. 1975. Skin-piercing blood-sucking moths. I. Ecological and ethological notes

on Calpe eustrigata (Lepid., Noctuidae). Acta Tropica, 32:125-144.

Bohart, R. M., and A. S. Menke. 1976. Sphecid Wasps of the World, a Generic Revision.

University Calif. Press, Berkeley, 695 pp.

Evans, H. E. 1958. Studies on the nesting behavior of digger wasps of the tribe Sphecini.

Part I: Genus Priononyx Dahlbom. Ann. Ent. Soc. Amer., 51:177-186.

Ferton, C. 1897. Nouvelles observations sur l’instinct des Pompilides. Actes Soc. Linn.,

Bordeaux, 52 [p. 123].

Ferton, C. 1899. Observations sur l’instinct des Bembex Fabr. Actes Soc. Linn., Bordeaux,

54:1-15.

Ferton, C. 1902. Notes detachees sur V instinct des hymenopteres melliferes et ravisseurs.

2 e Ser., Ann. Soc. Entom. Fr., 71:449-530.

Hartman, C. 1905. Observations on the habits of some solitary wasps of Texas. Bull. Univ.

Tex. (Scientif. Ser. 7), 65, 73 pp.

Howard, L. O., H. G. Dyar, and F. Knob. 1912. The mosquitoes of North and Central

America and the West Indies. Carnegie Inst., Washington, Pub. 159, 1064 pp.

Janvier, H. 1928. Recherches biologiques sur les predateurs du Chi li . Ann. Sci. Nat. Zool.,

(10)11:67-207.

Nielsen, E. T. 1945. Moeurs des Bembex. Spol. Zool. Mus. Haun., 7, 174 pp.

Peckham, D. J., and F. E. Kurczewski. 1978. Nesting behavior of Chlorion aerarium. Ann.

Ent. Soc. Amer., 71:758-761.

Piel, O. P. 1935. Recherches biologiques sur les Hymenopteres du Yang-Tse (Chine). Etude

sur les Sphegides. Ann. Soc. Entom. Fr., 104:273-306.

Powell, J. A. 1963. Biology and behavior of Nearctic wasps of the genus Xylocelia with

special reference to X. occidentalis (Fox) (Hym.: Sphecidae). Wasmann J. Biol.,

21:155-176.

Roth, P. 1925. Les Sphex de l’Afrique du Nord. Ann. Soc. Entom. Fr., 94:365-404.

Smith, C. E. 1935. Larra analis Fabricius, a parasite of the mole cricket Gryllotalpa hex-

adactyla Perty. Proc. Ent. Soc. Wash., 37:65-82.

Snoddy, E. L. 1968. Simuliidae, Ceratopogonidae, and Chloropidae as prey of Oxybelus

emarginatum. Ann. Ent. Soc. Amer., 61:1029-1030.

Steiner, A. L. 1962. Etude du comportement predateur d’un Hymenoptere sphegien: Lins

nigra V.d.L. (=Notogonia pompiliformis Panz.). Ann. Sci. Natur. Zool. Biol. Anim.,

Ser. 12, 4:1-126.

Steiner, A. L. 1971. Behavior of the hunting wasp Liris nigra V.d.L. (Hym., Larrinae) in

particular or in unusual situations. Can. J. Zool., 49:1401-1415.

Steiner, A. L. 1976. Digger wasp predatory behavior (Hym., Sphecidae). II. Comparative

study of closely related wasps (Larrinae: Liris nigra, Palearctic; L. argentata and L.

aequalis, Nearctic) that all paralyze crickets (Orthop., Gryllidae). Z. Tierpsychol., 42:

343-380.

Williams, F. X. 1928. Studies in tropical wasps—their hosts and associates (with descriptions

of new species). Bull. Exp. Sta. Hawaiian Sugar Planters’ Assoc. (Ent.), 19:1-179.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 135-138

PALTOTHEMIS CYANOSOMA, A NEW SPECIES OF

DRAGONFLY FROM MEXICO (ODONATA: LIBELLULIDAE)

Rosser W. Garrison

1030 Fondale Street,

Azusa, California 91702

The libellulid genus Paltothemis Karsch has long contained one species,

P. lineatipes Karsch, found commonly in the southwestern United States

south to Venezuela (Racenis, 1953) and Brazil (Calvert, 1899). While ex¬

amining the dragonfly collection at the University of California, Davis, I

found three male specimens of a large, pruinose blue dragonfly incorrectly

labeled “Scapanea frontalis.” Subsequent examination of this material shows

them to represent an undescribed species which keys to the genus Palto¬

themis in Borror (1945).

Paltothemis cyanosoma, new species

(Figs. 1-3,5)

Head. —Labium dark brown becoming pale yellow brown toward sides

and margin; labrum black; mandibles red brown with tips black, yellow

brown at base; ante- and postclypeus dark brown, becoming gray brown on

sides of postclypeus and about fronto-clypeal sulcus. Frons rugose, dark

metallic purple becoming red brown at sides; vesicle of vertex divided lon¬

gitudinally with prominent pointed cone forming on each side, vertex ru¬

gose, dark metallic purple becoming red brown above. Eye seam as long as

occiput, latter red brown; antennae black; rear of head black, red brown

immediately posterior to eyes. Rear margin of head sprinkled with long

white hairs.

Thorax. —Entirely dark brown covered by dark blue pruinosity; pro thorax

pale brown at margin of lobes, otherwise pruinose blue; posterior lobe small,

rounded, bent posteriorly, without hairs. Pterothorax with no discernible

pattern, carinae black; front and sides covered with dark hairs. Femora red

brown, black at tips; armature black. Tibiae and tarsi black. Wings as shown

in Fig. 1, hyaline, distal'half slightly infumated; anal margin in hind wing

with faint patch of pruinose white extending distally 2-3 cell rows; venation

black.

Abdomen. —Slightly swollen at base, narrowing at middle before expand¬

ing slightly at tip; segments 1-3 dark pruinose blue, becoming black in re¬

maining segments. In life, entire abdomen probably pruinose blue. Abdom¬

inal pattern largely obscured by pruinosity and postmortem changes in

136

PAN-PACIFIC ENTOMOLOGIST

I 5mm |

Figs. 1, 2. Paltothemis cyanosoma, n. sp., paratype, 3. Fig. 1. Wings. Fig. 2. Abdominal

segments 1-4.

holotype and one paratype. Other paratype probably the youngest of 3 with

abdominal pattern as shown in Fig. 2. Juvenile abdominal coloration prob¬

ably gray and black. Carinae and abdominal appendages black. Secondary

genitalia (Fig. 3) similar to P. lineatipes (Fig. 4), but with genital lobe round¬

ed and posterior margin of anterior lamina in lateral view almost straight or

slightly convex. Hamules similar to those of P. lineatipes. Penis (Fig. 5)

lacking prominent horn on ventral margin of glans present in P. lineatipes

(Fig. 6). Lateral lobe more developed than in P. lineatipes.

Hind wing length (measurements in parentheses are for holotype): 41-44

mm (44 mm); pterostigma: forewing: 3.5-3.8 mm (3.8 mm), hind wing: 3.5-

VOLUME 58, NUMBER 2

137

Figs. 3-6. Figs. 3,5. Paltothemis cyanosoma , paratype, n. sp. S . Figs. 4, 6. P. lineatipes 6 .

Figs. 3,4. Accessory genitalia. Figs. 5, 6. Penis: a, lateral view; b, dorsal view. L = lateral lobe.

3.9 mm (3.8 mm); antenodal crossveins: forewings: 15-17 (16 left wing, 17

right wing), hind wing: 10-11 (10 left wing, 10 right wing); postnodal cross¬

veins: forewing: 10-13 (13 left wing, 12 right wing), hind wing: 12-14 (14

left wing, 13 right wing).

Holotype. —Male: Mexico, state of Jalisco, 6 mi N of Guadalajara, 13

August 1970 (Baldomero Villegas). Two paratype males: same data as ho¬

lotype. Holotype and 1 paratype in collection of University of California,

Davis, 1 paratype in author’s collection.

In response to my inquiry about the locality of the captures, B. Villegas

(in litt., 24 Oct. 1978) responded: “As I remember, on August 13, 1970, I

collected about 6 miles north of Guadalajara near the town of Experiencia.

The stream that I collected at is within a mile or so from this town. Fur¬

thermore, this stream empties into a large canyon located north of Guada¬

lajara. This canyon is called ‘Barranca de Oblatos.’” This species probably

has habits similar to P. lineatipes. I have taken P. lineatipes on small streams

where adults generally perch on large, exposed rocks in the stream. Dunkle

(1978) gives further details of adult behavior.

138

PAN-PACIFIC ENTOMOLOGIST

Male P. cyanosoma are easily distinguished from P. lineatipes by overall

coloration. Male P. lineatipes are red and black, possess degrees of orange

at the base of the wings, especially around the cubito-anal and triangle

regions, and the basal half of the wing venation is red. No red is present on

the wings or body of P. cyanosoma. Three structural characters further

segregate the two: The genital lobe is rounded in P. cyanosoma (Fig. 3),

but truncate in P. lineatipes (Fig. 4); the hood of the penis lacks the prom¬

inent horn present in P. lineatipes, and the hind wings are relatively nar¬

rower than in P. lineatipes. Dunkle (1978) gives ratio of maximum hind wing

width to length as 1:2.7 in P. lineatipes. Measurements from other speci¬

mens in my collection show a range of 1:2.6 to 1:2.8. The same ratio for P.

cyanosoma ranges from 1:2.9-1:3. The number of cells bordering the midrib

of the anal loop from the ankle to toe (terminology after Needham and

Westfall, 1955) ranges from 3-4 in P. cyanosoma, 4-6 in P. lineatipes. The

ratio of sole to gaff length is 0.76 to 0.82 ( x = 0.79, N = 3) in P. cyanosoma,

and is 0.71-1.00 (x = 0.86, N = 29) in P. lineatipes, but the means are not

significantly different (F 05[130] = 3.28).

Acknowledgments

Special thanks are due to Mr. Robert O. Schuster, University of Califor¬

nia, Davis, for allowing me to examine the specimens; Drs. Dennis R. Paul¬

son, University of Washington, Seattle, and Minter J. Westfall, Jr., Uni¬

versity of Florida, Gainesville, for criticizing the manuscript; Dr. Baldomero

Villegas for providing information concerning the type locality; and to my

wife, Jo, who typed the manuscript and provided helpful editorial com¬

ments .

Literature Cited

Borror, D. J. 1945. A key to the New World genera of Libellulidae (Odonata). Ann. Entomol.

Soc. Amer., 38:168-194.

Calvert, P. P. 1899. Odonata from Tepic, Mexico, with supplementary notes on those of

Baja California. Proc. Calif. Acad. Sci., Zool., (3)1:371-418.

Dunkle, S. W. 1978. Notes on adult behavior and emergence of Paltothemis lineatipes Karsch,

1890 (Anisoptera: Libellulidae). Odonatologica, 7(3):277-279.

Needham, J. G., and M. J. Westfall, Jr. 1955. A Manual of the Dragonflies of North America

(Anisoptera). University of California Press, Berkeley and Los Angeles, xii + 615 pp.

Racenis, J. 1953. Contribucion al estudio de los Odonata de Venezuela. Anales Univ. Central

Venezuela, 35:31-96.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 139-152

LIFE HISTORY AND GENERAL BIONOMICS

OF TRIRHABDA SERICOTRACHYLA BLAKE

(COLEOPTERA: CHRYSOMELIDAE)

IN SOUTHERN CALIFORNIA

Patrick Y. O’Brien and Peter R. Atsatt

Department of Ecology and Evolutionary Biology,

University of California, Irvine 92717

This study examines life history characteristics of Trirhabda sericotra-

chyla Blake 1 in relation to its host plant, Artemisia californica Less. (As-

teraceae). The results of laboratory rearing studies were correlated with field

observations made at several locations within 5 km of the University of

California, Irvine, campus from 1977 to 1979. Host plant records for other

sympatric Trirhabda spp. observed during the field phase of the study are

also summarized.

Systematics and Host Plant Relationships

Other than the original description and host records (Blake, 1931; Hogue,

1970), biological information about T. sericotrachyla is lacking. The genus

was introduced by LeConte (1865) and its systematics have been reviewed

several times (Blake, 1931, 1951; Wilcox, 1965; Hogue, 1970). Thirty-seven

species, including three fossil forms, comprise the genus which is distributed

throughout North and Central America (Wilcox, 1971). According to field

observations, analysis of host records, and rearings, Hogue (1970) conclud¬

ed that species of Trirhabda were restricted to host plants in the Asteraceae

and Hydrophyllaceae. The available literature recently reviewed by Hogue

(1970) includes accounts of the life histories of T. canadensis (Kirby) (Bal-

duf, 1929), T. flavolimbata (Mannerheim) (Tilden, 1953), T. pilosa (Blake)

(Arnott, 1957; Pringle, 1960; Banham, 1961), and T. nitidicollis LeConte

(Massey and Pierce, 1960). A description of the larva of T. canadensis and

notes on several other species appeared in Boving (1929).

Four species of Trirhabda probably occur sympatrically in the San Joa¬

quin Hills of Orange County, California, where they segregate according to

differences in host plant preference and/or suitability. Three species, T.

sericotrachyla, T. luteocincta LeConte, and T. confusa Blake, have pre¬

viously been reported from this area (Hogue, 1970). Trirhabda confusa was

not located during this study but the senior author observed another species,

T. geminata Horn, on a previously unreported host, Encelia californica

Nutt. (Asteraceae). Older specimens in the University of California, Irvine,

140

PAN-PACIFIC ENTOMOLOGIST

Museum of Systematic Biology confirm this finding. Previous host records

(Blake, 1931; Hogue, 1970) indicate that T. geminata occurs on E. farinosa

Gray and E. virginensis A. Nels. in both high and low elevation deserts of

southern California.

On hillsides in the coastal sage scrub community near Irvine, California,

T. sericotrachyla, T. geminata, and T. luteocincta can be found in close

proximity. Trirhabda luteocincta occurs on Haplopappus venetus ssp.

vernonioides (Nutt.) Hall and H. palmeri Gray (Asteraceae), which can be

intertwined with A. californica branches supporting T. sericotrachyla.

“ Aplopappus” was recorded as a host for T. luteocincta (Blake, 1931),

but Hogue (1970) did not locate it on any Haplopappus species and reported

its host plant as A. californica. During the course of this study, we occa¬

sionally observed adult T. luteocincta on A. californica foliage, but could

not find larvae or adults feeding on this plant. Attempts to switch first instar

larvae of T. luteocincta from H. palmeri to A. californica were unsuc¬

cessful.

Trirhabda sericotrachyla is locally common in coastal sage scrub com¬

munities where it feeds exclusively on A. californica as both larvae and

adults (Blake, 1931; Hogue, 1970) and is the plant’s principal insect defoli¬

ator. With a few exceptions, the life history of T. sericotrachyla corresponds

with that known for other species in the genus. The insect ranges along

much of the coast of California and parts of Oregon and Washington (Hogue,

1970) where its distribution roughly coincides with that of A. californica

(Munz and Keck, 1959). It is univoltine and can attain high densities which

may cause severe defoliation of its host, a characteristic shared by other

Trirhabda species (Hogue, 1970; F. Messina, pers. comm.).

Extensive stands of A. californica occur in undeveloped areas of cismon-

tane southern California. The plant is a primary indicator of the coastal sage

scrub community (Epling and Lewis, 1942; Kirkpatrick and Hutchinson,

1977) and is frequently the dominant shrub in this association (Mooney,

1975; Axelrod, 1978). It is a drought deciduous perennial that sheds its

foliage in summer or fall depending on environmental conditions and flushes

shortly after the first winter rains.

The Egg

During laboratory rearing studies oviposition behavior was observed on

many occasions. Attempts to monitor oviposition in the field, however, were

virtually unsuccessful because females deposit eggs only in stem crevices

at soil level. Detailed examination of foliage and branches and screening of

soil and duff near several colonized plants yielded no eggs. In the lab most

eggs were laid in narrow crevices and folds of the rearing container or were

hidden beneath pieces of toweling provided for cover.

The eggs are deposited in May or June and hatch after A. californica

VOLUME 58, NUMBER 2

141

Table 1. Proportions of T. sericotrachyla eggs hatching under different temperature and

humidity conditions.

Humidity treatment

Temperature treatment

Lab conditions

Field conditions

Pre-cooled

Control

0.02

0.0

Wet-dry

0.02

0.0

0.12

Saturated

0.32

0.38

0.30

plants initiate new growth during the winter rainy season. They are laid in

clusters of 15-50 eggs, several days apart. The individual eggs are 1-2 mm

long with a sculptured chorion. When laid they are golden tan and slowly

darken, and the clusters are enclosed in an adhesive matrix that quickly

hardens.

Egg hatching .—Several experimental treatments were utilized to evaluate

the effects of humidity and temperature on the timing of eclosion and hatch¬

ing success. The galerucines, Diabrotica virgifera virgifera LeConte and D.

longicornis barbari Smith and Lawrence, the western and northern corn

rootworms, lay eggs that undergo winter-diapause (Chiang, 1973; Krysan et

al., 1977). Their embryonic dormancy is influenced by humidity (Krysan et

al., 1977), and the timing of eclosion is affected by temperature (George and

Ortman, 1965). Both of these environmental factors are incorporated in

rearing programs for corn rootworms (Branson et al., 1975).

The hatching experiment employed two different methods of humidifying

the environment combined with three temperature treatments. The eggs

were allowed to remain in bare petri dishes at normal laboratory conditions

for about 150 days (approximately until the time of the first fall rains in the

field) before the experiment was started. Two replicates for each treatment

of approximately 60 eggs (laid over a three week span in May 1978) were

placed on sterilized U.C. Mix potting soil in petri dishes. One moisturizing

treatment consisted of maintaining the eggs in an environment of continu¬

ously high humidity by saturating the soil weekly and keeping the petri dish

covered. Eggs in the second humidity treatment were also moistened weekly

but were allowed to dry out between waterings. Temperature conditions

were: (1) normal laboratory regime near a south facing window with light

augmented by fluorescent room lights; (2) cooling in a refrigerator at 5°C

for one month prior to exposure to the laboratory conditions mentioned

above; and (3) maintaining the eggs outdoors under shelter where temper¬

atures ranged from 1-30°C. Control (unmoistened) eggs were used with each

temperature treatment. Emerging first instar larvae climbed to the petri dish

lid and were easily counted.

The hatchability of eggs was markedly determined by humidity (Table 1),

142

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Timing of eclosion of T. sericotrachyla eggs following initial exposure to saturated

atmospheres and a, preliminary cooling at 5°C for 30 days, ▲; b, laboratory temperatures, 21°

± 2°C, •; c, field conditions, 1°-30°C, O.

with pronounced success only in chambers having saturated atmospheres.

The timing of eclosion was largely influenced by temperature (Fig. 1). One

month of chilling at 5°C resulted in a compressed hatch duration, while eggs

maintained outdoors under a widely fluctuating temperature regime hatched

over a prolonged period. Eggs exposed to the higher laboratory tempera¬

tures began hatching after 55 days, whereas eggs exposed to fluctuating

outdoor temperatures did not achieve an equivalent cumulative percent hatch

until 30 days later.

An abbreviated hatch duration is also seen with prechilled eggs of D.

virgifera (George and Ortman, 1965). Krysan (1972) showed that the em¬

bryonic rudiment develops to the diapause stage either before or during

chilling, which suppresses further development. Release from chill allows

rapid development and relative synchrony. Trirhabda sericotrachyla may

undergo a similar diapause, but hatch duration under field conditions lasts

much longer than when eggs have been cooled in the lab. Prolonged winter

chills are very uncommon in coastal southern California, and any synchro¬

nizing effects of low temperatures are probably negated by the combined

influences of extended egg laying by adults in late spring, daily temperature

fluctuations, and the possible sensitivity of the eggs to some stimulus cor¬

related with the initiation of shoot elongation by the plant.

VOLUME 58, NUMBER 2

143

Hatching success under field conditions is unstudied. Corn rootworm

rearing programs employ humidified environments throughout embryologi-

cal development with hatching success approaching 100% (Branson et al.,

1975). Coastal southern California receives very little rainfall from April to

October, and despite the presence of interstitial water in the soil or con¬

densation of atmospheric moisture at night, humidity in the microenviron¬

ment of field eggs would not be expected to remain near 100%. However,

the lengthy and well protected egg stage effectively passes the dry summer

and fall after the host foliage is shed. Embryonic diapause in D. virgifera

is thought to have originated as an adaptation to similar environmental con¬

ditions in Mexico and secondarily functions to protect the eggs during harsh

winters in temperate latitudes (Krysan et al., 1977; Branson et al., 1978).

Whether eggs remaining unhatched the first year will eclose at a future

time remains untested. Hatchability of western corn rootworm eggs is re¬

duced by 80% after one year’s storage at 5°C (Branson et al., 1975). Several

clusters of T. sericotrachyla eggs which were maintained under normal

laboratory conditions for one year were desiccated and lacking protoplasm.

Fungal contamination is difficult to avoid after several months in humidified

chambers, and these organisms may also invade eggs under field conditions.

As with other stenophagic insects, T. sericotrachyla is closely adapted to

the annual cycle of its host. At a time of year when few herbivorous insects

are evident, and the physical environment can cause flooding or freeze dam¬

age to A. californica, new larvae emerge from the egg following early leaf

development. The synchrony of insect developmental stages with the avail¬

ability of the most suitable part of the plant is essential to successful ex¬

ploitation by herbivores (Breedlove and Ehrlich, 1972; Kogan, 1975; La-

beyrie, 1978). Artemisia californica plants are variable in their time of

flushing, and T. sericotrachyla eggs appear to be correspondingly variable

in their embryonic duration. The population must complete larval devel¬

opment, metamorphose, and attain reproductive maturity within a period of

time that may be compressed from either end. With mild winters in southern

California, T. sericotrachyla can afford to emerge early from the egg stage

in most years in order to exploit the youngest foliage available.

The Larva

Monitoring of the field sites began in mid-January to detect the presence

of first instar larvae on A. californica foliage. The sites were visited regularly

at several day intervals thereafter as long as larvae or adults were present,

and detailed observations of insect behavior were made frequently.

Newly eclosed first instar larvae collected from the field were fed in the

lab on fresh host branches maintained in small jars of distilled water. A filter

paper barrier was used to prevent the larvae from entering the water, and

144

PAN-PACIFIC ENTOMOLOGIST

Table 2. Mean duration ± SD, ranges of length and wet weight, and mean head capsule

width ± SD of T. sericotrachyla larvae reared under laboratory conditions.

Larval

instar

Length

(mm)

Head capsule

width (mm)

Wet weight

(mg)

Duration

(days)

1-3

0.5 ± 0.1, N = 12

0.8-3.0

6.5

± 1.5, N = 40

3-7

0.8 ± 0.2, N = 15

2-8

6.4

± 1.6, N = 38

7-13

1.2 ± 0.2, N = 15

7-74

6.5

± 1.8, N = 38

the vial was placed in a plastic cottage cheese container to catch any larvae

dropping from the filter paper. Larval food was changed, and each larva was

weighed approximately every two days. Larvae could also be reared in petri

dishes containing moistened filter paper. Laboratory conditions were a south

facing window augmented by normal overhead fluorescent lighting while

people were present, temperature 20-24°C, relative humidity 40-50%.

First instar larvae appeared in early February in the field with the time

of emergence varying between sites and between years. Emergence lasted

over a two month span and varied both between and within plants. Gen¬

erally most of the larvae on a given plant emerged within a two to three

week period, but a few first instar larvae could be found on a plant even

after many of their predecessors had migrated to the soil to pupate.

First instar larvae were concentrated near the growing tips with 84% (N =

245) on the distal % of new shoots. A substantial proportion (25%) was

found within the terminal sheath of leaves at the shoot apex. The vast

majority (91%) were on the upper surface of a leaf, but whether this is

advantageous for early feeding or adhering to the leaf surface is unknown.

Older larvae were more widely distributed over the foliage. They appeared

to be more tolerant of mature foliage and could consequently disperse away

from crowded shoot tips. New growth was sometimes destroyed during

intense herbivory by the beetles, forcing larvae to seek food elsewhere.

Size ranges and the duration of the three larval instars are indicated in

Table 2. Four larval instars were reported for Trirhabda by Hogue (1970),

an unaccountable observation considering Boving’s (1929) monograph on

galerucine larvae and considerable work done on other insects of economic

importance in the group. First instar larvae are mostly piceous in color with

occasional hints of metallic luster, moderately pubescent, and highly mobile.

The second instar larvae change to a metallic blue or green, typical of many

galerucine larvae (Boving, 1929).

Ecdysis requires several hours and is preceded by a nonfeeding period of

undetermined length. Cast larval head capsules and thoracic remnants were

not restricted to the inner portions of the plant as Hogue (1970) reported

VOLUME 58, NUMBER 2

145

for the genus, but were found generally distributed about the foliage. Larvae

in the third instar reared in the lab were quite sedentary until they concluded

feeding prior to descending to the soil for pupation.

Before pupation, larvae would either climb downward or drop to the soil

surface where they would soon begin burrowing to a depth of 1-2 cm. Under

laboratory conditions larval development prior to entering the soil averaged

19 days (SD = 3, N = 40). By mid-May the vast majority of larvae had

disappeared.

The quiescent pre-pupal period lasted several days during which up to

half of the maximum larval weight was lost. The metallic color was retained

until shortly before pupation when the insect darkened and the luster dis¬

appeared. Most, but not all, of the larvae created a spherical cell of soil

particles bound by anal secretions, approximately 1 cm in diameter, in which

pupation took place. The larva assumed a C-shaped form with its head up

within a cell, or it rested on its side if no cell was constructed. These

observations on pre-pupal larvae conform to those of Hogue (1970).

With respect to eight neighboring perennial shrubs (O’Brien, 1980), T.

sericotrachyla larvae fed exclusively on A. californica. Host specificity ex¬

periments did not preclude potential conditioning influences as newly eclosed

larvae obtained from the field already feeding on A. calif ornica were used

for the tests, but the results correlate with field observations and prior host

records. Factors regulating host discrimination are untested, but could in¬

volve several types of secondary chemicals known from Asteraceae in gen¬

eral and Artemisia in particular. These components include monoterpenes

(Halligan, 1975, 1976), sesquiterpene lactones (Mabry and Bohlman, 1977),

flavenoids (Rodriguez et al., 1972), and coumarins (Shafizadeh and Melni-

koff, 1970).

Larval migration .—The spring of 1978 was exceptionally wet in southern

California, and virtually all A. calif ornica plants at one field site were se¬

verely defoliated, presumably because of prolonged surface flooding and

saturation of the root zone. Second and third instar larvae had dropped from

their host plants and were actively crawling over the surface, in contrast

to pre-pupal larvae which immediately burrow into the soil. The vast

majority of these migrating larvae sustained mortality from desiccation and/

or starvation while stranded in surface depressions from which they were

unable to escape.

To determine if such disenfranchised larvae would seek new host plants,

100 vigorously mobile, third instar larvae were collected from the soil sur¬

face and marked with water soluble, non-toxic poster paint. They were

released shortly thereafter in a slightly sloping but relatively flat area where

a number of small (0.5 m tall) A. calif ornica plants had not suffered appre¬

ciable damage. Many larvae were dropping at the time experimental insects

were obtained, so most of the insects used had probably only recently de-

146

PAN-PACIFIC ENTOMOLOGIST

parted their host plants. The presence of marked larvae in surrounding plants

and their distance traveled were recorded on several occasions during the

following 24 hours.

This experiment established that the probability of successful colonization

of a new host plant was low (7.0%) even with plant densities of 1.03/m 2 .

Only 4.0% of the larvae were consistently able to travel further than 2.0 m

in 24 hours, although most of them made an initial attempt to move. Most

larvae traveled downslope, but many were unable to avoid or climb out of

surface irregularities. In the lab, a less severe habitat than the outdoor soil

surface, second instar larvae died within 72 hours without food or moisture,

which provides a crude estimate of the time available for migrating larvae

to locate a new host.

The possibility that heavy feeding by T. s eric otr achy la larvae aggravated

the stress imposed on the plants by saturated soil could not be explored as

all defoliated plants had been infested by insects. Of 36 defoliated plants

which were being used to sample densities of beetle larvae in the field, 17

resprouted about 2 months later. Foliage on the other plants did not regen¬

erate, and they were dead the following spring. Artemisia californica under¬

goes similar die-back and regeneration in response to prolonged exposure

to freezing temperatures (Mooney, 1977). Defoliation in response to envi¬

ronmental stress of plants with the capacity for resprouting could confer an

added benefit of reduced herbivory. We have observed heavy mortality in

T. geminata populations on Encelia farinosa near Riverside, California,

which suffered extensive freeze damage in January 1979, and resprouted

several weeks later. Breedlove and Ehrlich (1972) postulated a similar mech¬

anism for high altitude lupines to rid themselves of flower-feeding lycaenid

butterflies. In the present case, the density of first instar T. sericotrachyla

larvae the following year on plants that survived defoliation was about half

that preceding leaf-drop.

The Pupa

Larvae were allowed to pupate in the bottom of plastic cups which con¬

tained about 2 cm of U.C. Mix potting soil and were placed in an emergence

cage. Later experiments have shown that the addition of soil is unnecessary

for successful pupation; the insects will simply pupate in the bottom of a

bare petri dish to which a small piece of moistened filter paper is added.

At pupation the last larval cuticle splits longitudinally along the dorsum,

revealing a typical exarate pupa, yellowish in color that darkens somewhat

over time. Larvae not constructing a pupation cell of soil particles meta¬

morphosed normally. Pupae observed in the field were concentrated within

40 cm of the base of a plant. Some of them occurred on the surface, but

most were found approximately 1 cm deep. Several Trirhabda species pu¬

pate near a depth of 1 cm, but others either burrow deeper or metamorphose

VOLUME 58, NUMBER 2

147

in the duff on the soil surface (Hogue, 1970). The average time from descent

to the soil to adult emergence in the lab was 13 days (SD = 2, N = 40).

The Adult

Adults appeared in the field in late April or early May. They are rather

cryptically colored with dusty blue to green elytra marginally bordered with

yellow. Females are significantly larger than males, and the sexes can be

readily distinguished according to the shape of the posterior margin of the

terminal abdominal sternite, a characteristic of the tribe Galerucini (Wilcox,

1965). In males this edge is deeply invaginate centrally while in females it

is shallowly concave. The cumulative sex ratio in the field over a one month

period following the appearance of the first adults approximated 1:1 (N =

260). Females were markedly more abundant than males during the first

several days, and males gradually caught up over the remainder of the month.

In the lab, adults climbed up a nearby plant after emergence where they

generally remained inconspicuous near its interior for several days while

they fed prior to mating. Hogue (1970) reported a pre-feeding period of 1-

2 days following emergence. When females were ready to mate in the held,

they frequently assumed positions toward the distal ends of branches, and

males approached them from below. Plants could occasionally be found with

almost every terminal shoot supporting a female with very few males pres¬

ent. The female would frequently face downward, and as a male approached

she would either wait for him to contact her or she would move downward

to meet him before she turned around and allowed him to mount in a manner

typical of the genus (Hogue, 1970). In Diabrotica virgifera, mating behavior

is influenced by sex attractant pheromones (Ball and Chaudbury, 1973; Guss,

1976; Bartelt and Chiang, 1977; Lew and Ball, 1978). This possibility has

not yet been explored for Trirhabda spp.

Upon emergence, one male and one female adult were placed in 2-quart

ice cream cartons containing host branches maintained in water. Adult food

was changed every two days, and filter paper barriers were used to block

access to the water. Factors necessary for oviposition were unknown, so

folded paper toweling, strips of paper toweling, and petri dishes containing

U.C. Mix were placed in the bottom of the cartons to provide cover, tight

crevices, and soil, respectively. The eggs were removed and counted every

two days.

Copulation was observed to take much longer than the 1-3 minutes men¬

tioned by Hogue (1970), occasionally lasting up to 15-20 minutes. The fe¬

male caused the male to withdraw by starting to crawl and by wiggling her

abdomen laterally until the male’s position was no longer secure and he fell

off. In the lab, polygamous matings also took place, which were usually

separated by several days with a clutch of eggs deposited in the intervening

period. However, in the presence of several males, a single female was

148

PAN-PACIFIC ENTOMOLOGIST

observed to accept at least two males in a four day period between depos¬

iting successive egg clusters. In addition, several females marked for iden¬

tification mated with at least two different males in the field during obser¬

vation periods a few days apart. Single matings are apparently the prevailing

pattern for other Trirhabda spp. (Hogue, 1970). Although capable of mating

several times, a female removed from the presence of males after she once

mated continued to lay eggs throughout her adult fife. The influence of

multiple matings on fecundity is still unclear because of inadequate sample

sizes. No information is available on the relative contribution of early and

late sperm to the genetic constitution of these eggs.

Adult females lived up to 50 days in the lab during which time they de¬

posited up to ten clusters totaling a maximum of 240 eggs. In general, fe¬

males which lived longest and were largest at emergence laid the most eggs.

However, there were enough exceptions to render statistically insignificant

the slight positive correlations between fecundity and adult female size and

longevity.

Adult dispersal .—Investigation of adult movement between plants in the

field was accomplished by marking all the insects present on 8 plants at one

field site with water soluble, non-toxic poster paint and recording their dis¬

tribution every two or three days for the following month. Copulating pairs

and females heavily laden with eggs were specially identified.

Of 312 marked insects, 15% were found on new hosts during the following

30 days. Males and females both disperse, although females have a tendency

to remain longer with the same plant prior to mating than do males. The

bulk of flight activity occurs in the afternoon while temperatures are ele¬

vated and breezes active. Most migrating insects move into or across the

prevailing direction of the wind.

Over 80% of the insects observed to have migrated dispersed no further

than 5.0 m from the plant where they were marked, with many moving only

one or two plants away. The maximum distance covered by a marked insect

(male) was about 60 m. The fate of lost marked insects is obviously uncer¬

tain, but based on observations of flying beetles, dispersal beyond 60 m is

probable. However, the possibility of finding such insects was strongly re¬

duced because of search time limitations. Several insects remained on a new

plant for up to 2 weeks and two individuals, known to have migrated, re¬

turned to the plant where they were first identified.

Several gravid females were found on either the same (N = 3) or a dif¬

ferent plant (N = 2) following a reduction in abdomen size, indicating that

more than one clutch is possible under field conditions and that eggs are

not necessarily deposited at the base of the same plant. One specially marked

female was observed mating twice, and three males engaged in more than

one copulation with different females. One male changed host plants 3 times

and was observed mating twice on different hosts. Both males (N = 7) and

VOLUME 58, NUMBER 2

149

females (N = 6) which were not mating when marked did so later, and 2 of

each sex copulated while on new host plants.

Natural Enemies

Trirhabda sericotrachyla has a complement of arthropod predators and

parasitoids similar to that of other Trirhabda species (Hogue, 1970). Lebia

cyanipennis Dejean, a diurnal arboreal carabid that specializes on leaf bee¬

tles (Madge, 1967), was infrequently observed searching A. californica fo¬

liage for prey, and occasionally captured one. A predaceous thrips, affixed

to the dorsal surface just behind the head capsule of a third instar larva, fed

on exudations seeping through the integument. The pentatomid, Perillus

splendidus (Uhler), was more numerous than L. cyanipennis, and it fed on

both larvae and adults. Other arthropod predators on adult T. sericotrachyla

included orb-weaving and jumping spiders, and one unidentified assassin

bug (Reduviidae).

Of several hundred third instar T. sericotrachyla larvae brought to the

lab for experimental purposes, several that did not complete development

assumed a barrel shape and eventually died. One individual yielded a single,

unidentified tachinid fly. Nothing emerged from the other carcasses. Several

presumably parasitized larvae were dissected, and parasite larvae (probably

tachinids) were found in two hosts. The tachinid, Aplomiopsis xylota (Cur¬

ran), caused substantial mortality among several Trirhabda species exam¬

ined by Hogue (1970).

When disturbed, first and second instar larvae elevated and waved their

abdomen in an apparent defensive reaction. Several Trirhabda species dis¬

played this behavior in the presence of a tachinid parasitoid (Hogue, 1970).

Other chrysomelid larvae are known to possess defensive glands which emit

noxious substances (Blum et al., 1972; Blum et al., 1978), but these are

presumed absent in the Galerucinae (Boving, 1929).

In the field several T. sericotrachyla larvae had suffered wounds from

which extruded a hardened mass of fluid. Similar damage was caused when

high densities of active larvae were kept in small containers in the lab.

Whether the field condition resulted from intraspecific aggression or unsuc¬

cessful predator attacks is unknown.

Vertebrate predation was not observed except for a single black phoebe

that was hawking airborne adult beetles in 1978. Although flocks of foliage

gleaning birds (bushtits, warblers, gnatcatchers) are active in the coastal

sage scrub habitat, birds do not appear to concentrate on the larvae, even

though they occur in high densities and are quite visible. Predation by lizards

was not observed, however, they have taken adult Trirhabda during labo¬

ratory feeding studies (Hogue, 1970). The possibility that T. sericotrachyla

larvae sequester noxious components from their hosts or produce deterrent

substances themselves has not been investigated.

150

PAN-PACIFIC ENTOMOLOGIST

No parasitoids emerged from several egg masses or 25 pupae taken to the

lab for observation. None were previously reported by Hogue (1970). Shrews

and insect eating rodents should be considered potential predators of the

pupae, although no such activity was observed.

Acknowledgments

Appreciation is expressed to Ann McGee and Gordon Marsh for review¬

ing an original draft of the manuscript, to David Wood and George Hunt

for their criticism of the Ph.D. dissertation, and to an anonymous reviewer

for many incisive comments.

Literature Cited

Araott, D. A. 1957. Occurrence of Trirhabda pilosa Blake (Coleoptera: Chrysomelidae) on

sagebrush in British Columbia, with notes on life history. Proc. Entomol. Soc. B.C.,

53:14-15.

Axelrod, D. I. 1978. The origin of coastal sage vegetation, Alta and Baja California. Amer.

J. Bot., 65:1117-1131.

Balduf, W. V. 1929. The life history of the goldenrod beetle, Trirhabda canadensis Kirby

(Coleoptera: Chrysomelidae). Entomol. News, 40:35-39.

Ball, H. G., and M. F. B. Chaudbury. 1973. A sex attractant of the western corn rootworm.

J. Econ. Entomol., 66:1051-1053.

Banham, F. L. 1961. Distribution of Trirhabda pilosa Blake (Coleoptera: Chrysomelidae)

attacking big sagebrush in the interior of British Columbia. Proc. Entomol. Soc. B.C.,

58:38-40.

Bartelt, R. J., and H. C. Chiang. 1977. Field studies involving the sex attractant pheromones

of western and northern com rootworm beetles. Environ. Entomol., 6:853-861.

Blake, D. H. 1931. Revision of the species of beetles of the genus Trirhabda north of Mexico.

Proc. U.S. Nat. Mus., 79:1-36.

Blake, D. H. 1951. New species of chrysomelid beetles of the genus Trirhabda and Diso-

nycha. J. Wash. Acad. Sci., 41:324-328.

Blum, M. S., J. M. Brand, J. B. Wallace, and H. M. Fales. 1972. Chemical characterization

of the defensive secretion of a chrysomelid larva. Life Sci., 11:525-531.

Blum, M. S., J. B. Wallace, R. M. Duffield, J. M. Brand, H. M. Fales, and E. A. Sololoski.

1978. Chrysomelidial in the defensive secretion of the leaf beetle, Gastrophysa cyanea

Melsheimer. J. Chem. Ecol., 4:47-53.

Boving, A. G. 1929. Beetle larvae of the subfamily Galerucinae. Proc. U.S. Nat. Mus., 75:

1-48.

Branson, T. F., P. L. Guss, and J. L. Krysan. 1978. Winter populations of some Diabrotica

in central Mexico: Voltinism and pheromone response. Ann. Entomol. Soc. Amer., 71:

165-166.

Branson, T. F., P. L. Guss, J. L. Krysan, and G. R. Sutter. 1975. Com rootworms: Labo¬

ratory rearing and manipulation. U.S. Dep. Agr., ARS-NC-28, 18 pp.

Breedlove, D. E., and P. R. Ehrlich. 1972. Coevolution: Patterns of legume predation by a

lycaenid butterfly. Oecologia, 10:99-104.

Chiang, H. C. 1973. Bionomics of the northern and western com rootworms. Annu. Rev.

Entomol., 18:47-72.

Epling, C., and H. Lewis. 1942. The centers of distribution of the chaparral and coastal sage

associations. Amer. Midi. Natur., 27:445^162.

VOLUME 58, NUMBER 2

151

George, B. W., and E. E. Ortman. 1965. Rearing the western corn rootworm in the labo¬

ratory. J. Econ. Entomol., 58:375-377.

Guss, P. L. 1976. The sex pheromone of the western corn rootworm (Diabrotica virgifera).

Environ. Entomol., 5:219-223.

Halligan, J. P. 1975. Toxic terpenes from Artemisia californica. Ecology, 56:999-1003.

Halligan, J. P. 1976. Toxicity of Artemisia californica to four associated herb species. Amer.

Midi. Natur., 95:406-421.

Hogue, S. M. 1970. Biosystematics of the genus Trirhabda LeConte of America north of

Mexico. Ph.D. Dissertation, University of Idaho. 212 pp.

Kirkpatrick, J. G., and C. F. Hutchinson. 1977. The community composition of California

coastal sage scrub. Vegetatio, 35:21-33.

Kogan, M. 1975. Plant resistance in pest management. Pp. 103-146 in R. L. Metcalf and W.

H. Luckman (eds.), Introduction to Insect Pest Management. J. Wiley and Sons, New

York.

Krysan, J. L. 1972. Embryonic stage of Diabrotica virgifera (Coleoptera: Chrysomelidae) at

diapause. Ann. Entomol. Soc. Amer., 65:768-769.

Krysan, J. L., T. F. Branson, and G. DiazCastro. 1977. Diapause in Diabrotica virgifera

(Coleoptera: Chrysomelidae): A comparison of eggs from temperate and subtropical

climates. Entomol. Exp. Appl., 22:81-89.

Labeyrie, V. 1978. The significance of the environment in the control of insect fecundity.

Annu. Rev. Entomol., 23:69-89.

LeConte, J. L. 1865. On the species of Galeruca and allied genera inhabiting North America.

Proc. Acad. Nat. Sci. Phila., 17:204-222.

Lew, A. C., and H. J. Ball. 1978. The structure of apparent pheromone secreting-cells in

female Diabrotica virgifera. Ann. Entomol. Soc. Amer., 71:685-688.

Mabry, T. J., and F. Bohlman. 1977. Summary of the chemistry of the Compositae. Pp.

1094-1104 in V. Heywood, B. L. Turner, and J. B. Harborne (eds.), The Biology and

Chemistry of the Compositae. Academic Press, New York.

Madge, R. B. 1967. A revision of the genus Lebia Latreille in America north of Mexico

(Coleoptera, Carabidae). Quaest. Entomol., 3:139-242.

Massey, C. L., and D. A. Pierce. 1960. Trirhabda nitidicollis, a pest of rabbit brush in New

Mexico. J. Range Manage., 13:216-217.

Mooney, H. A. 1975. Southern coastal scrub. Pp. 471^190 in M. Barbour and J. Major (eds.),

Terrestrial Vegetation of California. Wiley Interscience, New York.

Mooney, H. A. 1977. Frost sensitivity and resprouting behavior of analogous shrubs of

California and Chile. Madrono, 24:74-78.

Munz, P. A., and D. D. Keck. 1959. A California Flora. University of California, Berkeley,

1681 pp.

O’Brien, P. Y. 1980. Adaptive relations between a stenophagic herbivore, Trirhabda seri-

cotrachyla Blake (Coleoptera: Chrysomelidae), and its host plant, Artemisia californica

(Compositae), in southern California. Ph.D. Dissertation, University of California, Ir¬

vine. 125 pp.

Pringle, W. L. 1960. The effect of a leaf feeding beetle on big sagebrush in British Columbia.

J. Range Manage., 13:139-142.

Rodriguez, E., N. J. Carman, G. Vander Velde, J. H. McReynolds, T. J. Mabry, M. A. Irwin,

and T. A. Geissman. 1972. Methoxylated flavenoids from seven Artemisia species.

Phytochemistry, 11:3509-3514.

Shafizadeh, F., and A. B. Melnikoff. 1970. Coumarins of Artemisia tridentata ssp. vaseyana.

Phytochemistry, 9:1311-1316.

Tilden, J. W. 1953. Biological notes on Trirhabda flavolimbata. Coleopt. Bull., 7:21-23.

152

PAN-PACIFIC ENTOMOLOGIST

Wilcox, J. A. 1965. A synopsis of the North American Galerucinae (Coleoptera: Chryso-

melidae). N.Y. State Mus. Sci. Serv. Bull., 400:1-226.

Wilcox, J. A. 1971. Chrysomelidae: Galerucinae. Oidini, Galerucini, Metacyclini, Sermylini.

Coleopterorum Catalogus Supplementa, 78(1): 1-220.

Footnote

1 Taxonomic determinations of Trirhabda spp. were made by Gordon Marsh, Museum of

Systematic Biology, U.C. Irvine. The work fulfilled partial requirements for the Ph.D. by the

senior author at the University of California, Irvine. These studies were supported in part by

a Slossen Fund Grant from the University of California.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 153-158

THE IMMATURE STAGES OF TIPULA SIMPLEX DOANE

AND T. ACUTA DOANE (DIPTERA: TIPULIDAE)

Margaret J. Hartman

Department of Biology, California State University,

Los Angeles 90032

AND

C. Dennis Hynes

Biological Sciences Department, California Polytechnic

State University, San Luis Obispo 93407

Records show that there are several species of crane flies capable of de¬

stroying valued rangeland in California. Of these, Tipula simplex Doane, the

range crane fly, is apparently the most devastating. Although outbreaks have

been reported in several counties, Tulare, Tehama and Marin counties have

sustained the heaviest infestations and damage. Marin County reported heavy

infestations in 1927, 1956, and 1961. In 1961, 12,000 hectares were involved

and at one location, records show infestations as great as 4300 larvae per

square meter (Marin County Ag. Comm., pers. comm.). Tulare County

reported outbreaks in 1961, 1967, and in the winter of 1972-1973 (Tulare

County Ag. Comm., pers. comm.). According to the latest survey, during

the winter of 1973-1974, Tipula simplex has, in Tulare County alone, af¬

fected in excess of 13,000 hectares of pasture with density in some samples

as great as 3000 larvae per square meter. In the more acute infestations the

hills are denuded of all grass and other important forage, which makes the

ranchers understandably concerned. The production of cattle is reduced

two-thirds or more during such infestations. The adverse effect on the wa¬

tershed must also be considered as there are no roots to hold the soil to¬

gether. The top layer is simply washed away, leaving a characteristic slick.

It takes some years for the grasses to re-establish themselves on such hill¬

sides and even longer to reach their normal cattle carrying capacity (Hart¬

man and Hynes, 1977). In the winter of 1978, Tehama County reported crane

fly damage for the first time.

Tipula simplex, however, is not the only crane fly living in grasslands.

Reports from the literature indicate Tipula quaylii Doane and T. gramini-

vora Alexander also are involved in range destruction (Alexander, 1921;

Packard and Thompson, 1921). Tipula simplex appears to live in sympatry

with crane fly species that do no detectable damage. We have found T.

silvestra Doane in Marin County and T. acuta Doane in Tulare County as-

154

PAN-PACIFIC ENTOMOLOGIST

sociated with T. simplex. Several other species belonging to the subgenus

Triplicitipula may be involved. The relative importance of species other than

T. simplex in rangeland destruction has not been ascertained but appears to be

minimal.

At present, there is no paper available which contains descriptions of the

immature stages or gives a key to their identification. Agricultural workers

have no way of recognizing the different immature stages of the species.

This paper is primarily aimed at giving such workers descriptions for field

identifications of the larvae and pupae of Tipula simplex and Tipula acuta.

Alexander (1920), Hennig (1948), and Chiswell (1956) have given general

accounts of the taxonomic characters which have been useful in describing

larvae of the Tipulini.

Measurements of most larval structures give some idea of size but are

not critical in differentiating between the larvae. The critical measurement

separating various instars is the dextro-sinistral width at the base of the

mandibles. The measurement given in the larval descriptions represents the

extremes of dozens of specimens measured over several years. The ranges

given hold true regardless of year or weather conditions. This information

is given for instars of T. simplex, but is presently unavailable for all instars

of T. acuta. By far the most useful characters in separating the species are

those of the spiracular disc, as described below.

Description of Immature Stages

Tipula simplex

First instar larva .—Length, 3.1-4.0 mm; dextro-sinistral and dorsal-ven¬

tral width, 0.4-0.6 mm. Head capsule width at mandibles, 0.2 mm. Egg

tooth on frons heavily sclerotized. Integument whitish; setal pencils black,

one pencil on lateral margins of terga and sterna of each segment from

prothorax through abdominal segment seven. Surface of larva covered

sparsely with very short brown setae giving larva a dirty white color. Spi¬

racular disc with four elongate setae coming off the lateral lobes. Markings

of spiracular disc as in Fig. 1. Anal lobes much darker brown than remainder

of body.

Last instar larva .—Length, 23.0-29.0 mm; dorso-ventral and dextro-si¬

nistral width similar, 3.71-4.71 mm.

Head capsule .—Length, 1.9-2.34 mm (4th instar); dorso-ventral width at

base of mandibles, 0.68-0.86 mm; dextro-sinistral width at base of mandi¬

bles, 0.860-1.189 mm (4th instar), 0.582-0.784 mm (3rd instar), not available

(2nd instar), 0.202-0.253 mm (1st instar).

Cuticle unpigmented; tergal and sternal microsetae subequal, plural mi-

crosetae much shorter. Microsetae of thoracic segments forming continuous

brown band around segment, anterior portion of band darker; pattern mot¬

tled, especially at junction of pleuron with tergum and sternum. Abdominal

VOLUME 58, NUMBER 2

155

Figs. 1-3. Figs. 1, 2. Caudal view of Tipula simplex. Fig. 1. First instar larva. Fig. 2.

Fourth instar larva. Fig. 3. Caudal view of Tipula acuta, fourth instar larva.

156

PAN-PACIFIC ENTOMOLOGIST

segments 1-7 with tergum light brown, dark tufts of micro setae at bases of

macrosetae; sternum with anterior dark brown bands of microsetae, pattern

mottled.

Spiracular disc with fleshy, conical, subequal dorsal and lateral lobes.

Ventral lobes short, heavily sclerotized at blunt tip, directed dorsad. Spira¬

cles dark brown, face of disc patterned with brown sclerotized areas as in

Fig. 2. Anal lobes separated from remainder of abdomen by dark brown

band of cuticle; lateral pair conical, directed laterad; remainder of lobes

bulbous.

Pupa. —Length, 12.1 mm; dorso-ventral and dextro-sinistral width at base

of wing pad, 2.5 mm. Body reddish brown; base of antennal sheaths armed

with small spinous tubercle directed basad; breathing horns annulated, tip

spatulate, light yellow, becoming abruptly dark brown at base of horn. Pro-

notal median carina dark brown; mesonotum with midline dark brown, face

of dorsal crest rugose, one lateral and one medial tubercle on each side of

midline, heavily patterned with dark brown at lateral edges. A tubercle also

located on either side of midline at one-quarter distance between dorsal

crest and posterior margin. Wing pads light brown with wing veins lighter,

ending at anterior margin of second abdominal segment. Leg sheaths of male

ending midlength of third abdominal segment; outer sheaths longest, medial

sheaths and inner sheaths progressively shorter. Leg sheaths of female end¬

ing midlength of second abdominal segment; outer and medial sheaths sub¬

equal, inner sheaths shorter. Abdominal segments 2-7 reddish brown, each

segment divided into 2 rings; pleural region carinate, yellow, speckled with

dark brown spots. Anterior ring with forward area patterned with dark brown;

posterior ring armed with transverse row of spines. Posterior ring patterned

medially and laterally. Male and female cauda as shown in Figs. 4 and 5

respectively.

Tipula acuta

Last instar larva. —Length, 29.5-37.0 mm; dorso-ventral and dextro-si¬

nistral width, 4.0-5.5 mm. Cuticle unpigmented, covered with dense, very

short microsetae; dorsal and ventral microsetae subequal in length, pleural

shorter. Microsetae of thoracic segments and first abdominal segment form¬

ing continuous bands around segment, with slightly mottled pattern at junc¬

tions of pleural area with tergum and sternum. Abdominal segments 3-7

with setae of tergum forming 4 dark brown stripes, posterior darker, wider,

and mottled; setae of sternum patterned in 3 dark brown stripes, posterior

wider. Pleural junctions with sternum and tergum mottled. Spiracular disc

with six fleshy lobes, dorsal pair separate, elongate, sclerotized from base

to hardened sharp point, ventral pair short; entire face of disc patterned as

shown in Fig. 3. Anal field separated from remainder of abdomen by a dark

brown band of cuticle; lateral lobes conical, remaining lobes bulbous.

VOLUME 58, NUMBER 2

157

Figs. 4-7. Figs. 4, 5. Lateral view of cauda of pupa of Tipula simplex. Fig. 4. Male. Fig

5. Female. Figs. 6, 7. Lateral view of cauda of pupa of Tipula acuta. Fig. 6. Male. Fig. 7

Female.

158

PAN-PACIFIC ENTOMOLOGIST

Pupa. —Length, 19.7-24.0 mm; dorso-ventral and dextro-sinistral width,

3.5-3.9 mm. Body brown. Pronotal breathing horns brown, darker distally,

annulated, spatulate at tips, directed laterad. Mesonotum with anterior sur¬

face rugose, dorsal crest with pair of spinous tubercles dorsally and a pair

laterally. Wing pads dark brown, ending at posterior margin of second ab¬

dominal segment; veins not lighter. Leg sheaths dark brown, those of male

ending midlength of third abdominal segment, those of female ending at

posterior margin of second abdominal segment. Both sexes with outer and

medial leg sheaths subequal; inner sheath slightly shorter. Abdominal seg¬

ments 2-7 divided into 2 rings, with dorsal surface brown, darker ventrally;

each ring abruptly lighter along posterior margins. Posterior ring armed with

transverse row of spines. Lateral edge of each segment carinate, light in

color, speckled with dark brown spots. Male and female cauda as shown in

Figs. 6 and 7.

Key to Larvae of Crane Flies Living in Grasslands in Tulare County

1. Dorsal and lateral lobes of spiracular disc fleshy, conical, and sub¬

equal . T. simplex

Dorsal lobes of spiracular disc sclerotized from base to hardened

sharp point, longer than lateral lobes . T. acuta

Key to Pupae of Crane Flies Living in Grasslands in Tulare County

1. Wing pads ending at anterior margin of second abdominal segment;

outer leg sheaths longest, medial and inner sheaths progressively

shorter . T. simplex

Wing pads ending at posterior margin of second abdominal segment;

outer and medial leg sheaths subequal . T. acuta

Acknowledgments

We would like to acknowledge the cooperation of the Agricultural Com¬

mission of Tulare County, and the financial assistance of the J. G. Boswell

Company and the Gill Cattle Company.

Literature Cited

Alexander, C. P. 1920. The crane-flies of New York. Part II. Biology and phylogeny. Cornell

Univ. Agric. Exp. Stn. Mem., 38:695-1133.

Alexander, C. P. 1921. A new species of Tipula injurious to pasture lands (Tipulidae, Diptera).

Insecutor Inscitiae Menstruus, 9:135-137.

Chiswell, J. R. 1956. A taxonomic account of the last instar larvae of some British Tipulinae

(Diptera: Tipulidae). Trans. R. Entomol. Soc. Lond., 108:409^484.

Hartman, M. J., and C. D. Hynes. 1977. Biology of the range crane fly, Tipula simplex Doane

(Diptera: Tipulidae). Pan-Pac. Entomol., 53:118-123.

Hennig, W. 1948-1952. Die Larvenformen der Dipteren. Akademie-Verlag, Berlin, 1:1-185; 2:

i-vii, 1—458; 3:i-vii, 1-628.

Packard, C. M., and B. G. Thompson. 1921. The range crane-flies in California. U.S. Dep.

Agric. Dep. Circ., 172:1-8.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, pp. 159-161

THE PEACH BEETLE, COT1NIS MUTABILIS (GORY AND PERCHERON),

IN CALIFORNIA (COLEOPTERA: SCARABAEIDAE)

M. W. Stone 1

131 Sir Damas Dr., Riverside, California 92507

A scarabaeid, Cotinis mutabilis Gory and Percheron, also known as C.

texana Casey, or the peach or fig beetle, belonging to the subfamily

Cetoniinae (Ritcher, 1945), has for years been a destructive pest of peaches,

figs and grapes in southern California. 2 The larvae of Cotinis nitida (Lin¬

naeus), the green June beetle, severely damages young plants in tobacco

beds, lawns, and golf fairways by burrowing and forming mounds of dirt

on the surface (Allen and Creighton, 1962). Larvae of the genus Euphoria

have similar habits and the adults also damage fruit and corn in the eastern

states (Ting, 1934). The writer became interested in these insects because of

the habit of the larvae, when tunneling or on the soil, of crawling on their

backs, despite the presence of 3 pairs of thoracic legs. A brief description

of the various stages of C. mutabilis and notes on their periods of activity

in the Riverside area follows.

Egg (Fig. 1A).—Sixty-one C. mutabilis eggs deposited during the period

August 6 to 13, 1979 hatched between August 15 and 26, for an average

duration of the egg stage of 12.4 days. The time required for hatching out¬

doors varied from 9 to 17 days. The eggs are whitish and large (2.1 by 2.6

mm) and easily detectable in the soil.

Larva (Fig. IB, C).—Newly hatched larvae are whitish with brownish

head and legs. With plentiful food they develop rapidly and reach a size of

12 to 50 mm prior to pupation. The first two instars are usually completed

by fall and completion of the third stage occurs in the spring of the second

year. These larvae are known as “back crawlers.” Nichol (1935) describes the

movement of Cotinis larvae as an undulating motion of the entire body, the

propellent being the motion of transverse rows of stiff, short, stout bristles

on the dorsum of the thoracic region.

Pupa (Fig. ID, E).—The pupa may be found in an earthen case con¬

structed by the mature larva. The case may vary in size depending upon

the size of the pupa and whether male or female. Measurements of 7 pupae

showed an average size of 15 by 25 mm. In a group of reared larvae, nine

pupated in the period June 12 to July 4. The duration of the pupal period

ranged from 25 to 27 days. Pupation outdoors may occur from early May

up until August. Newly formed pupae are whitish, becoming cream colored

160

PAN-PACIFIC ENTOMOLOGIST

Fig. 1. Cotinis mutabilis. A, eggs. B, third stage larva. C, thoracic legs on larva, never

used for locomotion. D, pupa in earthen case. E, pupa. F, adult feeding on fig. G, clypeal

horn used to penetrate hard skinned fruits.

as they mature. Traces of green later appear on the elytra, head and other

exterior parts.

Adult (Fig. IF, G).—Except for an outer brownish edge the elytra of the

newly emerged beetle are greenish in color. The head is a more reddish green

and the legs a bright green. Female adults are larger, averaging 17 by 25

mm, as compared with 13 by 22 mm for males. The beetles are equipped

with a clypeal horn which is used for puncturing the skin of hard-skinned

fruits. Adults collected by the writer and in the U.C. Riverside collection

VOLUME 58, NUMBER 2

161

showed beetles present in the field from the latter part of June and until the

middle of November. The peak of emergence occurred in July-August.

The life of adults will vary depending upon the type of cages used and

food furnished. Those reared on figs in large glass jars remained alive

from 30 to 55 days, whereas when confined in 4 oz. salve tins in soil with

grapes some mortality occurred after 11 days. Egg-laying females are es¬

pecially attracted to compost and manure piles.

Literature Cited

Allen, N., and C. S. Creighton. 1962. Controlling green June beetle larvae in tobacco beds.

U.S. Dep. Agric. Leafl. 504:1-4.

Nichol, A. A. 1935. A study of the fig beetle, Cotinis texana Casey. Univ. Ariz. Agric. Exp.

Stn. Tech. Bull., 55:157-198.

Ritcher, P. O. 1945. North American Cetoniinae with descriptions of larvae and keys to genera

and species (Coleoptera: Scarabaeidae). Ky. Agric. Exp. Stn. Bull., 476:1-39.

Ting, P. C. 1934. Back-crawling scarabaeid grubs (Potosia affinis Anderesch) intercepted in

quarantine at San Francisco. Mon. Bull. Calif. Dep. Agric., 13:185-191.

Footnotes

1 Collaborator—USDA-SEA AR. Boyden Entomological Laboratory. Deceased March 28,

1982.

2 L. D. Anderson, U.C. Riverside, collected larvae in compost in April 1949. Adults in

U.C.R. Museum collection collected in August 1955 by A. L. Melander. Max Peterson and F.

E. Jorgenson, Riverside Lawn Bowling Club, collected adults in August-September 1977-1980.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, p. 162

SCIENTIFIC NOTE

A NEW ELEVATIONAL RECORD FOR PIERIS PROTODICE

IN CALIFORNIA (LEPIDOPTERA: PIERIDAE)

The checkered white, Pieris protodice Bdv. and LeC., is an extremely

vagile insect which undergoes regular altitudinal displacement throughout

western North America (Shapiro, 1979, J. Res. Lepid., 17:1-23). In New

Mexico and Arizona summer records above 3000 m are not uncommon. In

California it is replaced at high elevations by Pieris occidentalis Edw., with

which it is often confused (Shapiro, 1976, J. Lepid. Soc., 30:289-300). Most

California records above 2000 m are untrustworthy due to this confusion;

however Tilden (1959, Wasmann J. Biol., 17:249-271) seems to have kept

them straight. At Donner Pass (2100 m), where I have monitored the fauna

since 1973, P. protodice is a frequent visitor and occasional breeder, but

does not overwinter; it is absent on nearby Castle Peak (2700+ m) (Shapiro,

1978, Great Basin Nat., 73:443-452). The highest Sierran record I have

confirmed is a female taken by S. R. Sims near Sonora Pass, 11 August

1977, “above 3000 m.”

On 18 June 1980, H. V. Carey found a female P. protodice dead on a

snowbank near Barcroft Laboratory at about 3630 m in the White Moun¬

tains, Mono County. At this time the resident pierids, including P. occi¬

dentalis, were not yet flying. The specimen is in excellent condition; it is of

summer phenotype, shows no significant wear, and contained one fresh

spermatophore and many mature ova. Pierids have been reported before on

snowbanks: Erhard (1929, Proc. X Int. Cong. Zook, Budapest, 10:1356-

1371) and Stauder (1920, Z. Wiss. Insektenbiol., 95:263-264) found mass-

migratory species dead in large numbers high in the Alps. Mani (1962, In¬

troduction to High-Altitude Entomology) discusses the general phenomenon

of transport and migration of lowland insects to the nival zone of the Him¬

alaya. Pieris protodice is not a mass migrant, and the Barcroft specimen

must represent an individual displacement of at least 10 km and through

some 2000 m of elevation. This is, however, a much shorter minimum dis¬

tance than the 80 km traversed by a female Pieris occidentalis taken near

sea level in suburban Sacramento (Shapiro, 1977, J. Lepid. Soc., 31:202-

203). None of these feats by gravid females has much chance of establishing

a permanent population, but all can potentially contribute to introgressive

hybridization or the buildup of isolating mechanisms between these two

sibling species.

Arthur M. Shapiro, Department of Zoology, University of California,

Davis 95616.

Note added in proof. Ms. Carey found a second female under identical circumstances on 12

June 1982, suggesting that altitudinal dispersal is a fairly regular seasonal phenomenon in this

species and region.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, p. 163

SCIENTIFIC NOTE

NOTES ON A COLLECTION OF INTERTIDAL BEETLES FROM

THE FARALLON ISLANDS, CALIFORNIA

Observations and collections of Coleoptera were made in mid-October

1978 on Southeast Farallon Island, San Francisco County, California.

This small granitic island lies 32 km south of Point Reyes, Marin County.

Among the twenty beetle species collected were the following intertidal

rock forms: Staphylinidae— Liparocephalus cordicollis LeConte, Diaulota

densissima Casey, and D. vandykei Moore; Hydraenidae— Ochthebius

vandykei Knisch; Melyridae— Endeodes collaris (LeConte); and Salpingi-

dae— Aegialites subopacus (Van Dyke), A. fuchsi Horn. All seven are

flightless and inhabit similar habitats on the coast of California. Staphylini¬

dae were identified by Ian Moore, of Riverside, California.

Over half of the specimens of E. collaris lack the red coloration

possessed by mainland individuals, indicating long enough isolation to have

produced a dark form.

The species of Aegialites have been treated as strictly intertidal (Doyen,

J. T., 1976, Marine beetles (Coleoptera excluding Staphylinidae). Pp. 497-

519 in L. Cheng, Marine insects. North Holland Publishing Co.). While

specimens of A. subopacus seemed to be as restricted to the vicinity of

the high tide line as those that occur on the mainland, both adults and

larvae of A. fuchsi were found far above the influence of all wave action.

They occurred on the steep north slopes in a continuous distribution from

the splash zone to the roof of the lighthouse atop the island’s 109 m

summit. Greatest densities were found in the five to 20 m elevation range

in association with a mossy growth on north-facing rock surfaces. At

times, over 100 were seen within a 0.5 m 2 area. Mounted pairs were

numerous, though no actual copulations were seen.

The cement lighthouse is cylindrical in shape, about 5 m in both height

and diameter, with direction indicators dividing its top into eight equal

sections. The number present in each section during a morning of heavy

fog was as follows:

S SW W NW N NE E SE S

8 12 74 69 52 41 3 3

The distribution of A. fuchsi atop the lighthouse was representative of

that found on large boulders, rock outcrops, and the island as a whole.

The amount of moisture deposited by wind-driven fog appeared to be the

major factor in determining this distribution.—Derham Giuliani, 170 Flower

Alley, Big Pine, California 93513.

PAN-PACIFIC ENTOMOLOGIST

April 1982, Vol. 58, No. 2, p. 164

SCIENTIFIC NOTE

SHRIKE PREDATION ON THE SCORPION

ANUROCTONUS PHAIODACTYLUS (WOOD)

AND ON A SOLPUGID

(SCORPIONIDA: VAEJOVIDAE; SOLPUGIDA)

On 22 January 1976, WHC and DRF found an unidentified solpugid impaled

on a leaf tip of the Mohave yucca, Yucca schidigera Roezl ex Ortgies, 6 km

northwest of San Agustin, Baja California, Mexico (29°59'N, 114°56'W),

elevation 580 m. This area is dominated by Larrea and Ambrosia and is

considered to be part of the Sonoran Desert.

On 21 February 1978, an adult Anuroctonus phaiodactylus (Wood) was

found by WHC and PLC impaled on the leaf tip of a Joshua tree, Yucca

brevifolia Herbertii (J. M. Webber) Munz, 6.5 km south of Olancha, Inyo

County, California, at an elevation of 1164 m. The site is a transitional

community and is Upper Mojavean in nature, with Great Basin elements

present.

The shrike or butcherbird is known to impale its prey on sharp objects such

as shrub thorns and barbed wire (Craig, 1978, Auk, 95:221-234). The

loggerhead shrike, Lanius ludovicianus, occurs at both sites (Small, 1974,

The birds of California, xxiv + 310 pp., Winchester Press, New York; Grin-

nell, 1931, Univ. Calif. Publ. Zool., 32:1-300). We could find no literature

references concerning shrike predation on these arthropods. Muma (pers.

comm., 1980) reported that similar observations have been made concerning

solpugids by several workers but knew of no published records. Williams

(1966, Proc. Calif. Acad. Sci., (4)34:419-428) reported the following predators

of A. phaiodactylus : Notiosorex crawfordii (desert shrew); Stenopelmatus

sp. (Jerusalem cricket); Eleodes sp. (darkling beetle); and several species of

owls. We add the shrike to this list of predators. Because both the scorpion

and solpugid are normally nocturnal, they may be an uncommon prey item of

the shrike.

The voucher specimen of A. phaiodactylus and photographs of it as well as

the solpugid have been deposited in the Museum of Natural History, College

of Idaho, Caldwell.—William H. Clark and Donald R. Frohlich, Museum of

Natural History, College of Idaho, Caldwell, Idaho 83605 ; and Peter L.

Comanor, U.S. Department of Interior, Bureau of Land Management, 202-B,

Washington, D.C. 20240.

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Vol. 58

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No. 3

THE

Pan-Pacific Entomologist

HEPPNER—Revision of American Thaumatographa, with a New Species from Cuba

(Lepidoptera: Tortricidae: Chlidanotinae). 165

SOKOLOFF, WILSON, MULDER, and FAUSTINI—Observations on Populations of

Tribolium brevicornis LeConte (Coleoptera: Tenebrionidae). II. The Habitat Niche

of a Local Population in Southern California. 177

PINTO—A New Eupompha from Baja California with Additional Information on E.

decolorata (Horn) (Coleoptera: Meloidae). 184

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PARKER and TEPEDINO—Behavior of Osmia ( Nothosmia ) marginata Michener in

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PFAFFENBERGER—Description of and Phylogenetic Comments on the Final Larval

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SCIENTIFIC NOTES

PAGE—Polyandry in Brachymyrmex depilis Emery (Hymenoptera: Formicidae). 258

COVILLE—A Note on the Biology of Ancistrocerus waldenii flavidulus Bequaert (Hy¬

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PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 165-176

Published 16 December 1983

REVISION OF AMERICAN THAUMATOGRAPHA, WITH A

NEW SPECIES FROM CUBA

(LEPIDOPTERA: TORTRICIBAE: CHLIOANOTINAE)

John B. Heppner

Department of Entomology, Smithsonian Institution,

Washington, D.C. 20560

Hilarographa and several related genera, primarily of pantropical distri¬

bution, until recently were included in the Glyphipterigidae as formed by

Meyrick (1914), an assemblage now comprising several families. During

revisionary studies on North American Glyphipterigidae and Choreutidae,

it was determined that the three North American species of Hilarographa

belong in the family Tortricidae. Since these species will be excluded from

forthcoming revisions of the glyphipterigids and choreutids (Heppner, in

prep, a, in prep, b), they are treated here. Hilarographa, Thaumatographa,

and related genera were in part transferred to the Tortricidae by Diakonoff

(1977a) and by Heppner (1978). The tribe Hilarographini was described for

Hilarographa and Mictopsichia (Diakonolf, 1977a). It is one of four tribes

in the Chlidanotinae, along with Polyorthini, Chlidanotini, and Schoeno-

tenini.

The Hilarographini superficially appear like some Choreutidae, primarily

because of bright coloration often enhanced by metallic-iridescent markings.

However, they do not possess characters amenable to the family Choreutidae.

The lack of a basally scaled haustellum is the most obvious character ex¬

cluding them from Choreutidae. The note by Diakonoff (1977a), and again

in Diakonoff (1977b), that Hilarographa and relatives possess a scaled haus¬

tellum, is erroneous: the “scales” referred to are actually elongated sensory

setae, not true scales. Relationships to other Chlidanotinae among genera

in Hilarographini are remarkable, in particular the slit valvae of some species

in which valval pouches serve as repositories for corematal hairs. This apo-

morphic character is otherwise only recorded for the Polyorthini, in fact,

this was originally a main character for the proposal of this tribe. Additional

apomorphic characters among Hilarographini are an accessory bursa and

asteroid-like signum in most females, which are characteristic of typical

Chlidanotini. Recently evidence from larval characters has demonstrated

that the Chlidanotinae may again warrant elevation to family status in the

Tortricoidea (Diakonoff and Arita, 1981). The larva and pupa of Thau¬

matographa eremnotorna Diakonoff and Arita, a Japanese species, have

provided the first immature stages for study for all the Chlidanotinae. The

166

PAN-PACIFIC ENTOMOLOGIST

larva of this Thaumatographa shows a bisetose pre-spiracular setal group

on the pro thorax, an arrangement thus far not known among Tortricidae

where all larvae known are trisetose for this setal group. Larvae of other

Chlidanotinae are unknown.

Presently the tribe Hilarographini contains 9 genera worldwide: Chari-

tographa, Embolostoma, Hilarographa, Idiothauma, Irianassa, Mictocom-

mosis, Mictopsichia, Nexosa, and Thaumatographa. The tribe will be re¬

viewed on a world basis together with a species checklist in a forthcoming

paper (Heppner, 1982). Little is known of the biologies of these moths.

Tropical Hilarographa are day-fliers and most other Hilarographini that are

as colorful as these species probably are also diumally active. North Amer¬

ican Thaumatographa species may only be nocturnally active, however, since

they have only been collected at lights. The larvae of only three species have

been reared: Thaumatographa eremnotorna, Thaumatographa leucopyrga

Meyrick from Java, and Thaumatographa regalis (Walsingham) from Cal¬

ifornia. The larvae of Thaumatographa eremnotorna and T. regalis feed on

the cambium layer in the trunks of pines, while the larva of T. leucopyrga

has been reared from the shoots of Ardisia sieboldi Miquel (Myrsinaceae).

Thaumatographa Walsingham, 1897:52

Type-species.—Hilarographa zapyra Meyrick, 1886 (Type locality: New

Guinea), by original designation.

This genus comprises 32 species, including the new species described here.

Most are known from the Indo-Australian region, with a few occurring in

Japan. Only three species are known from North America, all closely related

to Japanese species, and only the new Cuban species penetrates the Neo¬

tropical region. Hilarographa, contrarily, is exclusively Neotropical as far

as is known.

Thaumatographa regalis (Walsingham), New Combination

Glyphipteryx regalis Walsingham, 1881:320.

This western North American species is very similar to the Japanese

species Thaumatographa decoris (Diakonoff and Arita), differing in having

larger silver spots between the black tornal spots of the forewings and having

more orange bordering this row of spots.

Forewing length 7.8-10.0 mm. Head: fuscous mixed with tan on frons;

labial palpus rather smooth scaled with white and brown scales on exterior

side of basal segment; antenna short with long ventral setae in male. Thorax:

fuscous with orange spots on posterior comers of mesothorax; patagia lus¬

trous gray-brown; legs whitish with fuscous basally and orange on tarsal

segments. Forewing: as in Figure 1, with ground color orange below fuscous

costal margin; silver striae at base to fascia continuing to dorsal margin;

VOLUME 58, NUMBER 3

167

1 > 2

Figs. 1-4. Fig. 1. Thaumatographa regalis (Walsingham), <3, Kemville, California (UCB).

Fig. 2. T. cubensis Heppner, <3 holotype, La Casimba, Cuba (BMNH). Fig. 3. T. youngiella

(Busck), 3, Victoria, Vancouver Island, British Columbia, Canada (USNM). Fig. 4. T. jonesi

(Brower), 2 allotype, Martha’s Vineyard, Massachusetts (USNM).

silver fascia at x h from base and oblique fascia from costal white mark near

apex; three white costal marks following silver fascia at Vi, each with silver

mesad; large buff-white field on distal half of wing, with numerous black

striae and dots; tornal margin with 4 black spots bordered mesally by orange

line and interrupted by silver spots; fringe fuscous; ventral side dull fuscous

with white apical marks repeated. Hindwing: fuscous with central lighter

area; fringe white and fuscous; ventral side mostly white. Abdomen: fuscous

with pale scales on posterior margin of each segment. Male genitalia: as in

Figure 6, with oblong setaceous valva having apex blunt and rounded along

ventral margin; without dorsal corematal slit; tegumen and vinculum fused

as sclerotized ring; saccus a short point; uncus elongate with slightly bulbous

and setaceous distal end; hamus long and pointed; socius small setaceous

stub; gnathos a small round band; transtilla a broad, relatively unsclerotized

flap; anellus horseshoe-shaped with tapered distal ends; aedeagus (Fig. 7)

168

PAN-PACIFIC ENTOMOLOGIST

Fig. 5. Distribution map of Thaumatographa species in North America: T. cubensis Heppner

(open circle), T. jonesi (Brower) (closed squares), T. regalis (Walsingham) (closed circles), and

T. youngiella (Busck) (triangles).

elongate with cornutus having a small spine; vesica with elongated hood.

Female genitalia: as in Figure 15, with ovipositor flattened; apophyses stout;

ostium a large funnel; ductus bursae long and straight to curved area near

bursa, little sclerotized; corpus bursae ovate with small accessory bursa on

long duct; signum (Fig. 16) a small asteroid-like mass of spines.

Types. — Holotype 2: Mt. Shasta [Siskiyou Co.], CALIFORNIA, 2 Aug-1

Sep 1871, Walsingham (BMNH, Walsingham Coll. 92037).

Specimens studied. —19 <3, 7 2. CALIFORNIA: Calaveras Co.: 4 mi. E

Murphy’s, 3000 ft, 17 Jul 1963 (1 <3), 2 Jul 1963 (1 <3), P. Quyle (UCB). El

Dorado Co.: 2 mi. W Lk. Edson, 27 May 1972 (1 2), J. Doyen (UCB). Kern

Co.: Kernville, 28 Apr 1964 (6 <3), J. A. Powell (UCB); Mt. Pinos, 6800 ft,

2 Jul 1965 (1 <3), D. J. Calvert (UCB); Wofford Hts., Lk. Isabella, 2700 ft,

14 May 1976 (2 <3), R. J. Ford (LACM). Los Angeles Co.: Los Angeles, 13

May 1931 (1 <3), Wolff (LACM). Placer Co.: Carnelian Bay, Lk. Tahoe, 24

VOLUME 58, NUMBER 3

169

Figs. 6-14. Male genitalia of North American Thaumatographa species. Fig. 6. T. regalis

(Walsingham), Kernville, California (JBF1 504) (UCB). Fig. 7. Same, detail of aedeagus. Fig. 8.

T. cubensis Heppner [damaged], La Casimba, Cuba (BMNH 20267, holotype). Fig. 9. Same,

detached vinculum. Fig. 10. Same, aedeagus detail. Fig. 11. T. jonesi (Brower), Open Pond

Cpgd., Alabama (JBH 624) (JBH). Fig. 12. Same, aedeagus detail. Fig. 13. T. youngiella (Busck),

Departure Bay, Vancouver Island, British Columbia, Canada (USNM 77128, paralectotype).

Fig. 14. Same, aedeagus detail.

Jun 1964 (1 6), D. F. Veirs (UCB); “Placer Co.,” 1 May 1919(12), H. Vachell

(USNM); Michigan Bluff, 21 Jun 1967 (1 2), R. P. Allen (CSDA). Riverside

Co.: Idyllwild, 13 Jun 1940 (1 <3), F. H. Rindge (AMNH); Santa Rosa Mtn.,

7100 ft., 6-7 Jul 1974 (1 2), D. C. Frack (DCF). Sacramento Co.: Folsom,

170

PAN-PACIFIC ENTOMOLOGIST

Figs. 15, 16. Fig. 15. Thaumatographa regalis (Walsingham), 9 genitalia, Placer Co., Cali¬

fornia (USNM 77127). Fig. 16. Same, signum detail.

20 Jul 1885 (1 <$), ex Pinus sabiniana, Koebele (BMNH); Folsom, 23 Jul

1885 (1 6) [Koebele] (USNM). , San Bernardino Co.: Barton Flats [San Ber¬

nardino Mts.], 31 Jul 1971 (1 9), D. C. Frack (DCF); Big Bear Lk., 6800 ft.

[San Bernardino Mts.], 7 Aug 1978 (1 6), R. H. Leuschner (RHL). San Diego

Co.: Laguna, 18 Jun 1965 (1 9), ex Pinus ponderosa, K. Baker (CSDA).

Shasta Co.: Hat Cr., 11 Jun 1960 (1 <$), F. Stehr (CSDA). Siskiyou Co.:

Shasta City, 22 Aug 1958 (1 6), J. A. Powell (UCB). Tuolumne Co.: Twain

Harte, 4000 ft., 17 Jul 1964 (1 9), M. R. & S. H. Lundgren (CAS).

Hosts.—Pinus ponderosa Douglas and Pinus sabiniana Douglas (Pinace-

ae).

Distribution. — Known only from California.

Remarks. — This species is remarkably similar superficially to the Japanese

species, Thaumatographa decoris. The genitalia, however, will distinguish

the two species. The Japanese species T. eremnotorna, a related species, has

been reared from the cambium of Pinus densiflora Siebold & Zuccerini.

Larvae of T. regalis also feed on the cambium layer of pines.

Thaumatographa cubensis, Heppner, new species

This species is superfically similar to T. regalis, although smaller, but

differs in characters of the male genitalia and has less orange on the forewings.

VOLUME 58, NUMBER 3

171

Figs. 17-20. Female genitalia of Thaumatographa species. Fig. 17. T. youngiella (Busck),

Victoria, Vancouver Island, British Columbia, Canada (USNM 77177). Fig. 18. Same, signum

detail. Fig. 19. T. jonesi (Brower), Martha’s Vineyard, Massachusetts (JBH 501, paratype) (CPK).

Fig. 20. Same, signum detail.

172

PAN-PACIFIC ENTOMOLOGIST

Forewing length 5.2 mm. Head: brown; labial palpus white, upturned with

short tuft on 2nd segment. Thorax: fuscous; legs tan, with fuscous on femora

and tarsal segments. Forewing: relatively pointed; termen oblique; macula-

tion as in Figure 2, ground color fuscous; orange scales basally bordering

pale silvery white fascia at % from base and a conspicuous orange area on

apical quarter; distal of pale silvery white fascia another 4 white marks on

costal margin; white speckled area interspersed among black striae beyond

mid-wing, with 4 black spots somewhat merged along tornal angle inter¬

spersed with silvery spots distally; distal two costal white marks also ex¬

tended as silver lines; fringe fuscous; ventral side dull fuscous with white

costal marks repeated. Hindwing: uniformly fuscous. Abdomen: fuscous with

silvery scales at posterior margin of each segment. Male genitalia: as in

Figures 8, 9 (damaged); valva elongate, setaceous, with rounded apex; tegu-

men and vinculum fused; saccus a sharp point; uncus elongated, with a thorn

on tip ventrally and setaceous; hamus long, narrow and upcurved near distal

end; socius % hamus length, setaceous; gnathos a membranous spatulate

structure, truncate apically; transtilla large, blunt rounded; anellus horse-

shoe-shaped with a triangular-shaped base; aedeagus (Fig. 10) elongate with

long cornutus having one spine; vesica with long hood. Female genitalia:

unknown.

Types.— Holotype 3: La Casimba [Camaguey], CUBA, 12 Dec 1969, ex

Pinus cubensis (BMNH, slide no. 20267).

Distribution.— Known only from Cuba.

Host.—Pinus cubensis Grisebach (Pinaceae).

Remarks. — Like the Japanese Thaumatographa decoris, this species ap¬

pears remarkably similar to T. regalis. The genitalia are very different and

the wing maculation is different in details from the other two species.

Thaumatographa youngiella (Busck), New Combination

Hilarographa youngiella Busck, 1922:278.

Hilarographa olympica Braun, 1923:118.

This species is very similar in maculation to Thaumatographa jonesi

(Brower) but lacks the distinct red-orange areas, having instead pale yellow

or buff areas and a general dark brown appearance.

Forewing length 4.8-6.4 mm. Head: fuscous and buff, becoming white on

frons; labial palpus smooth-scaled and white, with fuscous on basal segment;

antenna short with long ventral setae in males. Thorax: fuscous; patagia

lustrous gray fuscous; legs white with fuscous bands on tibiae and tarsal

segments. Forewing: as in Figure 3; fuscous ground color, with 5 oblique

costal white fascia and apical white spot; dorsal margin with 3 closely spaced

oblique fascia at x h from base and another white fascia at % from base; ends

of all fascia silver metallic; yellow-buff areas interspersed between white

VOLUME 58, NUMBER 3

173

fascia, each with black borders; termen with 4 black spots toward tornus;

mid-terminal margin with silver spot; fringe fuscous with subapical white

spot; ventral side fuscous with costal white marks repeated. Hindwing: fus¬

cous; fringe pale fuscous; ventral side dull fuscous with white apical border.

Abdomen: fuscous with pale silvery scales on posterior of each segment.

Male genitalia: as in Figure 13; oblong setaceous valva, with rounded apex

and antero-dorsad slit for corematal setae; tegumen and vinculum fused as

sclerotized ring; saccus shortly extended; uncus spatulate, with narrowed

point; hamus long; socius equally long, flattened and setaceous; gnathos

horseshoe-shaped, somewhat evenly thickened; transtilla quadrate with

thickened posterior end; anellus horseshoe-shaped with slight ventral point;

aedeagus (Fig. 14) elongate, with small cornutus; vesica with elongated hood.

Female genitalia: as in Figure 17; ovipositor flattened; apophyses stout;

ostium a simple cup; ductus bursae straight, posterior half sclerotized; corpus

bursae moderate in size with small accessory bursa on moderately long duct;

signum (Fig. 18) a small linear patch of scrobinations.

Types. — Lectotype 9 ( youngiella ): Biological Station, Departure Bay [Van¬

couver Island], BRITISH COLUMBIA, CANADA, 4 Aug 1908, C. H. Young

(CNC No. 1129), hereby designated. Holotype 9 ( olympica ): Olympic Hot

Springs trail [Clallam Co.], Washington, 27 Jul 1922 [E. L. Braun] (ANSP).

Specimens studied. — (38 < 3 , 10 9 ): Paralectotypes (3 < 3 , 1 9 ): same locality

as lectotype of T. youngiella, 2 Aug 1908 (1 < 3 , USNM), 4 Aug 1908 (1 3 ,

CNC; 1 < 3 , USNM), 9 Aug 1908 (1 9 , CNC), C. H. Young. CANADA.-

BRITISH COLUMBIA: Duncan [Vancouver Island], 4 Aug 1920 (2 < 3 ), 9

Aug 1933 (4 3 ), W. Downes (CNC); Fraser Mills, 30 Jun 1923 (1 < 3 ), 25 Jul

1922 (1 < 3 ), L. E. Marmont (USNM); Victoria [Vancouver Island], 2 Jul 1922

(1 < 3 ), 10 Jul 1923 (1 < 3 ), 15 Jul 1923 (1 < 3 ), 7 Aug 1922 (3 3 ), 10 Aug 1922

(1 9 ), W. R. Carter (USNM); Victoria, 31 Jul 1920 (1 9 ), W. Downes (CNC);

Victoria, 12 Jul 1923 (1 < 3 , 2 9 , ANSP), 14 Jul 1923 (1 6 , ANSP; 9 < 3 , USNM),

4 Aug 1923 (1 9 , USNM), W. H. Blackmore. USA.-CALIFORNIA: Lake

Co.: 1 mi. NW Cobb, 31 May-27 Jun 1979 (1 < 3 ), Powell and De Benedictis

(UCB). WASHINGTON: Clark Co.: Vancouver, 22 Aug 1930 (1 < 3 , 1 9 ), J.

F. G. Clarke (USNM). King Co.: Seattle, 29 Jul 1963 (2 < 3 ), T. Kincaid (UCB).

Pierce Co.: Puyallup, 20 Aug 1930 (1 6 , 1 9 ), T. C. Clarke (USNM); South

Tacoma, 20 Aug 1930 (1 9 ), J. F. G. Clarke (USNM). Thurston Co.: Lk. St.

Clair, 21 Aug 1930 (1 6), T. C. Clarke (USNM). Whatcom Co.: Bellingham,

19 Aug 1962 (1 < 3 , 1 9 ), J. F. G. Clarke (USNM); Squalicum Lk., 14 Aug

1930 (1 < 3 ), J. F. G. Clarke (USNM), and 14 Aug 1930 (1 < 3 ), T. C. Clarke

(USNM).

Distribution.— Southern British Columbia to northern California.

Host. — Unknown.

Remarks.— This species and the following species, Thaumatographa jonesi

(Brower), are related yet quite different in appearance and in genital char-

174

PAN-PACIFIC ENTOMOLOGIST

acters from T. regalis and T. cubensis. The Japanese species Thaumatogra-

pha aurosa (Diakonoff & Arita) is very similar in forewing pattern to T.

youngiella but is distinguished by the genitalia.

Thaumatographa jonesi (Brower), New Combination

Hilarographa jonesi Brower, 1953:96.

Superficially this species from the eastern United States is similar to T.

youngiella but differs in having much more red-orange on the forewings.

Forewing length 5.9-6.9 mm. Head: fuscous with central frons having

white line and white along lateral borders of frons; labial palpus white,

smooth-scaled with fuscous basal segment and some brown apically; antenna

short with long ventral setae in males. Thorax: fuscous; patagia fuscous with

some buff-white anteriorly; legs white with fuscous bands, especially on tarsal

segments. Forewing: as in Figure 4; ground color fuscous between numerous

oblique fascia; an orange mid-wing stria from base to Vi of wing; along costal

margin striae and marks begin with white and alternate with red-orange,

each bordered by black, with apical two orange fascia extending to tornal

angle; apical white fascia extended as silver; along anal margin first 3 white

fascia, then a yellow-orange fascia, then orange alternated with white until

tornus; termen with 5 black spots to tornus; silver mid-termen mark; fringe

fuscous except for subapical white spot on termen; ventral side fuscous with

white costal marks repeated. Hindwing: fuscous with pale white and fuscous

fringe; ventral side fuscous with subterminal and apical white border. Ab¬

domen: fuscous with pale silvery scales on posterior of each segment. Male

genitalia: as in Figure 11, with oblong setaceous valva antero-dorsally slit

for corematal setae insertion; tegumen and vinculum as fused and sclerotized

ring, with extended saccus; uncus spatulate with short knob on tip; hamus

long; socius equally long, flattened and setaceous; gnathos horseshoe-shaped,

with divergent pointed base ends; transtilla quadrate with thickened pos¬

terior end; anellus horseshoe-shaped; aedeagus (Fig. 12) elongate with thorn¬

like cornutus; vesica with elongated hood. Female genitalia: as in Figure 19;

flattened ovipositor; apophyses stout; ostium a simple cup; ductus bursae

straight, posterior half sclerotized; corpus bursae large, ovate with small

accessory bursa on moderately long duct; signum (Fig. 20) a small linear

patch of scrobinations.

Types. — Holotype < 5 : Martha’s Vineyard, Dukes Co., MASSACHUSETTS,

6 Aug 1941, F. M. Jones (AEB). Paratypes (4 9 ): same locality as holotype,

18 Jul 1947 (1 9 ), F. M. Jones (CPK), 6 Aug 1943 (1 9 ), F. M. Jones (USNM).

New Jersey.— Lakehurst [Ocean Co.], 4 Jul 1909 (1 9 ), W. D. Kearfott

(USNM). New York.-O rient [Suffolk Co.], Long Island, 2 Sep 1948 (1 9 ),

R. Latham (AMNH).

Additional specimens (5 < 5 , 1 9 ; 2).— Alabama. —Covington Co.: Open Pond

VOLUME 58, NUMBER 3

175

Cpgd., 20 mi. S Andalusia, 18 Apr 1976 (1 5), J. B. Heppner (JBH). Florida.—

Escambia Co.: Pensacola, 28 Sep 1961 (1), S. M. Hills (SMH). Highlands

Co.: Archbold Biol. Sta., 24 Nov 1971 (1), S. A. Frost (FSCA). Louisiana.—

St. Tammany Pa.: Slidell, 20 Sep 1974 (1 6), V. A. Brou (VAB). South

Carolina. —O conee Co.: Cherry Hill Rec. Area, 2000 ft., 5 Sep 1958 (1 <5),

7 Sep 1958 (1 6, 1 9), J. G. Franclemont (USNM). Texas. — Montgomery

Co.: Conroe, Camp Strake, 14 Sep 1977 (1 5), E. C. Knudson (FSCA).

Distribution. — Eastern coastal United States from Massachusetts to Texas.

Flo st. —Unknown.

Remarks. — This species has a forewing pattern similar to T. youngiella

but is more red-orange as opposed to the fuscous color of T. youngiella. The

larva of this species probably feeds on several species of pine in the coastal

areas and into the piedmont of the eastern United States.

Acknowledgments

Specimens were studied from the following institutions and private col¬

lections: A. E. Brower, Augusta, Maine (AEB); American Museum of Natural

History, New York (AMNH); British Museum (Natural History), London

(BMNH); Vernon A. Brou, Edgard, Louisiana (VAB); California Academy

of Sciences, San Francisco (CAS); California State Department of Agricul¬

ture, Sacramento (CSDA); Canadian National Collection, Ottawa (CNC);

Florida State Collection of Arthropods, Gainesville (FSCA); D. C. Frack,

Los Angeles, California (DCF); S. M. Hills, North Carolina (SMH); J. B.

Heppner, Washington, D. C. (JBH); C. P. Kimball, West Barnstable, Mas¬

sachusetts (CPK); R. H. Leuschner, Manhattan Beach, California (RHL);

Los Angeles County Museum of Natural History, Los Angeles (LACM);

United States National Museum of Natural History, Smithsonian Institu¬

tion, Washington, D.C. (USNM); and the University of California, Berkeley

(UCB). I wish to thank my colleagues at the Smithsonian Institution, J. F.

G. Clarke and D. R. Davis, for their comments on the manuscript.

Literature Cited

Braun, A. F. 1923. Microlepidoptera: notes and new species. Trans. Am. Entomol. Soc., 49:

115-127.

Brower, A. E. 1953. Three new species of microlepidoptera (Olethreutidae, Glyphipterygidae

and Yponomeutidae). Ann. Entomol. Soc. Am., 46:95-98.

Busck, A. 1922. Microlepidoptera from British Columbia. Can. Entomol., 53:276-280.

Diakonoff, A. 1977a. Description of Hilarographini, a new tribus in the Tortricidae (Lepi-

doptera). Entomol. Ber. (Amst.), 37:76-77.

-. 1977b. Rearrangement of certain Glyphipterygidae sensu Meyrick, 1913, with de¬

scriptions of new taxa (Lepidoptera). Zool. Verh. (Leiden), 158:1-55.

-, and Y. Arita. 1981. The early stages of Thaumatographa eremnotorna Diakonoff &

Arita with remarks on the status of the Hilarographini (Lepidoptera Tortricidae). Ento¬

mol. Ber. (Amst.), 41:56-60.

176

PAN-PACIFIC ENTOMOLOGIST

Heppner, J. B. 1978. Transfers of some Nearctic genera and species of Glyphipterigidae

( auctorum ) to Oecophoridae, Copromorphidae, Plutellidae, and Tortricidae (Lepidop-

tera). Pan-Pac. Entomol., 54:48-55.

-. 1982. A synopsis of the Hilarographini (Lepidoptera: Tortricidae) of the world. Proc.

Entomol. Soc. Wash., 84:704-715.

-. In prep. a. Copromorphoidea. In R. B. Dominick et al., The moths of America north

of Mexico. Fasc. 5. Classey, London.

-. In prep. b. Sesioidea: Choreutidae. In R. B. Dominick et al., The moths of America

north of Mexico. Fasc. 8.5. Classey, London.

Meyrick, E. 1886. Descriptions of Lepidoptera from the South Pacific. Trans. Entomol. Soc.

Lond., 1886:189-296.

-. 1914. Lepidoptera Heterocera: Fam. Glyphipterygidae. In M. P. Wytsman (ed.),

Genera Insectorum. Fasc. 164. Brussels, Belgium, 39 pp., 2 pis.

Walsingham, T. de Grey. 1881. On some North American Tineidae. Proc. Zool. Soc. Lond.,

1881:301-325, pis. 35-36.

-. 1897. Western equatorial African Micro-lepidoptera. Trans. Entomol. Soc. Lond.,

1897:33-67, pis. 2-3.

PAN-PACIFIC ENTOMOLOGIST

July 1982. Vol. 58, No. 3, pp. 177-183

Published 16 December 1983

OBSERVATIONS ON POPULATIONS OF

TRIBOLIUM BREVICORNIS

LECONTE (COLEOPTERA: TENEBRIONIDAE).

II. THE HABITAT NICHE OF A LOCAL

POPULATION IN SOUTHERN CALIFORNIA

Alexander Sokoloff, Ruth Wilson, and Gary Mulder

California State College San Bernardino,

San Bernardino 92407

AND

Daryl Faustini

Department of Entomology, University of Wisconsin,

Madison 53706

Despite the great economic importance of flour beetles of the genus Tri-

bolium MacLeay, little is known about their natural ecological niche. In the

past, their role in nature was fabricated partly from naturalists’ observations

of the behavior of tenebrionids in general, and partly from their behavior

under artificial conditions (in warehouses or in the laboratory). Thus, two

main hypotheses about the role Tribolium plays in biological communities

were proposed: 1. Because the beetles occur in the nests of bees, commonly

inhabit stored products, and have been reported from under the bark of

trees, some investigators (e.g., Magis, 1954) suggested that they are herbi¬

vores, feeding primarily on carbohydrates, fungi, or other materials of plant

origin; and 2. Because many tenebrionids are scavengers or predaceous under

natural conditions (e.g., species of Ulominae including several species of

Tribolium invade nests of social insects), and because cannibalism was ob¬

served in laboratory cultures, some investigators (e.g., Linsley, 1944) sug¬

gested that they must be omnivores, surviving in nature as scavengers or

semi-predators.

More recently, Park et al. (1965, 1970), and Sokoloff and Lerner (1967)

from laboratory studies of single- and mixed-species populations of Tribo¬

lium in various media established that cannibalism is common and exten¬

sive. Sokoloff and Lerner have further emphasized that the interaction be¬

tween T. castaneum (Herbst) and T. confusum du Val in laboratory cultures

must be considered a predator-prey interaction rather than one of compe¬

tition. (For a comprehensive review of the literature and an extensive dis¬

cussion of this problem see Sokoloff, 1974.) From these recent laboratory

observations, a third hypothesis can be advanced, namely, that in nature

178

PAN-PACIFIC ENTOMOLOGIST

Tribolium is a secondary or tertiary consumer, engaging in scavenging, pred¬

atory and cannibalistic activities.

Historically Tribolium castaneum and T. confusum are species long as¬

sociated with stored products (Good, 1933). Their distribution and survival

has been greatly influenced by man, through commerce and experimentation.

Hence, these synanthropic species cannot be used effectively to test the three

alternative hypotheses. However, other more primitive and less affected

species of the genus may provide the information to fill this gap.

This paper will report field and laboratory observations of organisms

associated with Tribolium brevicornis LeConte, a primitive species native

to North America. Although recorded as a minor pest of stored products in

other parts of California (Okumura and Strong, 1965; Strong, 1970), and as

a serious pest to commercial growers of Megachile pacifica (Panzer) (Polk,

1979) in Idaho, this species appears to be little influenced by human activities

in the area surveyed. Thus, the present information is relevant in establishing

the habitat niche of T. brevicornis, and may contribute toward understanding

other species whose habitat niche remains undefined.

Materials and Methods

Specimens for this investigation were collected from Waterman Canyon

at an altitude of approximately 365 m (1200 ft.) located at the southwestern

base of the San Bernardino Mountains, just outside the northern city limits

of San Bernardino, California.

A survey of the flora was carried out using standard quadrat plot tech¬

niques.

To determine the diversity of organisms associated with Tribolium brevi¬

cornis in the decaying log biocoenosis we removed three 1 m sections from

a downed alder (Alnus rhombifolia Nutt.) and one 1 m section from a dead

sycamore ( Platanus racemosa Nutt.) lying nearby which contained a hive

of Apis mellifera Linnaeus. In the laboratory the pieces of alder were sec¬

tioned with a bandsaw either into longitudinal pieces 3 x 3 x 100 cm, or

into cross-sections 3 cm thick. Organisms crawling on the surface were either

preserved in alcohol, or (as in the case of T. brevicornis) saved and placed

in standard flour beetle culture medium (19 parts wheat flour, 1 part brewer’s

yeast). To recover organisms from the galleries of carpenter bees the sections

of the log were tapped against each other, and any loose material was allowed

to fall on a sheet of white poster board. Sawdust produced from sectioning

was sifted through a coarse silk-bolting cloth sieve to recover additional

specimens.

The honeycomb from the beehive was removed from the log and frozen.

Later the cells of the honeycomb were examined with a microscope for the

presence of the various stages of the flour beetle.

VOLUME 58, NUMBER 3

179

Results and Conclusion

Habitat and flora.— The foothills adjacent to the Tribolium brevicornis

collection site are typified by a mixture of Chamise Chaparral and Southern

Oak Woodland plant communities (Munz and Keck, 1949) represented by

Adenostoma fasciculatum H. & A., Rhamnus californica Esch., Ceanothus

spp., Yucca whipplei Torr., Prunus ilicifolia (Nutt.) Walp., Cercocarpus bet-

uloides Nutt, ex T. & G., Heteromeles arbutifolia M. Roem., Rhus ovata

Wats., R. laurina Nutt, in T. & G., Styrax officinalis L., Quercus dumosa

Nutt., Q. crysolepis Liebm., Sambucus mexicana Presl., and Toxicodendron

diversilobum (T. & G.) Green. Naturalized escapes and native “weedy”

Coastal Sage Scrub species were noted growing in areas disturbed by settle¬

ment.

The habitat of Tribolium brevicornis appears to be restricted to the more

mesic streamside sites. Alnus rhombifolia is the dominant tree species at the

streamside. Other tree species associated with this mesic area include Acer

macrophyllum Pursh., Umbellularia californica (H. & A.) Nutt., Plat anus

racemosa, Quercus crysolepis, and Salix spp. Seventy-five Alnus rhombifolia

trees were recorded from an area of about 480 m 2 , ranging in basal area

from 2246 cm 2 to 121 cm 2 and averaging about 734 cm 2 .

Fauna. —The fauna found in the alder tree (Table 1) is typical of a decaying

log. Many of the organisms (such as isopods, collembolans, termites, etc.)

require mesic conditions for their survival. From Table 1 it is possible to

speculate on fairly safe grounds that various arachnids (spiders and pseudo¬

scorpions) and chilopods probably include the immature and adult stages

of Tribolium brevicornis among their prey since their stereotyped predaceous

and carnivorous habits are well known.

The reported feeding habits of Tenebrionidae, on the other hand, are

variable, ranging from herbivory to omnivory to carnivory, and cannibalism

(literature review in Sokololf, 1974). Thus, it is not safe to speculate about

the feeding habits of Tribolium on the basis of feeding habits of other genera

within the family as Linsley (1944), Hinton (1948), Butler (1949) and Magis

(1954) have done.

Surveys show that most of the wild species of Tribolium have been found

mainly under bark, and occasionally synanthropic species return to and are

captured in this same habitat. Hence, Good (1933, 1936), Linsley (1944),

Butler (1949) and Magis (1954) have assumed that the primitive and natural

habitat of Tribolium (and of the whole family) is under the bark or in decaying

logs. On the other hand, reports also show that Tribolium in the various

species groups have a tendency to become associated with other organisms,

particularly Hymenoptera; e.g., T. brevicornis in nests of Xylocopa and An-

thidium, T. confusum in nests of Anthophora, Clisodon and Osmia (Linsley

and MacSwain, 1942), T. destructor Uyttenboogaart in nests of Anthophora

Table 1. Species associated with Tribolium brevicornis in decaying Alnus rhombifolia logs.

Phylum/

class

Order

Family

Genus or species if known

MOLLUSCA

Gastropoda

Pulmonata

Limacidae

Limax marginatus Muller

ARTHROPODA

Arachnida

Pseudoscor-

—

pionida

Acari

Trombidiidae

Ceraneidae

—

Araneidae

Gnaphosidae

Harpyllus sp.

Thomisidae

_ 1

Theridiidae

Pardosa sp. 2

Crustacea

Isopoda

Oniscidae

Porcellio sp.

Armadillididae

Armadillium vulgare (Latreille)

Diplopoda

—

Chilopoda

Scolopendra sp.

Insecta

Collembola

Entomobryidae

—

Isoptera

Kalotermitidae

Incisitermes minor (Hagen)

Hodotermitidae

Zootermopsis angusticollis Hagen

Hemiptera

Miridae

—

Coleoptera

Carabidae

Axinopalpus biplagiatus (Dejean)

Buprestidae

Dicerca sp. (prob. horni Crotch)

Elateridae

_ 3

Dermestidae

Trogoderma orbatum (Say)

Bostrichidae

Amphicerus cornutus (Pallas)

Ostomidae

Tenebroides crassicornis (Horn)

Anthicidae

Vacusus confinus (LeConte)

Rhizophagidae

Rhizophagus sp.

Cucujidae

Brontes dubius truncatus

(Motschulsky)

Endomychidae

Symbiotes montanus (Casey)

Tenebrionidae

Blapstinus sp.

Metoponium convexicolle

(LeConte)

Tribolium brevicornis (LeConte)

Cerambycidae

Parandra marginicollis margini-

collis Schaeffer

Curculionidae

Rhyncolus sp. angulans LeConte?

Lepidoptera

Arctiidae

Arachnis picta Packard

Diptera

Bombyliidae

Anthrax tigrinus (De Geer)

Hymenoptera

Chrysididae

—

Formicidae

Camponotus sp.

Liometopum occidentale (Emery)

Tapinoma sessile (Say)

Sphecidae

Ectemnius sp.

Megachilidae

—

Apidae

Xylocopa tabaniformes orpifex

(Smith)

CHORDATA

Amphibia

Anura

Hylidae

Hyla sp.

1 Immature. 2 Female with young in egg sac. 3 Early instar larva.

VOLUME 58, NUMBER 3

181

Fig. 1. a, Waterman Canyon creek with Alnus rhombifolia. b, Dead A. rhombifolia which

served as nesting site for Xylocopa. c and d, Sections of A. rhombifolia showing galleries of

Xylocopa.

and Osmia (Linsley, 1944), T. anaphe Hinton from cocoons of Anaphe

moloneyi Druce (Hinton, 1948), T. castaneum from nests of Megachile, and

T. apiculum Neboiss from nests of Trigona carbonaria Smith (Neboiss,

1962). Tribolium madens Charpentier was found in beehives in Silesia, and

182

PAN-PACIFIC ENTOMOLOGIST

Haragsim (1965) states that this species and T. confusum occur in beehives

in Czechoslovakia. Tribolium audax Halstead has been obtained from cells

of the bee Megachile (Eutricharaea) rotundata (Fabricius) (Leech, 1943;

Halstead, 1969). Tribolium myrmecophilum Lea has been found in the nests

of the ant Iridomyrmex nitidus Mayr (Lea, 1905) and has been recorded

feeding on the pollen reserves of the Australian stingless bee Trigona (Ray-

ment, 1932). Tribolium antennatum Hinton may occupy a similar habitat

(Hinton, 1948). The literature fails to specify what kind of food the beetles

have taken in. The present study shows that T. brevicornis occurs in decaying

logs where the carpenter bee Xylocopa tabaniformes orpifex (Smith) has built

galleries as nesting sites. Tribolium brevicornis was found in the galleries

of Xylocopa and in other sites (but not in ant or termite nests). The species

of tree apparently is of no importance. In the present case T. brevicornis was

found in a decaying Alnus rhombifolia, but Linsley (1944) found it infesting

a nest of Xylocopa in a decaying Libocedrus decurrens Torr.

Interestingly, T. brevicornis was found in Xylocopa nests but did not occur

in a beehive of Apis mellifera found only three meters away, even though

the latter would provide a richer source of food than the nests of Xylocopa

(Sokoloff and Moore, unpublished). Over four dozen flour beetles were re¬

trieved from the portion of the tree we examined. Unfortunately the vibra¬

tion resulting from the sawing procedure caused the beetles to scatter, so

that there were no aggregations in any one site of the log. The discovery of

several large larvae of T. brevicornis verifies the fact that these beetles re¬

produce within the log. Some were found in galleries containing dead, dis¬

membered Xylocopa adults. In a later study Sokoloff and Moore (unpub¬

lished) found T. brevicornis larvae feeding on pupae of Xylocopa. Further

examination of T. brevicornis adults freshly captured in the field revealed

the remains of carpenter bee exoskeletons in their digestive tracts.

Hence, the evidence gathered so far suggests that T. brevicornis is probably

a secondary or tertiary consumer, engaging in scavenging, predatory, and

possibly cannibalistic activities within the decaying log biocoenosis.

Acknowledgments

We are grateful to the following specialists who classified the fauna as¬

sociated with Tribolium brevicornis. Dr. Fred Andrews, Dr. Thomas Eichlin,

Mr. Ray Gill, Dr. Alan Hardy, Mr. A. Toku, Miss Marjorie Moody, Mr.

Terry N. Seeno, Dr. M. K. Rust, Dr. Marius S. Wasbauer, and to Mr. Roy

Martin, California State College, San Bernardino, who identified the botan¬

ical material.

We thank Daniel L. Lopez, Elaine A. Sokoloff and Michael A. Sokoloff

for assistance in the field and Stan Ziegler and Frank L. Lootens for help in

the shop.

VOLUME 58, NUMBER 3

183

This investigation was supported by U.S. Army Research Office grant

DRXRO-CB-14447-L.

Literature Cited

Butler, P. M. 1949. Observations on the biology of Palorus ratzeburgi Wissman with com¬

parative notes of Tenebrionidae in general (Coleoptera). Trans. R. Entomol. Soc. Lond.,

100:249-273.

Good, N. E. 1933. Biology of the flour beetles, Tribolium confusum Duv. and T. ferrugineum

Fab. J. Agric. Res., 46:327-334.

-. 1936. The flour beetles of the genus Tribolium. U.S. Dep. Agric. Tech. Bull., 498:

1-58.

Halstead, D. G. H. 1969. Notes of the systematics and distribution of some Tribolium species

(Coleoptera: Tenebrionidae). J. Stored Prod. Res., 3:269-272.

Haragsim, O. 1965. Potemnici ( Tribolium sp.) jako skudci pylovych zasob. ( Tribolium species

as pests of pollen supplies). Ved. Pr. vysk. Ust. vcelar CSAZV, 4:61-65. (Rev. Appl.

Entomol. (A), 1967:55, 606.)

Hinton, H. E. 1948. A synopsis of the genus Tribolium MacLeay with some remarks on the

evolution of its species groups. Bull. Entomol. Res., 39:13-55.

Lea, A. M. 1905. On Nepharis and other ants’ nest beetles taken by Mr. J. C. Goudie at

Birchip. Proc. R. Soc. Victoria, 17:371-385.

Leech, H. B. 1943. Black flour beetle Tribolium madens Charp., in British Columbia. Can.

Entomol., 75:40.

Linsley, E. G. 1944. Natural sources, habitats, and reservoirs of insects associated with stored

food products. Hilgardia, 16:187-224.

-, and J. W. MacSwain. 1942. The parasites, predators and inquiline associates of

Anthophora linsleyi. Am. Midi. Nat., 27:402-417.

Magis, N. 1954. Apergu de l’histoire naturelle des complexes d’especes du genre Tribolium

(McLeay, 1825) (Coleoptera Tenebrionidae). Bull. Inst. R. Sci. Nat. Belg., 30:1-10.

Munz, P. A., and D. D. Keck. 1949. California plant communities. El Aliso, 2:87-105.

Neboiss, A. 1962. Notes on distribution and description of new species. Mem. Natl. Mus.

Victoria, 25:243-258.

Okumura, G. T., and R. G. Strong. 1965. Insects and mites associated with stored foods and

seeds in California. Bull. Calif. Dep. Agric., 54:13-23.

Park, T., D. B. Mertz, W. Grodzinski, and T. Prus. 1965. Cannibalistic predation in popu¬

lations of flour beetles. Physiol. Zool., 38:289-321.

-, M. Nathanson, J. R. Ziegler, and D. B. Mertz. 1970. Cannibalism of pupae by mixed-

species of adult Tribolium. Physiol. Zool., 43:166-184.

Polk, D. F. 1979. The life cycle and control of Tribolium brevicornis (LeConte) in leafcutting

bee boards. Unpublished MS Thesis, University of Idaho.

Rayment, T. 1932. The stingless bees of Australia. Victorian Nat., 48:183-189, 203-212,

246-254; 49:9-15, 39-43, 104-107.

Sokoloff, A. 1974. The biology of Tribolium with special emphasis on genetic aspects. Oxford

University Press, vol. 2.

-, and I. M. Lemer. 1967. Laboratory ecology and mutual predation of Tribolium species.

Am. Nat., 101:261-276.

-, and R. A. Moore. 1981. A search for methods of determining the type of food ingested

by Tribolium. Tribolium Inf. Bull., 22:139-140.

Strong, R. G. 1970. Distribution and relative abundance of stored-product insects in Cali¬

fornia: A method of obtaining sample populations. J. Econ. Entomol., 63:591-596.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 184-195

Published 16 December 1983

A NEW EUPOMPHA FROM BAJA CALIFORNIA WITH

ADDITIONAL INFORMATION ON E. DECOLORATA (HORN)

(COLEOPTERA: MELOIDAE ) 1

John D. Pinto

Department of Entomology, University of California, Riverside 92521

A recent examination of miscellaneous Meloidae in the California Insect

Survey Collection, University of California, Berkeley revealed eight indi¬

viduals of a new Eupompha, herein named E. vizcaina. This series was

collected by John T. Doyen in the spring of 1976 at Miller’s Landing in the

Vizcaino region of Baja California. A visit to the area in the spring of 1980

produced additional material. A description of the adult, first instar larva

and courtship behavior of E. vizcaina is given below. A second Baja Cali¬

fornia species, E. decolorata (Horn) also was found. Its first instar larva and

courtship behavior are also described.

Eupompha vizcaina and E. decolorata are the only Eupompha confined

to Baja California. They apparently are restricted to the Vizcaino Desert in

the central third of the peninsula. Adults of both species are synchronous.

However, the few collections so far suggest habitat and plant host differences.

E. vizcaina occurs on sand dunes where it feeds on flowers of Sphaeralcea

axillaris Watson. E. decolorata appears to be more generally distributed off

the dunes. Flowers of Malacothrix californica DC, Encelia palmeri Vasey

and Rose and Viguiera deltoidea Gray are known food sources.

Eupompha vizcaina and E. decolorata belong to Section I of Eupompha

as recently defined in the generic revision by Pinto (1979). However, they

are not close relatives. E. vizcaina is placed in the Viridis Group with E.

edmundsi (Selander) from northern Arizona and southern Utah and E. viridis

(Horn), a Chihuahuan Desert species. E. decolorata was placed in the Elegans

Group with two primarily southern California species, E. elegans (LeConte)

and E. imperialis (Wellman), on the basis of adult anatomy (Pinto, 1979).

Larval anatomy and courtship behavior (see below) corroborate this assign¬

ment.

Eupompha vizcaina Pinto, new species

(Figs. 1, 2, 5, 6, 8, 9)

Adult.— Moderately robust. Surface shining. Color uniformly dark, brassy

with virescent to purpurescent luster over most of body except abdominal

terga I-VI more distinctly virescent, and eyes and antennae black; head

without red spot on frons; wing membrane colorless to lightly infuscate.

VOLUME 58, NUMBER 3

185

Fig. 1. Eupompha vizcaina, male.

186

PAN-PACIFIC ENTOMOLOGIST

Pubescence cinereous, coarse, moderately dense to dense throughout, no¬

ticeably affecting body color. Length from frons (head in hypognathous po¬

sition) to apex of elytra 7.8 ± 0.2 (6-10) mm (N = 10). 2

Head 0.76 ± 0.01 (0.7-0.8) (N = 10) as long as wide, subrectangular,

widest at eyes; sides above eyes relatively straight; occiput straight to broadly

arcuate; moderately densely punctate, center of head typically impunctate,

or obsolescently so; surface between punctures microreticulate; male with a

very shallow, oval, setate sulcus on frons between eyes rarely extending to

epistomal suture, female with sulcus as well or almost as well developed;

frontal area of male densely set with very small cuticular pores (visible with

scanning electron microscope at 1000 x). Eyes large, bulged, angularly emar-

ginate anteriorly slightly above center, not noticeably narrower in dorsal

half, ca. % as wide as long, extending to a point 3 / 5 the distance from epistomal

suture to occiput. Antennae short, slightly compressed anteroposteriorly, not

tapering apically, segment III ca. % and 3 / 4 as long as I in males and females,

respectively; IV almost as wide as long, ca. 3 / 4 as long as III, V-X ca. 1.10

as wide as long. Labial palpi with segment III subequal in length and width

to II.

Pronotum usually very slightly longer than wide, 0.99 ± 0.01 (0.96-1.03)

(N = 10) as wide as long; sides evenly convergent anteriorly and posteriorly

from widest point in apical half; disk uneven, gradually declivent to posterior

margin, with a broad, shallow, horizontal impression across apical third

which often extends posteriorly at middle to center of disk, and a shallow,

subtriangular impression at base; punctures and surface as on head, less

densely punctate on either side of midline, setae directed posteriorly in apical

third, anteriorly in basal third, swirled (variously directed) at center.

Elytra densely, moderately coarsely scabropunctate; setae recumbent, dense

and partially obscuring cuticle. Venter with punctation similar to elytra

except surface along midline of metasternum smoother and impunctate, setae

longer than on dorsum. Legs with femora relatively slender, hind femora

only slightly wider than middle femora; tibial spurs on fore and middle legs

straight, spiniform, brown; spurs on hind tibiae pale yellow, inner spur

bladelike, outer spur spoon-shaped (Fig. 2); fore tarsi (Fig. 8) of male with

segments I-IV swollen, II-IV only slightly so, I almost half as wide as long,

slightly longer than II and III combined; ventral surface of I distinctly con¬

cave, glabrous and impunctate along midline, dorsal surface not sulcate, II-

V with moderately dense, short erect cinereous setae on venter; claws with

Figs. 2-7. Figs. 2-4. Apex of left tibia (ventral view) in species of Eupompha. Fig. 2. E.

vizcaina. Fig. 3. E. decolorata. Fig. 4. E. edmundsi. Figs. 5, 6. E. vizcaina, male genitalia. Fig.

5. Aedeagus. Fig. 6. Gonoforceps. Fig. 7. Hind claw of first instar larva of E. vizcaina.

187

VOLUME 58, NUMBER 3

188

PAN-PACIFIC ENTOMOLOGIST

VOLUME 58, NUMBER 3

189

curvature of ventral tooth slightly more abrupt than that of dorsal blade,

apex of tooth falling short of apex of dorsal blade; onychium with three

setae. Male genitalia (Figs. 5, 6) with posterior margin of basal piece slightly

concave; aedeagus with dorsal spines small, ventral spine elongate, slender

throughout. Female gonostyli moderately long.

First instar larva. — Light brown with head darker; venter with normal,

elongate, spiniform setae throughout; dorsum with both very short, stout

setae, and highly modified, elongate, clavacostate setae (Figs. 11,12); mod¬

ified setae somewhat shorter than unmodified homologues in other Eupom-

pha. Modified setae distributed as follows. Head capsule: widespread except

on labrum, also, lateral-most seta on first setal row behind labrum, seta

posteromedial to eye and those on occiput normal; thorax: 3 on lateral margin

of pronotum; on lateral and posterior margin of meso- and metanotum; a

single seta at base of meso- and metacoxae; abdomen: on posterior margin

of terga only. Unmodified dorsal setae stout, peg-like, much shorter (ca.

V 5 as long) than homologues in other species.

Head (Fig. 11) 30% wider than long, widest near level of eyes; sides

convergent to base; six setae between clypeus and frons (first row behind

labrum); gula about as wide as long, its setae inserted on anterior margin;

eyes subequal in diameter to mesothoracic spiracles. Antenna (Fig. 13) with

segment I ca. twice as wide as long; II as wide as long, ca. twice as long as

I; III twice as long as wide, subequal in length to II; sensory organ as wide

as long, 20% shorter than III; terminal seta moderately long, 60% longer

than III. Mandibles slender apically, abruptly widened at base; inner margin

entire; apical seta twice as long as basal seta. Maxillae with ca. 5 setae at

apex of mala; segment III of palpi ca. 40% longer than wide, asymmetrical,

lateral margin distinctly longer than medial margin, sensory area with ca.

25 papillae, two-segmented appendix not present. Labrum with setae of first

prementum long, seta of second prementum minute; segments I and II of

palpi subequal in length.

Thorax with line of dehiscence confined to pro- and mesonotum and

anterior fourth of metanotum; 26 setae on pronotal disk. Abdomen with

posterolateral margin of terga abutting against pleurites; spiracles placed in

membranous area between pleurites and anterolateral margin of terga; di¬

ameter of first spiracle ca. 20% less than that of mesothoracic spiracle and

75% greater than that of second spiracle, spiracles on segments II-VIII

subequal in size; terga with posterior marginal row of setae ca. half tergum

Figs. 8-11. Figs. 8-10. Fore tarsi (ventral view) of Eupompha spp. Fig. 8. E. vizcaina, male.

Fig. 9. E. vizcaina, female. Fig. 10. E. decolorata, male. Fig. 11. Head (320 x) of first instar

larva of E. vizcaina showing distribution of clavacostate setae (artifact at center of photo).

190

PAN-PACIFIC ENTOMOLOGIST

Figs. 12, 13. First instar larva of Eupompha vizcaina. Fig. 12. Detail of a clavacostate seta

from vertex of head (3200x). Fig. 13. Antenna (ventromedial view, 940x).

length; sternum with sclerites distinct on segments I-IX, those on I-VII

small, paired, increasing in size on posterior segments; VIII and IX each

with a single medial sclerite. Legs slender, hind claw (Fig. 7) long, broadly

curved, ca. 3 / 5 as long as hind tibia; claws with their two setae slightly but

distinctly separated at base, longer seta not approaching apex of claw. Body

length 1.25 mm, caudal setae 0.46 mm (N = 5).

Type information. — Holotype, adult male, from MEXICO, Baja California

Sur, Vizcaino Peninsula, ca. 27°24'N, 114°05'W; 51 road km E Rancho San

Jose Castro, 26 March 1980, dune association; on Sphaeralcea axillaris, J.

D. Pinto, J. M. Mathieu, and E. M. Fisher; deposited in the collection of

the California Academy of Sciences.

Geographic distribution. — Known from three locales in the Vizcaino Des¬

ert of central Baja California.

Records. —19 specimens as follows: MEXICO. Baja California Norte:

Guerrero Negro, 9 km N, 13; Miller’s Landing, 8. Baja California Sur:

Rancho San Jose Castro, 51 road km E, 58.

Larval material examined.— Larvae from eggs laid by females collected

at the type locality (see above). Five separate egg masses laid by different

females hatched in 9 days at 26°C.

Seasonal distribution. — Specimens from Miller’s Landing were collected

VOLUME 58, NUMBER 3

191

on 6 April 1976. Material from the two other locales was taken 24-26 March

1980.

Remarks. — Adults of E. vizcaina are distinguished from all other Eupom-

pha by body color and the moderately dense cinereous setation. Also, it is

the only Eupompha with males having inflated fore tarsi (Figs. 1, 8) but

lacking a distinct cephalic sulcus (Fig. 1).

Eupompha decolorata is the only other Eupompha occurring in the Viz¬

caino Desert. The two are quickly separated by coloration. E. decolorata is

bicolored. The elytra are orange, at least in part, and the rest of the body is

blue-black. E. vizcaina is dark throughout. Also, the fore tarsi of males of

E. decolorata are less modified (Fig. 10).

The first instar larva of E. vizcaina is almost identical to that of E. ed-

mundsi. They are the only meloids known with clavacostate body setae (Figs.

11, 12; also, see Pinto, 1975). The two are separated by rather subtle features.

In E. vizcaina the two claw setae (Fig. 7) do not arise from the same level

as they do in E. edmundsi (Pinto, 1975, Fig. 3). Also, the terminal seta of

the antenna (Fig. 13) is longer in E. vizcaina (60% longer than antennal

segment III) than in E. edmundsi (subequal in length), and abdominal spi¬

racles III-VIII are subequal in E. vizcaina but gradually decrease in size

posteriorly in E. edmundsi.

Eupompha decolorata (Horn)

Calospasta decolorata Horn, 1894:437.

Eupompha decolorata: Pinto, 1979:414.

First instar larva. — Body uniformly brown, with long, heavy spiniform

setae throughout. Head 30% wider than long, widest near level of eyes, sides

gradually convergent to base; six setae between clypeus and frons (1st row

behind labrum); gula about as wide as long, its setae inserted on anterior

margin. Eyes with diameter 20-30% less than that of mesothoracic spiracles.

Antennae with segment I ca. 75% wider than long; II slightly longer than

wide, ca. 50% longer than I; III twice as long as wide, ca. 20% shorter than

II; sensory organ slightly wider and longer than III; terminal seta twice as

long as III. Mandibles slender, distinctly widened basally, feebly crenulate

on inner margin; apical seta long, basal seta minute but usually distinct (ca.

V 5 as long as apical seta). Maxilla with ca. 6 setae on mala; segment III of

palpi ca. 35% longer than wide, asymmetrical, lateral margin distinctly longer

than medial margin, sensory area with ca. 20 papillae, two-segmented ap¬

pendix not evident and probably absent. Labium with setae of first pre-

mentum long, setae of second prementum x h as long; segment II of palpi

slightly longer than I.

Thorax with line of dehiscence distinct on pro- and mesonotum, weakly

expressed on metanotum; 24 setae on pronotal disk. Abdomen with pos-

192

PAN-PACIFIC ENTOMOLOGIST

terolateral margin of terga abutting against pleurites; spiracles placed in

membranous area between pleurites and anterolateral margin of terga; first

spiracle with diameter ca. 20% less than that of mesothoracic spiracle and

twice the diameter of second spiracle; spiracles on segments II-VIII subequal

in size; terga with posterior marginal row of setae ca. % tergal length. Sternum

with a pair of subtriangular sclerites on I-VIII, sclerites gradually increasing

in size on posterior segments, sclerites on VIII partially fused medially or

not; IX with a single, large sclerite. Legs slender, hind claw straight, curved

slightly at apex, ca. % as long as hind tibia; claws with their two setae

separated at base by ca. V 10 claw length, longer seta reaching slightly beyond

apical ‘/ 5 of claw. Body length 1.48 mm; caudal setae 0.62 mm (N = 5).

Material examined. — Larvae from eggs laid by a female collected in MEX¬

ICO, Baja California Sur, 39 km W San Ignacio, on Malacothrix californica.

Eggs hatched 9 days after oviposition at 26°C.

Remarks.— The larva of E. decolorata is closest to that of E. imperialis

and E. elegans and will key to the couplet separating these species in Pinto

(1979). The larva of E. elegans was described by MacSwain (1956) and that

of E. imperialis by Pinto (1975). E. decolorata is distinguished from E.

imperialis primarily by eye size. In E. imperialis the eyes are large. Their

diameter is subequal to that of the mesothoracic spiracles. In E. decolorata,

as in E. elegans, the diameter of the eyes is 20-30% less than that of the

mesothoracic spiracles.

Eupompha decolorata is best separated from E. elegans by structure of

the hind claw, size of the abdominal spiracles and the length of the terminal

seta of the antenna. In E. elegans the longest seta on the hind claw reaches

beyond the apical V 10 of the claw (Pinto, 1975, Fig. 4); in E. decolorata it

only approaches the apical V 5 . The abdominal spiracles decrease in diameter

posteriorly in E. elegans; in E. decolorata they are subequal on all segments

posterior to II. The terminal seta is ca. 3 x as long as segment III of the

antenna in E. elegans ; it is only 2 x as long in E. decolorata.

Additional records. — The recent treatment of E. decolorata (Pinto, 1979)

includes only five locality records, the type locality (Calimalli Mines), the

questionable inclusion of specimens from south of San Miguel Comondu,

and three locales within 25 mi. N of Punta Prieta. Additional records are

as follows: BAJA CALIFORNIA NORTE: Bahia de Los Angeles, 25 mi.

W; El Crucero. BAJA CALIFORNIA SUR: Guerrero Negro, 51 km SE;

Rancho San Jose Castro (Vizcaino Peninsula), 25 km E; San Ignacio, 39 km

W; Vizcaino, 56 km W. This additional material was collected between 24

March and 4 April.

Courtship Behavior in E. decolorata and E. vizcaina

The courtship of seven species of Eupompha was described earlier (Pinto,

1977). Courtship in E. decolorata and E. vizcaina is similar to that of most

VOLUME 58, NUMBER 3

193

Eupompha. The male performs all precopulatory behavior from a mounted

position directly above the female. Display consists of antennation and tarsal

rubbing. The male antennae direct those of the female onto the frontal area

of his head capsule and the fore tarsi stroke her maxillary palpi. Bouts of

display are highly variable in duration, lasting from one second to a minute

or more. Most bouts last less than 30 sec. Periods of display alternate with

genital insertion attempts, during which the male tries to copulate, or with

short periods of relative quiescence (dorsal riding). Although courtship dis¬

play is similar in both species the activities are performed differently.

Descriptions of behavior are based on laboratory studies and observations

in the held of very short duration. In E. vizcaina 10 pairs were observed for

a total of 3 hours; in E. decolorata 4 pairs were observed for a total of 1

hour.

E. decolorata.— Antennation dominates courtship display. Each male an¬

tenna is loosely curled around the corresponding antenna of the female and

both are kept in this position during the entire antennation bout. The male

rapidly shifts his body from side to side contacting first one and then the

other female antenna with the front of his head. In this species contact of

each female antenna with the male head is apparently made both by a slight

pulling by the antenna and, more importantly, by his head moving to meet

it. The mean rate that each antenna contacted the male head at 25°C in one

pair was 1.4 sec (1.3-1.5) (N = 3).

During antennation the fore tarsi are directed medially, venter up, and

are either held adjacent to one another or overlap slightly. They usually

remain motionless under the cervical area of the female. Most bouts of

display consist of antennation only. In a minority of bouts, however, the

tarsi quickly move forward in unison and brush the maxillary palpi of the

female (1-several times?) with the ventral surface of the tarsal segments.

Bouts of display are usually followed by genital insertion attempts. If

unsuccessful, the male either dismounts or continues display.

E. vizcaina. — Behavior in E. vizcaina is similar to that in E. decolorata

but with two important differences. Males of E. vizcaina do not move the

head from side to side to meet the female antenna. Instead, the head capsule

is stationary and each female antenna is pulled to the frontal area solely by

the corresponding male antenna as in E. fissiceps (Pinto, 1977, Fig. 5). The

second difference is that during tarsal rubbing the fore tarsi are not in contact

and do not stroke the female’s maxillary palpi in unison. Instead, each is

directed ventromedially, and moves independently and often alternately. As

in E. decolorata, however, it is the venter of the fore tarsus that contacts

the female’s palpi.

Antennation and tarsal rubbing are typically concurrent in E. vizcaina.

However, the rate of tarsal rubbing is ca. twice that of antennation. In a

single bout observed at 29°C each antenna was contacted by the male head

194

PAN-PACIFIC ENTOMOLOGIST

capsule 0.7 x/sec. Concurrent tarsal rubbing contacted each female palpus

1.4 x/sec. Rarely, antennation occurs without tarsal rubbing. This typically

occurs when the female is feeding.

The male genitalia are usually extruded during display and probe the

dorsum of the female. After most lengthy bouts of display the male moves

back slightly and attempts to insert regardless of female receptivity. If failing

to insert, the male either decamps or continues its display.

Three matings were observed. All followed a single bout of display lasting

30-40 sec. Males moved to a linear position within 2 min. after insertion

and remained coupled a mean of 2.17 (2.1-2.2) hours at 26°C.

Discussion

The placement of E. vizcaina in Section I of Eupompha is clearly indicated

by the modification of the ventral surface of the male fore tarsus, the elongate

ventral spine of the aedeagus, and the 6 setae on the frontoclypeus and the

asymmetrical segment III of the maxillary palpi in the first instar larva. The

explanate outer hind tibial spur in E. vizcaina is unique within Section I of

Eupompha (Fig. 2). In other species it is spiniform or stick-shaped and either

similar to the inner spur (Fig. 3) or only slightly wider (Fig. 4).

Assignment of this species to the Viridis Group is indicated by the sim¬

ilarity of its first instar larva to that of E. edmundsi. The only adult feature

suggesting relationship to E. edmundsi is the relatively dense body setation.

This character as well as the presence of clavacostate setae in the larva of

both are unique within Eupompha. The placement of E. edmundsi with E.

viridis, the only other member of the group, was based on the slightly clavate

ventral spine of the aedeagus (Pinto, 1979). In E. vizcaina the spine is not

obviously clavate (Fig. 6). The only features shared by E. vizcaina and E.

viridis are probably primitive within the genus (e.g., short, non-tapering

antennae; relatively subquadrate head; bulged eyes). Until the currently un¬

known larva of E. viridis is described, the relationship of this species to the

others in Section I will remain questionable.

Unlike the other members of the Viridis Group, E. vizcaina has a highly

developed courtship display. Both structure and associated behavior are

unique within Eupompha. Like members of the Elegans Group and unlike

those in Section II, it is the venter rather than the dorsum of the fore tarsus

that is used in tarsal rubbing. It is distinct from Elegans Group species,

however, in that the tarsi work independently and the ventral surface of

segment I is glabrous rather than setate (Figs. 8, 10). Although antennation

is well developed in E. vizcaina, males lack the distinct cephalic sulcus of

the Elegans Group. This shows that the absence of overt structural modi¬

fication is not always correlated with the absence of behavior. Furthermore,

the relationship of E. vizcaina to E. edmundsi, a species without well defined

VOLUME 58, NUMBER 3

195

display (Pinto, 1977), suggests that both tarsal rubbing and antennation

evolved independently within Section I and at least three times within the

genus.

Larval anatomy and courtship behavior show E. decolorata to be closely

related to E. elegans and E. imperialis as previously suggested by adult

structure (Pinto, 1979). Courtship behavior in all three species is basically

the same. The only difference in E. decolorata is the relative infrequency of

tarsal rubbing. Interestingly, the male fore tarsi are only slightly modified

in this species compared to those in E. elegans and E. imperialis (Fig. 10

below and Fig. 17 in Pinto, 1977).

Acknowledgments

I wish to thank the following individuals: John Chemsak and John Doyen

(University of California, Berkeley) for the loan of the first collection of E.

vizcaina ; Eric Fisher and Juan Mathieu for assistance in the held; Steve

Manweiler for helping in the laboratory and for operation of the scanning

electron microscope; Patricia Mote for preparation of Figures 1-7; and An¬

drew Sanders for plant identifications.

Literature Cited

Horn, G. H. 1894. The Coleoptera of Baja California. Proc. Calif. Acad. Sci., (2)4:302-449.

MacSwain, J. W. 1956. A classification of the first instar larvae of the Meloidae (Coleoptera).

Univ. Calif. Publ. Entomol., 12:1-182.

Pinto, J. D. 1975. A description of the first instar larva of Eupompha imperialis and E.

edmundsi (Coleoptera: Meloidae). Pan-Pac. Entomol., 51:134-139.

-. 1977. Comparative sexual behavior in blister beetles of the subtribe Eupomphina

(Coleoptera: Meloidae), and an evaluation of its taxonomic significance. Ann. Entomol.

Soc. Am., 70:937-951.

-. 1979. A classification of the genus Eupompha (Coleoptera: Meloidae). Trans. Am.

Entomol. Soc., 105:391-459.

Footnotes

1 This study was supported by grant DEB-7915307 from the National Science Foundation.

2 Mean ± standard error.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 196-201

Published 16 December 1983

A NEW APHID SPECIES, CINARA RADICIVORA

(HOMOPTERA: APHIDIDAE), LIVING ON WHITE FIR

David Voegtlin

Illinois Natural History Survey, Department of Energy and

Natural Resources, 607 East Peabody, Champaign 61820

The new species of Cinara described below was discovered during a study

of Camponotus spp. in the Giant Forest area of Sequoia National Park,

California. It was found feeding at the root crown on old growth white fir

{Abies concolor (Gord.) Lindl. ex Hildebr.).

Cinara radicivora Voegtlin, new species

(Figs. 1, 2)

Apterous viviparous females.— Color of living specimens: Entire body shiny,

dark green-black with scattered irregular patches of brown on dorsum of

abdomen. Appendages concolorous with body except for lighter areas as

follows: on antennae, proximal % of III, Vi of IV and l k of V; on legs, femora

lightening gradually on proximal Vi and proximal Vi of tibiae becoming

gradually lighter.

Color of mounted specimens: Degree of darkness of sclerotized areas quite

variable between specimens. In general dorsal sclerites lighter brown, lighter

than cauda, anal plate, subgenital plate and legs which vary from light to

dark brown. The sclerotization pattern is as follows: head, pro- and meso-

notum evenly sclerotic with paired, irregularly shaped, membranous patches

spinally on mesonotum (Fig. 2). Rest of dorsum membranous except for

paired sclerites on metanotum, and on abdominal tergites I and VIII. Some

specimens with a few sclerites on tergites VI and VII. Antennae concolorous

with head except as noted in living specimens. Extent of light areas on femora

and tibiae highly variable: both can be almost entirely pale except for joint

areas, or in some specimens, only proximal V 3 is pale with remainder quite

dark.

Measurements (mm).— Length of body, 4.4-6.1. Length of rostrum, 2.9-

3.8; ultimate rostral segment, 0.43-0.56. Antennal segments: III, 0.76-0.90;

IV, 0.31-0.46; V, 0.39-0.55; VI base, 0.20-0.25; process terminalis, 0.05-

0.08. Metatibiae, 2.6-3.2; tarsal I, 0.13-0.15; tarsal II, 0.36-0.46. Diameter

of base of siphuncular cone, 0.31-0.59. Length of setae: on tergite V, 0.13-

0.18; on tergite VIII, 0.15-0.23; longest on antennal III, 0.10-0.16; longest

dorsal surface mid-metatibiae, 0.09-0.14.

Morphology. — Sensoria on antennae: III, 0; IV, 0-2; V, 0-3. No meso-

VOLUME 58, NUMBER 3

197

0 0.2mm 0.4mm

i_i-1

0 0.1mm 0.2mm

I_I- 1

Fig. 1. Cinara radicivora, new species. A, First and second metatarsi; B, Subgenital plate

(Drawn from Holotype); C, Antennal VI; D, Ultimate rostral segment showing accessory setae.

sternal tubercle. Siphuncular cones with irregular anterior and often posterior

edges. Setae on siphunculi evenly scattered, dense (80-160) and gradually

decreasing in length and basal diameter towards top of cone. Subgenital plate

with 29-56 setae distributed as in Figure IB. Ultimate rostral segment with

26-54 accessory setae, those not aligned on either side of the stylet groove

are scattered about segment (Fig. ID). Antennal setae: II, 25-41; VI base,

19-33; process terminalis, 3-4 (Fig. 1C). Tergite V with 175-200 setae and

tergite VIII with 58-90 setae.

Alate viviparous females .—Color of living specimens: Head and thorax

black, abdomen dark green-black. Legs dark throughout except for proximal

x k of femora. Antennal II and proximal areas of III, IV and V light, remainder

black.

Color of mounted specimens: Head and prothorax medium amber, ptero-

198

PAN-PACIFIC ENTOMOLOGIST

Fig. 2. Cinara radicivora, new species. Apterous viviparous female. Drawing by Donna

Baron.

thorax dark brown. Abdomen membranous except for paired sclerites on

tergite VIII. Legs and antennae patterned as in life. Siphunculi medium

brown.

Measurements (mm). — Length of body, 4.1-6.1. Length of rostrum, 3.2-

3.9; ultimate rostral segment, 0.51-0.57. Antennal segments: III, 0.83-1.05;

IV, 0.41-0.51; V, 0.53-0.65; VI base, 0.24-0.29; process terminalis, 0.06-

0.08. Metatibiae, 3.0-4.1; tarsal I, 0.14-0.16; tarsal II, 0.42-0.51. Diameter

VOLUME 58, NUMBER 3

199

of base of siphuncular cone, 0.30-0.65. Length of setae: tergite V, 0.09-0.18;

tergite VIII, 0.13-0.23; longest on antennal III, 0.12-0.16; longest dorsal

surface mid-metatibiae, 0.13-0.19.

Morphology. — Secondary sensoria on antennal III, 1-5; IV, 2-5; V, 1-3.

Siphuncular cones with irregular anterior and often posterior edge. Setae on

siphunculi evenly distributed, dense (110-140) and gradually decreasing in

length and basal diameter towards top of cone. Subgenital plate shaped much

as in apterae (Fig. IB) but in some specimens narrower and longer, with

from 50-78 setae distributed as in apterae. Ultimate rostral segment with

52-67 accessory setae, those not aligned along stylet groove are scattered

about segment. Antennal setae: II, 39-50; base VI, 30-40; process terminalis,

4. Tergite V with 160-200 setae; tergite VIII with 53-80 setae.

Other forms. — No males, oviparae or fundatrices have been collected for

this species.

Types. — Holotype, apterous viviparous female on Abies concolor, Sequoia

National Park, CALIFORNIA, 19-VI-1977, David Tilles, slide #356-1.

Para types: Sequoia National Park, CALIFORNIA; 3 apterae, 19-VI-1977;

4 apterae, 21-VI-1977; 2 apterae, l-VII-1977; 1 aptera, 14-VII-1977; 1

aptera and 2 alatae, 29-VII-1977; 2 alatae, 8-VIII-1977; 1 alata, 14-VIII-

1977; 1 aptera and 1 alata, 15-VIII-1977, David Tilles. 19 apterae and 2

alatae, 11-VII-1979, Suzanne Paulaitis and David Voegtlin. Wildwood Camp,

Ochoco National Forest, OREGON, 2 apterae, 16-VII-1977, David Voegt¬

lin.

Type locality.— The Round Meadows area near Giant Forest in Sequoia

National Park, California.

Deposition of types. — Holotype and paratypes deposited in the collection

of the Illinois Natural History Survey. Other paratypes deposited in the

collections of the United States National Museum of Natural History, British

Museum (Natural History), Canadian National Collection, California Insect

Survey, Berkeley, and D. Hille Ris Lambers.

Diagnosis. —Of the approximately 200 species of Cinara there are only

six known with a rostrum of 3 mm or longer. All are associated with the

trunk, root crown, or root of their host. They are not closely related and the

elongate rostrum must have evolved independently in several species groups

within the genus. The absolute length of the rostrum will separate C. radi-

civora from the majority of the Cinara. C. radicivora can be separated from

the other 5 species with rostrum longer than 3 mm by the following key.

Key to Species of Cinara with Rostrum Longer than 3 mm

1(4). Dorsum of abdomen with very short setae (0.015 mm), each on

a very small sclerite only slightly larger than the base of the

setae.

2(3). Ultimate rostral segment 0.51-0.58 mm and with 22-30 acces-

200

PAN-PACIFIC ENTOMOLOGIST

sory setae. Antennal segment IV distinctly longer than anten¬

nal segment II. With 80+ setae on the siphuncular cones. On

Pinus halepensis Mill., on the bark. France.

. C. balachowskyi Ramaudiere (1974)

3(2). Ultimate rostral segment 0.64-0.74 mm and with 90+ accessory

setae. Antennal segment IV subequal to antennal segment II.

With 29-40 setae on siphuncular cones. On Pinus edulis En-

gelm., below ground level. Colorado. . . . C. puerca Hottes (1954)

4(1). Dorsum of abdomen densely covered with rather long setae, the

majority of which are not on sclerites.

5(6). With accessory setae on ultimate rostral segment in two rows

along the stylet groove, total 16-20 accessory setae. Subgenital

plate almost square in outline and densely covered with setae.

Process terminalis with seven subapical setae. On Picea glauca

(Moench) Voss, roots. Canada. C. saskensis Bradley (1962)

6(5). Accessory setae on the ultimate rostral segment always in more

than the two rows along the stylet groove (Fig. ID). More

than 25 accessory setae. Subgenital plate variable, usually

elongate oval (if it approximates a square then there are five

or fewer subapical setae on the process terminalis).

7(8). Ultimate rostral segment long and narrow, always greater than

0.60 mm. Siphunculi small with fewer than 30 setae. On Pinus

thunbergiana Franco and P. densiflora Siebold & Zucc., on

lower trunk. Japan. C. sorini Inouye (1970)

8(7). Ultimate rostral segment less than 0.56 mm long. Siphuncular

cones large and with 50 or more setae.

9(10). With a distinct mesosternal tubercle. Ultimate rostral segment

0.39-0.46 mm, with 22-26 accessory setae. Rostral length

2.4-3.5 mm. With less than 100 setae on the siphuncular

cones. On Pinus banksiana Lamb., on roots. Canada.

. C. piniradicis Bradley in Bradley and Wighton (1959)

10(9). Without a distinct mesosternal tubercle. Ultimate rostral seg¬

ment 0.43-0.56 mm, with 26-67 accessory setae. Rostral

length 2.9-3.9 mm. With 80-160 setae on the siphuncular

cones. On Abies concolor, root crown. California and Ore¬

gon.C. radicivora, new species

Biological notes. — This species is found beneath the ground level on the

root crown of old growth Abies concolor and in all collections it has been

attended by Camponotus spp. They are not easily collected as they are often

located in cavities formed under the outer layers of the bark. These are

probably naturally formed cavities but the shape and polished appearance

of them suggests that the ants may have assisted in cavity formation or

certainly were keeping the cavity free of debris.

VOLUME 58, NUMBER 3

201

Acknowledgments

I would like to thank David Tilles who provided many of the specimens

collected during his thesis research in Sequoia National Park. I would also

like to thank Donna Baron for the drawing of the whole aphid and George

Godfrey for his review of the manuscript.

Literature Cited

Bradley, G. A. 1962. Three new species of Cinara Curtis (Homoptera: Aphididae) from central

Canada. Can. Entomol., 94:1175-1182.

-, and D. C. Wighton. 1959. Description and life history of a new species of Cinara

(Homoptera: Aphididae) from Pinus banksiana Lamb. Can. Entomol., 91:453-456.

Hottes, F. C. 1954. Descriptions and notes on some species of Cinara (Aphidae). Proc. Biol.

Soc. Wash., 67:251-261.

Inouye, M. 1970. Revision of the conifer aphid fauna of Japan (Homoptera: Lachnidae). Bull.

Gov. For. Exp. Stn. (Tokyo), 228:57-102.

Remaudiere, G. 1974. Un nouveau Cinara des troncs de pins d’Alep (Horn. Aphididae). Ann.

Soc. Entomol. Fr., 10:527-534.

PUBLICATIONS RECEIVED:

Faunal affinities, systematics, and bionomics of the Orthoptera of the

California Channel Islands. By D. C. F. Rentz and David B. Weissman.

University of California Publications in Entomology, Vol. 94, xiv, 240 pp.,

337 figs., 11 tables, 1 map. Publication date: 9 March 1982. Published by

University of California Press, 2223 Fulton Street, Berkeley, CA 94720.

Price: $22.00 softbound. (Received by PCES at CAS 2 April 1982, P. H.

Arnaud, Jr., 1982-1).

The Pentatomoidea (Hemiptera) of northeastern North America with em¬

phasis on the fauna of Illinois. By J. E. McPherson. Southern Illinois Uni¬

versity Press, ix, 240 pp., 191 figs. Publication date given as 20 May 1982.

Available from Southern Illinois University Press, P.O. Box 3697, Carbon-

dale, Illinois 62901, telephone (618) 453-2281. Price: $30.00 hardbound.

ISBN 0-8093-1040-6. (Received by PCES at CAS 24 May 1982, P. H.

Arnaud, Jr., 1982-2).

The ecology of pests. Some Australian case histories. Edited by R. L.

Kitching and R. E. Jones, CSIRO Australia. Printed by CSIRO, Melbourne,

Beaverton, OR 97075, Exclusive Distributor. Price: $12.00 paperbound.

ISBN 0-643-00408-4. (Received by PCES at CAS 28 May 1982, P. H.

Arnaud, Jr., 1982-3).

PAN-PACIFIC ENTOMOLOGIST

July 1982. Vol. 58, No. 3, pp. 202-205

Published 16 December 1983

LIGHT TRAP COLLECTIONS OF THREE INTRODUCED

CONODERUS SPECIES (COLEOPTERA: ELATERIDAE)

IN SOUTHERN CALIFORNIA

M. W. Stone 1

131 Sir Damas Drive, Riverside, California 92507

AND

J. Wilcox 1

7551 Vista Del Sol, Anaheim, California 92807

This paper is a continuation and the final results of trapping studies

involving three introduced elaterids. All belong to the genus Conoderus,

namely exsul (Sharp) the sugarcane wireworm, falli (Lane) the southern

potato wireworm, and amplicollis (Gyllenhal) the Gulf wireworm. All are

destructive to various vegetable and field crops. Previous papers (Stone,

1975, 1976a, b; Stone and Wilcox, 1979a, b) presented data on the distri¬

bution of these pests in California, on preliminary trapping results, and on

life history studies conducted at Riverside.

The traps employed were 15 watt survey type fluorescent black light. One

was located at Riverside adjacent to a bare field formerly planted to citrus

and another in an avocado grove at Olive, Orange County, California 30

miles west of Riverside. The four year totals at Riverside (Table 1) show

that 57% of the sugarcane wireworm adults were trapped during July-August

and in the same period 65% at Olive. Totals for June to September at both

locations were lower but similar. These months usually have the highest

evening temperatures. At Riverside, C. exsul adults were trapped as early

as May 8 in 1978 and as late as November 27 in 1977. Early and late

emergence at Olive occurred on May 12, 1978 and on November 17, 1979.

Of interest was that the four year totals show 4500 more adults being collected

at Olive where the soil is not especially favorable for larval development.

The Gulf wireworm Conoderus amplicollis was recorded as a pest of veg¬

etables in Alabama in 1927 (Cockerham and Deen, 1936) and subsequently

discovered in Los Angeles County, California in 1938. Data regarding its

spread and life history studies have been reported on by Stone and Wilcox

(1979a). In previous trapping studies at Riverside and Olive in 1974-76 only

180 and 94 adults, respectively, were collected the entire season. As shown

in Table 2, low totals were also obtained in the four later years in Riverside

whereas at Olive, except in 1980, there was a substantial increase in numbers

in 1977-79. The earliest record of emergence, June 9, occurred in 1977 and

VOLUME 58, NUMBER 3

203

Table 1. Monthly and yearly totals of Conoderus exsul (Sharp) adults collected at black

light traps, Riverside and Olive, California.

May

June

July

Aug

Sept.

Oct.

Nov.

Total

Riverside

1977

313

899

1124

370

381

3100

1978

529

625

459

341

120

2143

1979

323

392

291

326

1387

1980

227

534

279

242

122

1414

Total

120

1392

2450

2153

1279

648

8044

Olive

1977

179

1106

1788

409

167

3675

1978

117

997

1158

700

649

3724

1979

371

888

1149

790

3261

1980

219

703

592

310

100

1930

Total

132

1766

3855

4229

2158

421

12,590

.02

Grand total

252

3158

6305

6382

3437

1069

20,634

.01

Table 2. Monthly and yearly totals of Conoderus amplicollis (Gyllenhal) adults collected at

black light traps, Riverside and Olive, California.

June

July

Aug.

Sept.

Total

Riverside

1977

1978

1979

1980

Total

193

Olive

1977

125

1978

142

202

1979

144

1980

Total

249

175

512

204

PAN-PACIFIC ENTOMOLOGIST

Table 3. Monthly and yearly totals of Conoderus falli (Lane) adults collected at black light

trap, Riverside, California.

April

May

June

July

Aug.

Sept.

Oct.

Nov.

Total

1977

196

588

602

339

510

2295

1978

413

234

588

334

507

214

2290

1979

281

345

177

557

130

1521

1980

113

186

105

453

333

1280

Total

113

514

783

1707

1218

1856

1187

7386

.01

1979 at both localities and the latest was September 26 at Riverside. It is

interesting that the totals for the four year period show that 83% of the adults

were collected in July-August at both localities. This elaterid species is not

considered to be highly attractive to light.

The southern potato wireworm Conoderus falli is an important pest of

vegetable crops in the southeastern United States (Day et al., 1971). It was

first collected in California in 1963 in Deep Canyon near Palm Desert and

in Riverside in 1966 (Stone, 1976b).

Over the four year period the highest adult catches were obtained in July

and in September (Table 3). August and October totals were exceptionally

high in 1977 but were lower in later years. Catches of adults were made as

early as April 10, 1980 and as late as November 27, 1977. During the six

year period of these studies no C. falli adults were collected in the trap

located in an avocado grove at Olive.

Of the 3 species, C. exsul and C. amplicollis have spread the most rapidly

and both have been found in 15 counties and as far north as Butte County.

Conoderus falli is now present in four counties, namely Riverside, San Diego,

Fresno, and Orange. In the absence of effective new soil insecticides or

fumigants or with the present restrictions on the use of the older highly

effective materials, these three species could become extremely hazardous

to California vegetable and held crops.

Literature Cited

Cockerham, K. L., and O. T. Deen. 1936. Notes on life history, habits and distribution of

Heteroderes laurentii (Guer.). J. Econ. Entomol., 29:288-296.

Day, A., F. P. Cuthbert, Jr., and W. J. Reid. 1971. The southern potato wireworm. Its biology

and economic importance in coastal South Carolina. USDA Tech. Bull., 1443:1-33.

Stone, M. W. 1975. Distribution of four introduced Conoderus species in California (Co-

leoptera: Elateridae). Coleopt. Bull., 29:163-166.

-. 1976a. Notes on the biology of the introduced elaterid Conoderus exsul (Sharp) (Co-

leoptera: Elateridae). Pan-Pac. Entomol., 52:304-310.

VOLUME 58, NUMBER 3

205

-. 1976b. The southern potato wireworm in California (Coleoptera: Elateridae). Coleopt.

Bull., 30:361-363.

-, and J. Wilcox. 1979a. The Gulf wireworm in California (Coleoptera: Elateridae). Pan-

Pac. Entomol., 55:235-238.

-, and-. 1979b. Population build-up of two introduced Conoderus elaterid species

in California (Coleoptera: Elateridae). Coleopt. Bull., 33:473-475.

Footnote

1 Collaborators—USD A, SEA, AR. Boyden Entomological Laboratory. M. W. Stone, de¬

ceased March 28, 1982. J. Wilcox, deceased December 22, 1982.

PUBLICATIONS RECEIVED:

Butterflies of the Rocky Mountain states. Edited by Clifford D. Ferris &

F. Martin Brown. Published by The University of Oklahoma Press, xix, 442

pp., 4 col. pis., 26 figs., 321 maps, & many unnumbered figs. Publication

date: October 1981. Published by The University of Oklahoma Press, 1005

Asp Avenue, Norman, Oklahoma 73019. Price: $35.00 hardbound, $15.95

softbound.

Community ecology of a coral cay. A study of One-Tree Island, Great

Barrier Reef, Australia. By Harold Heatwole, Terence Done & Elizabeth

Cameron. Published in Monographic Biologicae, Vol. 43, 400 pp., 99 figs.,

and 48 tables. Publication date: September 1981. Published by Dr. W. Junk

BV Publishers, P.O. Box 13713, 2501 ES The Hague, The Netherlands.

Price: DF1. 170/US $74.00 hardbound.

Ectoparasites of Hawaiian rodents (Siphonoptera, Anoplura and Acari).

By JoAnn M. Tenorio & M. Lee Goff. Bishop Museum, Special Publication

of the Department of Entomology, 32 pp., 19 figs. Publication date: Decem¬

ber 1980. Published by Bishop Museum Press, P.O. Box 19000-A, Honolulu,

Hawaii 96819. Price: $4.00 softbound.

Scientific and common names of insects and allied forms occurring in

Australia. By P. B. Carne, L. D. Crawford, M. J. Fletcher, I. D. Galloway,

& E. Highley. Commonwealth Scientific and Industrial Research Organi¬

zation, Australia, iii, 95 pp. Publication date: 1980. “Exclusive Distributor,”

ISBS, Inc., P.O. Box 1632, Beaverton, Oregon 97075. Price: $6.00 softbound.

Paul H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park,

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PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 206-215

Published 16 December 1983

NEW AND INTERESTING TRICHOPTERA FROM THE

WESTERN UNITED STATES

D. G. Denning

2016 Donald Drive, Moraga, California 94556

Six new species of western United States Trichoptera are described: Mi-

crasema oregoni, Tinodes schusteri, Farula wigginsi, Hydropsyche ander-

soni, Hydropsyche dorata, and Lepidostoma roemhildi. Several interesting

diagnostic characters of Amiocentrus aspilus Ross, Brachycentrus american-

us (Banks), Brachycentrus occidentalis (Banks) and Pedomoecus sierra Ross

are given. Unless stated otherwise types will be deposited in the California

Academy of Sciences, San Francisco.

Hy dropsy chi dae

Hydropsyche andersoni Denning, new species

The species is a member of the tana-abella species group. Characteristics

of the group are: crest-like profile of tergum 10, short membranous lateral

endothecal processes of the phallic apparatus, contour of segment 9 and the

translucent ovate area of dorsal tergum 9. It is closest to H. vanaca Denning.

Male. — Length 8.5 mm. Wings testaceous, head, thorax, appendages fus¬

cous, spurs prominent. Genitalia (Fig. 1). Segment 9 lateral lobe ovate,

demarcation of terga 9 and 10 indistinct. Tergum 9 translucent area outlined

by dark brown Y-shaped margin. Tergum 10 triangular crest prominent,

ventral margin arcuate, distolateral lobes wide, apex acute (Fig. 1); dorsal

surface with dense short acute setae, apices convergent (Fig. la). Claspers

(inferior appendages) distal segment short, obtuse. Phallic apparatus (Fig.

lb), typical of group, phallotheca sinuate, base large; endothecal processes

directed ventrocephalad, distal spine fuscous and short, directed laterad from

ventral aspect; phallotremal sclerite short, wide, fuscous, short internal scler-

ite directed cephalad; apex elongated, lateral ovoid area translucent, internal

sclerite bears 5 acute dark spines directed laterad, non-pigmented area with

dense spicules. Endophallus not discernible due to dark pigmentation.

Holotype, male. — OREGON: Lane County, H. J. Andrews Experimental

Forest, Willamette National Forest, 25 June 1977, David Voegtlin. Paratype,

male, same data as holotype, deposited in Entomology Department, Oregon

State University, Corvallis. Holotype will be deposited in California Acad¬

emy of Sciences.

This new species is named in honor of Dr. Norman H. Anderson, Oregon

VOLUME 58, NUMBER 3

207

Figs. 1-7. Fig. 1. Hydropsyche andersoni, male genitalia, lateral aspect; la, ninth and tenth

tergites, dorsal aspect; lb, phallic apparatus, lateral aspect. Fig. 2. Hydropsyche dorata, male

genitalia, lateral aspect; 2a, ninth and tenth tergites, dorsal aspect; 2b, phallic apparatus, lateral

aspect. Fig. 3. Tinodes schusteri, male genitalia, lateral aspect; 3a, clasper, ventral aspect; 3b,

phallic apparatus, lateral aspect. Fig. 4. Tinodes sigodana, male genitalia, lateral aspect; 4a,

phallic apparatus, lateral aspect, ae, aedeagus. Fig. 5. Tinodes sigodana, female genitalia, lateral

view, c, cercus; 5a, ventral aspect. Fig. 6. Farula wigginsi, male genitalia, lateral aspect; 6a,

dorsal aspect; 6b, ventral aspect. Fig. 7. Farula wigginsi, female genitalia, ventral aspect.

208

PAN-PACIFIC ENTOMOLOGIST

State University, who has made many noteworthy contributions to Tri-

chopterology.

Hydropsyche dorata Denning, new species

This species is a member of the tana-abella species group and is closely

related to H. tana Ross.

Male.— Length 9 mm. Wings, appendages fulvous; head, thorax fuscous,

setation sparse. Genitalia (Fig. 2). Segment 9 lateral lobes wide, apex obtuse;

from dorsal aspect (Fig. 2a), apices convergent, dark brown margin, median

translucent area subtriangular. Distal segment of clasper densely setose with

whitish setae. Phallic apparatus (Fig. 2b), base stocky, bulbous; endothecal

processes short, sessile except at apex, ventrolateral spur acuminate and

capable of lateral movement; phallotremal sclerite fuscous, short, narrow,

capable of only slight dorsoventral movement, apex bearing a membranous,

ventrally directed process, sessile except apex; slender brown internal spine

distally, apical ventral margin heavily pigmented; semicircular translucent

area at apex with brown extrusible sclerite with row of acute spicules; en-

dophallus narrowed abruptly near base.

Holotype, male. — OREGON: Marion County, Silver Falls State Park, Sil¬

ver Creek, 23 June 1974, P. H. Arnaud, Jr. Paratype, male, same data as

holotype.

Psychomyiidae

Genus Tinodes Stephens

The new species described herein is the twelfth North American species.

These described species are known only from the western United States and

Baja California, Mexico. Tinodes provo Ross and Merkley is widely distrib¬

uted, being known from Baja California to Utah; the remaining species have

an apparent limited distribution.

Tinodes consuetus McLachlan, 1871; California, Oregon.

Tinodes belisa Denning, 1950; Oregon, California.

Tinodes parvula Denning, 1950; California, Arizona.

Tinodes provo Ross and Merkley, 1950; Baja California, Utah, Nevada,

California, Arizona.

Tinodes sigodana Ross, 1950; California.

Tinodes siskiyou Denning, 1951; Oregon, California.

Tinodes cascadia Denning, 1956; California, Oregon.

Tinodes powelli Denning, 1964; Baja California, Utah, California.

Tinodes usilla Denning, 1966; California.

Tinodes gabriella Denning, 1973; California.

Tinodes twila Denning, 1975; California.

Tinodes schusteri Denning, n. sp.; California.

VOLUME 58, NUMBER 3

209

Tinodes schusteri Denning, new species

A member of the siskiyou-sigodana species group that is closely related

to Tinodes sigodana Ross. The new species differs from T. sigodana in the

shape of sternum 8, tergum 10, aedeagus, claspers and its mesal lobes. The

contour of the phallic apparatus and the setal ornamentation of the inter¬

mediate appendages also differ from T. sigodana.

Male. —Length 4.5 mm. Wings fuscous, maxillary palpi slightly longer

than forefemur. Genitalia (Fig. 3). Sternum 9 partially covers sternum 8;

tergum 9 acute distally. Tergum 10 reduced and semi-membranous. Inter¬

mediate appendages with 3 large, 1 small acute spines; apex obtuse and

densely setose; mesal surface concave. Phallic apparatus in repose may be

placed between the intermediate appendages; aedeagus (shaded in Fig. 3,

3b) curved dorsad, apices acute. Clasper basal segment short, almost quad¬

rate, dorsal margin convex, posterior mesal lobe narrow, erect, acute and

large; second lobe coalesced to mesal surface of basal segment, semicircular,

apex obtuse, curved ventrad; lobe is grooved and serves as a guide for phallic

apparatus as suggested by Schmid (1980); apical segment short, linear, apex

with a pair of short acute apices, lateral aspect (Fig. 3), convergent from

ventral aspect (Fig. 3a).

Holotype, male. — CALIFORNIA: Canada del Puerto, Santa Cruz Island,

Santa Barbara County, 20 June 1967, R. O. Schuster. Type deposited in the

Department of Entomology, University of California, Davis. I take pleasure

in naming this species in honor of the collector, Robert O. Schuster, who

has collected many interesting Trichoptera.

Tinodes sigodana Ross

Due to the similarity of T. schusteri to T. sigodana figures of T. sigodana

(Fig. 4) are presented for comparison to figures of T. schusteri (Fig. 3). In

neither figures are the preanal appendages shown.

Male. — Discernible major differences of T. sigodana from T. schusteri are:

heavier setation of intermediate appendages; longer basal segment of clasper,

the absence of the acute dorsal lobe near basal segment and acute prominent

apices of distal segment (Fig. 4). Phallic apparatus and aedeagus (Fig. 4a).

Female. — Genitalia (Fig. 5). Segment 8, terga 9, 10, 11 and cerci shown

(Fig. 5, 5a). General similarity apparent to T. provo (Schmid, 1980), except

the abrupt dorsal position of segment 11.

Male and female collected in Los Angeles County, California, Valley Forge,

San Gabriel Mountains, 11 July 1970, J. A. Honey. Specimens loaned by

Dr. Charles Hogue, Curator, Los Angeles County Museum of Natural His¬

tory. Type locality of T. sigodana Ross is “San Gabriel Mountains, Cali¬

fornia, 29 June, L. J. Milne.”

210

PAN-PACIFIC ENTOMOLOGIST

Limnephilidae

Genus Farula Milne

Members of the genus are known from Washington, Oregon and Califor¬

nia. The species are rare in collections with most known from only a few

specimens. Exceptions are F. malkini and F. honeyi which have been col¬

lected in large numbers. Larvae of two species have been described: F.

malkini and F.jewetti. The new species described herein is the eighth species

in Farula.

Farula rainieri Milne, 1936; Oregon, Washington.

Farula malkini Ross, 1950; Oregon.

Farula davisi Denning, 1958; Oregon.

Farula jewetti Denning, 1958; Oregon.

Farula reapiri Schmid, 1968; Oregon.

Farula honeyi Denning, 1973; California.

Farula petersoni Denning, 1973; California.

Farula wigginsi Denning, n. sp.; California.

In descriptions of all species except F. reapiri Schmid, the various com¬

ponents of the highly specialized male genitalia are designated by a letter (a,

b, etc.). Schmid (1980) recently named these morphological entities which

is followed here, but to avoid confusion with described species the same

lettering system initiated by Ross (1950b) is also used.

Farula wigginsi Denning, new species

Male. — Length 6 mm. Maxillary, labial palpi similar in length, segments

1 and 2 practically identical in length. Spurs 2-4-4, foreleg spurs inconspic¬

uous. Wings light brown. First antennal segment 4 times length of second.

Pronotum with pair of opaque ovate areas, apical portion not pigmented.

Genitalia (Fig. 6). Segment 8 caudal margin difficult to discern covering

about half of segment 9. Margin of segment 9 heavily sclerotized, dark brown.

Internal branch of tergum 10 (a) semi-membranous, light pigmentation,

directed dorsocaudad; external branch (b) undulating in lateral aspect, acu¬

minate, heavily sclerotized, dark brown; inferior branch (d) stout, heavily

sclerotized, apex oblique and projecting caudad; preanal appendage (c) short,

slender, non-pigmented; dorsal inferior branch (f) filamentous, directed cau¬

dad; lateral inferior branch (e) digitate, apex obtuse, curved caudoventral;

ventral inferior branch (cl) abruptly narrowed distally, apex acute. From

dorsal aspect (Fig. 6a), the fused internal branches of segment 10 distally

narrowed (a), apex bearing whitish setae; preanal appendages (c) slender,

directed laterally; external branch tergum 10 (b) base concave, acuminate

and convergent; inferior branch tergum 10 (d) with basal branch directed

slightly caudolaterad, apical branch bent laterad, structure fuscous, heavily

VOLUME 58, NUMBER 3

211

sclerotized. From ventral view (Fig. 6b), phallic apparatus (ae) lightly scler-

otized, trough-like, apical margin concave, aedeagus faintly visible in cleared

male.

Female.— Length 7 mm. Similar in coloration and general characteristics

to male. Ventral aspect of segment 9 (Fig. 7).

Holotype, male. — CALIFORNIA: Marin County, Point Reyes National

Seashore, 25 May 1975, D. G. Denning. Allotype, female, same data as

male.

Named in honor of Dr. G. B. Wiggins, Royal Ontario Museum, Toronto,

in recognition of his elucidation of the biology and immature stages of this

complex genus.

Pedomoecus sierra Ross

Although members of this monotypic genus are known from Alberta,

British Columbia, Washington, Oregon and California, they are rare in col¬

lections. Due to apparent variability in details of the male genitalia (Fig. 8

is presented) differing somewhat from the figures by Ross (1947) and Schmid

(1980). The phallic apparatus consists of a strongly sclerotized acute dorsal

prong, paired processes of 3 acute sclerotized prongs and an internal black

pigmented aedeagus near apex of the phallotheca. The spine shown in Ross’

(1947) figure at the base of the paired 3 branched structure is not present in

males examined from Washington and Oregon. Male used in Figure 8 was

collected at H. J. Andrews Experimental Forest, Willamette National Forest,

Lane County, Oregon, 22 August 1977, by David Voegtlin.

Brachycentridae

Genus Micrasema McLachlan

Currently there are 16 nearctic species in the genus but only five are from

the western region: M. bactro Ross, M. diteris Ross, M. onisca Ross, M.

alexanderi Denning, and M. etra Denning. There are probably several times

that number of undescribed species from the Rocky Mountains to the Pacific

Coastal regions.

Position and shape of lobes and contour of the mesodorsal margin of the

claspers are diagnostic. Existing figures of described species do not account

for variability which may be present. Figures should be shown from these

aspects: lateral, mesal, dorsal and dorsomesal. To avoid confusion in de¬

scriptions, these mesodorsal lobes of the clasper should be marked 1, 2, 3

in figures.

Micrasema oregoni Denning, new species

A member of the bactro species group with the characteristic configuration

of abdominal segments 6, 7, 8 and similar claspers, similar segment 9 and

tergum 10.

212

PAN-PACIFIC ENTOMOLOGIST

Male.— Length 5.5 mm. General coloration fuscous. Tergo-pleural ab¬

dominal segments 6, 7, 8 enclosed by heavily pigmented margin, remainder

opaque. Genitalia (Fig. 9). Segment 9 reduced dorsally to narrow strap.

Preanal appendages trianguloid, base wide and separated on meson. Tergum

10 dorsal aspect (Fig. 9d), narrowed distally, margin bilobed; near base an

elevated ridge with enlarged setose lobe discernible in dorsal or lateral view.

Phallic apparatus (Fig. 9e) contains a blackish furcate structure. Clasper

enlarged distally, mesodorsal margin subdivided into 3 lobes, lateral view

(Fig. 9). Mesal aspect (Fig. 9a), lobe 1 broad, of uniform width; lobe 2 (center

lobe) curved ventrally, acute, narrow; lobe 3 mesal surface appears as a

heavily pigmented ridge. Dorsal aspect (Fig. 9b), lobe 1 wide, triangular;

lobe 2 elongated, narrow, exceeds length of lobes 1, 3; lobe 3 twice length

lobe 1, acute apex. Dorsomesal view (Fig. 9c), lobe 1 wide, arcuate; lobe 2

pear-shaped, apex acute; lobe 3 same width throughout, apex rotundate.

Holotype, male.— OREGON: Lane County, H. J. Andrews Experimental

Forest, Willamette National Forest, Black Light Trap, 16 July 1977, David

Voegtlin. Type deposited in Department of Entomology, Oregon State Uni¬

versity, Corvallis.

Amiocentrus aspilus Ross

Widely distributed in British Columbia and western United States. The

phallic apparatus (Fig. 10) in cleared males includes brown pigmented and

opaque areas, a pair of brown internal sclerites, and the membranous faintly

folded endotheca. Male from Oregon, Lane County, H. J. Andrews Exper¬

imental Forest, Willamette National Forest, 19 June 1977, David Voegtlin.

In collections received by the author males and females of Brachycentrus

americanus and B. occidentalis are frequently misidentified. Therefore ad¬

ditional characters are given with figures pertinent to some of these char¬

acters.

Brachycentrus americanus (Banks)

Male.— Abdominal segments with no mesal lobes; 3-8 sterna fenestrate,

seventh with row of dense brown setae along margin. Lateral margin of

segment 9 with subacute lobe. Preanal appendages coalesced, V-shaped ex¬

cision of distal margin. Tergum 10 divided into a pair of elongate lobes, a

second pair of short branches arise dorsad. Phallic apparatus base dark

brown, ventral margin continues dark pigmentation to near apex, remainder

membranous; internal sclerite dark brown, dorsal branch subacute, ventral

branch apex truncate (Fig. 11), furcate from ventral view.

Female. — Tergum 10 somewhat quadrangular in lateral aspect (Fig. 1 la);

dorsally mesal excision short, narrow, anovaginal plate appears coalesced

VOLUME 58, NUMBER 3

213

Figs. 8-13. Fig. 8. Pedomoecus sierra, phallic apparatus, lateral aspect. Fig. 9. Micrasema

oregoni male genitalia, lateral view; 9a, clasper, mesal aspect; 9b, clasper, direct dorsal view;

9c, clasper, dorso-mesal aspect; 9d, tenth tergum, dorsal aspect; 9e, phallic apparatus, ventral

view. Fig. 10. Amiocentrus aspilus, phallic apparatus, lateral view, ae, internal sclerite shaded.

Fig. 11. Brachycentrus americanus, phallic apparatus, internal sclerite shaded; 11a, female

genitalia, lateral view, tergites 9, 10; lib, tergites 9, 10, dorsal aspect; 11c, bursa copulatrix,

ventral view. Fig. 12. Brachycentrus occidentalis, phallic apparatus, lateral aspect, internal

sclerite shaded; 12a, female genitalia, lateral view, tergites 9, 10; 12b, tergites 9, 10, dorsal

view; 12c, bursa copulatrix, ventral view. Fig. 13. Lepidostoma roemhildi, male genitalia, lateral

view; 13a, maxillary palpus, lateral view; 13b, antennal scape, first, second segments, lateral

view; 13c, antennal scape, first, second segments, dorsal view; 13d, phallic apparatus, lateral,

ventral view.

214

PAN-PACIFIC ENTOMOLOGIST

to tergum (Fig. lib). Bursa copulatrix as in Figure 11c. Male, female from

Box Elder County, Utah, One Mile Creek, 25 August 1978, R. W. Baumann.

Brachycentrus occidentalis (Banks)

Male. — Abdominal sternum 7 with large mesal lobe; the dark brown mar¬

ginal emargination of sternum 5 similar in both species. Segment 9 without

lateral lobe. Preanal appendages separated entire length. Tergum 10 entire,

ventral surface concave, distal margin with narrow mesal excision; base with

no dorsal branches. Base of phallic apparatus dark brown, dorsal and ventral

margins dark brown to apex, membranous area small; internal sclerite

brownish, apex of short dorsal branch ovoid, ventral branch apex acute,

prominent ventral lobe present (Fig. 12).

Female.— Tergum 10 dorsal and ventral margins with irregular lateral

aspect (Fig. 12a), distal margin with short excision; tergites widely separated

in dorsal view (Fig. 12b). Male, female from Wallowa County, Oregon,

Wallowa River, 19 May 1977, R. W. Baumann.

Lepidostomatidae

Lepidostoma roemhildi Denning, new species

This species is related to L. spicata Denning. Major differences from

L. spicata and other described species are present in the maxillary palpi,

claspers and phallic apparatus.

Male .—Length 9-10 mm. General color fulvous. Antennae carinate, seg¬

ments with dark brown marginal setal band. A dense pocket of long, slender,

black scale-like setae at base of forewings, wings without scales. Spurs 2-4-4.

Maxillary palpi one-segmented, 3 times length of first antennal segment,

appressed, slender, lightly pigmented, directed dorsad; mesal surface whitish,

no scales; lateral surface with fuscous setae progressively shorter distally

(Fig. 13a). Antennal scape lateral aspect (Fig. 13b), fuscous, massive, ex¬

panded dorsolaterally, thick whitish scales on mesal surface; first antennal

segment twice length of second, base curved laterally, mesal surface concave

with dense short black scales (Fig. 13c) dorsal view. Genitalia (Fig. 13).'

Tergum 10 confluent on meson, distal margin with rounded emargination

laterally (Fig. 13); dorsal and ventral margins arcuate, lateral surface bearing

minute tuberculate spinules. Clasper short, wide; baso-dorsal lobe digitate,

lateral lobe elongate, parallel to densely setose main structure curved dorsad;

a slender lobe present apically from concave mesal surface (Fig. 13). Phallic

apparatus short, apex rotundate laterally, bilobed ventrally; short pair flat

acuminate parameres closely appressed to phallotheca (Fig. 13d).

This species is named in honor of the collector, Dr. George Roemhild,

Montana State University, Bozeman, who has collected many interesting

Trichoptera.

VOLUME 58, NUMBER 3

215

Holotype, male. — MONTANA: Sanders County, 10 miles NE Thompson

Falls, West Fork Thompson River, 25 July 1975, George Roemhild. Para-

type, male. Glacier County, Montana, Many Glacier Area, small seeps along

Iceberg Lake Trail, 21 July 1979, R. W. Baumann, of Brigham Young Uni¬

versity, Provo, Utah.

Literature Cited

Ross, H. H. 1938. Descriptions of Nearctic caddis flies. Ill. Nat. Hist. Surv. Bull., 21:100—

183.

-. 1947. Descriptions and records of North American Trichoptera, with synoptic notes.

Trans. Am. Entomol. Soc., 73:125-168, pis. 2-8.

-. 1950a. The genus Tinodes in North America. J. Kans. Entomol. Soc., 23:64-67.

-. 1950b. Synoptic notes on some Nearctic limnephilid caddisflies. Amer. Midi. Nat.,

43:410-429.

Schmid, F. 1980. Genera des Trichopteres du Canada et des etats adjacents. The Insects and

Arachnids of Canada, 7:1-296.

PUBLICATIONS RECEIVED:

Wasps of the genus Trypoxylon subgenus Trypargilum in North America

(Hymenoptera: Sphecidae). By Rollin E. Coville. University of California

Publications in Entomology, Vol. 97, vi, 147 pp., 46 figs. Issue date on

review card given as “1 April 1982.” Published by University of California

Press, 2223 Fulton Street, Berkeley, CA 94720, telephone (415) 642-4562.

Price: $13.00 paperbound. ISBN 0-520-09651-7. (Received by PCES at CAS

7 June 1982, P. H. Arnaud, Jr., 1982-4).

Systematics of bees of the genus Eufriesea (Hymenoptera, Apidae). By

Lynn Siri Kimsey. University of California Publications in Entomology,

Vol. 95, ix, 125 pp., 129 figs. Issue date on review card given as “April,

1982.” Published by University of California Press, 2223 Fulton Street,

Berkeley, CA 94720, telephone (415) 642-4562. Price: $11.50 paperbound.

ISBN 0-520-09643-6. (Received by PCES at CAS 19 July 1982, P. H. Ar¬

naud, Jr., 1982-5).

A revision of the subfamily Coelidiinae (Homoptera: Cicadellidae). IV.

Tribe Coelidiini. By M. W. Nielson. Pacific Insects Monograph 38, i, 318

pp., 1103 figs. Publication date on review card given as “26 July 1982.”

Published by Department of Entomology, Bishop Museum. Price: $33.50

paperbound. Order from Bishop Museum Press, P.O. Box 19000-A, Ho¬

nolulu, Hawaii 96819. (Received by PCES at CAS 16 August 1982, P. H.

Arnaud, Jr., 1982-6).

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 216-222

Published 16 December 1983

PSEUDOPYGMEPHORUS ATYPOIDES RACK, NEW SPECIES

(ACARI: PYGMEPHORIDAE) ASSOCIATED WITH THE

FOSSORIAL MYGALOMORPH SPIDER

ATYPOIDES RIVERSI O. P.-CAMBRIDGE

(ARANEAE: ANTRODIAETIDAE) IN CALIFORNIA

Leonard S. Vincent 1

Department of Entomology, University of California, Berkeley 94720

AND

Gisela Rack

Zoologisches Institut und Zoologisches Museum, Universitat Hamburg,

Martin-Luther-King-Platz 3, 2000 Hamburg 13, West Germany

Mites of the family Pygmephoridae are often found in association with

insects and small mammals in soil and litter. Unlike other species of the

pygmephorid genus Pseudopygmephorus Cross, 1965, the host for the new

species of Pseudopygmephorus described here is a ground-loving spider,

Atypoides riversi O. P.-Cambridge (Araneae: Antrodiaetidae). A. riversi has

been recorded only in the coastal and Sierran mountain ranges of California

(Coyle, 1971). After overwintering in the parental burrow, spiderlings emerge

and establish their own permanent burrows. The burrow is enlarged as the

spider grows (Vincent, 1980). Details of the natural history and population

biology of A. riversi are found in Coyle (1971) and Vincent (1980). The

spiders and the mites found in association with them were collected by the

senior author at the University of California Blodgett Forest Research Sta¬

tion, which is located at an elevation of 1350 m. The forest is primarily

composed of Pinus ponderosa Dough ex Loud. The description of the mite

is to be attributed to the second author, Gisela Rack.

Although there are many reports of insect-mite associations (see Mac-

Naulty, 1971), there are few reports of nonparasitic spider-mite associations.

Mites parasitic on spiders have been recorded by Finnegan (1933), Lawrence

(1940) and Shiba(1969) (cited in Krantz, 1978), Parker (1962, 1963, 1965),

Parker and Roberts (1974), Domrow (1975), Cokendolpher et al. (1979),

and Forster and Forster (1973); the latter authors displayed a photograph

of a diplurid spider with parasitic mites on it. Some other brief associations

mentioned in the literature are as follows: Michael (1894) observed mites

(Bdellidae) on the web of the spider Amaurobius ferox (Walckenaer) under

stones. These mites may have been eating insects too small for the spider

to take. Vitzthum (1943) mentioned this observation as the only known case

VOLUME 58, NUMBER 3

217

of paraphagium of spiders and mites. Bristowe (1941) stated that spiders

rarely eat mites; however, Main (1976) briefly mentioned the frequent oc¬

currence of commensal relationships with regard to food between fossorial

spiders and mites. Parker (1963) referred to two species of mites in the

hypopial stage using spiders to disperse.

Methods and Materials

As part of a rearing study on A. riversi, 270 arbitrarily chosen spiders were

removed from their burrows and each was placed in a vial within a cooler

for later transport to the laboratory for measurement (Vincent, 1980). Nine

collections were made during seven different months over a three year period

(Table 1). All spiders were anesthetized before measurement: large spiders

were placed in shell vials suspended in ice; small spiders, which tended to

recover quickly from the ice bath, were anesthetized in a C0 2 chamber. The

carapace and sternum were measured to determine the age of each spider,

and each spider was examined under a dissecting microscope for parasitoids,

pathogens, and Pseudopygmephorus.

Results

Mites were found on 21 female and two adult male spiders (Table 1).

Among the females, mites were found on the cephalothorax, usually clustered

at the fovea, but on occasion also on one or both of the posterior lateral

bases of the cephalic region. They were found only on the area between the

labium and the sternum of male spiders. The number of mites on any

individual spider ranged from one to 15. The spiders were collected and

examined for mites from late March through late September. Mites were

present each time, and in all cases they were adult females.

Discussion

An adventitious dead-end relationship between A. riversi and P. atypoides

is suggested here rather than an established phoretic relationship since the

sedentary habit of A. riversi in general, and the lack of mites on spiderlings,

and the low frequency of mites on adults would seem to preclude successful

dispersal of P. atypoides (see Farish and Axtell, 1971, for a definition of

phoresy). Further, these mites have no apparent morphological adaptations

for a phoretic existence on A. riversi, and they were easily dislodged from

the spider with a camel’s hair brush. Parasitism seems unlikely since P.

atypoides was never found on membraneous areas of the spider, but only

on areas of thick integument, unsuitable for feeding; and there is no clear

evidence of parasitism among the Pygmephoridae in general (Krantz, per¬

sonal communication). Further, mites were not found on or in 10 spider egg

sacs removed from burrows. Commensalism also seems unlikely in that soil

and rejectamenta from spider burrows have never been observed to harbor

218

PAN-PACIFIC ENTOMOLOGIST

Table 1. Number of immature, mature female, and mature male Atypoides riversi examined

for Pseudopygmephorus atypoides.

Date

col¬

lected

Number of spiders collected and examined for mites

No mites found

Mites found

Total

spiders

Immatures Females

Males

Immatures Females

Males

April 77

—

May 77

—

June 77

—

Aug. 77

—

Sep. 77

Mar. 78

—

May 78

—

Sep. 78

—

June 79

—

Total

137

270

* Sub-adult.

mites. Furthermore, mites on an infested spider observed while feeding in

the laboratory did not leave the carapace to share in the food. Of course,

more observations of this type need to be made. Finally, only adult mites

were found on the spider. This indicates that the biology of the mite is not

intimately tied to that of the spider, and that the associated mites originated

somewhere outside of the spider’s burrow. Since A. riversi does not leave

its burrow to forage, the mite must come to it from the surrounding soil or

the burrow entrance.

Perhaps Pseudopygmephorus atypoides is normally associated phoretically

with one of the spider’s food sources, a food source too large for smaller

spiders to capture, since only adult and occasionally subadult spiders were

infested. It is conceivable that P. atypoides abandons its true host and climbs

aboard the feeding spider to avoid mastication or inundation by digestive

enzymes. Male spiders, which normally do not feed (Vincent, unpublished

observations), may have obtained P. atypoides while mating with infested

females.

Pseudopygmephorus atypoides Rack, new species

(Figs. 1-7)

Female.— Length (without gnathosoma) 260-300 ym, holotype 275 ym

(mean of 20 specimens 285 jtm); width 155-185 ym, holotype 155 ym (mean

of 20 specimens 170 jam); yellowish, dorsally and ventrally moderately cov¬

ered with punctations.

Dorsum (Figs. 1, 3). — Propodosoma free, longer than broad, with 2 pairs

of setae. Mediolateral prodorsal setae (pml) short (10-18 ym, holotype 14

VOLUME 58, NUMBER 3

219

jum, mean of 20 specimens 13 jiim), smooth, hidden behind posterior pro-

dorsal setae (pi), scarcely visible. Setae pi very long (127-144 y m, holotype

127 y m, mean of 20 specimens 130 ym), barbed. Sensillus with short pedicel,

round, covered with minute spines. Stigmata elliptical, far distant. Most

setae of hysterosoma strikingly long, barbed. Lengths of setae in ym (n =

20): cl = 140-160, holotype 145, mean 150; c2 = 130-165, holotype 145,

mean 151; dl = 140-160, holotype 145, mean 152; el = 127-140, holotype

130, mean 130; e2 = 115-130, holotype 127, mean 125; fl = 65-80, ho¬

lotype 70, mean 70; f2 = 70-85, holotype 80, mean 80.

Venter (Fig. 2). —Epimeres I and II each with 2 pairs of barbed setae. Setae

la shortest on venter. Apodemes 3 and 4 complete. Setae 3a arising more

closely than 3b. Setae 3a and 3b nearly of same length. Setae 4b longest on

venter. Distance between 4b much greater than between 4a. Setae hi and

h2 nearly of same length. Setae h3 arising in greater distance, longer than

hi and h2; lengths in ym (n = 20): hi = 25-30, holotype 29, mean 27; h2 =

29-35, holotype 30, mean 30; h3 = 37-45, holotype 40, mean 42. Venter

of opisthosoma laterally with fine striations.

Legs. — All legs nearly of same width. Leg I (Figs. 4, 5): Femur with 2

plumose setae and 1 hooked seta. Genu with 4 plumose setae, one of them

extremely long. Tibiotarsus somewhat broader than femur and genu. Claw

1 simple, without thumb, arising from distinct stalklike pedicel. In addition

to barbed setae, tibiotarsus I with 3 solenidia and 6 tactiles. Leg II (Fig. 6):

Femur with extremely long seta. Tibia with small solenidion, tarsus with

longer one, distally with 2 simple claws and large pulvillus. Leg III: Femur

with long seta like in leg II. Tibia with short solenidion. Claws and pulvillus

on tarsus as in leg II. Leg IV (Fig. 7): Trochanter only weakly constricted,

ventrally with 1 barbed seta. Proximal edge of trochanter nearly straight,

not bulbose. Femur with 2 barbed setae, one of them extremely long. Genu

ventrally with 1 short and barbed seta. Tibia proximal with tiny solenidion

and 4 plumose setae, one of them very long. Tarsus with 6 plumose setae,

2 simple claws and distinct pulvillus, shorter than at legs II and III.

Male. — Unknown.

Host and locality. — Female, holotype, and 9 females, paratypes, found on

the labium of an adult male (No. 269) of Atypoides riversi Cambridge, El

Dorado County, California, 13 September 1978, Leonard S. Vincent; 10

females, paratypes, from adult females (No. 276 and No. 277) of the same

species, locality, and date. Two females ex dorsal thoracic fova of Atypoides

riversi, Blodgett Forest Research Station, El Dorado County, California, 26

April 1977, L. S. Vincent.

Deposition of types. — Holotype and 17 paratypes in the California Acad¬

emy of Science, San Francisco. Two paratypes in Zoologisches Institut and

Zoologisches Museum of University Hamburg, D-2000 Hamburg 13, West

Germany.

220

PAN-PACIFIC ENTOMOLOGIST

VOLUME 58, NUMBER 3

221

Remarks.—Pseudopygmephorus atypoides is distinctive in having very

long dorsal setae pi, cl, c2, dl, el, and e2, until now not known in other

members of the genus Pseudopygmephorus. Only in the genus Bakerdania

are two species with such long setae known, B. longisetus (Mahunka, 1964)

and B. damboldti Mahunka, 1972. The new species differs from B. damboldti

in having shorter, and from B. longisetus in having somewhat longer, dorsal

setae. From both it differs in having setae pi and pml arising close together.

The shape and setation of tibiotarsus I and leg IV of P. atypoides also

distinguish it from Bakerdania species with long dorsal setae.

Acknowledgments

We wish to thank Drs. E. I. Schlinger and D. P. Furman, University of

California, Berkeley, Dr. B. J. Kaston, California State University, San Die¬

go, Dr. G. Krantz, Oregon State University, and Mr. I. A. Boussy, University

of California, Davis for reviewing this manuscript. Mr. I. Boussy assisted

in the field, and both I. Boussy and Ms. M. Buegler, University of California,

Berkeley, provided technical assistance.

Literature Cited

Bristowe, W. S. 1941. The comity of spiders. Vol. 2. Ray Society, London, pp. i-xiv, 229-

560, pis. 20-22.

Cokendolpher, J. C., N. V. Horner, and D. T. Jennings. 1979. Crab spiders of north-central

Texas (Araneae: Philodromidae and Thomisidae). J. Kans. Entomol. Soc., 52:723-734.

Coyle, F. A. 1971. Systematics and natural history of the mygalomorph spider genus Antro-

diaetus and related genera. Bull. Mus. Comp. Zool., 141:269-402.

Domrow, R. 1975. Ljunghia Oudemans (Acari: Dermanyssidae), a genus parasitic on mygalo¬

morph spiders. Rec. S. Aust. Mus. (Adelaide), 17:31-39.

Farish, D., and R. Axtell. 1971. Phoresy redefined and examined in Macrocheles muscae-

domestic ( Acarina: Macrochelidae). Acarologia, 13:16-29.

Finnegan, S. 1933. A new species of mite parasitic on the spider Liphistius malayanus Abra¬

ham from Malaya. Proc. Zool. Soc. Lond., 1933:413-417.

Forster, R. R., and L. M. Forster. 1973. New Zealand spiders, an introduction. Collins Brothers

and Company, Auckland and London, 254 pp.

Krantz, G. W. 1978. A manual of acarology. Second edition. Oregon State University Book

Stores, Inc., Corvallis, 509 pp.

Lawrence, R. F. 1940. New larval forms of South African mites from arthropod hosts. Ann.

Natal Mus., 9:401-408.

MacNulty, B. 1971. An introduction to the study of Acari—Insecta associations. Proc. Br.

Entomol. Nat. Hist. Soc., 4:46-70.

Main, B. Y. 1976. Spiders, the Australian naturalists library. W. Collins, Sydney and London,

296 pp.

Figs. 1-7. Pseudopygmephorus atypoides Rack, new species, holotype, female. Fig. 1. Dor¬

sum, Fig. 2. Venter. Fig. 3. Propodosoma with outstretched gnathosoma. Fig. 4. Left leg 1,

dorsal. Fig. 5. Left leg 1, ventral. Fig. 6. Left leg 2, dorsal. Fig. 7. Left leg 4, dorsal view.

222

PAN-PACIFIC ENTOMOLOGIST

Michael, A. D. 1894. The progress and present state of our knowledge of the Acari. J. R.

Microsc. Soc. Lond., 1894:18-33.

Parker, J. R. 1962. Ectoparasitic mites on spiders. Entomol. Mon. Mag., 98:64.

-. 1963. Ectoparasitic mites on spiders. Flatford Mill Spider Group Bull., 17:2-4.

-. 1965. More records of mites as ectoparasites on spiders. Brit. Spider Group Bull.,

25:6.

-, and M. S. Roberts. 1974. Internal and external parasites of the spider Pardosa hortensis

(Thorell) (Araneae: Lycosidae). Bull. Br. Arachnol. Soc., 3:82-84.

Shiba, M. 1969. Taxonomic investigations on free-living mites in the subalpine forest of

Shinga Heights IPB Area. II. Prostigmata. Bull. Nat. Sci. Mus. Tokyo, 12:65-115.

Vincent, L. S. 1980. The population biology of Atypoides riversi O. P.-Cambridge, a fossorial

mygalomorph spider (Araneae: Antrodiaetidae). Ph.D. Dissertation, University of Cal¬

ifornia, Berkeley, 142 pp.

Vitzthum, H. Graf. 1943. Acarina. In Bronn’s Klassen und Ordnungen des Tierreichs, 5(4)5:

1 - 1011 .

Footnote

1 Present address: Department of Biology, Georgia Southern College, Statesboro, Georgia

30460.

PUBLICATIONS RECEIVED:

Ecology of marine parasites. By Klaus Rohde. University of Queensland

Press, St. Lucia, Queensland, xvi, 245 pp., 6 colored pis., 71 figs. Publication

date on review card given as “October 14, 1982.” Price: $25.00 hardcover,

$12.95 paper. Order from University of Queensland Press, 5 South Union

Street, Lawrence, MA 01843, telephone (617) 685-3306. ISBN 0-7022-

1660-7. (Received by PCES at CAS 12 October 1982, P. H. Arnaud, Jr.,

1982-7).

Insect neurohormones. By Marie Raabe. Translated from French by Nis-

sim Marshall. Illustrated by Daisy Chervin. Plenum Press, New York, xiv,

poration, 233 Spring Street, New York, NY 10013. Price: hardbound $42.50

American price; prices 20% higher outside the U.S. and Canada. ISBN 0-306-

40782-5. (Received by PCES at CAS 29 November 1982, P. H. Arnaud, Jr.,

1982-9).

PAN-PACIFIC ENTOMOLOGIST

July 1982. Vol. 58. No. 3. pp. 223-230

Published 16 December 1983

PREY RECORDS FOR SOME NORTH AMERICAN SPIDER

WASPS (HYMENOPTERA: POMPILIDAE)

Marius S. Wasbauer

Laboratory Services/Entomology, California Department of

Food and Agriculture, Sacramento 95814

In the course of recent investigations on the Pompilidae of California, I

have had the opportunity to examine material in several western collections

of these wasps. Some of the specimens were pinned with or otherwise as¬

sociated with their spider prey. The records are presented here because, in

general, the biology of the North American Pompilidae is not well known

and because prey was previously unknown for some of the species. Deposito¬

ries for wasps and prey are as follows: California Academy of Sciences, San

Francisco (CAS); California Department of Food and Agriculture, Sacra¬

mento (CDFA); California Insect Survey, University of California, Berkeley

(CIS); Oregon State University, Corvallis (OSU); University of California,

Davis (UCD). Dr. Willis J. Gertsch, Portal, Arizona, kindly identified the

spiders, except for Anyphaena pacifica (Banks) which was determined by

Mr. L. Vincent, University of California, Berkeley. Dr. Gertsch also provided

comments on some of the spiders which have been included where appro¬

priate.

Cryptocheilus severini Banks

Prey.—Lycosa coloradensis Banks (Lycosidae). Female.

Collection data.— Arizona, Cochise Co., T17S-R31E-S3, October 20, 1958.

The specimen carries two additional labels—“Larrea” and “273” (CIS).

This represents the first prey record for C. severini and the first record

of Lycosa color adensis as prey of any North American species of Crypto¬

cheilus, although species of Lycosa are the only prey thus far recorded for

this genus in our fauna.

Priocnemis ( Priocnemis ) cornica (Say)

1. Prey.—Agroeca sp. (Clubionidae). Immature.

Collection data.— California, Sacramento Co., Sacramento (river levee

near Wheeler’s Landing), June 6, 1965 (M. Wasbauer, CDFA).

2. Prey.—Pardosa sp. (Lycosidae). Immature.

Collection data. — Missouri, Columbia, October 27, 1966 (F. D. Parker,

CDFA).

224

PAN-PACIFIC ENTOMOLOGIST

Host spiders of six families have been recorded for this rather nonselective

wasp (Evans and Yoshimoto, 1962:96; Kurczewski and Kurczewski, 1968:

3-4). Species of Pardosa have been reported as prey several times. Agroeca

is the third genus of Clubionidae known to be utilized.

Auplopus architectus metallicus (Banks)

Prey. —Metaphidippus aeneolus (Curtis) (Salticidae). Female.

Collection data. — California, Contra Costa Co., Orinda Village, San Pablo

Ridge below Eureka Peak, 1000-2000 feet, Oak-chaparral zone, May 25,

1972 (E. I. Schlinger, CIS).

Prey previously recorded for this subspecies include members of the Sal¬

ticidae and Clubionidae (Hurd and Wasbauer, 1956:169; Kurczewski, 1961:

23-24; Evans and Yoshimoto, 1962:109).

Auplopus nigrellus (Banks)

Prey. — Chiracanthium sp. (Clubionidae). Immature.

Collection data. — California, Stanislaus Co., La Grange, October 30, 1979

(R. P. Allen, CDFA).

This species of Auplopus occurs across the United States. It is characteristic

of woodland situations and has been recorded as provisioning with Saltici¬

dae, Anyphaenidae and Clubionidae in the eastern portion of its range.

Previous records of Clubionidae are Trachelas sp. prob. tranquillus (Hentz),

immature (Krombein, 1955:15) and Clubiona abboti L. Koch, immature

female (Kurczewski and Kurczewski, 1968a:369).

Ageniella ( Priophanes ) arizonica concolor Townes

Prey. — Oxyopes sp. near tridens Brady (Oxyopidae). Female.

Collection data. —Oklahoma, Lake Texoma, 2 miles east of Willis, July,

1965 (R. M. Bohart, UCD).

There are no previous prey records for this species of Ageniella, but the

related A. arcuata (Banks) and A. fuscipennis Townes have both been taken

with Oxyopes salticus Hentz (Evans and Yoshimoto, 1962:107; Hurd and

Wasbauer, 1956:169).

Aporus {Aporus) luxus (Banks)

A male specimen in the CIS collection taken in California at Point Reyes

National Seashore on January 18, 1976, by Paul Rude is pinned with a

cocoon and bears a label which reads “Ex. Apostichus (!) nest. Emerged 17-

II-1976.” The spider in this case is probably Aptostichus stanfor dianus Smith

(Ctenizidae) which is found in dune areas along the central California coast

and is also taken as prey by Aporus (Plectraporus) hirsutus (Banks) (Williams,

1928:136).

VOLUME 58, NUMBER 3

225

This record is of significance since it is the first indication of prey for any

North American species in the subgenus Aporus.

Agenioideus ( Agenioideus ) humilis (Cresson)

Prey.—Araneus bispinosus (Keyserling) (Araneidae). Immature female.

Collection data.— California, Marin Co., Mill Valley, October 20, 1960

(E. S. Ross, CAS).

Prey thus far recorded for A. humilis have all been araneid spiders (Evans

and Yoshimoto, 1962:88). Eberhard (1970:243) provided a record for Arane-

us cornutus Clerck in Florida.

Sericopompilus neotropicalis (Cameron)

Prey.—Misumenops sp. (Thomisidae). Female.

Collection data. — Arizona, Cochise Co., 2.5 miles southeast of Portal, July

31, 1960 (E. G. Linsley, CIS).

Prey have not been recorded previously for this common spider wasp,

although eleven genera of spiders in five families are known to be taken as

prey by the related S. apicalis (Say) of the eastern United States (Evans and

Yoshimoto, 1962:90).

Episyron quinquenotatus hurdi Evans

1. Prey.—Argiope trifasciata (Forskal) (Araneidae). Immature female.

Collection data.— Nevada, Lincoln Co., Caliente, July 21, 1958 (R. C.

Bechtel, UCD).

2. Prey.—Metepeira arizonica Chamberlin and Ivie (Araneidae). Female.

Collection data. — California, Imperial Co., Black Mountain, 4-5 miles

southeast Highway 78, 1300-1500 feet, March 28, 1978 (E. Fisher, CDFA).

Although there is some information on the prey and nesting behavior of

the nominate subspecies of Episyron quinquenotatus in the eastern U.S.

(Evans and Yoshimoto, 1962:91-92; Kurczewski and Kurczewski, 1968:9-

12, 1968a:370, 1973:67) there are no previous records of prey for the sub¬

species hurdi.

Tachypompilus ferrugineus burrus (Cresson)

Prey. — Cupiennius salei (Keyserling) (Ctenidae). Female.

Collection data. — Mexico, Veracruz, La Playa Escondida, 16 km north

Sontecomapan, Sierra de las Tuxtlas, August 2-9, 1980 (L. G. Bezark, C.

Y. Kitayama, CDFA).

Although the Nearctic subspecies of T. ferrugineus show a preference for

large lycosid spiders, Dolomedes (Pisauridae) has been recorded as prey

(Evans and Yoshimoto, 1962:87). Thus, it is not surprising that Ctenidae

would be taken by a Neotropical subspecies since they are wandering spiders

226

PAN-PACIFIC ENTOMOLOGIST

which hunt along the ground and over foliage, somewhat in the manner of

Lycosidae. According to Gertsch (in litt.) “The spider is common in Mexico,

Central America, West Indies and Brazil.”

Tachypompilus ferrugineus ferrugineus (Say)

Prey.—Lycosa helluo Walckenaer (Lycosidae). Immature female.

Collection data.— Kansas, Clearwater, August 9, 1971 (CDFA).

This species has been reported previously taking L. helluo as prey in

Kansas (Evans and Yoshimoto, 1962:87).

Anoplius ( Pompilinus ) insolens (Banks)

Prey. — Tibellus gertschi Chamberlin and Ivie (Philodromidae). Female.

Collection data.—\JXdh, Cache Co., Smithfield, July 18, 1958 (R. W. Thorp,

WJG, CIS).

There is one previous record of prey for this species, a female of the salticid

spider, Maevia vittata (Hentz) which Evans (1951:306) collected at East

Hartford, Connecticut.

Gertsch states (in litt.) that philodromid crab spiders of the genus Tibellus

are elongate, grassland species which frequently cling to stems. T. gertschi

is not a commonly encountered species.

Anoplius {Pompilinus) marginatus Complex

Prey.—Arctosa sp. (Lycosidae). Female.

Collection data.—North Dakota, Case Co., 10 miles east of Enderlin,

August 18, 1964 (J. R. Powers, CIS).

Females of A. {P.) bequaerti and townesi are not separable at present from

those of marginatus. Thus, the record given here may refer to any of the

three.

There are now many host records available for this complex (Evans and

Yoshimoto, 1962:79; Kurczewski and Kurczewski, 1968:19-20, 1968a:373-

374) and Arctosa sp. has been recorded previously.

Anoplius (Pompilinus) splendens (Dreisbach)

Prey. — Phidippus cardinalis (Hentz) (Salticidae). Male.

Collection data.— Missouri, Columbia, July 30, 1967 (F. D. Parker, CDFA).

Due mainly to the efforts of Frank and Edmund Kurczewski (1968, 1968 a,

1973), there have now accumulated a number of prey records for this species

which is common in the eastern United States. The range of prey is unusually

wide and encompasses the following families: Agelenidae, Amaurobiidae,

Araneidae, Clubionidae, Gnaphosidae, Lycosidae, Pisauridae, Salticidae and

Thomisidae.

In the Salticidae, the genera Habronattus, Pellenes, Marpissa and Phidip¬

pus are represented.

VOLUME 58, NUMBER 3

227

Anoplius {Anoplius) dreisbachi Evans

Prey.—Alopecosa kochi (Keyserling) (Lycosidae). Female.

Collection data.—\J\ah, Salt Lake City, October 15, 1914 (L. P. Rock-

wood, OSU).

There are no previous prey records for this wasp which is relatively com¬

mon in the west but Powell (1958:55) reports a female of Alopecosa gertschi

Schenkel as prey of the closely related Anoplius (A) toluca (Cameron), near

Pittsburg, Contra Costa County, California.

Anoplius {Anoplius) imbellis Banks

Prey.—Schizocosa mccooki (Montgomery) (Lycosidae). Immature.

Collection data.—\J\ah, Salt Lake City, October 14, 1914 (L. P. Rock-

wood, OSU).

Wasbauer (1957) studied the behavior of this wasp in two California

localities. In both areas, the prey was Pardosa ramulosa McCook. In pre¬

senting a record for an undetermined species of Pardosa as prey of A. imbellis,

Wasbauer and Powell (1962:399) suggested a narrow range of prey preference

because the wasp seems to show a restrictive hunting habitat selection.

Subsequent prey records (Evans and Yoshimoto, 1962:85; Kurczewski and

Kurczewski, 1968:22, 1973:72) although all Lycosidae, indicate a somewhat

wider range of preference than was suspected initially.

Anoplius {Anoplius) ithaca (Banks)

Prey.—Pardosa lowriei Kronestedt (Lycosidae). Female.

Collection data. — California, Placer Co., Chinquapin, 3 miles northeast

Tahoe City, June 25 to 29, 1979 (P. Adams, CDFA).

I am informed by Ms. Adams that the wasp was dragging the spider along

the ground between rocks on a boulder-strewn section of stream bank.

As far as existing knowledge goes, Anoplius ithaca seems to be restricted

in its prey preferences to members of the family Lycosidae. Only two genera

have been recorded, Arctosa and Pardosa (Evans and Yoshimoto, 1955:19,

1962:83; Krombein, 1956:42; Kurczewski, 1962:89, 1975:148; Kurczewski

and Kurczewski, 1968:22, 1973:72). Evans and Yoshimoto report an im¬

mature specimen of Lycosa sp. being fed upon by a female of ithaca.

Pompilus {Ammopshex) luctuosus luctuosus Cresson

Prey. —Anyphaena pacifica (Banks) (Anyphaenidae). Female.

Collection data. — California, Marin Co., Point Reyes National Seashore,

North Beach, June 2, 1979 (L. Vincent, CIS). Two wasps were taken at this

locality and date. Both were transporting adult females of A. pacifica.

There have been no records of prey previously reported for Pompilus l.

luctuosus. Anyphaenid spiders are not commonly recorded as prey of Pom-

228

PAN-PACIFIC ENTOMOLOGIST

pilidae. There are records of species of Anyphaena as prey of the mud-nest

building genus Auplopus and the Anoplius marginatus complex and of the

related spider, Aysha gracilis (Hentz) preyed upon by species of Sericopom-

pilus, Priocnemis and Auplopus (Evans and Yoshimoto, 1962:116; Kur-

czewski and Kurczewski, 1968:4, 6).

Pompilus (Archnospila ) arctus Cresson

1. Prey. — Gnaphosa muscorum (L. Koch) (Gnaphosidae). Female.

Collection data.— California, Nevada Co., Sagehen Creek, July 11, 1975

(B. Villegas, UCD).

2. Prey. — Cybaeus sp. (Agelenidae). Immature.

Collection data. — California, Contra Costa Co., Orinda Village, San Pablo

Ridge below Eureka Peak, 1000-2000 feet, Oak-chaparral zone, July 15,

1969 (E. I. Schlinger, CIS). Two wasps were taken at this locality and date,

both transporting immatures of Cybaeus.

Although Evans and Yoshimoto (1962:86) report the use of Gnaphosidae

(Orodrassus ) as prey of this pompilid, the genus Gnaphosa has not been

reported previously. Likewise, the use of Agelenidae has not been reported.

In fact, the genus Cybaeus was hitherto unknown as prey of any North

American spider wasp.

Pompilus ( Arachnospila ) fumipennis eureka (Banks)

1. Prey.—Lycosa sp. (Lycosidae). Immature.

Collection data.— California, Tuolumne Co., Sonora Pass, August 23, 1967

(R. W. Thorp, UCD).

2. Prey.—Alopecosa kochi (Keyserling) (Lycosidae). Female.

Collection data. — California, Areata, dunes, June 3, 1976 (D. M. Gordon,

CDFA).

All prey records to date are for Lycosidae. Wasbauer and Powell (1962:

400) list Alopecosa kochi (reported as Tarentula kochi) as prey from two

California localities and Evans (1951:268) reports Lycosa sp. from Cloud-

croft, New Mexico.

Pompilus {Arachnospila) scelestus Cresson

1. Prey.—Schizocosa pacifica (Banks) (Lycosidae). Immature male.

Collection data.— California, Santa Barbara Co., Santa Cruz Island, La

Cascada, May 8, 1968 (D. S. Homing, UCD).

2. Prey.—Schizocosa pacifica (Banks). Immature female.

Collection data. — California, Santa Barbara Co., Santa Cruz Island, Ca¬

nada del Medio, May 27, 1969 (J. E. Slansky, UCD).

This species is known to take spiders of three families: Lycosidae, Pi-

sauridae and Salticidae. Lycosid prey reported previously are Lycosa gulosa

VOLUME 58, NUMBER 3

229

Walckenaer, L. frondicola Emerton (Evans, 1951:264, 1970:483) and Geo-

lycosa raphaelana (Chamberlin) (Gwynne, 1979:685).

Aporinellus completus Banks

Prey. —Pellenes sp. (Salticidae). Female.

Collection data. — California, San Mateo Co., six miles northeast of Gazos

Creek Station, May 14, 1967 (J. Doyen, CIS).

There are several records of this species utilizing salticid spiders as prey.

Evans (1959:76) gives Pellenes oregonensis Peckham (female) at Sierraville,

California. Kurczewski and Kurczewski (1968a:377) give two records of

Pellenes viridipes (Hentz) at Presque Isle State Park, Pennsylvania.

Literature Cited

Eberhard, W. 1970. The predatory behavior of two wasps, Agenioideus humilis (Pompilidae)

and Sceliphron caementarium (Sphecidae), on the orb weaving spider Araneus cornutus

(Araneidae). Psyche, 77:243-251.

Evans, H. E. 1951. A taxonomic study of the Nearctic spider wasps belonging to the tribe

Pompilini (Hymenoptera: Pompilidae). Part 2. Am. Entomol. Soc. Trans., 76:207-361.

-. 1951a. A taxonomic study of the Nearctic spider wasps belonging to the tribe Pompilini

(Hymenoptera: Pompilidae). Part 3. Am. Entomol. Soc. Trans., 77:203-340.

-. 1959. Prey records for some midwestem and southwestern spider wasps (Hymenop¬

tera: Pompilidae). J. Kans. Entomol. Soc., 32:75-76.

-. 1964. Notes on the prey and nesting behavior of some solitary wasps of Mexico and

southwestern United States. J. Kans. Entomol. Soc., 37:302-307.

-. 1970. Ecological-behavior studies of the wasps of Jackson Hole, Wyoming. Bull. Mus.

Comp. Zool., 140:451-511.

-, and C. M. Yoshimoto. 1955. An annotated list of pompilid wasps taken at Blackjack

Creek, Pottawatomie County, Kansas (Hymenoptera). J. Kans. Entomol. Soc., 28:

16-19.

-, and-. 1962. The ecology and nesting behavior of the Pompilidae (Hymenop¬

tera) of the northeastern United States. Entomol. Soc. Am. Misc. Publ., 3:67-119.

Gwynne, D. T. 1979. Nesting biology of the spider wasps (Hymenoptera: Pompilidae) which

prey on burrowing wolf spiders (Araneae: Lycosidae, Geolycosa ). J. Nat. Hist., 13:681—

692.

Hurd, P. D., Jr., and M. S. Wasbauer. 1956. New host records for North American spider

wasps (Hymenoptera: Pompilidae). J. Kans. Entomol. Soc., 29:168-169.

Krombein, K. V. 1955. Miscellaneous prey records of solitary wasps. I. (Hymenoptera: Acu-

leata). Bull. Brooklyn Entomol. Soc., 50:13-17.

-. 1956. Miscellaneous prey records of solitary wasps. II. (Hymenoptera: Aculeata). Bull.

Brooklyn Entomol. Soc., 51:42-44.

Kurczewski, F. E. 1961. Some observations and prey records of Pompilidae (Hymenoptera)

from northeastern United States. Bull. Brooklyn Entomol. Soc., 56:23-24.

-. 1962. Observations, including new prey records of some Nearctic Pompilidae (Hy¬

menoptera). Bull. Brooklyn Entomol. Soc., 57:85-90.

-. 1975. Host records for some species of Pompilidae from southwestern United States

and Mexico (Hymenoptera). Pan-Pac. Entomol., 51:147-151.

-, and E. J. Kurczewski. 1968. Host records for some North American Pompilidae

230

PAN-PACIFIC ENTOMOLOGIST

(Hymenoptera) with a discussion of factors in prey selection. J. Kans. Entomol. Soc.,

41:1-33.

-, and-. 1968a. Host records for some North American Pompilidae (Hymenop¬

tera). First Supplement. J. Kans. Entomol. Soc., 41:367-382.

-, and-. 1973. Host records for some North American Pompilidae (Hymenoptera).

Third Supplement. Tribe Pompilini. J. Kans. Entomol. Soc., 46:65-81.

Powell, J. A. 1958. Biological notes on the burrow and prey of Anoplius ( Anoplius) ventralis

tarsatus (Banks) (Hymenoptera: Pompilidae). Pan-Pac. Entomol., 34:53-56.

Wasbauer, M. S. 1957. A biological study of Anoplius ( Anoplius ) imbellus Banks (Hymenop¬

tera: Pompilidae). Wasmann J. Biol., 15:81-97.

-, and J. A. Powell. 1962. Host records for some North American spider wasps with

notes on prey selection (Hymenoptera: Pompilidae). J. Kans. Entomol. Soc., 35:393—

401.

Williams, F. X. 1928. Studies in tropical wasps—their hosts and associates (with descriptions

of new species). Hawaii. Sugar Plant. Assoc. Exp. Sta. Bull. (Entomol. ser.), 19:1-179.

PUBLICATIONS RECEIVED:

South American species of the subgenus Anisotarsus Chaudoir (genus

Notiobia Perty:Carabidae:Coleoptera). Part I: Taxonomy and natural his¬

tory. By Gerald R. Noonan. Milwaukee Public Museum, Contributions in

Biology and Geology, Number 44, 84 pp., 70 figs. Publication date on cover

(confirmed by Mary Garity, Editor) cited as “November 2, 1981.” ISBN

0-89326-071-1. Price: $5.75 paper.

South American species of the subgenus Anisotarsus Chaudoir (genus

Notiobia Perty:Carabidae:Coleoptera). Part II: Evolution and biogeography.

By Gerald R. Noonan. Milwaukee Public Museum, Contributions in Biology

and Geology, Number 45, 117 pp., 21 figs. Publication date on cover (con¬

firmed by Mary Garity, Editor) cited as “December 1, 1981.” ISBN 0-89326-

072-X. Price: $6.75 paper,

Order from Publications Section, Milwaukee Public Museum, 800 West

Wells Street, Milwaukee, WI 53233, Telephone (414) 278-2700. Postage and

handling: $1.00 for first publication, 250 for each additional publication on

the same order. (Received by PCES at CAS 8 November 1982, P. H. Arnaud,

Jr., 1982-8).

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 231-235

Published 16 December 1983

BEHAVIOR OF OSMIA (NOTHOSMIA) MARGIN AT A MICHENER

IN THE NEST (HYMENOPTERA: M EG A CIIILI DA E)

F. D. Parker and V. J. Tepedino

Bee Biology and Systematics Laboratory,

Agricultural Research, Science and Education Administration,

USDA, Utah State University, UMC 53, Logan 84322

Osmia ( Nothosmia ) marginata Michener is a relatively uncommon, mega-

chilid bee restricted to the southwestern deserts of Arizona, California, Ne¬

vada and Utah (Parker and Bohart, 1966; Parker and Tepedino, in prep.).

Individuals have been recorded visiting a variety of plants in the spring and

the species appears to be polylectic (Hurd, 1979). Like other members of

the subgenus Nothosmia, O. marginata uses pre-existing holes in which to

nest. The species also nests readily in artificial domiciles (Parker and Bohart,

1966), and we took advantage of this behavior to observe the within-nest

activities of adult females in the greenhouse. Specifically we report on prep¬

aration of the cell, provisioning, egg-laying and construction of the cell

partition.

Methods

Individuals of O. marginata were obtained from elderberry trap-nests that

had been set out near Joshua Tree National Monument, Riverside County,

California, the previous year. Upon eclosion in late April, males and females

were released into a greenhouse (6 x 8 m) that contained species of Coreopis,

Papaver, Borago and Phacelia as pollen and nectar sources and Oenothera

as a source of leaf material. The greenhouse also contained a 1 x 1 x 2 m

observation box made of plywood with holes drilled in one wall. Elderberry

(Sambucus) twigs were inserted into these holes as nesting material. A 6 mm

diameter hole was drilled in each twig to a depth of about 23 cm. A transverse

cut was made halfway through each twig 20-30 mm below the entrance hole

and then continued lengthwise to the end of the twig (Fig. 1). This section

was then removed to expose the bore of the hole and replaced with a glass

plate which was taped to the twig. The behavior of bees using these nests

could be clearly observed and was recorded with a digital clock and tape

recorder.

Results

Detailed observations on two nesting females were conducted from 1015—

1520 hrs on 4 June about five weeks after the bees had been released. In the

232

PAN-PACIFIC ENTOMOLOGIST

observed nests, Bee 1 was provisioning her fifth cell and Bee 2 her second

cell. The behavior of nesting females is described in chronological sequence,

from preparation of the new cell to its completion.

Preparation of the new cell. — Complete preparation of a new cell was

observed for both bees, and was the same except for one important detail.

Upon completion of the partition of the cell below, Bee 2 appeared to roughly

equate the length of the cell to be constructed with the length of her body.

Facing outward, and with the tip of the abdomen close to the previously

finished partition, she applied masticated leaf material to the cell walls with

her mandibles and forelegs to form a raised rim or threshold around the

entire circumference of the hole. This threshold later served as the foundation

of the partition that sealed the cell. Bee 1 did not build a threshold at this

time but proceeded directly to the next phase in which the walls of the new

cell were partially lined with leaf pulp. The females made numerous trips

for leaf material and always entered the nest head-first carrying a ball of leaf

pulp in the mandibles. Before application to the walls, the ball was transferred

to the forelegs and chewed and flattened with the mandibles. Perhaps a

salivary secretion was introduced as well. This pulp was then applied to the

walls with mandibles and forelegs, beginning at the posterior end of the cell.

After each deposition of leaf pulp, she applied a salivary secretion to both

leaf material and bare pithy walls with her mouthparts. Both bees applied

their mouthparts to areas of the cell wall that were quite distant from the

area of leaf application. The secretion was clear and could not be discerned

when it dried (Fig. 2). Neither female applied leaf material to the glass wall

but did fill in spaces between glass and twig with leaf pulp.

When lining and shellacking of the cell walls was complete, Bee 2 im¬

mediately began to provision the cell since she had already constructed her

threshold. In contrast, Bee 1 constructed the threshold after lining the cell

and before provisioning. Her threshold was also more substantial and was

constructed differently from that of Bee 2. The partial partition in the nest

of Bee 2 was made by assuming a C-shape and pressing the abdomen against

the leaf ball while it was held against the cell wall and worked by mandibles

and forelegs into a flattened disc.

Twenty-four leaf collecting trips (2 hr 38 min) were required by Bee 1 to

complete the cell preparation process. Average duration of time periods spent

in the nest was 3 min 18 sec ± 1 min 32 sec (range 40 sec-7 min). Leaf¬

collecting trips averaged 3 min 36 sec ± 3 min 22 sec (range 1 min 15 sec-

17 min 35 sec).

Nectar and pollen deposition. — Females collected both nectar and pollen

on almost all foraging trips. A returning female entered the nest head first

and disgorged a nectar droplet upon the provision; this droplet was used to

wet the dry pollen deposited on the previous trip and to work it into the

provision with mandibles and forelegs. She then backed out of the nest,

VOLUME 58, NUMBER 3

233

Figs. 1-4. Fig. 1. Experimental nest constructed from an elderberry stem and fitted with a

glass plate. Fig. 2. A female Osmia marginata applying the salivary secretion to bare cell walls.

The closing partition, pollen ball and egg in the previous cell are clearly visible. Fig. 3. A female

Osmia marginata scraping pollen from the ventral abdomen onto the provision. Note the light-

colored layer of unincorporated pollen on the provision. Fig. 4. A female in the act of laying

the egg (arrow).

turned around and re-entered, abdomen first, and continued backwards to

the provision where the pollen in the abdominal scopa was kicked onto the

provision with the hind legs (Fig. 3). The unloading of pollen from the scopa

took only a few seconds; while unloading pollen, the female maintained

herself in rigid position supported by fore and mid legs with the antennae

stretched out rigidly in front. As she left the nest after unloading, she con¬

tinued to scrape the abdominal venter with the hind legs.

The completed provision typically measured 7-8 mm in length and 4 mm

wide. The bottom and sides conformed to the shape of the cell. The provision

was usually oval-shaped and somewhat moist or pasty.

A complete cell provisioning sequence was recorded for Bee 2. Twenty-

six foraging trips were made to provision the cell completely over a period

of slightly less than five hours. Nectar and pollen were collected on each

trip. Average duration of foraging trips was 8 min 38 sec ± 2 min 58 sec

(range 4 min 25 sec-13 min 25 sec). Little time was used to deposit nectar

234

PAN-PACIFIC ENTOMOLOGIST

and pollen in the nest; deposition time averaged only 57 sec ±12 sec (range

25-85 sec). Except for two in-nest resting periods of roughly 5 and 17 min¬

utes, this female worked constantly to complete provisioning.

Egg-laying. —Both bees were observed to lay eggs. Egg-laying in both cases

immediately followed deposition of the final nectar and pollen load. Bee 1

followed the usual sequence of nectar and then pollen deposition and then

turned around outside the nest and re-entered head first to incorporate the

dry pollen into the provision. Bee 2, however, deviated from this sequence;

she backed into the nest initially and deposited pollen first and then turned

around outside, re-entered, and incorporated her nectar load while molding

the provision. Bee 2 thus accomplished the final incorporation of pollen and

nectar into the provision with one less trip to the outside to turn around

than did Bee 1. This was the only time she exhibited this behavior.

In egg-laying the female backed down the burrow and worked the surface

of the provision with the tip of the abdomen for a few seconds before the

egg appeared. The female remained braced against the side walls for 90-120

sec with the abdomen curved slightly upward as the egg was extruded (Fig.

4). She then pulled herself away, and the egg remained with its posterior tip

stuck in the provision. The egg was arched with the anterior end bent towards,

but not touching, the provision. The female then moved away and groomed

herself for a brief period.

Construction of the cell partition. — Closing of the cell partition began im¬

mediately after egg-laying. The working of leaf material was as described

previously. The sides of the threshold were gradually thickened and extended

and the opening covered by flattened leaf pulp. The entire process took about

30 min and required only seven leaf-foraging trips.

Discussion

Certain aspects of the nesting behavior of Osmia marginata agree with

observations reported for other members of the subgenus Nothosmia. All

species for which there is information utilize pre-existing burrows and con¬

struct cell partitions and closing plugs of leaf material ( O. cordata Robertson:

Rau, 1937; O. albiventris Cresson: Medler, 1967; O. pumila Cresson: Krom-

bein, 1967; Medler, 1967). The provisions of all species also appear to be

somewhat moist and liquid.

The use of leaf material to partially line the cell walls and the construction

of a threshold appears variable among species. Medler (1967) reported that

O. albiventris lined the walls with leaf paste in one nest but not another, and

O. cordata (Rau, 1937) and O. pumila (Krombein, 1967; Medler, 1967) do

not appear to line the walls at all. In the two O. marginata nests observed

cells were only partially lined with leaf paste. In addition to O. marginata

only O. pumila has been reported to construct a threshold (Krombein, 1967).

VOLUME 58, NUMBER 3

235

Other species of Osmia in other subgenera are also known to construct

thresholds (Rust, 1974).

The application of a salivary secretion to bare cell walls has not been

observed for any other species of Osmia. Indeed, only one other twig-nesting

species in the family Megachilidae, Chelostoma minutum Crawford, is sus¬

pected to apply a salivary secretion to the bare cell wall (Stephen et al., 1969;

P. F. Torchio, pers. commun.), although the use of saliva to make mortar

nests has been reported for other megachilids (Eickwort, 1975). It is possible

that other twig-nesting species in the family engage in similar behavior, but

this can be confirmed only by direct observation because the secretion is

visually undetectable after drying.

Acknowledgments

We thank W. P. Nye for making the observation nests and G. C. Eickwort,

R. W. Matthews, and R. W. Rust for helpful comments on the manuscript.

Literature Cited

Eickwort, G. C. 1975. Nest-building behavior of the mason bee Hoplitis anthocopoides (Hy-

menoptera: Megachilidae). Z. Tierpsychol., 37:237-254.

Hurd, P. D., Jr. 1979. Superfamily Apoidea. Pp. 1741-2209 in K. V. Krombein, P. D. Hurd,

Jr., D. R. Smith, and B. D. Burks (eds.), Catalog of Hymenoptera in America north of

Mexico, Vol. 2. Smithsonian Institution Press, Washington, D.C., pp. 1199-2209.

Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests and associates.

Smithsonian Press, Washington, D.C., 570 pp.

Medler, J. T. 1967. Biology of Osmia in trap nests in Wisconsin (Hymenoptera: Megachilidae).

Ann. Entomol. Soc. Am., 60:338-344.

Parker, F. D., and R. M. Bohart. 1966. Host-parasite associations in some twig-nesting

Hymenoptera from western North America. Pan-Pac. Entomol., 42:91-98.

Rau, P. 1937. The life-history of Osmia lignaria and O. cordata, with notes on O. conjuncta.

Ann. Entomol. Soc. Am., 30:324-343.

Rust, R. W. 1974. The systematics and biology of the genus Osmia, subgenera Osmia, Chal-

cosmia, and Cephalosmia (Hymenoptera: Megachilidae). Wasmann J. Biol., 32:1-93.

Stephen, W. P., G. E. Bohart, and P. F. Torchio. 1969. The biology and external morphology

of bees, with a synopsis of the genera of northwestern America. Oregon State University,

Corvallis, 140 pp.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 236-239

Published 16 December 1983

NOTES ON THE SUBGENUS BOMBYLIUS {ZEPHYRECTES)

(DIPTERA: BOMBYLIIDAE)

Neal L. Evenhuis

Department of Entomology, Bernice P. Bishop Museum,

P.O. Box 19000-A, Honolulu, Hawaii 96819

Since the erection of the subgenus Bombylius ( Zephyrectes ) by Evenhuis

(1978), described to include the species B. anthophoroides Evenhuis and B.

montanus Johnson and Johnson and since data on these two species were

presented in Hall and Evenhuis (1980), new distributional data, ecological

observations and systematic studies have been made on this unique group

of flies. Four additional species of Bombylius ( B. incanus Johnson, B. ravus

Loew, B. cruciatus Fabricius, B. nicholsonae Hall and Evenhuis) are here

relegated to this subgenus. The inclusion of one Palearctic species ( cruciatus )

now gives Zephyrectes a Holarctic distribution.

Bombylius ( Zephyrectes) incanus Johnson

Types of this northeastern U.S. species deposited in the Museum of Com¬

parative Zoology, Harvard University (MCZ) have been recently examined,

with 8 and 2 genitalic studies made on paratypes. Data additional to those

presented in Hall and Evenhuis (1980) are as follows: 8 —Antennal segments

I and II with white scales intermixed with sparse black hairs. Mesonotum,

post alar callus and mesopleura with orange macrochaetae. Genitalia (Fig.

1) in lateral view with basistylus linear-lanceolate, length 3.5 x width, slightly

clawed basally; dististylus length 4 x width, linear, slightly tapering to point¬

ed apex; epiphallus broadly tapering to aedeagal tip, dorsal surface with well

pronounced club-shaped projection, projection larger than in B. anthopho¬

roides or B. montanus ; aedeagus long, thin, slightly tapering to truncate apex;

basal apodeme large, rounded; epandrium subrhomboid, with slightly pro¬

nounced posterior process; anterior process weak; cercus long, length 2x

width. 2—Genitalia: similar to that of B. anthophoroides.

Bombylius ( Zephyrectes) ravus Loew

The unique female holotype of ravus deposited in the MCZ has been

recently examined and the following data additional to that presented in

Hall and Evenhuis (1980) is noted: 2—Antennal segments I and II with white

scale-like hairs, basal Vi of antennal segment III with white scale-like hairs

laterally and dorsally, bare mesally. Anterior thoracic spiracle with dark

VOLUME 58, NUMBER 3

237

Figs. 1, 2. Fig. 1. Bombylius ( Zephyrectes) incanus Johnson, male genitalia: a) lateral view,

b) dorsal view. Fig. 2. B. (Z.) cruciatus Fabricius, female spermatheca.

brown bristles; macrochaetae and bristles on mesonotum, postalar callus,

mesopleuron and scutellum amber.

The “green pollinose” thorax mentioned in Hall and Evenhuis (1980) is

actually grayish ground color on the pleura as well as the lateral and anterior

portions of the mesonotum. The remainder of the thoracic ground color is

dark brown.

Bombylius ( Zephyrectes ) cruciatus Fabricius

Examination of both males and females of this species show it to possess

the characters typical of Zephyrectes (viz., white scale-like hairs on antennal

segments I and II and presence of amber-colored macrochaetae on the tho¬

rax). The female genitalia (Fig. 2) is also consistent with that of other Ze¬

phyrectes species in having the apical spermathecal duct slightly sclerotized

and the ejaculatory apparatus with many canaliculi.

It is very likely that other Palearctic species allied to B. cruciatus (e.g.,

cinerarius Pallas, androgynus Loew, armeniacus Paramonov, vlasovi Para-

monov, quadrifarius Loew, testaceiventris Paramonov) may also belong to

238

PAN-PACIFIC ENTOMOLOGIST

Fig. 3. Distribution of Bombylius ( Zephyrectes ) anthophoroides Evenhuis (small circles =

previous collecting records; large dots = new collecting records) and the plant Trichostema

lanceolatum Bentham (shading).

VOLUME 58, NUMBER 3

239

the subgenus Zephyrectes. Examination of both male and female genitalia

of those species will be necessary before any conclusions can be made as to

a subgeneric allocation. The male genitalia of those species listed above and

figured in Zaitzev (1966) fit the characters exhibited in other species be¬

longing to Zephyrectes.

Bombylius ( Zephyrectes ) anthophoroides Evenhuis

This species has previously been noted as a monolectic pollinator of the

labiate plant Trichostema lanceolatum Bentham (Hall and Evenhuis, 1980).

A collecting expedition in September 1980 was conducted by the author and

Mr. Thomas Plichta in northern and central California in an attempt to

extend the known distribution of this species which coincides remarkably

with the distribution of T. lanceolatum. The results of the expedition are

shown in Figure 3. New county records for B. (Z.) anthophoroides include

Calaveras, Madera, Mariposa, Sacramento, Solano and Yolo. The north¬

ernmost extension of this species was found in Yolo County, 1.6 mi [2.8

km] west of Winters. The finding of this species on the eastern side of the

central valley further emphasizes that B. (Z.) anthophoroides is restricted in

distribution to locations where Trichostema lanceolatum is present. Evi¬

dence at hand strongly supports the theory that this species of Bombylius

has co-evolved with, and restricted its distribution, emergence and flight

period to coincide with the location and blooming of T. lanceolatum. Future

collecting of this species will concentrate on more northern areas into Oregon

and southerly locations into Baja California. A forthcoming paper will de¬

scribe in detail the biological and ecological observations on this remarkable

species of bee fly and will include a discussion on the co-evolution of B.

anthophoroides and T. lanceolatum.

Bombylius ( Zephyrectes ) nicholsonae Hall and Evenhuis

The description of B. nicholsonae in Hall and Evenhuis (1980) failed to

state the subgeneric allocation of this species. Dissection of the female gen¬

italia and examination of other morphological characters show it to be a

member of the subgenus Zephyrectes.

Literature Cited

Evenhuis, N. L. 1978. New species and new subgenus of Bombylius (Diptera: Bombyliidae).

Entomol. News, 89:33-38.

Hall, J. C., and N. L. Evenhuis. 1980. Volume V. Homeodactyla and Asilomorpha, Part 13.

Bombyliidae, Number 1, pp. 1-96. In: G. C. D. Griffiths (ed.), Flies of the Nearctic

region. E. Schweizerbart’sche, Stuttgart.

Zaitzev, V. F. 1966. Paraziticheskie mukhi semeyestva Bombyliidae (Diptera) v faune zak-

avkaz’ya. Akademiia Nauk, Moscow and Leningrad, 375 pp.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 240-244

Published 16 December 1983

DESCRIPTION OF AND PHYLOGENETIC COMMENTS ON THE

FINAL LARVAL INSTAR OF CARYOBRUCHUS VESEYI

(HORN) (COLEOPTERA: BRUCHIDAE)

Gary S. Pfaffenberger

Department of Life Sciences, Eastern New Mexico University,

Portales 88130

Numerous species of the widely distributed and morphologically diverse

genus Caryobruchus Bridwell have been studied (e.g., Bridwell, 1929; Pfaf¬

fenberger, 1974; Pfaffenberger and Johnson, 1976). This is not so for C.

veseyi (Horn) which apparently is restricted to the range of Erythea bran-

degeei Purpus, its host. According to Bridwell (1929), this bruchid is found

in western Mexico and the Cape region of Baja California.

Adults oviposit on the seed coat of drupaceous fruits and like other bruchid

species, the ecluding larva bores into the underlying fruit to feed within the

oil-rich endosperm (Bridwell, 1929). Available information seems to indicate

that C. veseyi has a high degree of host specificity toward its geographically

restricted host.

To date there has only been one published larval description within this

genus (Pfaffenberger, 1974). Therefore, the foregoing description is deemed

important because of the recent active interest in larval taxonomy and the

subsequent support such information will lend to a reliable scheme of clas¬

sification among the Bruchidae.

Gratitude is expressed to D. H. Kavanaugh for supplying the larval spec¬

imen which is deposited in the collection of the California Academy of

Sciences.

Caryobruchus veseyi (Horn)

Body.—{ See Fig. 1 in Pfaffenberger, 1977.) Width-depth 4-5 mm by 9

mm long; C-shaped, robust; greatest width-depth in meso-metathoracic seg¬

ments tapering slightly toward anterior end; abdominal segments 1-5 smaller

and subequal in size, segments 6-10 with distinct posterior taper, segment

10 minute and nearly obscured by segment 9; cuticle white to yellowish,

without conspicuous pigmented or sclerotized areas, setae restricted pri¬

marily to thoracic and abdominal sternites (chaetotaxy similar to Fig. 1A

in Pfaffenberger, 1974), tergal setae sparse, found primarily on plical crests

of larger abdominal segments.

Head.—(See Figs. 3, 4 in Pfaffenberger, 1977.) Retractable, hypognathous,

VOLUME 58, NUMBER 3

241

dorso-ventrally flattened, invaginated portion of capsule tan in color, mouth-

parts heavily pigmented, particularly the mandibles.

Ocelli.— Absent.

Antenna (Fig. 1). —Located at base of mandible near distal curvature of

epicranial arm (Fig. 3 in Pfaffenberger, 1977); 2-segmented, frequently re¬

tracted into sclerotized antennal sheath, segments of subequal lengths, second

segment narrower, with crenulated distal margin; 5 well-developed, sharp,

stout, occasionally decurved, setae circumscribe distal, sensory cones; each

setal base partially encircled by scallop in sclerotized, crenulated antennal

margin; center of distal end occupied by 2 basiconic sensillae, median sen-

sillum enlarged, 3-4 times larger than lateral sensillum, lateral sensillum

with pointed, distal end.

Clypeolabrum (Fig. 2).—Clypeal portion 3 times as wide as long, with

convex basal border; 2 setae and sensory pore aligned along proximolateral

border, each spaced equidistantly by length of one seta, with sensory pore

located medial most; labral portion with flat proximal border and arcuate

distal border, distal margin concealed by several rows of elongated setal-like

structures forming a brush border; 10 asymmetrically arranged setae partially

obscured by brush border; setal pairs A, B, D, and E appear to form somewhat

of an arc with setal pair C located near center of arc; pair of sensory pores

located near proximolateral base of asymmetrically arranged setae; 2 addi¬

tional pairs of setae and sensory pores located laterally along proximal bor¬

der, all being equidistantly spaced by length of one of the setae, sensory pore

proximomedial compared to setae.

Epipharynx (Fig. 3). —Distal border supporting dense, spinous mat which

broadens laterally; 2 pairs of short, sharp, decurved setae located medially

along lateral borders of epipharyngeal groove; between anterolateral spinous

mat and medial, decurved setae is enlarged cluster of transversely aligned

rows of spinous projections in mini-clusters, sclerotized projections more

elongate and compact proximomedially, forming lateral borders for epi¬

pharyngeal groove.

Mandible .—(See Fig. 8 in Pfaffenberger, 1977.)

Maxilla (Fig. 4). —Cardo present (destroyed during preparation); stipes

entirely membranous, bearing 24, well-developed setae on distal, medio-

ventrolateral surface; sclerite of palpifer lightly sclerotized and bearing 5,

strongly decurved, elongate setae on medioventral surface; near base of

medial-most palpifer seta is small cluster (10) of apparent placoid-like sen¬

sillae; palpifer membrane bearing sensory pore and short, strongly decurved

setae on distoventral surface; palpus single-segmented, bearing seta, midway

between proximal and distal ends, on medial surface; lacinial sclerite, located

ventrolaterally, bearing single, ventral sensory pore, 5 spatulate setae bor¬

dering distolateral surface, distoventral surface bordered by 3, equidistantly

spaced, strongly decurved setae, distomedial area occupied by 5, transversely

242

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-7. Final larval instar Caryobruchus veseyi (Horn), scale line 0.1 mm. Fig. 1. Antenna.

Fig. 2. Clypeolabrum. Fig. 3. Epipharynx. Fig. 4. Left maxilla. Fig. 5. Labium. Fig. 6. Spiracle.

Fig. 7. Anal sulcus.

VOLUME 58, NUMBER 3

243

arranged setae, boomerang-shaped, spatulate-like seta near dorsomedial bor¬

der, 2 hair-like (possibly trichoid sensillae) setae on dorso-medial surface.

Labium (Fig. 5). —Entirely membranous; somewhat flattened proximally,

with concave skewed, arch-like, distal taper; submentum fleshy, bearing pair

of setae near each anterolateral border, anteromedial seta short, stout, half

as long as other seta; lateral margin of mentum bordered basally by stout

seta subtended by sensory pore, remainder of lateral margin bordered by

(from proximal to distal end) equidistantly spaced sensory pore and 2 stout,

sharp setae.

Leg. — (See Fig. 1H in Pfaffenberger, 1974.) Number of segments unde¬

termined; without sclerotization.

Spiracle (Fig. 6). —Oval-shaped; uniforous; mesothoracic pair enlarged;

atrial orifice nearly obscured by sclerotized, highly branched projections.

Anus (Fig. 7).—Y-shaped, terminal.

Host plants. —Erythea brandegeei Purpus.

Specimen examined. — One final instar (determined by association with

adults collected from host plant). Collected along trail between “Tapon” and

Rancho Poza Larga, 1100-1200 m, Arroyo de San Francisquito (from San

Jorge to San Francisquito and La Chuparosa), east side Sierra de la Victoria,

Baja California Sur, MEXICO, 13 April 1955 (Annetta Carter and Roxana

S. Ferris, 3375).

Significant characters. — Small size; absence of ocelli; crenulated distal

margin of second antennal segment; 5 setae circumscribing distal, antennal

sensory cones; presence of 2 basiconic sensillae on antenna; absence of con¬

spicuous sclerotized areas on clypeolabrum; brush-like border comprising

several rows of elongate setae, found along distal margin of labrum; 5 pairs

of asymmetrically arranged setae located anteromedially on labrum; en¬

larged, comma-shaped patch with transversely aligned rows of spinous pro¬

jections on epipharynx; absence of sclerite on stipes; 24 well-developed setae

on stipes; poorly sclerotized palpifer sclerite; presence of small cluster (10)

of apparent placoid-like sensillae; 5 strongly decurved setae along distal

margin of palpifer sclerite; single segmented palpus without visible sensory

structures on distal end; 5 spatulate setae bordering distolateral surface of

lacinia; 2 transverse rows of setae on distal end of lacinia; labium entirely

membranous; labium tapered anteriorly with skewed, concave lateral mar¬

gins; submentum indistinguishable from nonsclerotized mentum; labium

bordered laterally with 5 setae and 2 sensory pores; atrial orifice of spiracle

surrounded by rows of elongate, highly branched, sclerotized projections.

Discussion

According to Bridwell (1929), the genus Caryobruchus appears to consist

of a deviant C. veseyi and two well defined species groups. The differences

in arrangement, and/or presence/absence of chaetotaxy and particularly the

244

PAN-PACIFIC ENTOMOLOGIST

absence of ocelli in C. veseyi confirm his observations. Other major differ¬

ences involve the extent of sclerotization as well as the number of sclerites.

The latter may easily be interpreted on the basis of adaptation to seed

hardness. For example, seeds of Scheelea rostrata Burret (host for C. buscki )

and members of the genus Sabal (e.g., S. minor (Jacq.) Pers.; Woodruff,

1968) (host for C. gleditsiae (Linnaeus)) have a very hard seed whereas, the

seed of E. brandegeei (host for C. veseyi) is relatively soft.

Increased seed hardness translates into a greater demand for musculature

to enable the larva to penetrate and successfully excavate the seed endo¬

sperm. With increased musculature a commensurate need arises for enlarged

and more numerous sclerotized bases (sclerites) of attachment for the mus¬

cles. Therefore, larvae occupying harder seeds should theoretically possess

a greater sclerotized surface area than larval forms parasitizing seeds with

softer endosperm and more fragile seed coats. Such an hypothesis derives

support from comparative observations of C. buscki Bridwell (Pfaffenberger,

1974) and C. veseyi. To confirm this observation, however, would require

a comparative examination of the first (see C. gleditsiae. ; Pfaffenberger and

Johnson, 1976) and final larval instars of several larval species in addition

to those mentioned herein.

The extreme differences in presence/absence and degree of sclerotization

of the sclerites suggests the need for a review of the integrity of this divergent

genus. Such a recommendation is particularly evident when one observes

the presence of 3 pairs of ocelli in C. buscki and none in C. veseyi. Because

of the comparatively low adaptability of ocelli, they have been of distinct

importance in distinguishing genera (Prevett, 1971) and even subfamilies

(Boving, 1929) among larval Bruchidae.

Literature Cited

Boving, A. G. 1929. On the classification of beetles according to larval characters. Bull.

Brooklyn Entomol. Soc., 24:63.

Bridwell, J. C. 1929. A preliminary generic arrangement of the palm bruchids and allies

(Coleoptera) with descriptions of new species. Proc. Entomol. Soc. Wash., 31:141-160.

Pfaffenberger, G. S. 1974. Comparative morphology of the final larval instar of Caryobruchus

buscki and Pachymerus sp. (Coleoptera: Bruchidae: Pachymerinae). Ann. Entomol. Soc.

Am., 67:691-694.

-. 1977. Comparative descriptions of the final larval instar of Bruchus brachialis, B.

rufimanus, and B. pisorum (Coleoptera: Bruchidae). Coleopt. Bull., 31:133-142.

-, and C. D. Johnson. 1976. Biosystematics of the first-stage larvae of some North

American Bruchidae (Coleoptera). U.S.D.A. Tech. Bull., 1525:1-75.

Prevett, P. R. 1971. The larvae of some Nigerian Bruchidae (Coleoptera). Trans. R. Entomol.

Soc. Lond., 123:247-312.

Woodruff, R. E. 1968. The palm seed “weevil,” Caryobruchus gleditsiae (L.), in Florida

(Coleoptera: Bruchidae). Fla. Dept. Agric. Entomol. Circ., 73:1-2.

PAN-PACIFIC ENTOMOLOGIST

July 1982. Vol. 58. No. 3, pp. 245-249

Published 16 December 1983

AGGREGATION OF COELOPA (. NEOCOELOPA ) VANDUZEEI

CRESSON ON THE MONTEREY PENINSULA COAST,

CALIFORNIA, AND NOTES ON THE FAMILY

(DIPTERA: COELOPIDAE)

Paul H. Arnaud, Jr.

California Academy of Sciences, Golden Gate Park,

San Francisco 94118

The kelp fly, Coelopa ( Neocoelopa) vanduzeei Cresson is found along our

Nearctic Pacific coast beaches from Baja California north to Alaska. Poinar

(1977:81, 83) has observed that adults are more abundant on beaches in

southern California in the summer than in the winter. Adults may swarm

over stranded kelp on beaches and literally darken the adjacent sand with

their numbers. Observations on a natural aggregation of C. vanduzeei, in¬

dependent of a kelp or sand matrix, along the central California coast, during

the late spring of 1973, illustrated with photographs, are here documented,

and observations made by two colleagues are also recorded.

On June 9, 1973, at the Seal and Bird Rocks, located on the 17 Mile Drive

of the private Del Monte Properties Company, Monterey County, California,

at about 1600 hours daylight standard time, in clear weather, a large aggre¬

gation of C. vanduzeei was observed and photographed. The aggregation

was located on the then lee side of a granitic rock outcrop. The rock outcrop,

oriented somewhat at right angles to the incoming surf, measured about 15

meters in length and three meters in height, and even though separated from

the shore line, supported a limited vegetation that included the natives

Eriogonum parvifolium Sm. in Rees and Spergularia macrotheca (Hornem.)

Heynh., and the introduced Cakile maritima Scop. The seaward side of the

rock outcrop was bordered by an extensive wrack deposit. The kelp flies in

the aggregation were quiescent at the time of my observations.

When C. vanduzeei was described by Cresson (1914:457-458) from La

Jolla, California, only four specimens were known. Aldrich (1929:1-6) re¬

vised Coelopa for the Nearctic Region and knew of 119 museum specimens

of C. vanduzeei from the California coast. Kompfner (1974:44-51) described

and illustrated the third instar larva and puparium and provided biological

information on C. vanduzeei from the population at Pacific Grove on Mon¬

terey Bay. Poinar (1977:81-86) studied its biology on a San Diego County

beach at Solana Beach and reported differences in the utilization of beach

wracks by this southern population from those reported from Monterey Bay.

A review of this small family Coelopidae (with less than two dozen world

246

PAN-PACIFIC ENTOMOLOGIST

Figs. 1, 2. Fig. 1. Granitic rock outcrop, at Seal and Bird Rocks, site of aggregation of

Coelopa ( Neocoelopa ) vanduzeei Cresson. Fig. 2. Aggregation of C. ( N .) vanduzeei Cresson

located near center of granitic rock outcrop illustrated in Figure 1.

VOLUME 58, NUMBER 3

247

Fig. 3. Aggregation of Coelopa (Neocoelopa) vanduzeei Cresson located about a meter and

a half from left end of granitic rock outcrop illustrated in Figure 1.

248

PAN-PACIFIC ENTOMOLOGIST

species)—their great abundance, on occasions, and their biology—include

the following references. In the Palearctic region Ringdahl (1921:66-67)

discussed four species of Coelopidae in southern Sweden. Karl (1931:198)

in his paper on Diptera of Amrum and Helgoland Islands, of the North

Frisian Islands, Germany, discussed Coelopa frigida (Fabricius). Hennig

(1937:16) in his fine overview of the Coelopidae revised the Palearctic species,

and provided additional information on adult and larval morphology. Old-

royd (1954:198-202) has provided an excellent account of the exceptionally

abundant occurrence of C. frigida on the south coast of England and their

large numbers far inland, chiefly in the London area, and Taylor has dis¬

cussed their occurrences inland at Oxford (1955a:97) and along the coast at

Dawlish, South Devon (1955b: 107). Oldroyd reported that C. frigida is

irresistibly attracted to organic solvents such as trichloroethylene, chloro¬

form, and carbon tetrachloride, and certain paints and a few detergents as

well as odors from pharmacists shops. Egglishaw (1960:109-140) has pro¬

vided a detailed study of the larval stages and the biology of the five British

species of Coelopidae, and reported (1961:11-17) mass migrational flights

of C. frigida and C. pilipes Haliday. In the Australian region, Scotti, Gibbs,

and Wrigley (1976:1) report that on the South Australian Coast in New

South Wales adults of Chaetocoelopa Sydneyensis Schiner “. . . congregate

during winter in large numbers on the spindrift-moistened undersurfaces of

overhanging rocks near the high water level.”

The mechanisms by which the aggregation at the Seal and Bird Rocks

could have formed are not known. Possible mechanisms could consist of an

independent and individual response to an environmental gradient (or gra¬

dients) leading to aggregation in an environmentally optimum location, or

individual response to some stimulus (or stimuli) provided by other indi¬

viduals, leading to aggregation at a common location, or a combination of

both as suggested by Kavanaugh (1977:27-31) with an aggregation of the

carabid genus Scaphinotus. The length of time that such aggregations remain

intact is not known but due to the flight capabilities of the kelp flies they

could be rapidly decreased in size or terminated in response to some new

stimulus (or stimuli). The flies did not seem to be disturbed by my close

proximity in their observation or by the taking of the flash pictures. Con¬

siderable dispersal did occur, however, when a net was used to collect vouch¬

er specimens. J. R. Yockeroth (in a letter dated March 17, 1975) reported

that on July 4, 1973, at the base of low cliffs at Carmel, California that

several feet of the cliff were virtually black with C. vanduzeei, but in contrast

they all flew as he approached. V. F. Lee (in conversation) reports obser¬

vations of kelp flies thought to be C. vanduzeei, made near Bolinas Point,

Marin County, California, on October 5, 1979. He noted some individuals

along the beach cliff flying into the prevailing wind, others flying into the

eddy of the lee, and many present on the lee surface of the cliff, crawling

toward the windward edge.

VOLUME 58, NUMBER 3

249

Scotti, Gibbs, and Wrigley also report that adult Chaetocoelopa sydney-

ensis carry a virus, which they named kelp fly virus (KFV). It appears to

be distinct from other previously described viruses. As mentioned by Poinar

(1977:85), adults of C. vanduzeei in California can . . become a general

nuisance to people, alighting on their bodies and are sometimes seen around

the eyes of children.” A study of the kelp fly virus, relative to our kelp flies,

has not been made.

Acknowledgments

The author acknowledges with thanks the aid of: Brian H. Cogan and

James P. Dear, British Museum (Natural History), for furnishing a copy of

the Oldroyd reference; Maurice C. Giles, California Academy of Sciences,

for the photographic prints used in Figures 1 to 3, made from negatives on

Kodak Panatomic-X film; John T. Howell, California Academy of Sciences,

for plant identifications; Vincent F. Lee, California Academy of Sciences,

and J. R. Vockeroth, Canadian Department of Agriculture, for sharing in¬

formation on their observations of California kelp flies; George O. Poinar,

Jr., University of California, Berkeley, for supplying a copy of the Scotti et

al. reference and other information; George C. Steyskal, United States De¬

partment of Agriculture, for the confirmation of the identification of the

Coelopa and for references; and Madeline M. Arnaud and Nicole van Baal

for their interest and participation in this and other fieldwork.

Literature Cited

Egglishaw, H. J. 1960. Studies on the family Coelopidae (Diptera). Trans. R. Entomol. Soc.

Lond., 112:109-140.

-. 1961. Mass migrational flights of Coelopa frigida (Fabricius) and C. pilipes Haliday

(Diptera, Coelopidae) and Thoracochaeta zosterae Hal. (Diptera, Sphaeroceridae). Ento¬

mologist, 94:11-18.

Hennig, W. 1937. Coelopidae. [Fam.] 52, 39 pp., 40 figs., In E. Lindner (ed.), Die Fliegen

der palaearktischen Region, vol. 5, Stuttgart.

Karl, O. 1931. Fliegen von der Insel Amrum. Ein Beitrag zur Fliegenfauna der nordfriesischen

Inseln (Dipt.). Dtsch. Entomol. Z., Jahrgang 1930:193-206.

Kavanaugh, D. H. 1977. An example of aggregation in the Scaphinotus subgenus Brennus

Motschulsky (Coleoptera: Carabidae: Cychrini). Pan-Pac. Entomol., 53:27-31.

Kompfner, H. 1974. Larvae and pupae of some wrack dipterans on a California beach (Diptera:

Coelopidae, Anthomyiidae, Sphaeroceridae). Pan-Pac. Entomol., 50:44-52.

Oldroyd, H. 1954. The seaweed fly nuisance. Discovery, 15:198-202.

Poinar, G. O., Jr. 1977. Observations on the kelp fly, Coelopa vanduzeei Cresson in southern

California (Coelopidae: Diptera). Pan-Pac. Entomol., 53:81-86.

Ringdahl, O. 1921. Bidrag till kannedomen om de skfinska stranddynemas insektfauna. Ento¬

mol. Tidskr., 42(1):21—40; (2):65-92.

Scotti, P. D., A. J. Gibbs, and N. G. Wrigley. 1976. Kelp fly virus. J. Gen. Virol., 30:1-9.

Taylor, E. 1955a. Seaweed flies (Diptera, Coelopidae) at Oxford. Entomol. Mon. Mag., 91:

97.

-. 1955b. Abundance of seaweed flies (Dipt., Coelopidae) near Dawlish, South Devon.

Entomol. Mon. Mag., 91:107.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58. No. 3, pp. 250-257

Published 16 December 1983

DIURNAL VARIATION IN THE PREDATORY BEHAVIOR OF THE

GRASSLAND ROBBER FLY, PROCTACANTHELLA LEUCOPOGON

(WILLISTON) (DIPTERA: ASILIDAE)

Todd E. Shelly 1 and David L. Pearson

Department of Biology, 208 Mueller Laboratory,

The Pennsylvania State University, University Park 16802

Diurnal variability in robber fly foraging activity is most commonly stud¬

ied using a transect-census approach (Lavigne and Holland, 1969; Lavigne

and Dennis, 1975; Dennis and Lavigne, 1975, 1976a, 1976b; Hespenheide

and Rubke, 1977; Scarbrough and Norden, 1977, is an exception). The usual

procedure involves walking a transect throughout the day and counting the

number of actively foraging individuals or the number of individuals in

different microhabitats. When applied to asilids of the arid western United

States, this method often reveals a 4 stage activity pattern for clear, relatively

windless days (Lavigne and Holland, 1969; Lavigne and Dennis, 1975; Den¬

nis and Lavigne, 1975, 1976a, 1976b). As outlined by Lavigne and Holland

(1969), the 4 stages are: (1) an early morning warm-up, (2) a mid-morning

foraging peak, (3) late morning to mid-afternoon heat avoidance with low

foraging activity, and (4) a late afternoon increase in foraging activity.

Despite a general recognition of this pattern, the actual hourly variation

in flight activity (both foraging and relocation) has never been measured for

an arid grassland robber fly species. Consequently, precise statements re¬

garding the actual degree of foraging rate variability cannot be made, e.g. is

the peak foraging rate of mid-morning double, triple, etc. the minimum

foraging rate of mid-afternoon? The present study was thus undertaken to

provide a quantitative assessment of diurnal flight activity for the grassland

species Proctacanthella leucopogon (Williston). The results of a short-term

mark-recapture study are also presented.

Proctacanthella leucopogon is a medium sized asilid (12-15 mm) of the

grasslands of western United States and northern Mexico (Wilcox, 1965).

In Colorado, adults are active from late June to early September (Rogers

and Lavigne, 1972). Rogers and Lavigne (1972), Lavigne and Dennis (1980),

and Shelly and Pearson (1980) all recorded Homoptera (primarily Cicadel-

lidae), Diptera, and Hymenoptera as the major dietary components. Re¬

cently, Lavigne and Dennis (1980) described the foraging and courtship

behavior of this species. Their observations of diurnal activity, however, are

largely qualitative. The data presented here together with recent prey rec-

VOLUME 58, NUMBER 3

251

ognition tests (Shelly and Pearson, 1980) complement their observations

through a more quantitative interpretation of foraging behavior.

Materials and Methods

The foraging observations and mark-recapture study were conducted from

25 July to 6 August 1978, in Sulphur Springs Valley, Cochise Co., Arizona,

8 km southeast of Wilcox, Arizona (elev. 1273 m). The study site was a

shallow gully (approximately 100 m long and 6-11 m wide) in a moderately

used pasture. The predominant plant species were Distichlis stricta (Rybd.),

Prosopis velutina Woot., Suaeda torreyana S. Wats, and Atriplex griffithsii

(Strandl.).

Data on foraging were collected between 07:00 and 17:00 on sunny days

when winds were light. Observations were confined to females since (1)

females were more abundant and hence more easily located than males and

(2) potential complications arising from sexual differences were eliminated.

An individual was observed until lost from view or until 10 minutes elapsed.

The observations included: (1) number of foraging flights, (2) number of

relocation flights, (3) capture success (number of successful captures/number

of foraging flights), and (4) time spent perched in the sun on the ground, in

the shade on the ground and on vegetation above the ground. In addition,

visual estimates were made of distance for both foraging and relocation

flights.

In most cases, the potential prey that elicited a foraging flight could be

seen. However, even when the prey was not apparent, foraging and relocation

flights were easily distinguished. A foraging flight was typified by a rapid,

straight flight to an aerial point followed by a similar return to the ground.

In addition, foraging flights were usually made at angles approaching vertical.

In contrast, relocation flights were slower, less direct, and usually low and

parallel to the ground.

Data for all individuals observed during a given hour (07:00-08:00,

08:00-09:00, etc.) were combined, and both foraging and relocation activity

were expressed as flights per minute of observation time. Perch selection

was expressed as the percentage of the total observation time for a given

hour spent in each of the 3 perch categories. In these analyses, data for

12:00-13:00 and 13:00-14:00 were combined and treated as 1 time period

since few data were collected in either of these hours alone.

The mark-recapture study was conducted during a 3 day period following

completion of the foraging observations. Ten sections were established with¬

in the gully varying from 8.5 m to 10.0 m in length and from 6.25 m to

11.9 m in width; these latter values reflect variation in gully width. For its

entire length the gully was bordered on either side by approximately 2-3 m

of bare ground. The western end of the gully opened onto a dry pond bed,

252

PAN-PACIFIC ENTOMOLOGIST

Table 1. Perching positions of female P. leucopogon throughout the day. Ground surface

temperatures were recorded on July 30, 1978, at the start of each observation period.

Time

Minutes of

observation

Ground surface

temperature (°C)

% time on ground

% time on

vegetation

Sun

Shade

Sun

Shade

07:00-08:00

08:00-09:00

09:00-10:00

10:00-11:00

11:00-12:00

102

12:00-14:00

14:00-15:00

15:00-16:00

16:00-17:00

while at the eastern end several mesquite bushes (Prosopis) separated the

study gully from another gully which ran perpendicular to it.

The flies were marked by gluing numbered, circular bee tags (Poalith-

Zeichenplattcheb, Graze®) directly onto the center of the mesothorax. The

tag did not appear to inhibit flight. In the field, new individuals were marked

and previously marked individuals were identified during both the morning

(09:00-11:00) and afternoon (13:00-15:00). The location of each fly was

marked by a spike bearing a number corresponding to that fly’s tag number.

The exact position of each spike was obtained by triangulation. Most marked

flies were later identified without capture thus disturbing them as little as

possible. Minimum travel distances (straight lines) between captures/obser¬

vations were then calculated for all marked individuals seen more than once.

Results

Although P. leucopogon females perched almost exclusively on the ground,

whether they did so in sun or shade appears to be highly dependent upon

the time of day and hence the ground surface temperature (Table 1). From

07:00 to 11:00 the ground temperature in the sun increased from 24°C to

44°C, and the flies spent the great majority of this time perched in the sun.

However, from 10:00-11:00 to 11:00-12:00 the percent time spent in the

sun and in the shade changed from 78% time in sun and 20% time in shade

to 17% time in sun and 69% time in shade. Individuals remained in the

shade until late day when the ground surface temperature dropped from

48°C (16:00) to 42°C (17:00). Perching on vegetation was not observed to

any significant degree until the 11:00-12:00 period when 14% of the time

was spent perched on vegetation. Little vegetation perching occurred in the

NO. FLIGHTS/min.

VOLUME 58, NUMBER 3

253

HOUR OF DAY

Fig. 1. Foraging and relocation flight activity throughout the day for female P. leucopogon.

Total observation times for each.hour are given in Table 1.

afternoon although this behavior was slightly more common than in early

to mid-morning.

Flight activity showed a similar dependence upon temperature (Fig. 1).

Low foraging flight activity between 07:00 and 08:00 was followed by a

foraging peak from 08:00 to 11:00 when 0.39 to 0.45 foraging flights/minute

were made. As the temperature increased to 45°-49°C by mid-day, foraging

flights/minute declined to 0.12 for the 12:00-14:00 period. From 14:00 to

254

PAN-PACIFIC ENTOMOLOGIST

Table 2. Mean minimum distance traveled by marked P. leucopogon females as a function

of daylight hours since last capture. Standard deviation is given in parentheses.

Daylight hours since

last capture

No. of

observations

Mean minimum distance

traveled (m)

9.0 (7.7)

18.8(12.5)

17.1 (7.9)

28.8

5.4 (1.6)

28.1 (8.3)

1.5 (0.1)

17:00, however, foraging flight frequency increased, although it never reached

the peak morning levels. Relocation flight behavior exhibited a similar hourly

trend (Fig. 1). Unlike the morning plateau of foraging flight frquency, how¬

ever, the frequency of relocation flights continued to increase and peaked

during the 10:00-11:00 period. Similarly, in late afternoon relocation activity

increased dramatically rather than leveling off as did foraging flight activity.

Based on visual estimates, individual foraging flights ranged from 5 cm

to approximately 1.8 m with the majority being between 10 cm and 60 cm.

Foraging flights were rarely more than 60 cm above ground. Capture success

(number successful captures/number foraging flights) was 6%. Relocation

flights ranged from 5 cm to approximately 1.2 m with the majority being

between 30 cm and 90 cm.

In the mark-recapture study, the majority (67%) of the 64 females marked

were not recaptured (or re-sighted) again. Fourteen individuals were recap¬

tured once, 5 individuals were recaptured twice, 1 individual was recaptured

3 times, and another individual was recaptured 4 times. The mean distances

traveled between captures for individuals recaptured once and twice were

15.4 m (SD 12.9) and 14.4 m (SD 10.6), respectively. For the individual

recaptured 3 times, the minimum distances traveled between capture sites

were 2.0 m, 16.1 m, and 18.0 m, and for the individual recaptured 4 times,

the corresponding distances were 5.0 m, 5.5 m, 9.8 m, and 11.7 m.

The high mobility of P. leucopogon females is also demonstrated in a plot

of minimum distance traveled as a function of daylight hours since last

capture or sighting (Table 2). Following marking in the morning, the 15

females recaptured in the afternoon of the same day had traveled an average

of 9.0 m. Seven and 11 daylight hours after the previous capture, females

traveled an average of 18.8 m and 17.1 m, respectively. Further relations

between distance travelled and hours since last capture were obscured by

apparent movement of individuals in and out of the study site. For example,

VOLUME 58, NUMBER 3

255

after 28 daylight hours, 3 females had moved a minimum of 23 m, but 3

females were found within 3 m of their marking site (Table 2). Similarly,

after 24 hours, 1 female had traveled 28.8 m, whereas 3 females were found

between 4.1 m and 6.9 m of their marking site (Table 2).

Discussion

Compared to the transect-census approach, measurement of actual flight

activity more precisely describes diurnal variability of robber fly behavior.

During the present study period, female P. leucopogon exhibited 4 behavioral

stages:

1. Early morning warm-up period (07:00-08:00). Females perched in the

sun almost exclusively (99% observation time) but exhibited relatively

low foraging and relocation rates (47% and 31% of peak rates, respec¬

tively).

2. Mid-morning foraging period (08:00-11:00). Within this period, females

perched in the sun at least 78% observation time for any 1 hour and

exhibited the peak foraging rate for the day and a very high relocation

rate (89% of the peak rate).

3. Mid-day shade seeking period (11:00-16:00). Within this period, females

perched in the shade a minimum of 63% observation time for any 1 hour

and exhibited the lowest foraging and relocation rates for the day (27%

and 30% of peak rates, respectively).

4. Late afternoon foraging period (16:00-17:00). Females perched in the

sun 63% observation time and exhibited a relatively high foraging rate

(67% of the peak rate) and peak relocation rate of the day.

Shade seeking behavior is apparently a thermoregulatory response to ex¬

cessively high ground surface temperatures. The critical temperature range

prompting such behavior was approximately 43°-45°C. The dramatic late

morning decrease in sun perching occurred when the ground surface tem¬

perature in the sun exceeded 44°C at 11:00. Conversely, the late afternoon

increase in sun perching occurred when the ground surface temperature in

the sun dropped from 46°C to 42°C from 16:00 to 17:00.

Although shade seeking appears to be a thermoregulatory response, it is

less clear whether or not the lowered foraging rate from shaded perches also

results from thermoregulatory constraints. Two lines of indirect evidence,

however, suggest that high mid-day temperatures do not limit foraging flight

frequency. First, ground surface temperatures in the shade ranged from 36°-

42°C during the shade seeking period. Thus, for the majority of the mid¬

day period ground surface temperatures in the shade were not appreciably

higher than the ground surface temperatures in the sun during the mid¬

morning period of peak foraging activity. Second, as shown for a neotropical

256

PAN-PACIFIC ENTOMOLOGIST

robber fly (Shelly, in prep.), foraging rate in asilids may be strongly correlated

with the encounter rate of potential prey items. For western grasslands,

Lavigne and Holland (1969) and Hespenheide and Rubke (1977) have both

suggested that prey availability decreases during the hot mid-day hours.

Consequently, the low mid-day foraging rate of P. leucopogon may simply

reflect this reduced encounter rate.

The present observations agree in large part with those of Lavigne and

Dennis (1980) for a population of P. leucopogon in Mexico. In the Mexican

population, individuals were described as “sluggish” in the early morning

when the ground surface temperature was less than 29.5°C. As temperature

increased, individuals foraged actively until seeking shade when the ground

surface temperature in the sun exceeded 45.5°-47.5°C. Although slightly

higher, this estimate is similar to the corresponding estimate of 43°-45°C

for the present observations. Perhaps the major difference between the 2

populations involves the frequency with which individuals perch on vege¬

tation to avoid high ground temperatures. Lavigne and Dennis (1980) report

perching upon vegetation as an intermediate step between sun and shade

perching, whereas in the present study the shift from sun to shade perching

was direct. In Arizona, individuals rarely perched on vegetation (maximum

value: 14% observation time for 11:00-12:00).

Regarding the mark-recapture study, 2 results indicate that P. leucopogon

females forage over considerable areas. First, the majority of the individuals

marked were not recaptured. Second, those recaptured had traveled relatively

large distances. Two hours after being marked, recaptured females had trav¬

eled an average of 9.0 m, and after 7 and 11 daylight hours the corresponding

values were 18.8 m and 17.1 m, respectively. Rather than reflecting a small

home range, the small travel distances noted for several individuals after 24

and 28 daylight hours probably indicate that these individuals left the study

site soon after being marked only to re-enter at approximately the same

location. Support for this interpretation comes from the fact that these in¬

dividuals were all recaptured only once and not repeatedly as would be

expected if their movements were confined to a small area. Thus, whereas

male P. leucopogon may set up territories (Lavigne and Dennis, 1980), P.

leucopogon females do not appear to exhibit strong site fidelity.

Summary

Fluctuations in ambient temperature impose severe constraints upon the

foraging activity of female Proctacanthella leucopogon. The diurnal activity

rhythm consisted of 4 periods: (1) an early morning warming up period, (2)

a mid-morning peak in foraging flight frequency, (3) a late morning to late

afternoon shade seeking period, and (4) a late afternoon increase in foraging

activity. Peak foraging and relocation rates were 0.45 flights/minute and 0.97

flights/minute, respectively. The mid-day shade seeking behavior reduced

VOLUME 58, NUMBER 3

257

foraging and relocation activity to 29% and 31% of peak levels, respectively.

A preliminary mark-recapture study showed that 2 hours after being marked

females had traveled an average distance of 9 m and showed no indication

of territoriality.

Acknowledgments

T. Shultz, J. Brown, and F. Virrazzi aided with the field work. J. Wilcox

kindly confirmed our species identification. We also thank D. S. Dennis, S.

Juliano, and R. J. Lavigne for their comments on the manuscript. This

research was supported by National Science Foundation grant DEB 78-

OS 172 to Pearson.

Literature Cited

Dennis, D. S., and R. J. Lavigne. 1975. Comparative behavior of Wyoming robber flies. II

(Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr., 30:1-68.

-, and-. 1976a. Ethology of Leptogaster parvoclava in Wyoming (Diptera: Asili¬

dae). Proc. Entomol. Soc. Wash., 78:208-222.

-, and-. 1976b. Ethology of Efferia varipes with comments on species coexistence

(Diptera: Asilidae). J. Kans. Entomol. Soc., 49:48-62.

Hespenheide, H. A., and M. A. Rubke. 1977. Prey, predatory behavior, and the daily cycle

of Holopogon wilcoxi Martin. Pan-Pac. Entomol., 53:277-285.

Lavigne, R. J., and D. S. Dennis. 1975. Ethology of Efferia frewingi (Diptera: Asilidae). Ann.

Entomol. Soc. Am., 68:992-996.

-, and-. 1980. Ethology of Proctacanthella leucopogon in Mexico (Diptera: Asi¬

lidae). Proc. Entomol. Soc. Wash., 82:260-268.

-, and F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming robber

flies (Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr., 18:1-61.

Rogers, L. E., and R. J. Lavigne. 1972. Asilidae of the Pawnee National Grasslands, in

northeastern Colorado. Univ. Wyo. Agric. Exp. Stn. Sci. Monogr., 25:1-35.

Scarbrough, A. G., and A. Norden. 1977. Ethology of Cerotainia albipilosa Curran (Diptera:

Asilidae) in Maryland: Diurnal activity rhythm and seasonal distribution. Proc. Entomol.

Soc, Wash., 79:538-554.

Shelly, T. E., and D. L. Pearson. 1980. Predatory behavior of Proctacanthella leucopogon

(Diptera: Asilidae): Prey recognition and prey records. Environ. Entomol., 9:7-9.

Wilcox, J. 1965. Proctacanthella Bromley (Diptera: Asilidae). J. Kans. Entomol. Soc., 38:

106-110.

Footnote

1 Present address: Department of Biology, University of California, Los Angeles, California

90024.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, p. 258

Published 16 December 1983

SCIENTIFIC NOTE

POLYANDRY IN BRACHYMYRMEX DEPILIS EMERY

(HYMENOPTERA: FORMICIDAE)

A large swarm of Brachymyrmex depilis Emery was observed at 1830

hours, September 1, 1980. This swarm consisted of two large column shaped

clouds of predominantly male ants and extended from about three or four

meters off the ground to an elevation of about ten meters. It occurred in an

open area of the central quad at the University of California, Davis.

Twenty-five to 30 males and three reproductive females were observed

and collected on a white sheet that was placed on the ground under the

swarm. One female alighted on the sheet with three males attached to her

abdomen, apparently in copula. These three males were motionless, attached

only by their everted genitalia and were being dragged around by the walking

female. Another female alighted with two males attached in the same man¬

ner; the third female was not observed copulating.

Multiple mating behavior by reproductive female ants has been reported

many times in the literature. These include: Acromyrmex landolti Forel

(Kerr, 1969, Evol. Biol., 3:119-175); Atta sexdens (Linnaeus) (Kerr, 1961,

Rev. Bras. Biol., 21:45-48); Ection burchelli (Westwood) (Rettenmyer, 1963,

Univ. Kans. Sci. Bull., 44:281-465); Formica aquilonia Yarrow (Pamilo,

Rosengren, Vepsalainen, Varvio-Aho, and Pisarski, 1978, Hereditas, 89:

233-248); F. montana Emery (Kannowski, 1963, Symp. Genet. Biol. Ital.,

12:74-102); F. opaciventris Emery (Scherba, 1961, J. N.Y. Entomol. Soc.,

69:71-87); F. rufa Linnaeus (Marikovsky, 1961, Insectes Soc., 8:23-30); F.

subintegra Emery (Kannowski, 1963, loc. cit.); Mycocepurus goeldii Forel

(Kerr, 1961, loc. cit.); Prenolepis imparis (Say) (Talbot, 1945, Am. Midi.

Nat., 34:504-510); Pogonomyrmex barbatus Smith, P. desertorum Wheeler,

and P. maricopa Wheeler (Holldobler, 1976, Behav. Ecol. Sociobiol., 1:405-

423); P. occidentalis Cresson (Nagel and Rettenmyer, 1973, J. Kans. Ento¬

mol. Soc., 46:82-101); and P. rugosus Emery (Holldobler, 1976, loc. cit.).

Multiple copulation does not automatically imply multiple insemination.

Other methods such as dissection and the use of genetic markers are needed

to provide conclusive evidence. However, observational data are necessary

to derive a comprehensive understanding of insect mating behavior.

Robert E. Page, Jr., Department of Entomology, University of Wisconsin,

Madison 53706.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, pp. 259-260

Published 16 December 1983

SCIENTIFIC NOTE

A NOTE ON THE BIOLOGY OF ANCISTROCERUS WALDENII

FLAVIDULUS BEQUAERT (HYMENOPTERA: EUMENIDAE)

Ancistrocerus waldenii flavidulus Bequaert occurs in the Pacific Coastal

regions of northern California and Oregon (Bequaert, 1943, Entomol. Am.,

23:225-295). Previous biological information on A. w. flavidulus is based

on two nests reported by O. W. Richards (1962, Pan-Pac. Entomol., 38:145-

146) from Marin County, California. The nests were located within rock

crevices and consisted of several cells plastered over with mud. Richards

also reported that prey were larval Tortricidae ( Cnephasia and Archips). The

data presented here are based on a nest found on a west facing road cut at

Shelter Cove, Humboldt County, California, on 21 May 1976. The mud

nest (Fig. 1) was in a shallow depression in the slightly overhanging side of

a protruding rock. The female wasp (identification confirmed by J. R. Car¬

penter) was plastering mud onto the nest surface when she was captured.

The rock was brought into a field laboratory and the nest walls were removed.

Three cells provisioned with caterpillars (identified by Dr. J. A. Powell) and

a small empty cell in the upper left hand corner (Fig. 2) were exposed.

Presumably the first cell constructed was at the bottom and the last at the

top. The uppermost provisioned cell contained two larvae of Chionodes sp.

(Gelechiidae) and 9 of Cnephasia sp. (Tortricidae). The wasp egg (about 2

mm long) was suspended by a silk thread from the upper side of the cell

near the presumed inner end. The middle cell contained 9 larvae of Cne¬

phasia sp. and six undetermined small dark caterpillars. A small chrysidid

wasp larva and a shriveled host wasp egg were also present. The bottom cell

had 12 larvae of Cnephasia sp., a slightly larger chrysidid larva, and a

Figs. 1, 2. Laboratory photographs of mud nest of Ancistrocerus waldenii flavidulus. Fig. 1..

Nest on side of rock, scale equals 1 cm. Fig. 2. Nest with outer wall removed to expose the

cells, scale equals 0.5 cm.

260

PAN-PACIFIC ENTOMOLOGIST

shriveled wasp egg. The egg in the uppermost cell failed to develop, but the

chrysidid larvae were successfully reared to the adult stage within gelatin

capsules supplied with some of the caterpillars. Dr. R. M. Bohart identified

the chrysidids as Chrysis coerulans Fabricius. Chrysis coerulans is a common

widespread enemy of many eumenid wasps, but this is the first record of it

from A. w. flavidulus.

Rollin E. Coville, Division of Entomology and Parasitology, 201 Wellman

Hall, University of California, Berkeley 94720.

PAN-PACIFIC ENTOMOLOGIST

July 1982, Vol. 58, No. 3, p. 260

Published 16 December 1983

SCIENTIFIC NOTE

ADDITIONAL RECORDS OF THYMELICUS LINEOLA

(OCHSENHEIMER) IN BRITISH COLUMBIA

(LEPIDOPTERA: HESPERIIDAE)

Bums (1966, Can. Entomol., 98:859-866) summarized the currently known

distribution of Thymelicus lineola (Ochsenheimer) in North America. This

Palearctic species was first found in North America in 1910. Since its first

introduction at London, Ontario, it has spread to much of temperate eastern

North America.

The only western North American population was collected at Terrace,

British Columbia in 1960. Since then repeated efforts by resident collectors

to find this species, especially at Smithers, British Columbia, have failed.

This last summer D. Threatful sent me a series of eleven males of T.

lineola from 2 miles east of Sicamous, British Columbia, collected June 19,

1980.1 encouraged him to return to the locale where on July 12 he observed

“several hundred specimens” of both sexes. Further collecting on July 12

showed that the species was present in ever decreasing numbers east from

Sicamous to Solsqua, Cambie, Malakwa, Craigellachie, Taft, and Three Val¬

ley. At Revelstoke, where Mr. Threatful lives, the species has not been seen.

When Mr. Threatful last collected at Sicamous in 1970 he did not observe

this species.

Thus it would appear that this hay and pasture lands pest is well established

in western North America and can be expected to eventually spread over a

wide area.

Jon H. Shepard, R.R. no. 2, Nelson, British Columbia, Canada V1L 5P5.

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ciety, Memoir No. 4. Published August 1972.

This unique book, written by one of the outstanding entomologists of our time,

relates personal experiences and episodes from boyhood through his professional

career. Memories spanning a life of service to the University of California cover a

period of over thirty years, and these memories also include incidents from World

War II, details of research expeditions around the world, a fascinating account of the

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Vol. 58

OCTOBER 1982

No. 4

THE

Pan-Pacific Entomologist

In Memoriam

PAUL DAVID HURD, JR.

1921-1982

KROMBEIN and LINSLEY-Paul David Hurd, Jr., 1921-1982 . 262

HAFERNIK—Mimetic Selection and Subspecific Variation in the Spicebush Swallowtail,

Papilio troilus Linnaeus (Lepidoptera: Papilionidae). 278

PARKER and GRISWOLD—Biological Notes on Andrena ( Callandrena ) haynesi

Viereck and Cockerell (Hymenoptera: Andrenidae). 284

O’BRIEN— Trypargilum tridentatum (Packard) in Trap Nests in Oregon (Hymenoptera:

Sphecidae: Trypoxylinae). 288

EHLER—Observations on Trialeurodes packardi (Morrill) in a Community Garden

(Homoptera: Aleyrodidae). 291

WHARTON—The Biology of Apiocera haruspex Osten Sacken (Diptera: Apioceridae)

in Central California, and Comparison with Other Asiloidea. 296

PARRELLA—A Review of the History and Taxonomy of Economically Important

Serpentine Leafminers ( Liriomyza spp.) in California (Diptera: Agromyzidae). 302

LANHAM—A New Species of Andrena (Micrandrena ) from Colorado, with Other Notes

on the Group (Hymenoptera: Apoidea). 309

DOWELL—Biology of Tetraleurodes acaciae (Quaintance) (Homoptera: Aleyrodi¬

dae). 312

RUST and HANKS—Notes on the Biology of Aegialia hardyi Gordon and Cartwright

(Coleoptera: Scarabaeidae). 319

(continued on back cover)

SAN FRANCISCO, CALIFORNIA • 1982

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

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G. A. Marsh, Editor

C. Pleines, Editorial Assistant

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Manuscripts, proofs, and all correspondence concerning editorial matters should

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

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Pacific Coast Entomological Society

OFFICERS FOR 1982

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E. L. Smith, President-Elect V. F. Lee, Secretary

Statement of Ownership

Title of Publication: The Pan-Pacific Entomologist.

Frequency of Issue: Quarterly (January, April, July, October).

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

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This issue mailed 30 December 1983

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The Pan-Pacific Entomologist (ISSN 0031-0603)

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

Paul David Hurd, Jr., 1921-1982

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 262-277

Published 30 December 1983

Paul David Hurd, Jr.

1921-1982

Karl V. Krombein 1 and E. Gorton Linsley 2

Paul David Hurd, Jr., son of Paul David Hurd and Ruth Dorothea Bick,

was born in Chicago, Illinois, on April 2, 1921, and died of a heart attack

on March 12, 1982, at George Washington University Hospital, Washington,

D.C.

In his early days his family moved to an extremely modest home on the

Mojave Desert in California. Neither of his parents had had much formal

education and were unable to encourage Paul’s early interest in natural

history nor to provide any financial support for his education. Nevertheless,

he attended Colton Union High School, Colton, California, transferring to

Newport Harbor Union High School, Newport Beach, California. One of

his teachers took a great interest in Paul and helped foster his interest in

science. During this period he began collecting insects and plants and study¬

ing birds. It was through his study of birds that he met Josephine R. Mich-

ener, mother of the distinguished entomologist-biologist, Charles D. Mich-

ener, at meetings of the Western Bird-banding Association and he stopped

at her home when he passed through Pasadena, once or twice a year. She

obviously enjoyed his visits, frequently wrote or told Charles about them,

and regarded him as an unusually pleasant and interesting young man. They

discussed ornithological matters, birds that they had seen or banded, etc.

She encouraged him and tried successfully to influence him to continue his

interest in natural history. She thought he might go on to a scientific career

in vertebrate zoology (ornithology), but she told Charles that she thought

he was even more interested in a career in entomology. However, his first

published paper, reporting the results of a bird census of Newport Upper

Bay, appeared in Audubon Magazine in 1941, a later paper in 1947. In the

late 1940’s he conducted bird-banding studies in Berkeley, on the Univer¬

sity’s Oxford Tract, in collaboration with a fellow student, H. E. Childs, Jr.

They coauthored a short paper describing a new multiple-catch bird-trap.

Although Paul subsequently devoted his primary efforts to entomology, he

maintained an interest in birds throughout his life. It is interesting to note

that although Charles Michener had been hearing about Paul from his mother

for years, he has no clear personal memory of him until he turned up at

Berkeley among entomology students when he, Michener, was a graduate

assistant.

264

PAN-PACIFIC ENTOMOLOGIST

Paul entered the University of California at Berkeley in 1940 with a major

in entomology, paying his way from whatever work he could find locally.

In 1942, he withdrew for war service, joining the United States Navy. He

served in the South Pacific as a Chief Pharmacist’s Mate, using periods of

shore leave for collecting insects in the Solomon Islands. He was wounded

in one of the naval engagements in the area and was awarded the Purple

Heart.

Upon returning to Berkeley in 1946, he resumed work toward the Bachelor

of Science degree and served briefly as a preparator for the departmental

insect collections. As a graduate student working for his M.S. degree (1948)

and a Ph.D. (1950) he held various teaching and research assistantships. In

1950 he was added to the staff as Senior Museum Entomologist. There, his

enthusiastic drive and technical skills enabled him to make substantial im¬

provements in the departmental collection.

During the summers of 1952 and 1953, he worked at Barrow, Alaska, as

a member of a research team led by Professor Frank A. Pitelka and sponsored

by the Office of Naval Research. Here Paul’s expertise in bird-banding proved

critical in studies of breeding and molt of two passerines, the Lapland Long-

spur and Snow Bunting. But his main duties were entomological—moni¬

toring soil surface insect populations to define seasonal trend of change;

examining stomach contents of longspurs, buntings and phalaropes to iden¬

tify prey species and estimate their relative importance; and updating the

knowledge of local insect fauna. This last service proved of general value,

as even into the years of the International Biological Program for tundra

(1969-1973), his faunal summary helped to guide field work in the Barrow

area.

While at Barrow, much of Paul’s sampling was done by means of an

aspirator. About two months after returning from his last trip he became ill

and during the next week living Coleoptera, Collembola, Diptera and Hy-

menoptera were passed from the left antrum of the sinus. An account of this

remarkable and unpleasant experience was reported in Science (No. 20 of

the appended bibliography).

In 1954, Paul was appointed Junior Entomologist in the California Ag¬

ricultural Experiment Station to be responsible for the Berkeley project en¬

titled, “The California Insect Survey” (now housed in the “Essig Museum

of Entomology”). Paul later was given teaching as well as research titles and

in 1965, before leaving Berkeley he had attained the rank of Professor of

Entomology and Entomologist in the Experiment Station.

Early in this period (1956) Paul and Ray F. Smith of the Department of

Entomology and Parasitology, Berkeley, proposed to the University’s As¬

sociates in Tropical Biology, a program of investigation of fossiliferous amber

in Chiapas, Mexico. The site had been reported to Smith by Franz Blom

VOLUME 58, NUMBER 4

265

and subsequently visited by graduate students. As a result, Paul and J. Wyatt

Durham, Professor of Paleontology, visited the amber sites on a mule trip

of 10 days that year under the guidance of Gertrude Blom, since Franz was

ill at the time. Accounts of this trip were published by Mrs. Blom in Pacific

Discovery, Vol. 10, pp. 8-14, under the title “On the Amber Trail in Chiapas”

and the scientific results were summarized by Hurd, Smith and Durham

(No. 55) as well as elsewhere. Smith had made a visit to Chiapas with Paul

shortly after the first trip with Durham. According to Durham, Paul super¬

vised much of the preparation of the contained fossil insects for study,

assigned them to their proper taxonomic categories, and distributed them

to specialists for study. The collection is maintained in the Museum of

Paleontology, University of California, Berkeley.

Hurd’s contributions to the academic program at Berkeley were many and

varied. The courses he taught or participated in at various times ranged

from an introduction to the natural history and classification of insects to

graduate research and seminars, as well as contributions to interdepartmental

courses in biological sciences. Graduate students who received the Ph.D.

degree under his supervision were Clarence D. Johnson, Evert E. Lindquist,

Lois B. O’Brien, Gerald I. Stage, Wallace A. Steffan, and Marius S. Wasbauer.

An important aspect of Paul’s entomological activities at Berkeley was his

work with students in the field. For many years he was in charge of the

summer field course in entomology and frequently took collecting parties to

various parts of the state in connection with his duties as head of The

Agricultural Experiment Station California Insect Survey Project. He also

led field trips of graduate and undergraduate students during the spring

season to collect insects from various areas, particularly in the Mojave and

Colorado deserts. These trips not only supplied excellent experience for the

students but helped to provide the California Insect Survey with one of the

finest collections of southwestern desert insects in existence and the best

collection of California material housed anywhere. His own research projects

not only involved these areas but took him into Mexico, Central and South

America, frequently with support from the National Science Foundation.

On campus, Paul was active on a large number of administrative com¬

mittees both in the department and elsewhere, among the most significant

of which was the Chancellor’s Advisory Committee on Landscape Planning,

which he chaired, and which had a profound influence in preserving natural

areas and siting new buildings with a minimum of environmental impact.

He also served on the statewide faculty committee advisory to the University

Press. As a member of the 75th Anniversary Committee of the University,

he introduced novel exhibit ideas, many of which are still evident.

His first sabbatical leave (1959-1960) was spent at Curitiba, Brazil, where

he worked with Padre I. S. Moure, with the support of awards from the

Guggenheim Foundation and Fulbright Commission. This experience not

266

PAN-PACIFIC ENTOMOLOGIST

only cemented a strong friendship between the two but initiated cooperative

research programs which continued until his death.

In 1967, Paul took leave from the University to serve for two years as

Associate Program Director, Systematic Biology, Division of Biological and

Medical Sciences, National Science Foundation. Harve Carlson, who was

then Director of the Division at the Foundation, recalls that Paul was soon

integrated into the work patterns of the program and took over immediate

evaluation of proposals in his area of expertise. His quiet and easy manner

allowed him to enjoy the respect of the Division staff immediately. According

to Dr. Carlson, Paul did an outstanding job during his period of service.

The Program Directors and Division office tried hard but unsuccessfully to

recruit him on a permanent basis, but being a dedicated scientist whose first

love was research, he declined although tempted.

In August, 1970, Paul left Berkeley to accept an appointment as Curator

of Apoidea in the Department of Entomology, National Museum of Natural

History, Smithsonian Institution, Washington, D.C. In July of the following

year he was selected as the third chairman of the department for a 5-year

term. Although departmental affairs always had first priority on his time,

he continued his research program vigorously, spending available free time

at the Museum and many hours at home in connection with the apoid section

of the Hymenoptera catalog (No. 91). Working closely and cooperatively

with the departmental scientific staff, he effected a reorganization which

improved departmental operations. The first was the elimination of major

divisions, such as Lepidoptera, Coleoptera and Neuropteroid orders. This

freed those division leaders of subsidiary administrative duties and allowed

them more time for research, curation and other Museum responsibilities.

Another improvement was the transformation of the existing technicians’

pool into a collections management unit to which were assigned a collections

manager and several permanent and part-time technicians. This resulted in

a reduction of the large backlog of unprocessed lots of insects as well as an

improved capability for handling new incoming accessions. Paul suggested

and helped develop an insect zoo as a temporary museum installation early

in 1971. Later, during his chairmanship but under the supervision of T. L.

Erwin, the zoo evolved into a permanent exhibit, one of the most popular

within the Museum.

Paul’s systematic and biological interests were broad, as reflected in the

appended bibliography. While a student at Berkeley, his early entomological

concern was primarily with wasps and his doctoral dissertation was on the

California species of Pepsis, an interest that he pursued to a culmination in

a revision of the Nearctic species. As his studies of aculeate Hymenoptera

expanded, he gave more and more attention to bees.

In the field of systematics, if one had to select a most consistent focus, it

would be on the carpenter bees (Xylocopinae). From a 1954 paper on a

VOLUME 58, NUMBER 4

267

polytypic interpretation of the California carpenter bee (No. 21), followed

a year later by a treatment of the carpenter bees of California (No. 23), he

proceeded to publish 12 more papers on this group of bees, significantly

correlating biological characters with more traditional methods, before join¬

ing Padre Moure to produce in 1963 the classical book entitled, “A Clas¬

sification of the Large Carpenter Bees ” (No. 60). This publication is not only

a landmark in the study of this group of bees, but a model of thoroughness

and meticulous attention to detail that has not been equalled in any major

apoid group. This treatment of the higher classification of the carpenter bees

of the world was followed in 1978 by an annotated catalog of the carpenter

bees of the Western Hemisphere (No. 89) —again a classic model for an

insect group, which brought together in general form for the genus, and

annotated form for the species, all the references and known information

about the systematics, biology and morphology of the group. Although his

shortened life did not permit him to complete career-long interests in the

parasitic bees and megachilids, he was able to complete with colleagues

shorter but equally comprehensive studies of smaller groups, such as the

Oxaeidae (No. 88).

Paul’s longtime interest in the flower relationships of bees was brought to

focus by a National Science Foundation grant in support of a research pro¬

gram on the systematics, intrafloral ecology and evolution of squash and

gourd bees ( Peponapis and Xenoglossa ) and their host plants of the genus

Cucurbita. With the assistance of colleagues he pursued this study throughout

their home areas in the Western Hemisphere south to Argentina, the results

of which helped clarify the origins of domestic squash and the evolutionary

history of wild species. In an attempt to introduce some of these bees to the

Hawaiian Islands, he and A. E. and M. M. Michelbacher flew chilled adults

to the islands of Oahu and Hawaii (No. 79), and provided larvae and pupae

for introductions to Australia. The results of these efforts are not yet known.

Other important studies involving pollination ecology, included the creo¬

sote bush ( Larrea trident at a) (No. 85, 86), now assuming significance in the

management and development of resources in the arid southwest, and sun¬

flower pollinators (No. 95), critical in the rapidly expanding crop production

of the cultivated forms of Helianthus. All of these publications have been

widely quoted in the literature of general biology as well as that of agriculture.

Late in 1970 Hurd and K. V. Krombein discussed the desirability of

producing a new edition of the synoptic catalog of Hymenoptera in America

north of Mexico (No. 9). They decided that the data base should be entered

into the computer for several reasons. The catalog could be produced on the

GPO Linotron from a special magnetic tape, and the data base could be

manipulated to produce indexes in alphabetical order. Computerization would

also permit manipulation of the data base to answer specific inquiries, and

it could be expanded and updated continually to provide revised editions

268

PAN-PACIFIC ENTOMOLOGIST

of the catalog or parts thereof as needed. Hurd, together with Krombein and

B. D. Burks, U.S. Department of Agriculture (later replaced by D. R. Smith,

USD A) constituted the Editorial Board which established the protocol for

the catalog and edited the sections prepared by the combined Smithsonian-

Agriculture hymenopterists. Their editorial duties began in mid-1971 and

finally terminated in 1979 with completion of the third volume of indexes.

Paul was responsible for preparation of the section on Apoidea (No. 91)

which required a large part of his available research time through 1976. His

section has received critical acclaim from other bee specialists and from

botanists working on pollination ecology. Subsequently, the Apoidea data

base provided much useful information for the large work on sunflower bees

(No. 95) undertaken jointly by Hurd, W. E. LaBerge and E. G. Linsley.

Because of his expertise in computer techniques, Paul began a series of

annotated catalogs on the New World bees. The first on carpenter bees (No.

89) was by him alone. The second part with J. S. Moure on Halictidae has

been completed except for the indexes which will be produced by computer

manipulation. Hurd and Moure planned to complete the entire catalog in

ten additional parts, but Paul had made no progress on these at the time of

his death.

Several other projects are in press or nearing completion. Among these is

a study of the bee pollinators of selected trees and shrubs of the southwestern

deserts undertaken with E. G. Linsley and T. J. Zavortink. He also had in

progress a revision of the cleptoparasitic bee genus Triepeolus. He had ex¬

amined all readily available material, sorted it to species with names applied

to all new taxa, and had a number of illustrations prepared by a staff artist.

Paul had not begun to write up this study, but it is hoped that it can be

completed by one of his colleagues.

Many other aspects of his research deserve mention, but space will not

permit a fuller analysis, and the reader is referred to the appended bibli¬

ography for items of interest.

Paul belonged to a large number of professional organizations, many of

which he served in an appointed or elected capacity. For example, as a

member of the Pacific Coast Entomological Society, he edited the Pan-Pacific

Entomologist for several years. In the Entomological Society of America,

among other functions, he served on the Governing Board and chaired the

important Advisory Committee for Systematics Resources in Entomology.

He was President of the Association for Tropical Biology in 1969-70, and

co-Chairman of the Program Committee for the First International Congress

of Systematic and Evolutionary Biology for which he received the Congress

Medal.

As might be expected, Paul was a member of several honor societies

including Sigma Xi. He was also a Fellow of the American Association for

the Advancement of Science and of the California Academy of Sciences.

VOLUME 58, NUMBER 4

269

Among his numerous professional activities, he served as Section Editor

(Hymenoptera) for Biological Abstracts.

The Smithsonian Institution recognized his stature as an entomologist of

world renown because of the superb quality of his scientific contributions.

He was promoted to supergrade status in the Federal Civil Service in 1978,

the third entomologist on the Smithsonian staff to be so honored. In 1980

he was appointed a Senior Scientist, one of only five in the Museum of

Natural History.

Paul was elected to membership in the prestigious and exclusive Cosmos

Club shortly after moving to Washington, and later he served for several

years on the Admissions Committee. He greatly enjoyed his membership in

this organization of men who have done meritorious original work in science,

literature or the arts. Numerous small and even more exclusive groups of

members have formed within the Cosmos Club. One such is the Friday

Morning Cheese Group with membership limited to the first three depart¬

mental chairmen (J. F. G. Clarke, Krombein and Hurd). Every Friday those

of us in residence went to the club for cocktails, cheese and crackers, followed

by an advisedly light lunch!

He was also a member of the much smaller Washington Biologists’ Field

Club with headquarters on Plummers Island in the Potomac River. He and

Krombein collected there during 1971 and 1972, anticipating eventually that

they would publish an annotated list of the bees, comparing the recent fauna

with that collected more than 50 years earlier. Regrettably, this project was

put in abeyance because of Hurd’s many other commitments.

Paul suffered a severe heart attack in 1980, and was hospitalized for several

months. After an extended period of testing various drugs, his cardiologist

eventually found a combination that reduced his pressure to acceptable

limits. Unfortunately the drug therapy had some adverse side effects, so that

Paul became moody and very withdrawn during his last months. The side

effects fortunately did not diminish his research interests or capability, and

he drove himself relentlessly and incessantly, perhaps realizing that his time

was limited. His fatal heart attack, from which he never regained conscious¬

ness, came at 5:20 a.m., just after he entered the Museum on Friday March

12 .

He is survived by his wife, Grace Isabelle, a son, Philip James, and a

daughter, Mrs. Katherine Lee Hartfield, all now of Austin, Texas, and by a

son from an earlier marriage, Rodney Wayne of Berkeley, California.

We are grateful to Charles D. Michener for contributing reminiscences of

Paul’s early work with birds, to Frank A. Pitelka for information on Paul’s

work at Barrow, Alaska, to J. Wyatt Durham and Ray F. Smith for infor¬

mation about the Chiapas Amber Project, and to Harve J. Carlson for

recollections of Paul’s service with the National Science Foundation. We

270

PAN-PACIFIC ENTOMOLOGIST

are also indebted to Michener and Jerome G. Rozen for their helpful com¬

ments on a draft of this biographical sketch.

List of Taxa Named for Paul D. Hurd, Jr.

Paul was honored by a number of systematists who gave new taxa a specific

or subspecific name based on his last or first name. Many of these taxa had

been collected by Hurd himself. Such patronyms known to Paul are as

follows:

COPEPODA

Parastenocaris hurdi Jakobi and de Loyola e Silva, 1962.

ACARINA

Amphicalvolia hurdi Turk, 1963.

Bdellodes hurdi Atyeo, 1960.

Charadracarus hurdi Newell, 1960.

Oppia hurdi Woolley, 1971.

HEMIPTERA

Dysmicoccus hurdi McKenzie, 1962.

Phenacoccus hurdi McKenzie, 1964.

COLEOPTERA

Anthaxia hurdi Cobos, 1949.

Cenocephalus hurdi Schedl, 1962.

Crossidius hurdi Chemsak and Linsley, 1959.

Cryptorhynchus hurdi Zimmerman, 1971.

Dactylozodes hurdi Cobos, 1976.

Nemognatha hurdi MacSwain, 1951.

Rhipiphorus luteipennis hurdi Linsley and MacSwain, 1952.

DIPTERA

Aphantorhapha hurdi Reinhard, 1959.

Apiocera ammophila hurdi Cazier, 1982.

Brunnettia hurdi Quate, 1950.

Cophura hurdi Hull, 1960.

Forcipomyia hurdi Wirth, 1952.

Lordotus hurdi Hall, 1957.

Metapogon hurdi Wilcox, 1964.

Pholeomyia hurdi Sabrosky, 1959.

Phyllolabis hurdi Alexander, 1964.

Procolobstema hurdi Cook, 1971.

VOLUME 58, NUMBER 4

271

HYMENOPTERA

Agathilla hurdi Townes, 1967.

Ammophila hurdi Menke, 1963.

Andrena hurdi Lanham, 1949.

Anthocopa hurdiana Michener, 1954.

Brachycistis lacustris hurdi Wasbauer, 1966.

Calliopsis hurdi Shinn, 1967.

Ceratina hurdi Daly, 1974.

Cerceris hurdi Scullen, 1972.

Chelonus hurdi McComb, 1968 (1967).

Dipogon hurdi Evans, 1974.

Episyron quinquenotatus hurdi Evans, 1950.

Heterostelis hurdi Thorp, 1966.

Hylaeus hurdi Snelling, 1966.

Larropsis hurdi Bohart and Bohart, 1962.

Melissodes hurdi LaBerge, 1961.

Melissodes paulula LaBerge, 1961.

Microdynerus hurdi Parker, 1970.

Neolarra hurdi Shanks, 1978.

Nomada hurdi Evans, 1972.

Nomadopsis cincta hurdi Rozen, 1958.

Osmia hurdi White, 1952.

Oxybelus hurdi Bohart and Schlinger, 1956.

Paratiphia hurdi Allen, 1965.

Perdita hurdi Timberlake, 1956.

Plenoculus hurdi Williams, 1960.

Priocnessus hurdi Dreisbach, 1960.

Protandrena hurdi Timberlake, 1976.

Pseudisobrachium hurdi Evans, 1961.

Psorthaspis macronotum hurdi Evans, 1954.

Pterocheilus hurdi Bohart, 1950.

Scolia ribbei hurdi Krombein, 1963.

Synhalonia hurdi Timberlake, 1969.

Tachysphex hurdi Bohart, 1962.

Tiphia hurdi Allen, 1965.

Bibliography of Scientific Contributions by

Paul D. Hurd, Jr.

Joint authorship is indicated parenthetically following the complete ci¬

tation, e.g. (With W. F. Barr.) as in No. 2.

1. 1941. Forty-first Christmas bird census. Newport Upper Bay. Audubon Mag.,

43:145.

272

PAN-PACIFIC ENTOMOLOGIST

2. 1947a.

3. 1947b.

4. 1947c.

5. 1948.

6. 1950a.

7. 1950b.

8. 1950c.

9. 1951a.

10. 1951b.

11. 1951c.

12. 195 Id.

13. 195 le.

14. 1952a.

15. 1952b.

16. 1953.

17. 1954a.

18. 1954b.

19. 1954c.

20. 1954d.

21. 1954e.

22. 1955a.

Notes on the Dasymutilla of the Palo Verde Valley, California with the

description of a new species (Hymenoptera: Mutillidae). Pan-Pac. Ento-

mol., 23:85-90. (With W. F. Barr.)

Redescription of Agenioideus humilis (Cresson) with notes on its biology

(Hymenoptera: Pompilidae). Pan-Pac. Entomol., 23:132-134.

Unusual winter visitants to Berkeley, California. Condor, 49:173.

Systematics of the California species of the genus Pep sis Fabricius (Hy¬

menoptera: Pompilidae). Univ. Calif. Publ. Entomol., 8:123-150, 4 figs.

Claude “I” Smith, 1922-1949. Pan-Pac. Entomol., 26:132-133.

Nomenclatorial notes on the genus Pepsis (Hymenoptera: Pompilidae).

Pan-Pac. Entomol., 26:132-133.

Some insects associated with nests of Dianthidium dubium dilectum

Timberlake, with a list of the recorded parasites and inquilines of Dian¬

thidium in North America. J. N.Y. Entomol. Soc., 58:247-250. (With

E. G. Linsley.)

Genus Pepsis Fabricius. In Muesebeck, Krombein and Townes, Hy¬

menoptera of America north of Mexico-Synoptic catalog. U.S. Dept.

Agric., Agric. Monogr., 2:909-910.

The history of the Pacific Coast Entomological Society. Pan-Pac. Ento¬

mol., 27:97-119, 4 figs. (With E. O. Essig et al.)

The California velvet ants of the genus Dasymutilla Ashmead (Hyme¬

noptera: Mutillidae). Bull. Calif. Insect Surv., 1:89-118, 1 pi.

The female of Pseudomethoca anthracina (Fox) (Hymenoptera: Mutil¬

lidae). Pan-Pac. Entomol., 27:156.

The melectine bees of California (Hymenoptera: Anthophoridae). Bull.

Calif. Insect Surv., 1:119-135, 1 pi., 5 maps. (With E. G. Linsley.)

The Scoliidae of California (Hymenoptera: Aculeata). Bull. Calif. Insect

Surv., 1:141-152, 2 pis.

Revision of the Nearctic species of the pompilid genus Pepsis (Hymen¬

optera: Pompilidae). Bull. Am. Mus. Nat. Hist., 98:257-334, 49 figs., 2

tabs.

Notes on the melectine bees of Mexico (Hymenoptera: Anthophoridae).

J. Kans. Entomol. Soc., 26:35-37, 1 fig.

Distributional notes on Eutricharaea, a Palearctic subgenus of Mega¬

chile, which has become established in the United States (Hymenoptera:

Megachilidae). Entomol. News, 65:93-95.

Monodontomerus montivagus Ashmead, a parasite of Megachile cen-

tuncularis (Linnaeus) (Hymenoptera). Pan-Pac. Entomol., 30:146. (With

A. E. Michelbacher.)

California insects—Survey provides accurate data for study of state’s

insect problems. Calif. Agric., 8(5):3, 1 map.

“Myiasis” resulting from the use of the aspirator method in the collection

of insects. Science, 119:814-815.

A polytypic interpretation of the California carpenter bee Xylocopa cal-

ifornica with the description of a new subspecies and notes on a possible

polytopic form. Pan-Pac. Entomol., 30:199-202, 1 fig.

The megachiline bees of California (Hymenoptera: Megachilidae). Bull.

VOLUME 58, NUMBER 4

273

Calif. Insect Surv., 3:1-248, 141 figs., 1 tab., 112 maps. (With C. D.

Michener.)

23. 1955b. The carpenter bees of California (Hymenoptera: Apoidea). Bull. Calif.

Insect Surv., 4:35-72, 4 pis., 4 maps.

24. 1955c. The Aculeate wasps. Pp. 573-575. In: A Century of Progress in the

Natural Sciences, 1853-1953. California Academy of Sciences, San

Francisco, pp. i-x, 1-807.

25. 1955d. Carpenter bees as a pest in California. Pest Control Rev., December,

p. 2.

26. 1956a. Xylocopa rufina utilizing Mexican cedar timbers for nesting purposes

(Hymenoptera: Apoidea). Pan-Pac. Entomol., 32:28.

27. 1956b. Packaging and shipping of specimens for identification. Pest Control

Rev., April, pp. 4-5.

28. 1956c. Notes on the subgenera of the New World carpenter bees of the genus

Xylocopa (Hymenoptera: Apoidea). Am. Mus. Novit., 1776:1-7, 1 fig.

29. 1956d. A new multiple-catch bird trap. News from the Bird Banders, 31:52-

53. (With H. E. Childs.)

30. 1956e. New host records for North American spider-wasps (Hymenoptera:

Pompilidae). J. Kans. Entomol. Soc., 29:168-169. (With M. S. Was-

bauer.)

31. 1957a. Notes on the autumnal emergence of the vernal desert bee, Hesperapis

fulvipes Crawford (Hymenoptera: Apoidea). J. Kans. Entomol. Soc., 30:

10 .

32. 1957b. The meaning of Mexico’s amber. Pac. Discovery, 10(2):6-7, illustr. (With

R. F. Smith.)

33. 1957c. Fossiliferous amber of Chiapas, Mexico. Bull. Geol. Soc. Am., 68:1824.

(With J. W. Durham.)

34. 1957d. Stack beetles at southern California cement plants. Pit and Quarry,

49(11): 168, 208. (With E. G. Linsley.)

35. 1957e. Melanophila beetles at cement plants in southern California (Coleoptera,

Buprestidae). Coleopt. Bull., 11:9-11. (With E. G. Linsley.)

36. 1957f The conopid flies of California (Diptera). Bull. Calif. Insect Surv., 6:19-

50, 4 figs., 25 maps. (With S. Camras.)

37. 1958a. New fossil find. Nat. Hist., 67:223-224, illustr.

38. 1958b. American bees of the genus Dioxys Lepeletier and Serville (Hymenop¬

tera: Megachilidae). Univ. Calif. Publ. Entomol., 14:275-302, 28 figs.,

4 maps.

39. 1958c. Observations on the nesting habits of Colletes stepheni Timberlake (Hy¬

menoptera: Apoidea). Pan-Pac. Entomol., 34:147-153, 1 fig. (With J.

A. Powell.)

40. 1958d. Observations on the nesting habits of some New World carpenter bees

with remarks on their importance in the problems of species formation.

Ann. Entomol. Soc. Am., 51:365-375, 5 figs.

41. 1958e. The carpenter bees of the Eastern Pacific Oceanic Islands (Hymenoptera:

Apoidea). J. Kans. Entomol. Soc., 31:249-255, 4 figs.

274

PAN-PACIFIC ENTOMOLOGIST

42. 1958f.

43. 1959a.

44. 1959b.

45. 1959c.

46. 1959d.

47. 1959e.

48. 1960a.

49. 1960b.

50. 1961a.

51. 1961b.

52. 1961c.

53. 1961d.

54. 1961e.

55. 1962.

56. 1963a.

57. 1963b.

Cretaceous and Tertiary insects in Arctic and Mexican amber. Proc.

Tenth Int. Congr. Entomol. (1956), 1:856.

The larval habits of Plinthocoelium suaveolens plicatum (LeConte) (Co-

leoptera: Cerambycidae). Bull. South. Calif. Acad. Sci., 58:27-33, 2 pis.

(With E. G. Linsley.)

Ethological observations on some bees of southeastern Arizona and New

Mexico (Hymenoptera: Apoidea). Entomol. News, 70:63-68. (With E.

G. Linsley.)

Beefly parasitism of the American carpenter bees belonging to the genus

Xylocopa Latreille (Hymenoptera: Apoidea). I. Kans. Entomol. Soc., 32:

53-58, 1 hg.

Observations on the nest-site behavior of Melissodes composita Tucker

and its parasites, with notes on the communal use of nest entrances

(Hymenoptera: Apoidea). Entomol. News, 70:141-146. (With E. G.

Linsley.)

Some nomenclatorial problems in the genus Xylocopa Latreille (Hy¬

menoptera: Apoidea). Pan-Pac. Entomol., 35:135-148.

On the systematic position of three anthidiine bees described by Adolfo

Ducke (Hymenoptera: Apoidea). Bol. Mus. Para. Emilio Goeldi (n. s.),

28:1-13, 5 figs. (With J. S. Moure.)

A New World subgenus of bamboo-nesting carpenter bees belonging to

the genus Xylocopa Latreille (Hymenoptera: Apoidea). Ann. Entomol.

Soc. Am., 53:809-821, 14 figs., 2 tabs. (With J. S. Moure.)

Some notes on sapygid parasitism in the nests of carpenter bees belonging

to the genus Xylocopa Latreille (Hymenoptera: Aculeata). J. Kans. Ento¬

mol. Soc., 34:19-22, 1 fig. (With J. S. Moure.)

Lestis Lepeletier and Serville, 1828 (Insecta, Hymenoptera); proposed

designation of a type-species under the plenary powers. Z. N. (S.) 1383.

Bull. Zool. Nomencl., 18:201-202. (With C. D. Michener.)

Systematics of the carpenter bee types (Genus Xylocopa Latreille) con¬

tained in the collections of the Museo Argentino de Ciencias Naturales

“Bernardino Rivadavia,” Buenos Aires (Hymenoptera: Apoidea). J. Kans.

Entomol. Soc., 34:181-195, 12 figs. (With J. S. Moure.)

Some problems of species formation in the carpenter bee genus Xylocopa

Latreille (Hymenoptera: Apoidea). Act. Trabajos Primer Congr. Suda-

mer. Zool., 3:77-82. (With J. S. Moure.)

A synopsis of the carpenter bees belonging to the subgenus Xylocopoides

Michener (Hymenoptera: Apoidea). Trans. Am. Entomol. Soc., 87:247-

257, 2 pis.

The fossiliferous amber of Chiapas, Mexico. Ciencia, 21:107-118, 2 pis.

(With R. F. Smith and J. W. Durham.)

Xilocopa Latreille, 1802 (Insects, Hymenoptera): proposed suppression

under the plenary powers. Z. N. (S.) 1527. Bull. Zool. Nomencl., 20:

227-228.

A new subspecies of Xylocopa tabaniformis Smith from Mexico (Hy-

VOLUME 58, NUMBER 4

275

menoptera: Apoidea). Pan-Pac. Entomol., 39:275-276. (With L. B.

O’Brien.)

58. 1963c. Pollination of the unicorn plant (Martyniaceae) by an oligolectic, corolla¬

cutting bee (Hymenoptera: Apoidea). J. Kans. Entomol. Soc., 36:248-

252. (With E. G. Linsley.)

59. 1963d. A contribution to the systematics of the bee genus Protepeolus Linsley

and Michener (Hymenoptera: Apoidea). J. Kans. Entomol. Soc., 36:

253-255. (With E. G. Linsley.)

60. 1963e. A classification of the large carpenter bees (Xylocopini) (Hymenoptera:

Apoidea). Univ. Calif. Publ. Entomol., 29:vi + 365 pp., 244 figs., fron-

tisp. (With J. S. Moure.)

61. 1964a. Pollination of squashes, gourds and pumpkins. Calif. Agric., 18(5):2—4,

3 figs. (With A. E. Michelbacher and R. F. Smith.)

62. 1964b. The squash and gourd bees—genera Peponapis Robertson and Xeno-

glossa Smith—inhabiting America north of Mexico (Hymenoptera:

Apoidea). Hilgardia, 35:375-477, frontisp., 18 figs. (With E. G. Linsley.)

63. 1965a. Carpenter bees of the subgenus Notoxylocopa (Hymenoptera: Apoidea).

Ann. Entomol. Soc. Am., 58:175-196, 38 figs. (With L. B. O’Brien.)

64. 1965b. A new subgenus and species of matinal Andrena from the flowers of

Sicyos (Cucurbitaceae) in Mexico (Hymenoptera: Apoidea). Pan-Pac.

Entomol., 41:186-193, 5 figs. (With W. E. LaBerge.)

65. 1966a. Problems of bee distribution in Western North America. In: Distribu¬

tional patterns of selected Western North American Insects. Bull. Ento¬

mol. Soc. Am., 12:110-111.

66. 1966b. The Mexican squash and gourd bees of the genus Peponapis (Hyme¬

noptera: Apoidea). Ann. Entomol. Soc. Am., 59:835-851, 12 figs., 8

maps. (With E. G. Linsley.)

67. 1966c. The pollination of pumpkins, gourds and squashes (Genus Cucurbita).

Bee World, 47(Suppl.):97-98.

68. 1967a. The identity of Megachile rotundata (Fabricius) and M. argentata (Fa-

bricius) (Hymenoptera: Apoidea). Entomol. Medd., 35:3-10.

69. 1967b. South American squash and gourd bees of the genus Peponapis (Hy¬

menoptera: Apoidea). Ann. Entomol. Soc. Am., 60:647-661, 6 figs., 2

tabs. (With E. G. Linsley.)

70. 1967c. Squash and gourd bees of the genus Xenoglossa (Hymenoptera: Apoi¬

dea). Ann. Entomol. Soc. Am., 60:988-1007, 19 figs., 5 maps, 2 tabs.

(With E. G. Linsley.)

71. 1967d. The Mexican squash and gourd bees of the genus Peponapis (Hyme¬

noptera: Apoidea). Folia Entomol. Mex., 15/16:40-41.

72. 1968a. Intrafloral ecology. Annu. Rev. Entomol., 13:385-414. (With H. G.

Baker.)

73. 1968b. Late season foraging activities of Xenoglossa gabbii crawfordi. Pan-Pac.

Entomol., 44:58-68. (With A. E. Michelbacher.)

74. 1968c. The feasibility of introducing squash bees (. Peponapis and Xenoglossa )

into the Old World. Bee World, 49:159-167. (With A. E. Michelbacher

and E. G. Linsley.)

75. 1968d. Notes on the ecology and distribution of the squash bee, Xenoglossa

276

PAN-PACIFIC ENTOMOLOGIST

76. 1970a.

77. 1970b.

78. 1971a.

79. 1971b.

80. 1972a.

81. 1972b.

82. 1974a.

83. 1974b.

84. 1974c.

85. 1975a.

86. 1975b.

87. 1975c.

88. 1976.

89. 1978a.

90. 1978b.

gabbii crawfordi. Folia Entomol. Mex., 18/19:110-111. (With A. E.

Michelbacher.)

A classification of the squash and gourd bees Peponapis and Xenoglossa

(Hymenoptera: Apoidea). Univ. Calif. Publ. Entomol., 62:1-39, 11 figs.,

3 maps, 4 tabs. (With E. G. Linsley.)

A new, narrowly polylectic, autumnal species of Dialictus from the flow¬

ers of Jepsonia heterandra, an endemic California Saxifrage (Hymenop¬

tera: Apoidea). Pan-Pac. Entomol., 46:209-212.

Squash and gourd bees (. Peponapis, Xenoglossa) and the origin of the

cultivated Cucurbita. Evolution, 25:218-234, 5 figs., 3 tabs. (With E.

G. Linsley and T. W. Whitaker.)

Experimental introduction of squash bees ( Peponapis ) to improve yield

of squashes, gourds, and pumpkins. Bee World, 52:156-166, 4 figs., 2

tabs. (With A. E. Michelbacher and E. G. Linsley.)

Parasitic bees of the genus Holcopasites Ashmead (Hymenoptera: Apoi¬

dea). Smithson. Contrib. Zool., 114:1-41, 16 figs., 1 tab. (With E. G.

Linsley.)

A new edition of the catalog of North American Hymenoptera. Taxon,

21:361-362.

Ecology of the squash and gourd bee, Peponapis pruinosa, on cultivated

cucurbits in California (Hymenoptera: Apoidea). Smithson. Contrib.

Zool., 168:1-17, 4 figs., 8 tabs. (With E. G. Linsley and A. E. Michel¬

bacher.)

The status of Nomia mesillensis Cockerell (Hymenoptera: Halictidae).

Proc. Entomol. Soc. Wash., 76:198-199. (With E. G. Linsley.)

Report of the Advisory Committee for Systematics Resources in Ento¬

mology. Bull. Entomol. Soc. Am., 20:237-242, 1 fig., 1 tab. (With R.

L. Fischer et al.)

Some insects other than bees associated with Larrea tridentata in the

southwestern United States. Proc. Entomol. Soc. Wash., 77:100-120.

(With E. G. Linsley.)

The principal Larrea bees of the southwestern United States (Hyme¬

noptera: Apoidea). Smithson. Contrib. Zool., 193:1-74, 18 figs., 15 tabs.

(With E. G. Linsley.)

Report of the Advisory Committee for Systematics Resources in Ento¬

mology. Part II: The current status of entomological collections in North

America. Bull. Entomol. Soc. Am., 21:209-212. (With R. L. Fischer et

al.)

The bee family Oxaeidae with a revision of the North American species

(Hymenoptera: Apoidea). Smithson. Contrib. Zool., 220:1-75, 68 figs.,

3 pis., 3 maps, 2 tabs. (With E. G. Linsley.)

An annotated catalog of the carpenter bees (genus Xylocopa Latreille)

of the Western Hemisphere (Hymenoptera: Anthophoridae). Smithson¬

ian Institution Press, 106 pp., frontisp.

Bamboo-nesting carpenter bees (genus Xylocopa Latreille) of the sub¬

genus Stenoxylocopa Hurd and Moure (Hymenoptera: Anthophoridae).

J. Kans. Entomol. Soc., 51:746-764, 12 figs.

VOLUME 58, NUMBER 4

277

91. 1979a. Apoidea. In: Krombein et al., Catalog of Hymenoptera in America north

of Mexico, 3 vols., Smithsonian Institution Press, pp. 1741-2209. (Hurd

also was a joint author of introductions to each volume, and to the

section Hymenoptera.)

92. 1979b. Pollination of Opuntia lindheimeri and related species. Plant Syst. Evol.,

132:313-320, 2 tabs. (With V. and K. A. Grant.)

93. 1979c. Pollination of the southwestern opuntias. Plant Syst. Evol., 133:15-28,

2 tabs. (With V. Grant.)

94. 1980a. Skeletal parts of the sting apparatus of selected species in the family

Andrenidae. Proc. Entomol. Soc. Wash., 82:562-567, 5 figs. (With I. T.

Radovic.)

95. 1980b. Principal sunflower bees of North America with emphasis on the south¬

western United States (Hymenoptera: Apoidea). Smithson. Contrib. Zool.,

310:1-158, frontisp., 11 figs., tabs. 1-3, tabs. A-N, 5 pis. (With W. E.

LaBerge and E. G. Linsley.)

96. 1981. In-nest behavior of the carpenter bee, Xylocopa pubescens Spinola (Hy¬

menoptera: Anthophoridae). J. Kans. Entomol. Soc., 54:209-218, 8 figs.

(With D. Gerling and A. Hefetz.)

97. 1983a. [In Memoriam], Philip Hunter Timberlake, 1883-1981. Pan-Pac. Ento¬

mol., 58:2-18. (With E. G. Linsley and J. C. Hall.)

98. 1983b. Comparative behavioral biology of two Middle East species of carpenter

bees (Xylocopa Latreille) (Hymenoptera: Apoidea). Smithson. Contrib.

Zool., 369:1-33, frontisp., 30 figs., 7 tabs. (With D. Gerling and A.

Hefetz.)

Footnotes

1 Senior Scientist, Department of Entomology, Smithsonian Institution, Washington, D.C.

20560.

2 Professor Emeritus, Department of Entomological Sciences, University of California, Berke¬

ley, California 94720.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 278-283

Published 30 December 1983

MIMETIC SELECTION AND SUBSPECIFIC VARIATION IN THE

SPICEBUSH SWALLOWTAIL PAPILIO TROILUS LINNAEUS

(LEPIDOPTERA: PAPILIONIDAE)

John E. Hafernik, Jr.

Department of Biology, San Francisco State University,

San Francisco, California 94132

Butterfly color patterns are determined by the interaction of a number of

selective factors. Among these are sexual selection (Magnus, 1957, 1963;

Tinbergen et al., 1942; Silberglied and Taylor, 1973; Hafernik, 1982), se¬

lection for thermoregulatory efficiency (Watt, 1968), crypsis (Wickler, 1968),

aposematic coloration (Rothschild, 1972) and mimicry (Brower, 1958a, b;

Platt et al., 1971; Platt and Brower, 1968; Wickler, 1968; Punnett, 1915).

Perhaps the most interesting phenomena involved with selection for various

wing patterns are those resulting from predator avoidance strategies. This

paper discusses one such factor, mimetic selection, and proposes a new role

for it in influencing subspecific wing pattern variation in the spicebush swal¬

lowtail Papilio troilus Linnaeus.

The Battus philenor Mimicry Complex

Larvae of the pipevine swallowtail Battus philenor (Linnaeus) feed on

leaves of Aristolochia, a genus known to contain substantial concentrations

of toxic aristolochic acids (Rothschild et al., 1970; Euw et al., 1968). As a

result adults of B. philenor are distasteful to at least two species of birds and

presumably to a wide array of other vertebrates (Brower, 1958a; Platt et al.,

1971). In the eastern United States B. philenor generally is recognized as the

model for a series of palatable mimics. This Batesian association includes

males and females of P. troilus troilus, females of P. polyxenes Fabricius,

and the dark female form of P. glaucus Linnaeus (all Papilionidae); and two

nymphalid butterflies Limenitis arthemis astyanax (Fabricius) and females

of Speyeria diana (Cramer); as well as males of the saturniid moth Callo-

samia promethia (Drury) and related species (Brower and Brower, 1962;

Waldbauer and Sternburg, 1975).

Battus philenor ranges from New England south into Central America but

is absent or rare in Florida (Klots, 1951; Brower and Brower, 1962; Kimball,

1965). In the eastern United States most B. philenor mimics surpass the

distribution of their model. Two of these L. arthemis (Drury) and P. glaucus

show well-documented phenotypic variation correlated with increasing rar¬

ity or absence of B. philenor at the northern or southern limits of its distri-

VOLUME 58, NUMBER 4

279

bution (Platt and Brower, 1968; Burns, 1966). In the eastern United States,

L. arthemis occurs as two subspecies: L. a. astyanax, which mimics B.

philenor, and L. a. arthemis, which is disruptively colored. In New England

and the Great Lakes States, as B. philenor becomes increasingly rare, selec¬

tion apparently favors a switch from a mimetic strategy to one of camouflage

(Platt and Brower, 1968). Similarly, north of the range of B. philenor mimetic

females of P. glaucus are replaced by a disruptively colored form similar to

the male in coloration (Platt and Brower, 1968; Burns, 1966). In southern

Florida the frequency of mimetic morphs of P. glaucus is also reduced to

six to eight per cent (Brower and Brower, 1962).

Battus polydamus as a Model for Papilio troilus in Florida

Nominate P. troilus is characterized by blue or greenish spots on the dorsal

hindwing and small light spots on the dorsal forewing. This pattern is very

similar to that of B. philenor (Fig. la, c). In Florida P. troilus greatly out¬

numbers B. philenor, its usual model further north. In fact, Brower and

Brower (1962) found that the troilus to philenor ratio in southern Florida

was about 1000 to 1 and that B. philenor was only slightly more common

in northern Florida. In southern Georgia and northern Florida P. t. troilus

intergrades with its floridian subspecies P. t. ilioneus Abbot and Smith, a

butterfly characterized by a generally brighter, more yellowish green color

pattern with well-developed forewing spots (Fig. Id). As with P. glaucus,

mentioned above, this switch in phenotypes parallels the decline in frequency

of B. philenor.

Two hypotheses have been suggested to explain the switch over to the

floridian phenotype in P. troilus. Brower and Brower (1962) suggest that the

phenotypic change results from a relaxation of selection for a mimetic color

pattern allowing the expression of deviant color patterns. This explanation

would predict a similar deviation, although not necessarily in the same

direction, in northern populations of P. troilus which exceed the range of

their model, yet no such phenotypic change occurs in northern populations.

Moreover, I believe it likely, as is apparently the case in L. arthemis and P.

glaucus, that if selection for mimicry were relaxed other selective factors

would increase in importance in determining color patterns. There is no

indication that the P. t. ilioneus pattern is more cryptic than the P. t. troilus

pattern nor have other selective reasons been suggested. The ilioneus phe¬

notype is, if anything, more conspicuous than the normal morph and perhaps

more easily spotted by predators.

The second explanation is that of Remington (1968) who believed that

the ilioneus phenotype is the result of previous isolation of Florida P. troilus

populations. Remington believed that a dense band of forest across northern

Florida provided a barrier that until recently maintained geographic isolation

280

PAN-PACIFIC ENTOMOLOGIST

Fig. la-d. Battus-Papilio troilus mimicry complex. Fig. la, Battusphilenor, St. Louis, Mis¬

souri. Fig. lb, Battus polydamus, Sontecomapan, Veracruz, Mexico. Fig. lc, Papilio troilus

troilus, Bear Mountain, New York. Fig. Id, Papilio troilus ilioneus, Lutz, Florida.

between the two subspecies. He explains the area of intergradation in north¬

ern Florida and southern Georgia as a region of secondary contact which

may eventually result in the evolution of prezygotic isolating mechanisms

between the two taxa. I do not find this scenario compelling since P. troilus

is a forest butterfly (Klots, 1951; Emmel, 1975). It occurs commonly in such

heavily forested areas as the Big Thicket area of east Texas (personal ob¬

servation). This suggests that a dense forest would not significantly restrict

gene flow between Florida and Georgia populations as it might have for

other non-forest butterflies which show a similar distribution, but no evi¬

dence of subspecific differentiation.

In place of the above explanations, I propose a third, although not mutually

exclusive, hypothesis. Study of spread specimens of P. t. ilioneus suggests

the pattern of another Aristolochia-feeding swallowtail B. polydamus (Lin¬

naeus) (Fig. lb, d). Aristolochic acids have been isolated from the tissues of

B. polydamus (Rothschild et al., 1970; Rothschild, 1972) and it is likely that

VOLUME 58, NUMBER 4

281

B. polydamus is unpalatable to a wide range of vertebrate predators although

feeding trials have not been done. Waldbauer and Sternburg (1975) suggest

that B. polydamus serves as the model for several Batesian mimics in the

neotropics among them females of Papilio androgeus Cramer and P. lyco-

phron Hiibner. Battus polydamus is common in Florida and ranges north

to southern Georgia where it is infrequent (Harris, 1972). No quantitative

data are available on the relative proportions of B. polydamus and P. troilus

in Florida. However, Kimball (1965) states that B. polydamus is at times

more common than P. troilus. Thus there is circumstantial evidence that

the change from the typical troilus morph to the ilioneus morph results from

a change in frequency of suitable models.

Some workers group P. troilus populations in Texas and the Gulf States

under the subspecies ilioneus (=texanus Ehrman) (Klots, 1951; Emmel, 1975).

I am unable to distinguish differences among Texas populations and more

northern P. t. troilus. B. polydamus occurs periodically in central and north¬

ern Texas and the Gulf States. If any tendencies toward ilioneus- like patterns

exist in some populations, they could result from selection for patterns that

provide mimetic advantage from both B. philenor and B. polydamus.

Discussion

The influence of availability and abundance of appropriate models on

geographic variation of mimetic butterflies is well-known, especially for

tropical species. Perhaps the best known and most intricate example of this

kind of geographic pattern is that found in the African swallowtail Papilio

dardanus Brown. This species occurs as a complex array of races whose

females mimic several members of the unpalatable genera Danaus and

Amauris. Frequencies of mimetic morphs are related to the abundance of

particular models in each area (Clarke and Sheppard, 1960). In the United

States subspecific variation in the well-known viceroy butterfly Limenitis

archippus (Cramer) is also related to a change in abundance of models.

Throughout most of its range L. archippus mimics the monarch Danaus

plexippus (Linnaeus), but in the southwestern United States it mimics the

related D. gilippus strigosus (Bates). In northern Florida and southern Geor¬

gia L. archippus populations switch from a D. plexippus-like pattern to one

resembling D. gilippus berenice (Cramer). As pointed out by Remington

(1968), this situation closely parallels the one outlined above for Papilio

troilus.

The phenotypic similarity of P. t. ilioneus to Battus polydamus, the co¬

incidence in range of the two, and the rarity of B. philenor in Florida offer

strong circumstantial evidence of color pattern selection in Florida P. troilus

for mimicry of B. polydamus. This explanation does not mutually exclude

previous hypotheses. Indeed, the explanations of Remington (1968) and

Brower and Brower (1962) may be partly correct. Previous isolation of

282

PAN-PACIFIC ENTOMOLOGIST

Florida P. troilus, as postulated by Remington, probably would have accel¬

erated evolution of new color patterns using B. polydamus as a model.

Furthermore, the absence of B. philenor from Florida, emphasized by Brower

and Brower, would have been important in releasing P. troilus populations

from stabilizing selection for B. philenor-\ik.e patterns.

Research into the following areas is needed to clarify the factors influencing

variation among the southern populations of P. troilus : 1. Laboratory studies,

of the type pioneered by the Brower group, utilizing caged predators in order

to ascertain the unpalatability of B. polydamus and the efficiency of P. troilus

ilioneus vis-a-vis P. t. troilus as a B. polydamus mimic. 2. Field studies of

the relative abundance of B. polydamus and P. troilus in various parts of

Florida and Georgia in conjunction with a quantitative analysis of geographic

variation in P. troilus wing patterns in the southeastern United States.

Summary

The similarity in wing pattern of Papilio troilus ilioneus Abbot and Smith

to that of Battus polydamus (Linnaeus), the coincidence in range of the two,

and the rarity in Florida of Battus philenor (Linnaeus) the presumed model

for P. t. troilus Linnaeus suggest that the P. t. ilioneus pattern is the result

of selection for mimicry of B. polydamus. Previous explanations of the causes

of subspecific variation in P. troilus are discussed in light of this new hy¬

pothesis.

Acknowledgments

J. T. Doyen, R. W. Garrison, and J. A. Powell read an earlier draft of this

manuscript and provided helpful suggestions. Specimens figured are from

the California Insect Survey, University of California, Berkeley, except for

the B. polydamus specimen which is from the collection of the author.

Literature Cited

Brower, J. V. Z. 1958a. Experimental studies of mimicry in some North American butterflies.

Part II. Battus philenor and Papilio troilus, P. polyxenes, and P. glaucus. Evolution, 12:

123-136.

-. 1958b. Experimental studies of mimicry in some North American butterflies. Part

III. Danaus gilippus berenice and Limenitis floridensis. Evolution, 12:273-285.

Brower, L. P., and J. V. Z. Brower. 1962. The relative abundance of model and mimic

butterflies in natural populations of the Battus philenor mimicry complex. Ecology, 43:

154-158.

Bums, J. M. 1966. Preferential mating versus mimicry: Disruptive selection and sex-limited

mimicry in Papilio glaucus. Science, 153:551-553.

Clarke, C. A., and P. M. Sheppard. 1960. The evolution of mimicry in the butterfly Papilio

dardanus. Heredity, 14:163-173.

Emmel, J. F. 1975. Subfamily Papilioninae. Pp. 390-402, in W. H. Howe (ed.), The butterflies

of North America, Doubleday and Company, New York.

VOLUME 58, NUMBER 4

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Euw, J. von, T. Reichstein, and M. Rothschild. 1968. Aristolochic acid—I. In the swallowtail

butterfly Pachlioptera aristolochiae (Fabr.). Isr. J. Chem., 6:659-670.

Hafemik, J. 1982. Phenetics and ecology of hybridization in buckeye butterflies (Lepidoptera:

Nymphalidae). Univ. Calif. Publ. Entomol., 96:i-vii, 1-109.

Harris, L. 1972. Butterflies of Georgia. University of Oklahoma Press, Norman, 326 pp.

Kimball, C. P. 1965. Arthropods of Florida and neighboring states. Vol. 1. Lepidoptera of

Florida. Florida Department of Agriculture, Gainesville, 363 pp.

Klots, A. B. 1951. A field guide to the butterflies. Houghton Mifflin Co., Boston, 349 pp.

Magnus, D. B. 1957. Experimentelle Untersuchungen zur Bionomie und Ethologie des Kais-

ermantels Argynnis paphia L. (Lepidoptera: Nymphalidae) I. Uber optische Ausloser

von Anfliegereaktionen und ihre Bedeutung fur das Sichfinden der Geschlechter. Z.

Tierpsychol., 15:397-426.

-. 1963. Sex-limited mimicry. II. Visual selection in the mate choice of butterflies. Proc.

16 Int. Congr. Zool., Washington, D.C. 4:179-183.

Platt, A. P., and L. P. Brower. 1968. Mimetic versus disruptive coloration in intergrading

populations of Limenitis arthemis and astyanax butterflies. Evolution, 22:699-718.

-, R. P. Coppinger, and L. P. Brower. 1971. Demonstration of the selective advantage

of mimetic Limenitis butterflies presented to caged avian predators. Evolution, 25:692-

701.

Punnett, R. C. 1915. Mimicry in butterflies. Cambridge University Press, Cambridge, 188

pp.

Remington, C. L. 1968. Suture-zones of hybrid interaction between recently formed biotas.

Evol. Biol., 2:321-428.

Rothschild, M. 1972. Secondary plant substances and warning coloration in insects. In H. F.

van Emden (ed.), Insect/plant relationships. Symposia Royal Entomological Society,

London 6. John Wiley and Sons, New York.

-, T. Reichstein, J. von Euw, R. Alpin, and R. R. Harman. 1970. Toxic Lepidoptera.

Toxicon, 8:293-299.

Silberglied, R. E., and O. Taylor. 1973. Ultraviolet difference between sulfur butterflies, Colias

eurytheme and C. philodice, and a possible isolating mechanism. Nature, 241:406-408.

Tinbergen, N. B., B. J. D. Meuse, L. L. Boerma, and W. W. Variosseau. 1942. Die Balz des

Samtfalters, Eumenis ( Satyrus) semele (L.). Z. Tierpsychol., 5:182-226.

Waldbauer, G. P., and J. G. Stemburg. 1975. Satumiid moths as mimics: An alternative

interpretation of attempts to demonstrate mimetic advantage in nature. Evolution, 29:

650-658.

Watt, W. B. 1968. Adaptive significance of pigment polymorphism in Colias butterflies. I.

Variation of melanin pigment in relation to thermoregulation. Evolution, 22:437-458.

Wickler, W. 1968. Mimicry in plants and animals. McGraw-Hill Book Co., New York,

255 pp.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 284-287

Published 30 December 1983

BIOLOGICAL NOTES ON ANDRENA ( CALLANDRENA ) HA YNESI

VIERECK AND COCKERELL (HYMENOPTERA: ANDRENIDAE)

F. D. Parker and T. Griswold

Bee Biology & Systematics Laboratory, Agricultural Research, USD A,

Utah State University, UMC 53, Logan 84322

In the summer of 1980, both sexes of Andrena haynesi Viereck and Cock¬

erell were collected from sunflower that grew on sand dunes in southeastern

Utah’s San Rafael Desert. The recorded distribution area of this large, dark-

colored andrenid bee includes the western great plains (LaBerge 1967), but

specimens from northeastern Arizona are in the USD A Bee Laboratory

collection at Logan, Utah. The specimens reported in this study are the first

records of this bee from Utah.

During August and September, we found many newly constructed nest

mounds of A. haynesi on the sand dunes near the sunflower patches where

the bees foraged. Thus, we were able to observe and record some aspects of

the biology of this sunflower bee. There is little biological information about

Callendrena, the subgenus where A. haynesi is placed. For example, nests

of only 2 of the 52 United States species have been described (Rozen, 1973;

Parker and Bohart, 1983).

Nesting Site

We found many of the nest mounds on the bare, windward side of crescent

dunes located on both sides of Highway 24, approximately 20 miles north

of Hanksville, Utah. Another nest concentration was found on the north

slope of a hill that borders a wash about one mile northwest of Goblin Valley

State Park, Utah, a location approximately seven miles from the first one.

Most of the nests at this latter site were scattered among small shrubs of

Eriogonum where the soil surface was firmer than that of the dunes. We

also found some nests in hard-packed sand along an old road. Adjacent to

both nesting sites were large patches of the sunflower, Helianthus anomalus

Blake.

Nest Architecture

Tumulus. — The nest entrances were characterized by large mounds of

excavated dirt surrounding the openings (Fig. 1). These tumuli were much

larger than nest mounds of other mining bees we have seen. The steep-sided,

cone-shaped mounds measured 8.9-10.2 cm wide and 3.8-4.4 cm high.

Entrances of most nests were in the center of the tumulus, but they were at

VOLUME 58, NUMBER 4

285

Figs. 1, 3. Andrena haynesi. Fig. 1. Nest tumulus. Fig. 3. Cell with loose pollen and partial

pollen ball.

the side in older nests, probably because rain had weakened the turrets,

causing them to fall on their sides. All new nests had a turret within and

extending to the base of the tumulus that the bees had made by cementing

8-15 mm of soil around the nest entrance (Fig. 2). We collected several

females that were in the turret just below the plug in the nest entrance by

Fig. 2. Andrena haynesi nest architecture: A = entire nest diagram; B = details of tumulus,

turret, and lateral burrow; C = cell.

286

PAN-PACIFIC ENTOMOLOGIST

quickly brushing the turret away from the main burrow. The 8-mm wide

nest entrance was closed by a 5-9 mm thick plug of moist sand except when

the female was actively foraging.

Burrow.— We excavated only one burrow because of their extraordinary

depths. The main burrow was the same diameter as the entrance, and the

walls were smooth. It descended vertically for 2.35 m and then sloped to

the east in a 45° spiral for 20 cm. At this point, the main burrow was 2.7

m below the soil surface. From the main burrow, a lateral burrow extended

eastward for 43 cm and ended in a single cell.

While the first nest was being excavated, which took about 7 hours, we

estimated the depth of the main burrows of other nests by removing the

tumulus and lowering a weighted string down the large hole. All of the nests

we measured had been plugged at the entrance. The main burrows of five

nests descended vertically 2.1-2.2 m below the surface. Other nests were

considerably shorter, only .5-1.3 m deep. After the bees replugged the en¬

trance of the shallower nests, we remeasured the burrows. In all cases, the

string dropped to a greater depth the second time, suggesting that the main

burrow is plugged at various intervals, and the bees use the sands from these

plugs to reseal the nest entrance.

Cell. — A single horizontal, but incomplete cell was found; it measured 23-

mm long and 11 -mm wide (Fig. 3). The basal two-thirds of the cell was coated

with a thick waxy material. This cell, which was in the process of being

provisioned (the female was captured as she returned to the nest with a

pollen load), contained several loads of loose pollen and a spherical pollen-

nectar ball that was 7 mm in diameter (Fig. 2). The pollen was 100% He-

lianthus. The soil temperature at the cell depth was 67°F (19°C).

Observations on Adults

Five female Andrena were observed making foraging trips. During the

foraging period, the bees always left the nest entrance open. The temperature

when some bees began foraging was only 47°F (8°C) because they opened

their nests as early as 6:27 a.m. (September 16). (It was still too dark for us

to see without the aid of a lantern.) We also observed an undescribed species

of Nomada (Pachynomada ) leaving an unplugged nest of A. haynesi early

in the morning. This parasite has been reared from cells of Andrena helianthi

Robertson, at Logan, Utah (Parker and Bohart, 1983).

The time the bee spent out of the nest to gather a pollen load averaged

12 min 6 sec (10 min 24 sec to 14 min 51 sec), and the time spent within

nests between trips averaged 2 min 50 sec (2 min 5 sec to 3 min 42 sec).

The time the bees spent at the sunflower head averaged 32.1 sec (2-88 sec,

n = 27). The female landed directly on the disk flowers, near the margin,

and then worked the ring of newly dehisced anthers in a circular pattern,

with some bees making more than one circuit. The foraging period ended

VOLUME 58, NUMBER 4

287

between 8:30 and 9:15 a.m. when the nest entrances were replugged; no

further activity was seen at these nests.

Discussion

The nests of A. haynesi are deeper than nests of any other recorded North

American bee, probably an adaptation of this species to their environment.

First, these bees nest in shifting sand dunes that are several feet above the

surrounding soil surface. Deep nests would protect the bee cells when the

dunes shifted from the nesting site. For example, 12 Andrena nests were

marked in August, but when we returned in September, all of the nest markers

and 7 of the nest turrets had been blown away; nevertheless, 5 of the original

nests were still active. Second, these bees are matinal oligoleges of sunflower

and forage during the coolest time of the day. Perhaps the higher soil tem¬

perature in these deep nests enables the bees to maintain their body tem¬

perature at the flight threshold so that they can forage early in the day at

cold temperatures. It is also possible that there are soil moisture requirements

for bee development and that this depth places the cells below the maximum

level of dry sand even in drought years.

The basic pattern of A. haynesi nest construction is similar to that of A.

helianthi, a related Callandrena that is also an oligolege of sunflower (Parker

and Bohart, 1983). The nests of A. helianthi were not as deep and the lateral

burrows were shorter than those of A. haynesi. Andrena accepta Viereck,

the only other biologically known Callandrena, has communal nests (Rozen,

1973).

Acknowledgments

We would like to thank D. Veirs of this laboratory for his assistance in

the field work and for the illustrations. Drs. N. Youssef, Department of

Biology, Utah State University, and W. LaBerge, Illinois Natural History

Survey, Natural Resources Building, Urbana, offered helpful suggestions for

improving this manuscript.

Literature Cited

LaBerge, W. E. 1967. A revision of the bees of the genus Andrena of the Western Hemisphere.

Part I. Callandrena. Bull. Univ. Neb. State Mus., 7:1-318.

Rozen, J. G. 1973. Biological notes on the bee Andrena accepta Viereck. J. N.Y. Entomol.

Soc., 81:54-61.

Parker, F. D., and G. E. Bohart. 1983. Notes on the biology of Andrena {Callandrena) helianthi

Robertson. Pan-Pac. Entomol. (1982), 58:111-116.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 288-290

Published 30 December 1983

TRYPARGILUM TRIDENTATUM (PACKARD) IN TRAP NESTS IN

OREGON (HYMENOPTERA: SPHECIDAE: TRYPOXYLINAE)

Mark F. O’Brien

Insect Division, Museum of Zoology,

The University of Michigan, Ann Arbor 48109

Trypargilum tridentatum (Packard), found from Connecticut to Florida

and west to British Columbia and California, is a member of the nitidum

species group which preys exclusively on web-building spiders (Krombein

et al., 1979). T. tridentatum has been found to nest in old borings in wood,

elderberry ( Sambucus ) stems, trap nests, and old Sceliphron (Sphecidae)

nests (Rau and Rau, 1918; Blackman and Stage, 1924; Krombein, 1967;

Krombein et al., 1979). According to Krombein (1967), T. tridentatum

usually provisioned cells with two to four species of spiders per cell in Arizona

and North Carolina. The prey within a single trap nest often consisted of

two or three spider families. Published prey records include members of the

families Theridiidae (4 spp.), Mimetidae (1 sp.), Araneidae (9 spp.), and

Dictynidae (1 sp.) (Krombein, 1967), but no nesting data is available for

populations from the northwestern United States.

The study area was located about 30 miles south of Burns, Harney Co.,

Oregon at the Malheur Field Station. Rabbitbrush ( Chrysothamnus sp.) and

Sagebrush (Artimesia sp.) was the dominant vegetation in the area.

Trap nests were similar to those used by Krombein (1967); 2.6 x 2.6 x

18 cm blocks of white pine with 6 and 10 mm borings 14 cm deep, and

were placed horizontally on a building 1.5 m above the ground during the

first week of July 1979. Four completed nests were retrieved on July 20, and

sent by mail to Syracuse, New York, where they were split open and the

contents removed, examined, and identified. Nests were taped back together

after measurements were taken and placed in an outdoor insectary for over¬

wintering. They were retrieved in March 1980, and cocoons along with

parasites were transferred to gelatine capsules and coded with cell numbers.

These were kept inside until the adults eclosed.

Four trap nests (J-5, 6, 20, 21) contained a total of 31 cells. All nests had

been fully provisioned and closed. Mud cell partitions were approximately

1 mm thick, with closing plugs 2.0, 2.1, and 3.5 mm thick. Three nests

contained vestibular cells: J-21, two, 2.3 and 2.5 cm long; J-20, 1.0 cm long;

and J-5, 1.1 cm long. Dimensions for the nests and the number of prey are

given in Table 1.

Mean cell length (both sexes) was 1.38 ± .312 cm (N = 31, 1.1-2.7). Male

VOLUME 58, NUMBER 4

289

Table 1. Nest data of Trypargilum tridentatum from Oregon.

Nest J-5 Cell

Cell length (cm)

1.6

1.3

1.5

1.35

1.6

1.25

1.1

Cocoon length

—

1.05

—

1.05

No. of prey

Sex

—

Nest J-6

Cell length

1.3

1.8

1.6

1.4

1.6

1.4

1.5

1.1

Cocoon length

1.1

1.3

1.15

1.2

1.0

—

No. of prey

—

Sex

—

Nest J-20

Cell length

1.25

1.3

1.1

1.2

1.1

1.5

1.2

Cocoon length

1.1

1.2

1.05

1.1

—

No. of prey

—

Sex

—

Nest J-21 b

Cell length

1.4

1.1

1.2

2.7

No. of prey

a Parasitized cells are designated with a “P.”

b All cell contents were removed and preserved in alcohol.

cocoons ranged from 1.0-1.3 cm (x = 1.11 ± .114, N = 5), whereas female

cocoons ranged from 1.05-1.3 cm long (x = 1.15 ± .097, N = 6). Male cells

varied from 1.1-1.8 cm long (x = 1.4 ± .292, N = 5), and female cell lengths

ranged from 1.3-1.6cm(x= 1.5 ± .126, N = 6). Krombein (1967) reported

populations in Arizona and North Carolina having male cocoons and cells

slightly longer and less variable than those of females.

Whereas the wasps in Arizona and North Carolina studied by Krombein

(1967) used several genera and families of spiders per nest, I found that

Metepeira grandiosa Chamberlin and Ivie (Araneidae), a new prey record,

was used exclusively in the 21 cells that still contained prey. The mean

number of spiders per cell was 8.7 ±1.7 (7-12). The use of only one species

of spider may reflect a local abundance of prey, or conditioning on the part

of the provisioning female.

Adult wasps emerged between May 21 and 30, 1980. Three cells in nest

J-5 had been parasitized by miltogrammine flies (Sarcophagidae) (Table 1).

Cell 6 contained one puparium, cells 7 and 8 each contained three. Unfor-

290

PAN-PACIFIC ENTOMOLOGIST

tunately, no adult flies were reared, but other workers (Parker and Bohart,

1966; Krombein, 1967) have recorded Amobia floridensis (Townsend) from

nests of T. tridentatum.

Acknowledgments

I thank Roy A. Norton, State University of New York College of Envi¬

ronmental Science and Forestry, Syracuse, for identifying the prey spiders

and for his helpful comments on the manuscript; and also John Hayden, for

setting out the traps in Oregon and sending them back to Syracuse.

Literature Cited

Blackman, M. W., and H. H. Stage. 1924. On the succession of insects living in the bark and

wood of dying, dead, and decaying hickory. N.Y. State Coll. For. Tech. Publ., 17:196-

197.

Krombein, K. V. 1967. Trap-nesting wasps and bees: Life histories, nests, and associates.

Smithsonian Press, Washington, D.C, 570 pp.

-. 1979. Superfamily Sphecoidea. Pp. 1573-1740, In: Krombein, K. V., P. D. Hurd,

Jr., D. R. Smith, and B. D. Burks, Catalog of Hymenoptera in America north of Mexico.

Vol. 2, Apocrita (Aculeata). Smithsonian Institution Press, Washington, D.C., pp. i-xvi,

1199-2209.

Parker, F. D., and R. M. Bohart. 1966. Host-parasite associations in some twig-nesting

Hymenoptera from Western North America. Pan-Pac. Entomol., 42:91-98.

Rau, P., and N. Rau. 1918. Wasp studies afield. Princeton University Press, Princeton, 372

pp.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 291-295

Published 30 December 1983

OBSERVATIONS ON TRIALEURODES PACKARDI (MORRILL) IN

A COMMUNITY GARDEN (HOMOPTERA: ALEYRODIDAE)

L. E. Ehler

Department of Entomology, University of California, Davis 95616

Strawberry whitefly, Trialeurodes packardi (Morrill), is commonly asso¬

ciated with strawberry ( Fragaria ananassa ) in urban areas of the Sacramento

Valley of California. In certain community gardens, strawberry is grown as

either a “monoculture” (i.e., a given garden plot planted entirely in straw¬

berry) or as part of a polyculture (i.e., a few strawberry plants mixed with

other crops). Thus, T. packardi occurs on strawberry plants grown in both

monocultures and polycultures. The purpose of the present study was to

assess the influence of such crop-plant diversity on abundance of T. packardi

in a community garden.

Materials and Methods

The study site was a community garden on the Davis Campus of the

University of California. This site was previously an experimental orchard

and was converted into a community garden during the early 1970’s. The

garden was composed of ca. 450, 21 m 2 plots. The experimental work was

conducted from April 20 through July 6, 1977. Subsequent destruction of

the gardens precluded further observations.

Fourteen gardens (referred to herein as plots) were chosen for study. Plots

1-7 were polycultures whereas plots 8-14 were strawberry monocultures.

Crops grown in the polycultures during the course of the experiment are

given in Table 1. Of these crops, apparently only strawberry is a suitable

host for T. packardi (Russell, 1948, 1963). All plots were well cared for by

the respective gardeners and none was treated with a chemical pesticide

during the course of the study.

Plots were sampled weekly from April 20-July 6. A sample consisted of

a careful inspection of the undersides of 10 randomly chosen leaves per plot.

The number of adults and immatures of T. packardi was recorded. This

procedure was carried out such that there was minimal disruption of the

plants in question. All of the specimens collected from strawberry and sub¬

mitted for determination were T. packardi. Greenhouse whitefly, T. vapora-

riorum (Westwood), infested numerous plants in the gardens but was never

collected from strawberry.

292

PAN-PACIFIC ENTOMOLOGIST

Table 1. Crops grown in polycultural plots.

Crops

Plots

Number strawberry plants

English pea

X a

Tomato

Egg plant

X b

Pepper

X b

X a

Swiss chard

X a

Onion

Beet

Carrot

Sweet corn

Squash

X b

x b

X b

Artichoke

Green bean

X b

Lettuce

Peanut

X b

Cucumber

X b

Broccoli

X b

Lima bean

X b

a Removed during the experiment.

b Added during the experiment.

Results and Discussion

Population density of T. packardi in the polycultures contrasted markedly

with that in the monocultures (Fig. 1). During the study, mean densities (i.e.,

whiteflies/10 leaves) in the poly cultures ranged from 1-90, 0-19, 2-34, 0-

18, 0-32, 0-29 and 0-39 in plots 1-7, respectively. In the 7 monocultures,

mean whitefly density during the study ranged from 0-7, 0-2, 0-1, 0-5, 0-

4, 0-3 and 0-2, respectively. As the ranges in mean whitefly density were

generally similar for both monocultural and polycultural plots, average den¬

sities (N = 7) for each set of plots were computed and these are plotted in

Figure 1. Mean density for the season was 12.74 whiteflies/10 leaves in the

polycultures compared to 0.69 in the monocultures. Mean percent leaves

infested during the season was 34.5% in the polycultures compared to 5%

in the monocultures. These results are also consistent with previous obser¬

vations made in the same community garden during 1976 (see Ehler, 1978).

In that study, mean whitefly density (adults + immatures) was 21.9/10 leaves

(55.2% leaves infested) in polycultures compared to 1.7/10 leaves (9.8%

leaves infested) in monocultures. Furthermore, whitefly populations exhib¬

ited instability in the polycultures and stability in the monocultures.

VOLUME 58, NUMBER 4

293

Fig. 1. Population trends of Trialeurodes packardi on strawberry grown in polyculture vs.

monoculture.

A number of hypotheses can be postulated to account for the empirical

dichotomy documented here. Perhaps the most obvious hypothesis would

be that, in polycultures, whiteflies build up on other crop plants and sub¬

sequently invade the strawberry plants, thus resulting in higher population

densities than in monocultures. However, this does not appear likely because,

according to Russell (1948, 1963), strawberry whitefly has not been recorded

from any of the crops (other than strawberry) utilized in the present study.

Furthermore, the whiteflies which infest these crops are characteristically T.

vaporariorum. Another hypothesis would involve differential action by nat¬

ural enemies; however, I was unable to detect such activity by natural ene¬

mies of strawberry whitefly in these plots. Some additional research regarding

the role of natural enemies is warranted however. A more plausible hy¬

pothesis at present involves differential habitat suitability. That is, straw¬

berry whitefly apparently prefers a more open habitat such as that provided

by a few, rather widely spaced strawberry plants in the mixed plantings.

Conversely, the more densely planted monocultures are less suitable habitats.

294

PAN-PACIFIC ENTOMOLOGIST

The following empirical evidence is submitted in support of the latter

hypothesis. On June 2, 1978,1 sampled 9 densely planted strawberry mono¬

cultures which also had a few, widely spaced plants along the edges. In each

case, 20 leaves were examined: 10 each from the interior plants and from

the marginal plants. Samples from the interior of these monocultures yielded

an average of 1.1 ± 0.4 whiteflies/10 leaves (7.8% leaves infested) compared

to 13.4 ± 3.3 whiteflies/10 leaves (56.7% leaves infested) among more widely

spaced plants along the edges. Thus, it appears that strawberry plants grown

in poly culture are quite suitable for strawberry whitefly such that “boom

and bust” cycles in population density result; in strawberry monocultures,

the habitat is apparently much less suitable for the insect and only a marginal

amount of reproductive success occurs. In this regard, only 5.2% of all the

individuals observed in the 7 monocultures were nymphs compared to 46.4%

in the 7 polycultures.

The view that diversity begets stability or lessens pest problems has been

critically assessed by numerous recent authors; these include theoretical

treatments (e.g., May, 1973; Gilpin et al., 1976), more practical accounts

(e.g., van Emden and Williams, 1974; Litsinger and Moody, 1976) and

combinations of both (e.g., Murdoch, 1975; Way, 1977). From such ac¬

counts, it becomes apparent that (1) the value of crop diversity in agroeco¬

systems is open to question and (2) the view that diversity begets stability

or lessens pest problems is an excessive generalization. The results of the

present study attest to the latter conclusion—i.e., the results were precisely

the opposite of what the theory predicts. In many ways, the diversity in the

polycultures was irrelevant; the more important factor apparently was hab¬

itat suitability as influenced by plant density and spacing, etc. However, this

hypothesis is in need of additional empirical verification.

Acknowledgments

I thank R. Gill and M. B. Stoetzel for taxonomic assistance.

Literature Cited

Ehler, L. E. 1978. Some aspects of urban agriculture. Pp. 349-57, In G. W. Frankie and

C. S. Koehler (eds.), Perspectives in urban entomology. Academic Press, New York,

417 pp.

Gilpin, M. E., G. A. H. McClelland, and J. W. Pearson. 1976. Space, time, and the stability

of laboratory mosquito populations. Am. Nat., 110:1107-1 111.

Litsinger, J. A., and K. Moody. 1976. Integrated pest management in multiple cropping

systems. Ch. 15, In R. I. Papendick, P. A. Sanchez, and G. B. Triplet (eds.), Multiple

cropping. Special Publ. No. 27, Am. Soc. Agronomy, Madison, WI.

May, R. M. 1973. Stability and complexity in model ecosystems. Princeton University Press,

Princeton, New Jersey, 235 pp.

Murdoch, W. W. 1975. Diversity, complexity, stability and pest control. J. Appl. Ecol., 12:

795-807.

VOLUME 58, NUMBER 4

295

Russell, L. M. 1948. The North American species of whiteflies of the genus Trialeurodes.

U.S.D.A. Misc. Publ., 635:1-85.

-. 1963. Hosts and distribution of five species of Trialeurodes (Homoptera: Aleyrodidae).

Ann. Entomol. Soc. Am., 56:149-153.

van Emden, H. F., and G. F. Williams. 1974. Insect stability and diversity in agro-ecosystems.

Annu. Rev. Entomol., 19:455-475.

Way, M. J. 1977. Pest and disease status in mixed stands vs. monocultures; the relevance of

ecosystem stability. Pp. 127-138, In J. M. Cherrett and G. R. Sagar (eds.), Origins of

pest, parasite, disease and weed problems. Blackwell Scientific Publications, Oxford.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 296-301

Published 30 December 1983

THE BIOLOGY OF APIOCERA HARUSPEX OSTEN SACKEN

(DIPTERA: APIOCERIDAE) IN CENTRAL CALIFORNIA, AND

COMPARISON WITH OTHER ASILOIDEA

Robert A. Wharton

Department of Entomology, Texas A & M University, College Station 77843

Relatively little has been published on the biology of the Apioceridae. 1 The

most complete study is that by Cazier (1963) on the bionomics of Apiocera

painteri Cazier. Lavigne (1975) noted similarities between A. painteri and

A. clavator Painter while discussing the general behavior of the latter. English

(1947) and Irwin and Stuckenberg (1972) emphasized larval morphology in

their works on Southern Hemisphere species; while Norris (1936) and Pa-

ramonov (1953) briefly reported on oviposition and adult feeding respec¬

tively. Observations on Apiocera haruspex Osten Sacken are recorded here

for comparison, both with other apiocerids and with related members of the

Asiloidea.

Materials and Methods

A population of A. haruspex was first observed along the sandy banks of

the middle fork of the Cosumnes River in August 1976. The site is ap¬

proximately 5 km southwest of Somerset, El Dorado County, California at

an elevation of 650 m. Subsequent observations were made in August and

September of 1977 and 1978 along the river and on grassy hillsides above

the river canyon. Observations on asilids were made in the same area.

Voucher specimens on which observations were made are deposited in the

collections of Mont Cazier and the author.

One A. haruspex egg clutch was excavated immediately following ovi¬

position, and the eggs kept with sand in a glass jar. The single hatched larva

is that on which the following description is based.

Results

Larva.— The single larva attained a length of 5.5 mm in 2 months before

it died, and moulted only once. Because it was not preserved immediately

after death, some details are difficult to discern. The larva of A. haruspex is

very similar to that of A. maritima Hardy (English, 1947) and completely

different from that of Tongamya miranda Stuckenberg (Irwin and Stuck¬

enberg, 1972).

All segments subequal in length except terminal segment noticeably short¬

er. Intersegmental constrictions of abdomen similar to those of A. maritima

VOLUME 58, NUMBER 4

297

mandibla

I f

l0.

1 ^r~^ \

^ (err

jnaxilla

palp

lateral keel

antenna

dorsal rod

-capsule rod

Figs. 1, 2. Apiocera haruspex. Fig. 1. Posterior spiracle and terminal portion of tracheal

trunk in penultimate abdominal segment, second instar larva. Fig. 2. Dorsal view of head

capsule of same larva.

(English, 1947: Fig. 1) most evident on abdominal segments 1-5. Apical

segment broadly rounded as in A. maritima and A. painteri, and bearing at

least one pair of long, subapical setae. Posterior spiracle opening on pen¬

ultimate segment, funnel-shaped, with single opening (Fig. 1). Head (Fig. 2)

very similar to A. maritima, consisting of well-developed anterior and pos¬

terior portions. Posterior portion dorsally with rounded antennal discs an-

terolaterad, each disc bearing a raised tubercle; pair of median dorsal rods

extending further posteriorly than in A. maritima ; posterolateral lobes and

capsule rod present and similar in shape to those of A. maritima ; posterior

portion with 2 pairs of stout setae ventrolaterad. Anterior portion arched

dorsally; mandibles, maxillae, and maxillary palps as in A. maritima, prom¬

inent; lateral keel and setae less well-developed.

Phenology.—A. haruspex adults were found only during August 1976 and

1978, and September 1977. Adult activity was restricted to a 2-3 week period

in each population, with slight variation among populations and years. The

298

PAN-PACIFIC ENTOMOLOGIST

Table 1. Foray behavior by Apiocera haruspex males (6 individuals).

Individuals

Totals

Interspecific encounters

Intraspecific encounters

with other males

No other animals visible

ratio of females to males became greater as the season advanced, as was

noted for A. painteri by Cazier (1963). Males and females were active

throughout the day, generally initiating activity between 0900 and 1000

PDT. Individuals sitting in sunny positions usually became active before

those in shaded positions, suggesting that ambient temperatures influence

initiation of activity (see also Lavigne, 1975).

Mating. — Mating behavior in A. haruspex is similar to that described by

Cazier (1963) and Lavigne (1975) for other Apiocera species. Four pairs of

A. haruspex were observed in copulo, all between 1130 and 1715 PDT.

Initiation of coupling was not seen; and when first observed, all couples were

on the ground. Pairs remained in copulo up to 23 min in a tail-to-tail position.

As in A. painteri, the flies became active following an initial stationary

period. While still in copulo they rotated in a clockwise and/or counter¬

clockwise direction and flew short distances. Final separation occurred after

several jerks of the male abdomen. During copulation, males were more

active than females.

Male behavior.—As noted by Cole (1969:179), A. haruspex closely resem¬

bles asilids of the genus Ejferia. In addition to similarity in shape and color

pattern, males are behavioral mimics as well. Males occupy sunny perches

(e.g., twigs, grass stems), or sit in a clearing, and make forays identical to

prey-capture flights of a sympatric Ejferia sp. In 66% of the A. haruspex

forays, no other insects were in the vicinity (Table 1). While such forays

may serve for visual orientation, this species also flew at other insects passing

nearby. Both inter- and intraspecific encounters were observed, but never

any case of predation. Species of the genus Apiocera are, in fact, not pre¬

daceous (Paramonov, 1953; Lavigne, 1975), and have only sponging-type

mouth parts.

Oviposition and fecundity.—The oviposition behavior of six females was

recorded. The following sequence was noted: female stood with body at steep

angle to substrate; abdomen pushed from side to side into substrate; complete

burial of abdomen and part of thorax accomplished in 5-10 sec; female

remained buried for 1.5-2.0 min; abdomen pulled out of soil in 5-10 sec;

VOLUME 58, NUMBER 4

299

tip of abdomen usually wriggled briefly over surface; fly moved quickly to

one side and quickly kicked sand over the oviposition site with hind legs;

female moved to new oviposition site. Total oviposition time was between

2 and 3 min. Similar to findings of Cazier (1963) with other Apiocera species,

females oviposit in sandy soils in shaded situations, most often in leaf litter

or near the base of small bushes. False starts were commonly noted, with

females inserting only the apical 1-3 segments into the substrate before

withdrawing without ovipositing. Cazier (1963) suggested that failure to

oviposit in such cases was most probably due to the fly encountering an

obstacle or harder subsurface soils before the abdomen was fully distended.

All observations on oviposition were made in the afternoon.

One cluster of five eggs was uncovered in the field. Two adult females

were dissected, one of which had eight follicles per ovariole and contained

200 eggs in various stages of development. The second female had only 16

very large eggs filling the abdominal cavity.

Discussion

Available information suggests that larvae of Apiocera species are very

similar to each other. Major features are the shape of the posterior spiracles,

head, and caudal segment; presence of a lateral keel, dorsal rod, and capsule

rod; shape and placement of mandibles, maxilla, and maxillary palp. All of

the above features differ in the first instar larva of Tongamya miranda (Irwin

and Stuckenberg, 1972). Although such lack of agreement between genera

may merely reflect differences among instars, the magnitude of the dissim¬

ilarities suggests that the various character states may be representative of

distinct phylogenetic lines. It will thus be of interest to discover if other

members of the Megascelinae have larvae similar to that of Tongamya-, and

if Rhaphiomydas has larvae similar to Apiocera.

Combining the observations on A. haruspex with those of Cazier (1963)

on A. painteri and Lavigne (1975) on A. clavator, mating in Apiocera consists

of an initial pairing, followed by a stationary period in a tail-to-tail position,

and finally an active uncoupling period which may include short flights.

Similar behavior has been reported for some asilid species (Lavigne and

Holland, 1969; Dennis and Lavigne, 1975). Known mydid species, however,

mate in a male over female position (Wharton, 1981).

Male forays probably serve a dual purpose. They may function in behav¬

ioral mimicry of their aggressive asilid models and they may also serve to

increase intraspecific encounters, which may be important for initial meeting

of the sexes. Toft and Kimsey (personal communication) are working on the

role of forays and other behavioral traits in aggregation behavior. Estab¬

lishment of temporary territories in conjunction with forays was also noted

for the asilid Proctacanthella leucopogon (Williston) by Lavigne and Dennis

(1980), and for My das by Wilcox and Papavero (1971).

300

PAN-PACIFIC ENTOMOLOGIST

Oviposition in sandy substrates is noticeably similar in some asilids (Mei-

lin, 1923), mydids (Hesse, 1974; Wilcox and Papavero, 1971; Wharton, 1981),

apiocerids, and the Thereva-growp in therevids (Irwin, 1976). In these groups,

species that oviposit in sandy soils possess acanthophorites. Many other

therevids also possess acanthophorites, but their oviposition behavior is

more complex (Irwin, 1976). The long, telescopic abdomen and burial of

abdomen (and often part of the thorax) allow for deep penetration and egg

deposition away from surface predators and parasitoids; and in hot, arid

regions, where most such species occur, provide a cooler environment for

developing eggs.

Acanthophorites serve in initial excavation (Meilin, 1923), in creating a

temporary oviposition chamber by rapid twirling immediately prior to egg

deposition (Cazier, 1963; personal observation), and in scraping the ovi¬

position site afterwards (Lavigne and Holland, 1969; Dennis and Lavigne,

1975). Placement of the stout, modified setae on the acanthophorite reflects

their use in oviposition. In A. haruspex, setae occur in lateral rows, one row

on each side. As the abdomen is inserted into the soil during oviposition,

it is moved from side to side—for maximum efficiency in use of the setae

as excavators. In Proctacanthus, an asilid, setae are concentrated in a cluster

posteriorly. As the abdomen is inserted into the soil, it is moved back and

forth rather than from side to side—again for maximum efficiency during

excavation.

Irwin (1976) discussed plesiomorphic and apomorphic states for the acan¬

thophorite. The laterally placed rows of thickened setae, while undoubtedly

an apomorphic state, are nevertheless found throughout the Asiloidea. This

condition is thus a relatively stable one, and indicative of common ancestry.

Similarities in the oviposition behavior of apiocerids, certain mydids, and

some asilids suggest that such characteristics are of a plesiomorphic nature.

This supports the findings of Irwin (1976) that similar behavior in the The-

reva- group is the plesiomorphic state within the Therevidae.

Clutch size and duration of oviposition are similar in Apiocera, several

mydids (Wharton, 1981), and asilids which oviposit in the soil (Dennis and

Lavigne, 1975). Clutch size is considerably larger in Tongamya, however

(Irwin and Stuckenberg, 1972). The striking differences between Tongamya

and Apiocera, both biologically and morphologically, further suggest that

the two should be regarded as only distantly related.

Acknowledgments

I am most grateful to Dr. M. Cazier for verifying the identity of A. ha¬

ruspex', and to Drs. M. Irwin and M. Cazier for suggesting improvements.

Dr. C. Toft kindly made available her unpublished work on apiocerids,

preventing unneeded duplication.

VOLUME 58, NUMBER 4

301

Literature Cited

Cazier, M. A. 1963. The description and bionomics of a new species of Apiocera, with notes

on other species (Diptera: Apioceridae). Wasmann J. Biol., 21:205-234.

Cole, F. R. 1969. The flies of western North America. University of California Press, Berkeley

and Los Angeles, 693 pp.

Dennis, D. S., and R. J. Lavigne. 1975. Comparative behavior of Wyoming robber flies II.

(Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr., 30:1-68.

English, K. M. I. 1947. Notes on the morphology and biology of Apiocera maritima Hardy

(Diptera, Apioceridae). Proc. Linn. Soc. N. S. W., 71:296-302.

Hesse, A. J. 1974. A new South African representative of the South West African genus

Namibimydas Hesse (Diptera: Mydaidae), with some ecological notes on the habits of

the species. Ann. South Afr. Mus., 66:25-34.

Irwin, M. E. 1976. Morphology of the terminalia and known ovipositing behaviour of female

Therevidae (Diptera: Asiloidea), with an account of correlated adaptations and comments

on phylogenetic relationships. Ann. Natal Mus., 22:913-935.

-, and B. R. Stuckenberg. 1972. A description of the female, egg and first-instar larva

of Tongamya miranda, with notes on oviposition and the habitat of the species (Diptera:

Apioceridae). Ann. Natal Mus., 21:439-453.

Lavigne, R. J. 1975. Redescription of Apiocera clavator with notes on its behavior (Diptera:

Apioceridae). Ann. Entomol. Soc. Am., 68:673-676.

-, and D. S. Dennis. 1980. Ethology of Proctacanthella leucopogon in Mexico (Diptera:

Asilidae). Proc. Entomol. Soc. Wash., 82:260-268.

-, and F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming robber

flies (Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr., 18:1-61.

Meilin, D. 1923. Contributions to the knowledge of the biology, metamorphosis and distri¬

bution of the Swedish asilids—in relation to the whole family of asilids. Zool. Bidr.

Upps., 8:1-317.

Norris, K. R. 1936. New species of Apioceridae (Diptera) from Western Australia. J. R. Soc.

West. Aust., 22:49-70.

Paramonov, S. J. 1953. A review of the Australian Apioceridae (Diptera). Aust. J. Zook, 1:

449-537.

Wharton, R. A. 1981. Mydas fly populations in the central Namib Desert (Diptera: Mydidae).

Ann. Transvaal Mus., 33:145-151.

Wilcox, J., and N. Papavero. 1971. The American genera of Mydidae (Diptera), with the

description of three new genera and two new species. Arq. Zool. (Sao Paulo), 21:41-119.

Footnote

1 The observations of Toft and Kimsey (1982, J. Kans. Entomol. Soc., 55:177-186) were

published while this paper was in press and are not incorporated herein.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 302-308

Published 30 December 1983

A REVIEW OF THE HISTORY AND TAXONOMY OF

ECONOMICALLY IMPORTANT SERPENTINE LEAFMINERS

{LIRIOMYZA SPP.) IN CALIFORNIA

(DIPTERA: AGROMYZIDAE)

Michael P. Parrella

Department of Entomology, University of California, Riverside 92521

Liriomyza trifolii (Burgess) is a common serpentine leafminer on cut and

pot chrysanthemums in California. This insect has recently become a serious

threat to the chrysanthemum industry, presumably due to the development

of resistance to many commercially registered insecticides. Research nec¬

essary to develop a pest management program for L. trifolii on chrysanthe¬

mums is currently in progress. Part of this research effort has involved a

search of the literature for previous work done on other economically im¬

portant members of the genus Liriomyza Mik. According to Spencer (1973),

there are four species of economic importance in California: L. brassicae

(Riley), L. huidobrensis (Blanchard), L. sativae Blanchard, and L. trifol-

iearum Spencer. A fifth can be added to this list, L. trifolii, which was

introduced into California after 1973.

There has been considerable taxonomic confusion in the past with regard

to the polyphagous Agromyzidae. This has been particularly true with mem¬

bers of the genus Liriomyza, due to their wide, overlapping host ranges and

general morphological similarity. Although only five economically important

Liriomyza spp. are currently recognized as occurring in California, the lit¬

erature records 15 species. This is a result of numerous misidentifications

and synonymies that have been determined since these early works. Spencer

(1973) prepared the definitive work on the systematics of the Agromyzidae

and in this he clarifies the present status of most of the California species.

Steyskal (1973) summarized Spencer’s clarification of the status of Liriomyza

spp. in the United States and his list of species is identical to the list for

California presented here.

This paper examines: (1) the taxonomic confusion that has occurred with

the above five species in California including citations not covered by Spen¬

cer (1973); and (2) the host plants from which these species have been reared

in California including recent surveys of economically important hosts for

several of these flies completed in 1980. The purpose is to provide a concise

chronological reference for researchers working with this important group

of leafmining flies.

VOLUME 58, NUMBER 4

303

Liriomyza brassicae (Riley)

This species is the most cosmopolitan of all the Agromyzidae and occurs

primarily on Cruciferae. There are limited records of L. brassicae in Cali¬

fornia. Oatman and Platner (1969) examined the population trends and

parasitization of this species on cabbage in southern California. They con¬

cluded that L. brassicae was of minor economic importance.

Liriomyza huidobrensis (Blanchard)

Liriomyza huidobrensis, the pea leafminer (Sutherland, 1978), is Nearctic

and Neotropical in distribution. This species was first cited in California as

a pest of peas and spinach by Lange (1945) (as L. orbona (Meigen)). Sub¬

sequent work by Smith and Lange (1946) and Lange and Smith (1947)

involved control of L. huidobrensis (as L. orbona (Meigen)) on peas. Lange

(1949) discussed the occurrence of leafmining flies in California and stated

that the species causing damage along the California coast, in coastal valleys

and in areas having a coastal influence was L. huidobrensis (as Agromyza

(. Liriomyza ) flaveola Fallen). This species was considered a pest of cruciferous

crops, lettuce, melons, peas, sugar beets, tomatoes and cultivated flowers.

In 1957, Lange et al. discussed L. huidobrensis (as L. langei Frick) as a

serious pest of spinach. They indicated this species also attacked peas, pep¬

pers, lettuce, carrots, onions, asters, celery, beans, cineraria, zinnia, stock,

guayule, cabbage, brussel sprouts, kohlrabi, cauliflower, turnip, broccoli,

okra, parsnip, radish, dandelion, endive, chicory, and rutabagas.

Control measures for L. huidobrensis as a pest of asters were explored by

Jefferson and Pence (1948) and Jefferson and Eads (1949) (as L. flaveola

(Fallen)) and by Jefferson and Eads (1952) (as L. langei Frick).

Frick (1951) described a new species of serpentine leafminer, L. langei,

from peas, sugar beets, spinach, celery and aster. Later, Frick (1958) de¬

scribed a new species of leafminer from carnation, L. dianthi, which he

(Frick, 1964) synonymized with L. langei. L. langei was then synonymized

with L. huidobrensis (Blanchard) by Spencer (1973).

Elmore and Ranney (1954) described the injury to seedling pepper plants

by L. huidobrensis (as L. langei Frick), and Wilcox and Howland (1955)

examined control measures for L. huidobrensis (as L. langei Frick) on sugar

beets. Pritchard (1957) discussed a new leafmining pest of carnations (as

Liriomyza sp.) which was probably L. huidobrensis.

During 1980, L. huidobrensis was reared from gypsophila grown in the

San Diego area.

Liriomyza sativae Blanchard

Liriomyza sativae is Nearctic and Neotropical in distribution, occurring

on a wide range of plants. In the San Joaquin and Sacramento Valleys Lange

304

PAN-PACIFIC ENTOMOLOGIST

(1949) indicated L. sativae (as Agromyza ( Liriomyza) subpusilla Frost) was

the most common species causing damage to alfalfa, beans, melons and

tomatoes. In southern California, Lange (1949) found L. sativae (as Agro¬

myza ( Liriomyza) pusilla Meigen) damaging tomato.

Tilden (1950) reported on the oviposition and behavior of L. sativae (as

L. pusilla (Meigen)) on Baccharis pilularis D.C. This reference indicates that

larvae have the ability to leave one leaf and to enter another of the same

host species.

Michelbacheretal. (1949, 1951, 1952, 1953, 1955) discussed the chemical

control of Liriomyza spp. as a pest of tomatoes (probably L. sativae ) and

L. sativae as a pest of melons (as L. subpusilla Frost). Wilcox and Howland

(1952) reported on the control of Liriomyza spp. (probably L. sativae ) on

tomatoes in southern California.

In 1957, Frick synonymized Agromyza {Liriomyza) pusilla (Meigen),

Agromyza ( Liriomyza) subpusilla Frost and Liriomyza subpusilla (Frost)

with his new species L. munda. Spencer (1973) synonymized L. munda

Frick with L. sativae Blanchard, the vegetable leafminer (Sutherland, 1978).

Frick (1957) discussed a new combination within the species L. pictella

(Thomson) and subsequently identified the “melon leafminer” discussed by

Oatman and Michelbacher (1958, 1959) and Oatman (1959a, 1959b, 1960a,

1960b, 1961). The present status of L. pictella in these studies is unclear

and as indicated by Oatman (1961) may be a sibling species of L. sativae

from sympatric natural populations.

Shorey et al. (1962) and Shorey and Hall (1963) reported on the toxicities

of insecticides to Liriomyza spp. (probably L. sativae) on poled tomatoes

in southern California.

Jensen (1969) and Jensen and Koehler (1970) discussed the status of L.

sativae (as L. munda) on alfalfa. In these papers the authors indicate that

two species of leafminers occur on alfalfa, L. munda and L. pictella. It is

interesting that they state that the papers published by Oatman dealt with

L. munda, not L. pictella, but that in alfalfa both species were present.

Spencer (1973) identified the L. pictella of Jensen (1969) and Jensen and

Koehler (1970) as a new species, L. trifoliearum (discussed below).

In 1976, Oatman and Kennedy demonstrated that outbreaks of L. sativae

on tomatoes could be caused by applications of methomyl. This was followed

by the recent work of Johnson (1979) and Johnson et al. (1980a, 1980b,

1980c, 1980d) with L. sativae as a pest of tomatoes where the parasite

complex, effect of pesticides, and sampling plans were examined. Also in

1979, Hoskinson completed a study on the spatial and temporal distribution

of L. sativae on pole tomatoes in southern California. Even though research¬

ers working with leafmining Diptera on tomatoes in southern California

since 1976 have cited only one species, it is probable that mixed populations

of L. sativae and L. trifolii were present. During 1980, L. sativae was reared

VOLUME 58, NUMBER 4

305

from tomato and squash grown in southern California. Selected chemicals

were evaluated for control of L. sativae as a pest of summer squash (Sharma

et al., 1980). They reported that several materials provided good control

and significantly increased yields.

Liriomyza trifoliearum Spencer

Liriomyza trifoliearum is Nearctic in distribution and has a rather narrow

host range compared to the other economically important Liriomyza spp.

in California. This leafminer has been cited as a pest of alfalfa in California

(as Liriomyza pictella Thomson) (Jensen, 1969; Jensen and Koehler, 1970).

Damage to alfalfa was caused by the tendency of mined leaflets to drop from

the plant before or during harvest. This species was not collected during

1980.

Liriomyza trifolii (Burgess)

This species is Nearctic and Neotropical in distribution and enjoys a wide

host range. Frick (1959) described specimens from California, Oregon and

Washington as L. trifolii. Those from the latter state were described as new

species by Spencer (1965) and the flies from California and Oregon were

misidentifications. The true L. trifolii is a recent introduction into California

(1975 or 1976) probably originating on chrysanthemum cuttings from Flor¬

ida. This species was reared from chrysanthemums, verbena, cineraria, ca¬

lendula, gypsophila, gerbera, snapdragon, sugar beans, tomato and celery in

southern California during 1980. Bivins and McCloskey (1978) discussed

chemical control of this species and Oetting et al. (1981) examined chry¬

santhemum varietal susceptibility to this leafminer species. Liriomyza trifolii

is currently a very serious pest of chrysanthemums and is becoming more

important as a pest of tomato and celery.

Discussion

The probable specific determinations of Liriomyza spp. in California were

made on the assumption that there are five economically important species

in California. The actual flies worked with by these researchers were not

examined. Identifications were made using Spencer (1973) and Frick (1951,

1957, 1958, 1964) and by examining how each researcher viewed the iden¬

tifications of flies from previous related work.

True polyphagy in the Agromyzidae is rare (Spencer, 1964) with only 10

polyphagous species recorded throughout the world (Spencer, 1977). It is

probable that few, if any, further dominant and widespread polyphagous

species will be found. California is in the unenviable position of having at

least four polyphagous Liriomyza species. However, as stated by Spencer

(1973) and demonstrated by Oatman (1961) the process of speciation among

the polyphagous Agromyzidae is probably being reversed and further spe-

306

PAN-PACIFIC ENTOMOLOGIST

ciation can be expected. Thus, flies identified as the same species from

different parts of California may not necessarily exhibit similar character¬

istics, even though they may be from the same host.

Acknowledgments

Appreciation is extended to Drs. John T. Trumble and Robert A. Van

Steenwyk, Department of Entomology, University of California at Riverside

for reviewing earlier drafts of this manuscript. The comments by two anon¬

ymous reviewers are also greatly appreciated.

Literature Cited

Bivens, J. L., and W. McCloskey. 1978. New compound effective against leafminer fly on

greenhouse chrysanthemums—Progress report. Flower and Nursery Report. Coop. Ext.,

Univ. Calif. Fall: 1-2.

Elmore, J. C., and C. A. Ranney. 1954. Injury to pepper plants by the pea leafminer. J. Econ.

Entomol., 47:357-358.

Frick, K. E. 1951. Liriomyza langei, a new species of leafminer of economic importance in

California. Pan-Pac. Entomol., 27:81-88.

-. 1957. Nearctic species in the Liriomyza pusilla complex. No. 2. L. munda and two

other species attacking crops in California. Pan-Pac. Entomol., 33:59-70.

-. 1958. Liriomyza dianthi n. sp., a new pest of carnations in California. Proc. Entomol.

Soc. Wash., 60:1-5.

-. 1959. Synopsis of the species of agromyzid leaf miners described from North America.

Proc. U.S. Nat. Mus., 108:347-465.

-. 1964. Liriomyza dianthi, a new synonym of L. langei Frick (Diptera: Agromyzidae).

Pan-Pac. Entomol., 40:11-12.

Hoskinson, S. E. 1979. Spatial and temporal distribution of L. sativae Blanchard (Diptera:

Agromyzidae) on fall tomatoes, with implications in pest management. M.S. Thesis, San

Diego State University, 82 pp.

Jefferson, R. N., and C. O. Eads. 1949. Control of leaf miners on field-grown asters. Univ.

Calif. (Los Angeles) News Letter No. 41.

-, and-. 1952. Control of leaf miners and other insect pests of aster. J. Econ.

Entomol., 45:476-481.

-, and R. J. Pence. 1948. Preliminary experiments on the control of the leafminer

Liriomyza flaveola on asters. J. Econ. Entomol., 41:653-654.

Jensen, G. L. 1969. Investigations of the economic importance of certain Liriomyza species

on alfalfa in north and central California (Diptera: Agromyzidae). Thesis, University of

California, Berkeley, 125 pp.

-, and C. S. Koehler. 1970. Seasonal and distributional abundance and parasites of leaf

miners of alfalfa in California. J. Econ. Entomol., 63:1623-1628.

Johnson, M. W. 1979. Insect pest management strategies for control of Liriomyza sativae

Blanchard (Diptera: Agromyzidae) on pole tomatoes in southern California. Dissertation,

University of California, Riverside, 126 pp.

-, E. R. Oatman, and J. A. Wyman. 1980a. Effects of insecticides on populations of the

vegetable leafminer and associated parasites on summer pole tomatoes. J. Econ. Ento¬

mol., 73:61-66.

-,-, and-. 1980b. Effects of insecticides on populations of the vegetable

leafminer and associated parasites on fall pole tomatoes. J. Econ. Entomol., 73:67-71.

VOLUME 58, NUMBER 4

307

-,-, and-. 1980c. Natural control of Liriomyza sativae (Dip.: Agromyzidae)

in pole tomatoes in southern California. Entomophaga, 25:193-198.

-,-,-, and R. A. Van Steenwyk. 1980d. A technique for monitoring Lir¬

iomyza sativae in fresh market tomatoes. J. Econ. Entomol., 73:552-555.

Lange, W. H., Jr. 1945. Investigations with DDT in California, 1944. Univ. Calif. Agr. Exp.

Stn. : 18-22.

-. 1949. Notes on the occurrence of agromyzid flies during 1948, and a record of two

unreported species in California. Pan-Pac. Entomol., 25:91-92.

-, A. A. Grigarick, and E. C. Carlson. 1957. Serpentine leafminer damage. Calif. Agr.,

11:3-5.

-, and L. M. Smith. 1947. Control of a leafminer on peas. J. Econ. Entomol., 40:496-

499.

Michelbacher, A. E., O. G. Bacon, and J. Underhill. 1953. Leaf miner on tomato, control by

dieldrin. Studied for northern California conditions. J. Econ. Entomol., 7:15.

-, W. W. Middlekauf, O. G. Bacon, and J. E. Swift. 1955. Controlling melon insects

and spider mites. Calif. Agr. Exp. Stn. Bull., 749:1-46.

-,-, and L. C. Glover. 1951. Studies with aldrin and dieldrin against melon insects.

J. Econ. Entomol., 44:390-393.

-,-, and-. 1952. Aldrin, dieldrin and heptachlor to control California melon

insects. J. Econ. Entomol., 44:470-475.

-,-, F. C. Lamb, and N. B. Akesson. 1949. Further investigations of control of

tomato insects in northern California. J. Econ. Entomol., 42:666-674.

Oatman, E. R. 1959a. Host range studies of the melon leaf miner, Liriomyza pictella (Thom¬

son) (Diptera: Agromyzidae). Ann. Entomol. Soc. Am., 52:739-741.

-. 1959b. Natural control studies of the melon leaf miner, Liriomyza pictella (Thomson)

(Diptera: Agromyzidae). J. Econ. Entomol., 52:759-762.

-. 1960a. Intraspecific competition studies of the melon leaf miner, Liriomyza pictella

(Thomson) (Diptera: Agromyzidae). Ann. Entomol. Soc. Am., 53:130-131.

-. 1960b. Parasitism of the overwintering pupae of the melon leaf miner, Liriomyza

pictella. J. Econ. Entomol., 53:682.

-. 1961. Crossbreeding studies with two closely related species of Liriomyza. Pan-Pac.

Entomol., 37:53-57.

-, and G. G. Kennedy. 1976. Methomyl induced outbreak of Liriomyza sativae on

tomato. J. Econ. Entomol., 69:667-668.

-, and A. E. Michelbacher. 1958. The melon leaf miner Liriomyza pictella (Thomson)

(Diptera: Agromyzidae). I. Life history studies. Ann. Entomol. Soc. Am., 51:557-566.

-, and -. 1959. The melon leaf miner, Liriomyza pictella (Thomson) (Diptera:

Agromyzidae). II. Ecological studies. Ann. Entomol. Soc. Am., 52:83-89.

-, and G. R. Platner. 1969. An ecological study of insect populations on cabbage in

southern California. Hilgardia, 40:1-40.

Oetting, R. D., F. S. Morishita, and A. L. Helmkamp. 1981. Susceptibility of selected chry¬

santhemum cultivars to leafminer damage. Flower and Nursery Rep., Coop. Ext., Univ.

Calif., Winter, p. 5.

Pritchard, E. A. 1957. New carnation pests. Bud mite and leaf miner found in California may

cause serious problems. Calif. Agr., 11:5.

Sharma, R. K., A. Durazo, and K. S. Mayberry. 1980. Leafminer control increases summer

squash yields. Calif. Agr., 34:21-22.

Shorey, H. H., and I. M. Hall. 1963. Toxicity of chemical and microbial insecticides to pest

and beneficial insects on poled tomatoes. J. Econ. Entomol., 56:813-817.

-, H. T. Reynolds, and L. D. Anderson. 1962. Effect of Zectran, Sevin, and other new

carbamate insecticides upon insect populations found on vegetables and field crops in

southern California. J. Econ. Entomol., 55:5-11.

308

PAN-PACIFIC ENTOMOLOGIST

Smith, L. M., and W. H. Lange, Jr. 1946. Investigations with DDT and other new insecticides—

Experiments with DDT and lead arsenate for controlling the pea leafminer and other

pea insects. Calif. Agr. Exp. Stn. Circ., 365:52-54.

Spencer, K. A. 1964. The species-host relationship in the Agromyzidae as an aid to taxonomy.

Int. Cong. Entomol. Proc., 12:101-102.

-. 1965. A clarification of the status of Liriomyza trifolii (Burgess) and some related

species. Proc. Entomol. Soc. Wash., 67:32-40.

-. 1973. Agromyzidae (Diptera) of economic importance. Dr. W. Junk B. V., The Hague,

Netherlands, 418 pp.

-. 1977. A revision of the Australian Agromyzidae. West. Aust. Mus. Special Pub.,

8 : 21 .

Steyskal, G. C. 1973. The strange fate of the “serpentine leaf miner” (Liriomyza spp., Agro¬

myzidae, Diptera). U.S.D.A. Coop. Econ. Ins. Rep., 23:735-736.

Sutherland, D. W. S. 1978. Common names of insects and related organisms (1978 revision).

Special Publication, Entomological Society of America, 132 pp.

Tilden, J. W. 1950. Oviposition and behavior of Liriomyza pusilla (Meigen). Pan-Pac. Ento¬

mol., 26:119-121.

Wilcox, J., and A. F. Howland. 1952. Control of dipterous leaf miner on tomatoes in California.

J. Econ. Entomol., 45:634-639.

-. 1955. Control of the pea leaf miner in southern California. J. Econ. Entomol., 48:

579-581.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 309-311

Published 30 December 1983

A NEW SPECIES OF ANDRENA (MICRANDRENA ) FROM

COLORADO, WITH OTHER NOTES ON THE GROUP

(HYMENOPTERA: APOIBEA)

U. N. Lanham

University of Colorado Museum, University of Colorado, Boulder 80309

The new species described below, so far known only in the female, closely

resembles A. melanochroa Cockerell, and runs to that species in the key in

Ribble’s monograph of the subgenus Micrandrena (1968). Both species are

included in Ribble’s species group of A. piperi Viereck. The key can be

modified to include the new species by the following change in the vicinity

of couplet 24:

24. Enclosure of the propodeum with at least dorsal surface very

granular, so as to appear beaded, contrasting with smoother

surface of rest of propodeum .24a

Enclosure of propodeum not especially granular, little contrast

between enclosure and rest of propodeum. 25

24a. Facial fovea abruptly narrowed in lower third to half or less the

width of upper portion, integument between eye and narrow

part of fovea widened to at least half width of adjacent fovea;

clypeus with a few shallow, wide irregular wrinkles, lower por¬

tion with few very irregularly spaced strong punctures, upper

half reticulate, more evenly and finely punctate, somewhat flat¬

tened; process of labrum narrowly triangular, pointed.

. kristina, n. sp.

Facial fovea normal, only gradually narrowed below, integumen-

tal strip between narrow portion and eye not widened; clypeus

evenly convex, lower shining portion rather evenly and finely

punctured, becoming impunctate toward upper margin; process

of labrum transverse, broadly rounded or subrectangular ....

. melanochroa Cockerell

25. piperi, etc.

In the Boulder area, melanochroa is the common Micrandrena of Poten-

tilla fissa Nutt., while kristina is found on Physaria bellii G. A. Mulligan, a

local endemic crucifer.

Andrena kristina Lanham, new species

Female. — Length 7 mm, forewing 6 mm. Integument black, except for

hyaline amber apical margins of abdominal terga and reddish brown flagella.

310

PAN-PACIFIC ENTOMOLOGIST

Head having width/length ratio 4.8/4.0, measuring between outer eye orbits

and from vertex to bottom of clypeus, hairs white. Clypeus with a few

irregular shallow wrinkles, especially above; lower half shining, with a few

coarse and irregularly spaced punctures, upper half flattened, reticulate, more

regularly punctured. Face above clypeus vertically striate, above ocelli be¬

coming coarsely granular, with polished impunctate area at dorsal ends of

compound eyes. Facial foveae with dorsal ends extending slightly above

eyes, width here slightly less than V 2 distance between eye and lateral ocellus,

extending below beyond antennal insertions but not as far as upper margin

of clypeus, lower third abruptly narrowed to V2-V3 width of upper portion

by incurving inner margin of fovea and increasing width of integument

between fovea and eye, which becomes more than V2 as wide as fovea.

Antennae with scape plus pedicel reaching to middle of lower ocellus; 1 st

segment of flagellum longer than 2 4- 3 on outer margins. Process of labrum

narrowly triangular. Mouthparts normal for the piperi group, 1 st 3 maxillary

segments about equal to length of galea. Thorax. Pubescence on the thorax

long for the group, sparse and erect dorsally, denser and decumbent on sides,

color white tending to brown on median posterior surfaces of dorsum. Prono-

tum without shoulders, but with strong groove running anterio-medially

from base of pronotal lobe. Mesonotum shining but strongly reticulate, punc¬

tures small, sparse, 4-5 puncture-widths apart, mesoscutum about the same;

metanotum coarsely reticulate; propodeum coarsely beaded, with delicate

but distinct ridges on dorsal face, sculpture at edges of enclosure about same

as that of adjacent part of propodeum. Corbicula without anterior fringe,

dorsal fringe poorly developed, interior with a few long simple hairs widely

scattered over most of surface, integument of interior shining, rather coarsely

reticulate. Hind leg with trochanteral floccus complete but sparse; hairs of

tibial scopa simple, long, the hairs on posterior margin about as long as

width of tibia at its widest point, entire scopa moderately loose and sparse;

all hairs white, except for dark tuft at tibio-femoral junction; tarsal claw with

tooth fully developed, of size normal for the piperi group. Wing membranes

clear, veins amber, 1st transverse cubital vein ending 1 to 3 vein widths

distad from pterostigma, basal vein falling well distad of transverse medial

nervure. Abdomen with terga semi-shining, reticulation moderately coarse.

Tergum 1 without apical hair band, 2 with widely interrupted weak white

hair band, 3 with band narrowly interrupted, 4 with band entire, caudal

fimbria entirely brown tinged with orange; terga anterior to fimbria without

long hairs dorsally; pygidial plate finely granular, narrowly rounded, with

weakly developed raised central triangular area.

Type material.— Holotype, female, 4 miles north Boulder, COLORADO,

5500 feet, 23 May 1980, collected at flowers Physaria bellii (U. N. Lanham).

Seventeen paratypes, females: 2 with same data as holotype; 4 same except

24 and 25 May 1980 (K. Neff); 8 from 10 miles SW Loveland, COLORADO,

VOLUME 58, NUMBER 4

311

26 May 1980, collected at flowers Physaria bellii (U. N. Lanham); 1 from

3 miles east Lyons, COLORADO, 7 June 1980 (U. N. Lanham and K. Neff);

2 from 12 miles north Ft. Collins, COLORADO, 7 June 1980 (U. N. Lanham

and K. Neff).

The name is pronounced with the second syllable accented and the “i” as

in mile.

All localities named in the type description are on or very near the outcrop

of Cretaceous Niobrara limestone and shale that lies at the eastern base of

the foothills of the Front Range. This habitat is described in “Some Colorado

Andrena of the subgenus Scaphandrena of presumed hybrid origin . . . ,”

Lanham, 1981.

Of the 11 species of the piperi group of Micrandrena only three, including

piperi, show a strong preference for Cruciferae.

The rather remarkable primulifrons group of Micrandrena is apparently

oligolectic on crucifers, according to Ribble. The group is distinctive on

account of the heavy and coarsely punctate integument. The two species of

the group, primulifrons Casad and trapezoidea Viereck have previously been

known only from the Southwest. It is therefore of interest to find a species

that matches Ribble’s description of trapezoidea on crucifers in northeastern

Colorado. I collected two females and a male at Briggsdale, Weld Co., 19

and 21 May 1979 from flowers of Descuraina and 2 females, 10 miles SW

Loveland, 26 May 1980 from Physaria bellii. In his Micrandrena paper

Ribble suggests that the primulifrons group is annectant between that sub¬

genus and Scaphandrena. His monograph of Scaphandrena (1974) contains

the statement that the group should be transferred to Scaphandrena, ap¬

parently as an afterthought, since the group is not included in the key to

species.

Literature Cited

Lanham, U. N. 1981. Some Colorado Andrena of the subgenus Scaphandrena of presumed

hybrid origin, with special reference to the tarsal claws (Hymenoptera: Apoidea). J. Kans.

Entomol. Soc., 54:537-546.

Ribble, D. W. 1968. Revisions of two subgenera of Andrena: Micrandrena Ashmead and

Derandrena, new subgenus (Hymenoptera: Apoidea). Bull. Univ. Nebr. State Mus., 8:

237-394.

-. 1974. A revision of the bees of the genus Andrena of the Western Hemisphere subgenus

Scaphandrena. Trans. Am. Entomol. Soc., 100:101-189.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 312-318

Published 30 December 1983

BIOLOGY OF TETRALEURODES ACACIAE (QUAINTANCE)

(HOMOPTERA: ALEYRODIDAE ) 1

Robert Y. Dowell 2

Agricultural Research Center, University of Florida,

3205 S. W. 70 Avenue, Fort Lauderdale 33314

Tetraleurodes acaciae (Quaintance) is a legume-feeding whitefly found in

California, Cuba, Florida, Jamaica, and Mexico (Quaintance, 1900; Hamon,

1978; Mound and Halsey, 1978). First described as Aleyrodes acaciae by

Quaintance in 1900, it was placed in the genus Tetraleurodes by Quaintance

and Baker in 1914. Despite its being an occasional pest of powder puff plants

('Calliandra haematocephala Hassk.) in Florida (Hamon, 1978), little is known

of its life history or biology. Information on the developmental stages, pred¬

ators, parasitoids, pathogens, and adult behavior of T. acaciae is provided

here.

Methods and Materials

Samples of 20 leaves from powder puff plants in Plantation and Davie,

Broward County, Florida were taken at irregular intervals between January

1978 and August 1980. Five leaflets/sample were held at 26-28°C and 50-

70% RH for parasitoid emergence. The remainder were examined for in¬

formation on size, shape, color, density and distribution of larval stages,

and the presence of pathogenic fungi. Adult whiteflies and predators were

collected from ten 25 cm long branch tips by dislodging them into plastic

bags. The bags and their contents were frozen and later examined for predator

species and number, and whitefly sex ratio. The behavior of T. acaciae adults

and associated insects was observed for thirty minutes/sample date.

Several other whitefly species have demonstrated a preference for yellow

sticky traps (Dowell et al., 1979; Mound, 1962, Vaishampayan et ah, 1975),

and Fitzpatrick et al. (1979) effectively used a yellow sticky trap to monitor

population trends of citrus blackfly, Aleurocanthus woglumi Ashby. To de¬

termine if T. acaciae prefers yellow traps, the response of T. acaciae adults

to sticky traps of different colors was determined over two T. acaciae gen¬

erations between March and June 1978. Translucent yellow coffee can lids

were tested against the same lids painted green, black, red, and white (Dowell

et al., 1979b). The traps were covered with Tanglefoot 1 and one trap of each

color hung on 5 visually isolated powder puff plants for 48 hours after which

the number of T. acaciae/Xmp was counted. This procedure was repeated 5

times for a total of 25 traps exposed/color. The data were pooled by date

VOLUME 58, NUMBER 4

313

prior to analysis of variance and least significant difference tests. The re¬

sponse of T. acaciae adults to yellow traps of varying sizes (7.5-136 cm 2 )

was determined in the same manner over the same time period.

Concurrent held work on other projects (Dowell, 1979; Dowell et al.,

1979a; Dowell and Cherry, 1981) provided the opportunity to look for T.

acaciae infestations on many plant species.

Results and Discussion

The lengths and widths of the eggs and the four larval instars are shown

in Figure 1. Like the citrus blackfly, the immature female is markedly larger

than the male in the 3rd and 4th instars (Fig. 1) (Dietz and Zetek, 1920;

Dowell et al., 1981).

The newly emerged first instar is translucent green with short antennae

and clearly visible eyes. Like other whiteflies it is the only mobile stage

besides the adult. Unlike most other whiteflies, the first instar shows a distinct

tendency to migrate from the lower surface of the leaf (site of oviposition)

and settle on the upper surface. Eggs are deposited singly on the underside

of host plant leaves. Only one of 420 eggs was laid on the upper surface of

25 leaflets examined; 120 of 396 nymphs had settled on the upper surface

of the same leaflets.

The 2nd-4th instar larvae are black with a pronounced white fringe around

the lateral margin. The instars are most easily separated by size (Figs. 1, 2).

Hamon (1978) gives a detailed description of the 4th instar larva (pupa).

The adults are uniform light yellow in color and of typical aleyrodid shape.

Both sexes are small with the female (0.83 ± 0.032 mm long) being larger

than the male (0.70 ± 0.031 mm long). Both sexes have immaculate pow¬

dery-white wings and incompletely separated red eyes. Unlike the citrus

blackfly, or cloudy-winged white fly [ Dialeurodes citrifolii (Morgan)], both

sexes are very active flyers. When populations are large, thousands of adults

are constantly in flight around infested plants.

There are ~8 generations/year with all stages continuously present

throughout the year in south Florida. A total of 2449 adults were sexed, of

which 1226 were males and 1223 were females giving a 1:1 sex ratio. T.

acaciae host plants are generally in the Leguminosae. In addition to the

plants listed in Bemis (1904), Hamon (1978), Mound and Halsey (1978),

and Quaintance (1900), T. acaciae has been observed infesting plants in the

following genera: Albizzia, Citrus, Coccoloba, and Lysiloma in Florida and

Albizzia, Amorpha, Brachysema, Ceratonia, Dolichos, Hardenbergia, Poin-

ciana, Pueraria, Robinia, Sophora, and Wisteria in California (Ray Gill,

CDFA personal communication). There are no data indicating which of the

recorded host plants support complete development of T. acaciae. High

densities of T. acaciae on C. haematocephala result in extensive sooty mold

314

PAN-PACIFIC ENTOMOLOGIST

LENGTH(mm)

Fig. 1. Relationship between developmental stages and their length and width. The bars

represent one standard deviation and the numbers in () indicate sample size. It is possible to

sex the 3rd and 4th instar nymphs. M = male, F = female.

growth on the leaf surfaces coated by honeydew excreted by the whitefly.

Premature leaf drop then gives the plants a “spindly” appearance with black

leaves. Sooty mold growth followed by defoliation are undesirable (Dowell

et al., 1979a) for ornamental plants which constitute the majority of T.

acaciae host plants.

The following predators were observed feeding upon various stages of T.

acaciae : Chilocorus stigma (Say), Cycloneda sp Delphastuspusillus LeConte

(Coleoptera: Coccinellidae), Chrysopa spp. (Neuroptera: Chrysopidae), Con-

dylostylus chrysoprasi (Walker) (Diptera: Dolichopodidae) and an uniden¬

tified ant species was observed carrying off newly emerged adults. Ants were

Figs. 2, 3. Fig. 2. Egg (a), 1st (b), 2nd (c), 3rd (d) and 4th instar (e) T. acaciae (42 x). Line

represents 1 mm. Note the white fringe around later 2nd-4th instars. Fig. 3. Underside of leaflet

showing T. acaciae nymphs (a) and Aschersonia aleyrodis growth (b) (12x).

VOLUME 58, NUMBER 4

315

316

PAN-PACIFIC ENTOMOLOGIST

0 _

I-

TRAP COLOR

Fig. 4. Relationship between trap color and number of T. acaciae adults caught. Yellow

traps (Y) caught significantly more whiteflies than the other colors tested [F = 57.25, P(5,24) <

0.05, 1 SD ± 68.14] (Red = R, Green = G, White = W, Blue = B, Black = Bl). Different letters

within the bars represent significantly different means at P = 0.05. All traps were 58 cm 2 /side.

Note the Log 10 scale on the y-axis.

never observed tending the larvae for honey dew. Numerous adult T. acaciae

were caught in the webs of various spiders. Three species of parasitic wasps

emerged from 4th instar T. acaciae. They were identified by Dr. L. A. Stange

(Florida State Collection of Arthropods, Gainesville, Florida) and James

Woolley (Entomology, University of California; Riverside) as Eretmocerus

Iportoricensis Dozier, Encarsia sp. (Hymenoptera: Aphelinidae), and Sig-

niphora sp. (Signiphoridae). The Signiphora sp. was the most abundant and

represented >60% of all parasitoids that emerged from sample material.

Specimens are held in the Florida State Collection of Arthropods in Gaines-

VOLUME 58, NUMBER 4

317

ville, Florida. Parasitization rates/sample date range from 1-28% with an

overall average of 19.6% of T. acaciae larvae being parasitized (249/1269)

by all species.

Each year (1978-1980) during August-November a pathogenic fungus (red

aschersonia) (Fig. 3) identified as Aschersonia aleyrodis Webb (Deutero-

mycotina) by Dr. J. J. McRitchie (Division of Plant Industry, Gainesville,

Florida) reduced T. acaciae populations by 96% (from 24/leaflet to 0.8/

leaflet in 1978) on the sample plants. T. acaciae populations rebounded the

following spring to original levels of 20-40 live larvae/leaflet. A. aleyrodis

has been previously reported attacking D. citrifolii and Dialeurodes citri

(Ashmead) (Berger, 1910, 1917) and appears to be a critical factor in reducing

high population densities of T. acaciae.

Yellow colored sticky traps caught significantly (P < 0.05) more T. acaciae

adults than the other colors tested (Fig. 4) and trap size had no measurable

influence (P > 0.05) on the number of adult whitefly caught per cm 2 trap

surface.

A small number of parasitoids of all three species were caught on the

yellow sticky traps. Dowell and Cherry (1981) have shown that such traps

can be used to monitor changes in the numbers of citrus blackfly parasitoids

and coccinellid predators.

Acknowledgments

I thank Dr. L. A. Strange and James Woolley for identification of the

parasitoids, Dr. J. J. McRitchie for identification of the fungus, and Ray

Gill for the information on California host plants.

Literature Cited

Bemis, F. E. 1904. The aleyrodids or mealy-winged flies of California with reference to other

American species. Proc. U.S. Nat. Mus., 27:471-537.

Berger, E. W. 1910. Whitefly control. Univ. Fla. Agric. Exp. Stn. Bull., 103:1-28.

-. 1917. Whiteflies of citrus. Month. Bull. Calif. State Comm. Hortic., 4:297-306.

Dietz, H. F., and J. Zetek. 1920. The blackfly of citrus and other sub-tropical plants. U.S.D. A.

Bull., 885:1-55.

Dowell, R. V. 1979. Host selection by the citrus blackfly, Aleurocanthus woglumi (Homoptera:

Aleyrodidae). Entomol. Exp. Appl., 25:289-296.

-, R. H. Cherry, and G. E. Fitzpatrick. 1979a. Citrus pests in an urban environment.

Fla. Sci., 42:196-200.

-, F. W. Howard, R. H. Cherry, and G. E. Fitzpatrick. 1979b. Field studies of the host

range of the citrus blackfly Aleurocanthus woglumi (Homoptera: Aleyrodidae). Can.

Entomol., 111:1-6.

-, and R. H. Cherry. 1981. Development of a survey trap for predators and parasitoids

of the citrus blackfly, Aleurocanthus woglumi Ashby. Entomol. Exp. Appl., 29:357-362.

-,-, G. E. Fitzpatrick, J. A. Reinert, and J. L. Knapp. 1981. Biology, plant-insect

relations and control of the citrus blackfly, Aleurocanthus woglumi Ashby (Homoptera:

Aleyrodidae). Univ. Fla. Tech. Bull., 818:1-49.

318

PAN-PACIFIC ENTOMOLOGIST

Fitzpatrick, G. E., R. H. Cherry, and R. V. Dowell. 1979. Effects of Florida citrus pest control

practices on the citrus blackfly (Homoptera: Aleyrodidae) and its associated natural

enemies. Can. Entomol., 111:731-734.

Hamon, A. G. 1978. Acacia whitefly, Tetraleurodes acaciae (Quaintance) (Homoptera: Al¬

eyrodidae). Fla. Dept. Agric. Consumer Serv. Entomol. Circ., 190:1-2.

Mound, L. A. 1968. Studies on the olfaction and color sensitivity of Bemisia tabaci (Genn.)

(Homoptera: Aleyrodidae). Entomol. Exp. Appl., 5:99-104.

-, and S. H. Halsey. 1978. Whitefly of the world. J. Wiley & Sons, 340 pp.

Quaintance, A. L. 1900. Contributions towards a monograph of the American Aleyrodidae.

U.S.D.A., Tech. Ser., Div. Entomol., 8:1-64.

-, and A. C. Baker. 1914. Classification of the Aleyrodidae. Part II. U.S.D.A., Tech.

Ser., Bur. Ent., 27(2):95-109.

Vaishampayan, S. M., M. Kogan, G. P. Waldbauer, and J. T. Wooley. 1975. Spectral specific

response in the visual behavior of the greenhouse whitefly, Trialeurodes vaporariorum

(Homoptera: Aleyrodidae). Entomol. Exp. Appl., 18:344-356.

Footnotes

1 Florida Agricultural Experiment Station Journal Series No. 2666. Received for publication

March 10, 1981. Mention of a trademark does not imply endorsement by the University of

Florida.

2 Present address: Department of Food and Agriculture, Analysis and Identification Unit,

1220 N Street, Room 340, Sacramento, California 95814.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 319-325

Published 30 December 1983

NOTES ON THE BIOLOGY OF AEGIALIA HARDYI GORDON AND

CARTWRIGHT (COLEOPTERA: SCARABAEIDAE)

R. W. Rust and L. M. Hanks 1

Department of Biology, University of Nevada-Reno, Reno 89557

Aegialia hardyi Gordon and Cartwright is a large (4.5 mm, 2.5 mg) species

in the genus and is endemic to Sand Mountain and Blow Sand Mountains,

Nevada. This species is flightless, a detritivore and winter active in both the

adult and larval stages. The present study details portions of the life history

and biological characteristics of this sand obligate Aegialia. We also compare

its known life history characteristics to other species of Aegialia.

The genus Aegialia contains 24 species in North America (Stebnicka,

1977). Species of Aegialia (s. str.) are associated with sand habitats (Jerath

and Ritcher, 1959; Gordon and Cartwright, 1977). Jerath and Ritcher (1959)

reported that A. blanchardi Horn was found by sifting sand around and

beneath the roots of vegetation on sand dunes near Waldport, Oregon. They

also found the adults of A. lacustris LeConte, A. latispina LeConte and A.

conferta Horn from sand dunes near Corvallis, Oregon. Jerath (1960a) re¬

ported A. crassa LeConte and A. punctata Brown from Oregon sand dunes

as well. Jerath (1960b) described the larvae of A. blanchardi and A. lacustris.

Recently, Gordon and Cartwright (1977) described some new Aegialia (s.

str.) adding four new species of which three are endemic to sand dunes in

Nevada (A. hardyi endemic to Sand Mountain, A. magnifica Gordon and

Cartwright endemic to Big Dune, and A. crescenta Gordon and Cartwright

endemic to Crescent Dune), and a fourth from California.

Study Areas

Two Nevada sand dunes were sampled from June 1979 through July 1980

for arthropods. Sand Mountain is approximately 46 km ESE of Fallon,

Churchill County, Nevada (39°20'N, 118°20'W) at 1250 m elevation. Blow

Sand Mountains are approximately 52 km SE of Fallon, Nevada (39°10'N,

118°35'W) and are at about 1400 m elevation. The two dunes are separated

by approximately 25 air kilometers. Sand Mountain is a star dune of ap¬

proximately 3.2 km 2 while Blow Sand Mountains are complex star and linear

dunes of approximately 9.2 km. Both dunes result from the same eolian

sand deposited during the Turupah and Fallon formations of about 4000

years B.P. (Morrison and Frye, 1965). Sand Mountain has easy access and

is a popular recreational area while Blow Sand Mountains have extremely

320

PAN-PACIFIC ENTOMOLOGIST

limited access and little recreational use; three-fourths of the dune is located

in a restricted travel area under the jurisdiction of the U.S. Navy.

The floras of the two dunes are similar. The dominant vegetation is Atriplex

confertifolia (Torr. and Frem.), Tetradymia tetrameres (Blake), Chrysotham-

nus viscidiflorus (Hook.), Astragalus lentiginosus Dougl. and Psoralea lan-

ceolata Pursh, and at Sand Mountain only, Eriogonum kearneyi Tidestr.

and Psorothamnus polydenius (Torr. ex. S. Wats.). The dominant grass is

Oryzopsis hymenoides (R. and S.).

Materials and Methods

Field sampling. —Adults and larvae of Aegialia hardyi were collected by

sifting sand. Surface sand to a depth of 0.4-0.5 m, both from beneath dune

vegetation and from non-vegetated areas, was sifted through 12 x 12 mm

and 1.5 x 1.5 mm mesh screens. Approximately 0.005 m 3 of sand were

sifted at a time. The dunes were sampled twice monthly from October to

May for adults and larvae, except in December and January when they were

sampled only once. Blow Sand Mountains were not sampled in November

because of severe weather conditions. Within one sample period four or five

different areas on the dunes were sampled and in any one area sand beneath

all vegetation types present were sampled as well as pure sand (no vegetation).

The presence of beetles in each vegetation type was recorded. Direct obser¬

vation of surface active beetles was also attempted during both days and

nights of a sampling period. Pitfall traps of both 24 hr and 30 day types also

were employed.

Reproductive and food analyses. — During each sampling period at Sand

Mountain 20-30 adult beetles were preserved in ethyl alcohol and acetic

acid (~ 10:1). These individuals were sexed, the females dissected and their

ovarian development and the number of ova present recorded. The digestive

tracts of both males and females were crushed and mounted on microscope

slides and examined for food content.

Laboratory rearing.—Adults and larvae of A. hardyi were returned to the

laboratory for rearing. The individuals used were from Sand Mountain and

were obtained by sifting. Adults were obtained in November, December and

January and the larvae in April. Four or five beetles were placed in 95 cc

plastic cartons 9/10th filled with Sand Mountain sand and detritus. A glass

tube (9 mm dia.) was placed in the center of the sand and functioned as a

watering tube. Cartons were watered at about two week intervals with about

20-25 ml water. This kept the bottom 3-4 cm of sand moist. Cartons were

kept at 24° ± 4°C and under (45 cm) growlux fluorescent tubes on natural

day length.

Climatological data. — Mean monthly temperatures were obtained from

the closest reporting station at Fallon, Churchill County, Nevada, approx¬

imately 50 km distance.

VOLUME 58, NUMBER 4 321

100 _

Fig. 1A-C. Seasonal distribution of Aegialia hardyi Gordon and Cartwright from Nevada

sand dunes. Fig. 1 A. Mean monthly temperature and precipitation from Fallon for the survey

period. Fig. IB. Aegialia hardyi distribution from Blow Sand Mountains (*Blow Sand Moun¬

tains not sampled). Fig. 1C. Aegialia hardyi distribution from Sand Mountain.

Results

Seasonal distribution. —Aegialia hardyi adults were first found in Febru¬

ary, March and April of 1979 during preliminary sampling of the dunes.

Adults were next collected in late October 1979. The abundance of adults

was fairly constant during the next six months (Fig. IB, C) with a slight

322

PAN-PACIFIC ENTOMOLOGIST

Table 1. Reproductive conditions of female Aegialia hardyi Gordon and Cartwright from

Sand Mountain, Churchill County, Nevada, collected from November 1979 to May 1980.

Ovarian development

Eggs present

Number examined

November

December

yes

January

yes

February

yes—no

5 (2-3)

March

yes—no

5 (2-3)

April

yes—no

4 (1-3)

May

increase in January at both dunes. Beetles were found as late as early May

at Sand Mountain. Larvae were found only at Sand Mountain in March,

April and May (Fig. 1C). In March several instars were present, the first

instar larvae are most likely present in February but simply were overlooked

or lost in the sifting process. No pupae were found. Adults and larvae were

found in months having a mean monthly temperature near to or below 10°C

(Fig. 1A).

After the sand was sifted away and the remaining detritus spread out on

a tray for examination, adult beetles present began crawling through the

detritus and so were easily collected. This activity was observed even when

the air temperature was around 5°C. Larvae showed the same activity.

Ova were present from December through April (Table 1). All females

examined from December and January contained ova and there was a steady

decline to a low of 25% in April. No ovarian development was observed in

the females from November and May. These ova were approximately 0.7

mm long by 0.3 mm wide. Twice, 3-0 and 3-2 ova were the observed number.

Additional developing ova were observed in all females with “mature” ova

present. The sex ratio was slanted towards males (1:0.73).

Dune distribution. — Both adults and larvae of A. hardyi were found in

sand removed from the bases of emergent-shrub and grass vegetation on the

dunes. Beetles were taken most often from beneath Tetradymia tetrameres

(56 samples: 41 with and 15 without) followed by Chrysothamnus viscidi-

florus (28/13) and Eriogonum kearneyi (26/15). They were rarely taken from

beneath Atriplex confertifolia (12/25), Oryzopsis hymenoides (12/37) and

Psoralea lanceolata (11/21) and never were found in sand associated with

Psorothamnus polydenius (0/17) or in non-vegetated sand (0/89). Both the

larvae and the adults showed extremely clumped distributions, larvae more

so than adults. For example, several times sand was removed from three

sides of a Tetradymia tetrameres bush with beetles collected only in one of

the three samples, or sparingly in the other two. Three samples from beneath

VOLUME 58, NUMBER 4

323

one Chrysothamnus viscidiflorus bush produced 27, 0, and 0 Aegialia adults;

all three samples were taken within a 4 m 2 area about the bush. From an

area of approximately 0.2 m 2 beneath a Tetradymia bush, 100+ larvae were

taken and sampling from beneath the three sides of the bush produced no

additional larvae. Single adults were rarely found in a sample.

Adult beetles were less commonly found at Blow Sand Mountains with

no adults unearthed in April and May 1980 nor were larvae ever found. We

suspect that this was due to the portion of the dunes available to us for

sampling, the linear dune type or to both conditions.

Adults and larvae were always found in moist sand and neither were seen

on the surface of the sand. Only one adult was obtained by pitfall sampling

of the dunes and it was taken in one 30 day pitfall trap.

Laboratory rearing.— Adult beetles, after a short adjustment period of

about a week during which individuals injured in capture died, lived for an

average of 120 days in the laboratory (longest recorded, November 2 to

March 24; 142 days). All adults were dead in June and no larvae were found

in the cartons with captive adults. Larval beetles lived less than one week

in the laboratory and most died within 24 hr after collecting. No larvae lived

long enough to pupate. Adults were observed moving about on the surface

of the sand between 2000 and 2200 hr on several occasions during January

and February. On 23 January, 7 of 23 cartons with adults had one or more

beetles active at 2100 hr. All 23 cartons were watered on the 24th; no more

beetle activity was observed. This same event occurred on February 6 and

7. Twice during the 6 months of attempted rearing adults were active during

daylight hours (0900 and 1000 hr), on 16 and 19 November.

Examination of cartons indicated that adult beetles were found within a

centimeter or two of the moist sand-dry sand interface. The exception being

the surface active beetles in ‘almost’ dry sand cartons. Dead larvae were

found throughout the moist sand and only rarely in dry sand. This along

with their high mortality rate suggests that soil moisture is much more critical

to the larvae than adults.

Food habits. —Aegialia hardyi adults and larvae feed on detritus trapped

in the sand that accumulates around emergent vegetation. Gut analyses

showed plant fibers from stems, sclereids from stem pith, spongy parenchyma

from leaf tissue and tracheids from stems. The cells and tissue clumps mea¬

sured up to a maximum of 25 g. The fungi Cladosporium, Stemphylium,

Alternaria and Torula were common in gut contents. These fungi are as¬

sociated with decay of plant materials. Whether or not these fungi serve as

food for the beetles is open for speculation. It is possible that the plant

material is the carrier of the actual food, the fungal mycelium; a situation

similar to that found in freshwater detritovores (Cummins, 1974). Clado¬

sporium and Alternaria were very abundant outnumbering the plant tissue

fragments in the guts of several adult beetles. Some of the beetles carried

pollen grains in their gut contents. One was identified as Pinus monophylla

324

PAN-PACIFIC ENTOMOLOGIST

Torr. and Frem., Pinyon Pine; the closest Pinyon in a windward direction

is over 50 km away. One gut contained insect integument, setae easily visible;

perhaps pupal or larval exoskeleton. There was no apparent difference in

the food selected by adults and larvae. Adults from November had empty

digestive tracts while all others were full.

Discussion

Aegialia hardyi is similar to some species of Aegialia (s. str.) in its asso¬

ciation with sand habitats. However, these sand habitats may be wet sand,

as along streams. It is different in having a winter activity period for both

adults and larvae. Aegialia blanchardi adults were found year round and the

larvae in the early summer (May to July) (Jerath, 1960b). Aegialia lacustris,

A. latispina and A. conferta were found during the summer months and their

larvae were found in the early summer (Jerath and Ritcher, 1959). These

researchers found adults, pupae and larvae just below the surface (“6 to 8

inches”) by sifting sand. They also reported the larvae to feed only on

decaying organic matter in the sand.

The winter activity period of A. hardyi may result from the limited pre¬

cipitation and hot summer temperatures found in these Great Basin sand

dunes. Although there are summer rain storms, the seasonal distribution of

the 10 to 15 cm of precipitation in the Fallon area is principally winter

(Houghton et al., 1975). Also, there is over an eighty percent chance of no

measurable precipitation during early August in the Sand Mountain and

Blow Sand Mountains areas. Of the 15.8 cm of precipitation falling during

the survey period, 76.6% fell between October 1979 and May 1980 (Fig.

1 A). The 30 mm of rain in August represents an unusual climatological event

not typical over extended years. The addition of new litter to the sand also

would be at a maximum rate during the fall months at the onset of the adult

activity period. Competition with other summer active detritovores also

may have narrowed the activity period or at least shortened it. Finally, all

three factors may be important in the sequence of steps resulting in the

winter activity period.

Acknowledgments

We would like to thank Bob Bechtel for his assistance with the project

and his review of the manuscript. Dr. Don Prusso identified the fungi and

plant parts. Also, the review comments of Drs. Fred Andrews, Bob Gordon

and Alan Hardy were most helpful. Bob Gordon identified the Aegialia

specimens. We would like to thank Dave Goicoechea, BLM State of Nevada,

for his support in making the study possible.

Literature Cited

Brown, W. J. 1931. Revision of the North American Aegialiinae (Coleoptera). Can. Entomol.,

63:9-19, 42-49.

VOLUME 58, NUMBER 4

325

Cummins, K. W. 1974. Structure and function of stream ecosystems. BioScience, 24:631-

641,

Grodon, R. D., and O. L. Cartwright. 1977. Four new species of Aegialia (s. str.) (Coleoptera:

Scarabaeidae) from California and Nevada Sand Dunes. J. Wash. Acad. Sci., 67:42-48.

Houghton, J. G., C. M. Sakamoto, and R. O. Gilford. 1975. Nevada’s weather and climate.

Nev. Bur. Mines Geol. Spec. Publ., 2:1-78.

Jerath, M. L. 1960a. Distribution of Aphodiinae in Oregon. Pan-Pac. Entomol., 36:183-188.

-. 1960b. Notes on larvae on nine genera of Aphodiinae in the United States (Coleoptera:

Scarabaeidae). Proc. U.S. Nat. Mus., 111(3425):43-94.

-, and P. O. Ritcher. 1959. Biology of Aphodiinae with special reference to Oregon.

Pan-Pac. Entomol., 35:169-175.

Morrison, R. B., and J. C. Frye. 1965. Correlation of the middle and late quartemary succes¬

sions of the Lake Lahonton, Lake Bonneville, Rocky Mountains (Wasatch Range), south¬

ern Great Plains, and eastern midwest areas. Nev. Bur. Mines Rep., 9:1-45.

Stebnicka, Z. 1977. A revision of the world species of the tribe Aegialiini. Acta Zool. Cracov.,

22:397-506.

Footnote

1 Present address: Department of Entomology, University of Maryland, College Park, Mary¬

land.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 326-329

Published 30 December 1983

NOTES ON NEOTROPICAL NAUCORIDAE II. A NEW SPECIES OF

AMBRYSUS AND REVIEW OF THE GENUS

POTAMOCORIS (HEMIPTERA ) 1

John T. Polhemus

3115 S. York Street, Englewood, Colorado 80110

AND

Dan A. Polhemus

Department of Biology, University of Utah, Salt Lake City 84112

This is the second in a series intended to revise the naucorid fauna of the

Neotropical region.

Specimens of all species treated here are held in the Polhemus Collection

(JTP); the disposition of other specimens is given in the text. Unless oth¬

erwise noted, for all measurements 40 units = 1 mm.

Ambrysus plautus Polhemus and Polhemus, new species

General appearance.—Of moderate size, ovate, shape as in Figure 1A.

Ground color yellowish-brown; head and pronotum shallowly rugose, mot¬

tled with areas of chocolate brown; scutellum rugulose; hemelytra light um¬

ber, minutely punctate, embolium yellowish brown. Venter light brown,

abdomen clothed with fine golden hairs; legs dark yellow, with light brown

spines.

Structure. — Head with eyes raised slightly above surface dorsally; vertex

moderately produced behind eyes; labrum shallowly triangular in shape,

apex rounded, width/length = 23/12; eyes convergent anteriorly, posterior/

anterior interocular space = 60/48. Pronotum with lateral margins broadly

convex, smooth; posterolateral angles rounded; width/length = 212/65. Scu¬

tellum wider than long, weakly sinuate along lateral margins, width/length =

130/63. Hemelytra fully developed; embolium expanded medially, broadly

rounded, not sinuate posteriorly; entire surface set with minute round pits

appearing white against darker background. Connexival margins weakly ser¬

rate; posterolateral angles strongly spinose. Fore leg femur length/width =

88/50; tarsus slightly exceeding adjacent proximal part of femur. Middle

and posterior femora with row of tiny, barely visible spines posterodorsally

and posteroventrally; middle and posterior tibiae set with numerous stout

spines; distally with two transverse rows of small spines. Propleura not

contiguous with prosternum; propleura barely contiguous medially, not con¬

tiguous there with mesosternum.

VOLUME 58, NUMBER 4

327

Fig. 1. Ambrysus plautus. A, Female, dorsal view. B, Female subgenital plate, ventral view.

C, Male genital segments, dorsal view, showing male genital process at left of center (in figure,

right side of insect).

Male genital process as in Figure 1C; female subgenital plate shape as

shown in Figure IB.

Total length of males, in mm; N = 7, x = 8.44, s = .39; total width of

males, in mm; N = 7, x = 6.23, s = .25. Total length of females, in mm;

N = 5, x = 9.36, s = .24; total width of females, in mm; N = 5, x = 6.9, s =

.14.

Type material.— Holotype, male, and allotype, female, MEXICO, Duran¬

go, 20 mi. E of La Ciudad, 8700 feet (2652 m), CL 726, VI-6-1975, J. T.

and D. A. Polhemus (JTP). Paratypes (all collected by J. T. and D. A.

Polhemus in J. T. Polhemus collection): 6 males, 4 females, 2 nymphs, same

data as holotype; 24 males, 24 females, 16 nymphs, same locality as holotype,

IV-22-1981; 1 male, 6 females, MEXICO, Chihuahua, Rio Concho, 9 mi.

N of Creel, IV-28-1982; 11 males, 14 females, Chihuahua, 7 mi. NE of

Rancho Basaseachic, IV-27-1982; 2 females, Chihuahua, 26 mi. NE of Ran¬

cho Basaseachic, IV-27-1982.

Etymology .—The name plautus (L.; broad; masculine) refers to the shape

of this insect.

Discussion.—Ambrysus plautus, n. sp. resembles Ambrysus drakei La-

328

PAN-PACIFIC ENTOMOLOGIST

Rivers in general facies, but is much smaller, with the embolium more

expanded medially and not sinuate posteriorly, the pronotum more rugose,

the body shape more rotund, and the male and female pregenital structures

differently shaped. The male process in both species is long and slender; in

A. plautus it is sharply bent basally (Fig. 1C) but in A. drakei it is upright

and gently curved basally, bent at or beyond the middle (see figure in LaRivers,

1957). The female subgenital plate of A. plautus is trispinose distally (Fig.

IB) whereas in A. drakei the subgenital plate is rounded posterolaterally

with the lateral spinose processes in a more medial position, and the median

process is blunt and sinuate (see LaRivers, 1957). The expanded, broadly

rounded embolium and the strongly spinose posterolateral connexival angles

are distinctive attributes of this species.

A Review of the Genus Potamocoris Hungerford

We have examined the males and females of all three known species of

Potamocoris and find that a revision of the LaRivers (1969) key is necessary.

P. robustus LaRivers has a small incised notch apically on the female subgen¬

ital plate, difficult to see because of the hairy vestiture; LaRivers’ key uses

the supposed absence of such a notch as a key character for robustus. The

heavy patch of yellow hairs on the posterior edge of the mesofemur (and

mesotibia) is not a key character but is sexually dimorphic and present in

males of all three species. The median rugulose V-shaped depression on

abdominal sternum V is also sexually dimorphic, occurring only in females,

and not found in other naucorid genera. The genus furthermore is unique

among naucorids in that the species commonly fly; all records for beckeri

LaRivers, and by far the longest series of robustus are from lights.

Nothing is known about the ecology or habitat of this interesting genus.

Hungerford’s (1941) original description of Potamocoris and only known

species at that time, parvus Hungerford, was based on material collected by

Alberto Schulze on the East Paraguay River at or near Horqueta, Paraguay

and sent to Hungerford by John Lutz of Philadelphia. If Lutz had any notes

on the collection method or habitat they were not furnished.

Twelve specimens of P. beckeri were taken by Clark, Murray, Hart, and

Schaffner at light, on July 5, 1972, 36 miles E of Acuyacan, Veracruz, Mexico.

This extends the known range of this species far north of the type locality

in Honduras, where the single female holotype was taken in a light trap

(LaRivers, 1950).

Many specimens of P. robustus were taken by R. T. and J. C. Schuh at

light, December 17, 1971, km 3, Tournavista Road, 34 km W of Pucallpa,

300 m, Loreto, Peru; the type locality is also in Peru. This splendid series

should permit morphological studies that will hopefully elucidate the rela¬

tionship of this annectant genus to the remainder of the Nepomorpha.

For their kindness in allowing the authors to study specimens, we are

VOLUME 58, NUMBER 4

329

indebted to P. D. Ashlock, University of Kansas; J. C. Schaffner, Texas

A&M University; R. T. Schuh, American Museum of Natural History.

Key to Species of Potamocoris Hungerford

1. Inner margins of eyes parallel; V-shaped depression on female fifth

sternum occupying almost entire length of segment (Mesoamerica)

. beckeri LaRivers

Inner margins of eyes distinctly convergent anteriorly; V-shaped

depression in female fifth sternum occupying half or a little more

of length of segment . 2

2. Interocular space more than 1.5 x eye width; female subgenital plate

sinuate posteriorly, with a small sharply incised notch apically

(Peru) . robustus LaRivers

Interocular space less than 1.3 x eye width; female subgenital plate

smoothly rounded posteriorly, not sinuate, not notched apically

(Paraguay) . parvus Hungerford

Literature Cited

Hungerford, H. B. 1941. A remarkable new naucorid waterbug (Hemiptera). Ann. Entomol.

Soc. Am., 34:1-4.

LaRivers, I. 1950. A new species of the genus Potamocoris from Honduras (Hemiptera;

Naucoridae). Proc. Entomol. Soc. Wash., 52:301-304.

-. 1957. A new Ambrysus from Mexico (Hemiptera; Naucoridae). Entomol. News 68:

232-237.

-. 1969. New naucorid taxa (Hemiptera). Biol. Soc. Nev., Occas. Pap., 20:

1 - 12 .

Footnote

1 Contribution from the University of Colorado Museum, Boulder, Colorado 80309.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 330-335

Published 30 December 1983

THE GENUS ANOPLOSIAGUM BLANCHARD

(COLEOPTERA: SCARABAEIDAE) IN CUBA.

PART I. DESCRIPTIONS OF FIVE NEW SPECIES

Miguel Garcia-Vidal

351 Grove Street, #1, San Francisco, California 94102

The subfamily Melolonthinae has only two genera reported from Cuba:

Anoplosiagum and Phyllophaga. I have been working on the latter for several

years (Garcia-Vidal, 1975, 1978). At the request of some Cuban parasitol¬

ogists I began to work on Anoplosiagum, since this genus was found as a

host for some parasites.

The first writer to describe a Cuban species of Anoplosiagum was E.

Blanchard, who published his catalogue of the entomological collections of

the Paris Museum in 1850 (Chapin, 1932). Blanchard established this genus

for four species, three from Brazil and the other (A. pallidum) from Cuba.

In 1865, Chevrolat described another (A. variabile). E. Chapin described six

more (A. flavicolle, A. peltatum, A. rufum, A. scabrosum, A. scaramuzzai,

and A simplicipes) and redescribed another species (A. rutilus ) that originally

was placed in the genus Clavipalpus (Chapin, 1932).

During my research I found specimens that did not fit the descriptions of

the previously described species of Anoplosiagum. The principal differences

were in the claws, the clypeus, and the male genitalia. Figures of these

distinguishing characters are presented in this paper.

Until now, only nine species have been reported from Cuba (Blackwelder,

1944). Adding the present five the total number of species of Anoplosiagum

from Cuba grows to fourteen.

The figures of the male genitalia were drawn by the author. I wish to

sincerely thank Heriberto Maza who made the figures of the claws and heads

and Ing. Fernando de Zayas who lent me the specimens from his collection

for my studies. Acronyms used in this paper: FZC—Fernando de Zayas

Collection, Havana; CAS—California Academy of Sciences, San Francisco.

Anoplosiagum cubensis Garcia-Vidal, new species

(Figs. 5, 7)

Shining. Elytra testaceous; underparts, legs testaceous to yellowish-brown;

head castaneous; disc of pronotum with piceous spot which extends to head

(as A. turquinensis)-, posterior, lateral margins yellowish-brown. Clypeus

densely punctured; anterior margin slightly, narrowly reflexed; clypeo-frontal

suture obliterate. Frons flat except medially, base punctate; punctures as on

Figs. 1-5. Heads (A) and claws (B) of Anoplosiagum spp. Fig. 1. A. zayasi. Fig. 2. A.

turquinensis. Fig. 3. A. swezeyi. Fig. 4. A. oteroi. Fig. 5. A. cubensis.

clypeus. Head pubescent with thick hairs. Pronotum rather sparsely punc¬

tured; punctures smaller than those of head, with thick, long, straight hair

mainly on lateral margins; anterior, basal margins entire; sides broadly di¬

lated medially; lateral margins finely denticulate; anterior angles slightly

acute, basal obtuse. Scutellum triangular, punctures as on elytra. Elytra more

densely punctured than pronotum but less than head; punctures smaller than

those of pronotum; sutural margins tumid but only slightly elevated, with

some punctures. Pygidium punctured as pronotum, with long, straight hairs;

apex broadly rounded. Antennal club 3-segmented, about as long as stem.

332

PAN-PACIFIC ENTOMOLOGIST

Upper tooth of protibia obsolete. Coxal plates with thick, straight hairs.

Sternites densely, regularly punctured; punctures shallow, with long, straight,

fine hairs. Urosternites less densely punctured than sternites; sparsely pu¬

bescent. Longer calcar of metatibia slender, acuminate, about as long as first

tarsal segment. Claws slightly curved, with split near base. Length, 8-9 mm.

Holotype.— Male, Matanzas, Central CUBA, June 12, 1940 (Scaramuzza)

(FZC).

Paratypes.— Five with same data as holotype (3, FZC; 2, CAS).

This species is similar in color to A. peltatum, but the spot on the pronotum

is more irregular, with its sides not well defined. In addition the claws of A.

peltatum are stout, bent, and split near middle and the upper tooth of the

protibia is present. It is a smaller species.

Anoplosiagum oteroi Garcia-Vidal, new species

(Figs. 4, 8)

Shining. Disc of elytra, sides, basal margins of pronotum, anterior margin

of clypeus yellowish-brown; sides of elytra, disc of pronotum, frons, posterior

half of clypeus, underparts piceous; legs variegated. Frons densely, coarsely

punctured, flat except at base, where it is impunctate; clypeo-frontal suture

obliterated. Clypeus less densely, more regularly punctured than frons; mar¬

gins narrowly reflexed. Pronotum less densely, more regularly punctured

than head; anterior, basal margins entire; lateral margins finely denticulate;

sides broadly dilated medially; anterior angles slightly acute; posterior ob¬

tuse; with spot, as in A. peltatum which continues to frons. Scutellum cas-

taneous, sparsely punctured. Elytra slightly less punctured than pronotum;

sutural margins slightly tumid; lateral, posterior margins bearing long, straight

hairs. Pygidium castaneous, coarsely punctured, apex broadly rounded. An¬

tennal club 3-segmented, about as long as stem. Protibia tridentate; upper

tooth very weakly developed, distal very long. Coxal plates bearing thick,

long, straight hairs. Sternites densely punctured; punctures fine, from which

arise fine, long, straight hairs. Urosternites with sides yellowish-brown, less

densely punctured than sternites. Longer calcar of metatibia acuminate,

longer than first tarsal segment. Claws little curved, without split. Length,

6-8 mm.

Holotype. — Male, Estacion Experimental Agronomica, Finca Coca, Prov.

Habana, CUBA, May 1961 (I. Garcia) (FZC).

Paratypes.—Two, Santiago de las Vegas, CUBA, May 20, 1933 (A. Otero),

with a label from Estacion Experimental Agronomica, Cuba, Ento. No.

10111 (FZC); one, Santiago de las Vegas, CUBA, May 20, 1933 (A. Otero),

with a label “Abundant in pasture fields,” Estacion Experimental Agrono¬

mica, Cuba, Ento. No. 10111 (FCZ). Four with same data as holotype (2,

FZC; 2, CAS).

This species is dedicated to A. Otero, who worked in the Entomology

VOLUME 58, NUMBER 4

333

Figs. 6-10. Male genitalia of Anoplosiagum spp., frontal view (A) and lateral view (B). Fig.

6. A. zayasi. Fig. 7. A. cubensis. Fig. 8. A. oteroi. Fig. 9. A. swezeyi. Fig. 10. A. turquinensis.

Department of the formerly Estacion Experimental Agronomica, Santiago

de las Vegas, Habana, for many years.

Anoplosiagum swezeyi Garcia-Vidal, new species

(Figs. 3, 9)

Elytra, head castaneous; pronotum castaneous with yellowish-brown band

on both sides; underparts, legs yellowish-brown. Head densely, coarsely

punctured with irregular punctures, absent at base, with thick, short, straight

hairs; clypeo-frontal suture obliterate; margins of clypeus abruptly reflexed.

Pronotum sparsely punctured; punctures shallow; anterior, basal margins

entire; sides dilated a little before subangulated middle; lateral margins finely

334

PAN-PACIFIC ENTOMOLOGIST

denticulate, with thick, long, reflexed hairs. Scutellum with punctures present

laterally, absent on disc. Elytra more densely punctured than pronotum, but

less than head; punctures deeper, bigger than those of pronotum; sutural

margins slightly tumid; lateral, posterior margins with short, thick, bent

hairs. Pygidium pubescent, with thick, long, straight hairs; apex broadly

rounded. Antennal club 3-segmented, about as long as four preceding seg¬

ments combined. Upper tooth of protibia almost obsolete; median rather

acute, distal long, acuminate. Coxal plates with long, thick hairs. Sternites

densely punctured with punctures fine, shallow, regularly distributed, from

which arise fine, short hairs. Urosternites punctured about as sternites; fifth

segment laterally bearing a tuft of long, fine hairs. Longer calcar of metatibia

acuminate, about as long as first tarsal segment with its apical spines. Claws

little curved; without split. Length, 6-9 mm.

Holotype.— Male, Las Martinas, Pinar del Rio, CUBA, June 24, 1940 (J.

Acuna) (FZC).

Paratypes.— One, Las Martinas, Pinar del Rio, CUBA, June 24, 1940 (J.

Acuna) (CAS); one, San Julian, Pinar del Rio, CUBA, January 8, 1938 (J.

Acuna), with a label from Estacion Experimental Agronomica, Cuba, Ento.

No. 10972 (FZC).

This species is dedicated to Dr. Sean Swezey, University of California at

Berkeley.

Anoplosiagum turquinensis Garcia-Vidal, new species

(Figs. 2, 10)

Very narrowly oval, long, shining; yellowish-brown to castaneous; with

two rather long piceous spots on pronotum; sutural margins from yellowish-

brown to piceous. Frons densely, coarsely punctured; punctures irregular;

with a depression on each side of the median line; impunctate toward base

and sides, which are darker than disc; clypeo-frontal suture obliterated.

Clypeus colored as disc of frons, similarly punctured, but punctures more

regular; margin abruptly reflexed. Pronotum less densely punctured than

head; punctures smaller but more regularly distributed; spots on both sides

of median line of disc; margins entire; sides dilated, rounded at middle; with

long, thick hairs on lateral margins. Scutellum almost triangular, with punc¬

tures toward margins; same color as sutural margins. Elytra more densely

punctured than pronotum but less than head; punctures rather bigger; sutural

margins tumid; lateral, posterior margins bearing long, thick, straight hairs.

Pygidium sparsely punctured; punctures fine; some fine hairs, mainly on

margins; apex broadly rounded. Protibia tridentate; upper tooth tiny; teeth

almost equidistant. Antennal club 3-segmented, about as long as stem. Coxal

plates with thick, long, straight hairs arising from fine punctures. Urosternites

less densely punctured than sternites, but with similar punctures; some fine

hairs. Longer calcar of metatibia acuminate, longer than first tarsal segment.

Claws little curved, split medially. Length, 7.5-9 mm.

VOLUME 58, NUMBER 4

335

Holotype. — Male, Pico Turquino, Sierra Maestra, Oriente, CUBA, June

1963 (F. de Zayas) (FZC).

Paratypes .—Three with same data as holotype (FZC). Two, Pico Turquino,

CUBA, 3750 feet altitude, June 10-29, 1930 (J. Acuna) (FZC); one, Palma

Mocha to Pico Joaquin, Sierra Maestra, CUBA, May 18, 1948, 3900-6300

feet altitude (collector unknown); one, Pico Turquino, CUBA, July 29, 1922,

4500-5000 feet altitude (S. C. Bruner and C. H. Ballou) (CAS).

Anoplosiagum zayasi Garcia-Vidal, new species

(Figs. 1, 6)

Shining. Head piceous; pronotum yellowish-brown; sides of elytra cas-

taneous; disc, underpart, legs yellowish-brown. Head densely, coarsely punc¬

tured; punctures irregular, absent at base; clypeo-frontal suture obliterated.

Pronotum less densely but more regularly punctured than head; punctures

shallow, more regular in shape; anterior, basal margins entire; sides medially,

broadly dilated; lateral margins finely denticulate; anterior angles almost

right, basal obtuse. Scutellum colored as pronotum, with punctures toward

margins. Elytra more densely punctured than pronotum but less than head;

punctures bigger, deeper; sutural margins conspicuous but not elevated; lat¬

eral, posterior margins bearing thick, straight hairs. Pygidium densely, coarsely

punctured; apex broadly rounded. Antennal club 4-segmented about as long

as rest of segments. Upper tooth of protibia obsolete; distal very long. Coxal

plates with thick, long hairs. Sternites pubescent, with fine, long, straight

hairs, arising from fine punctures. Urosternites less pubescent than sternites.

Longer calcar of metatibia slender, acuminate, longer than first segment.

Claws gently curved, without split. Length, 6-7 mm.

Holotype.— Male, La Glorieta, Guantanamo, Prov. Oriente, CUBA, June

1959 (F. de Zayas) (FZC).

Paratypes .—Four with same data as holotype (2, FZC; 2, CAS).

This species is characterized by its 4-segmented antennal club and the

absence of the upper tooth of the protibia. It is dedicated to my friend and

professor Ing. Fernando de Zayas.

Literature Cited

Blackwelder, R. E. 1944. Checklist of coleopterous insects of Mexico, Central America, the

West Indies, and South America. Bull. U.S. Nat. Mus., 185:189-341.

Chapin, E. A. 1932. Revision of the pleurostic Scarabaeidae of Cuba and the Isle of Pines. I.

The Melolonthinae. Ann. Entomol. Soc. Am., 25:173-209.

-. 1935. New Cuban pleurostict Scarabaeidae. Mem. Soc. Cubana Hist. Nat., 9:67-75.

Garcia-Vidal, M. 1975. Compilation sobre los principales aspectos relativos al genero Phyl-

lophaga Harris. Rev. Agrotecnia Cuba, 7:17-22.

-. 1978. El genero Phyllophaga Harris, 1826 (Coleoptera: Scarabaeidae) en Cuba. I.

Description de cinco nuevas especies. Poeyana, 182:1-14.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 336-351

Published 30 December 1983

NESTING BIOLOGY AND FLOWER RELATIONSHIPS OF

XYLOCOPA SONORINA SMITH IN HAWAII

(HYMENOPTERA: ANTHOPHORIDAE ) 1

Dan Gerling

The George S. Wise Faculty of Life Sciences,

Tel Aviv University, Israel

Carpenter bees have been the object of numerous studies during recent

years (Anzenberger, 1977; Barrows, 1980; Bonelli, 1976; Gerling and Her¬

mann, 1978; and Hurd, 1978). As a result of these and other works, much

information was added to our knowledge of the biology of these bees and

new insight has been gained of their biology, especially with respect to the

social relationships.

During my sabbatical leave, I spent 9 months at the Department of Ento¬

mology of the University of Hawaii at Manoa and had a chance to observe

the Hawaiian carpenter bee, Xylocopa sonorina Smith, to study some of its

biological attributes, and to compare them with some of the results men¬

tioned in other recent works. The results of these studies are reported herein.

Materials and Methods

Observations were conducted of both unmarked and marked bees. Mark¬

ing was carried out by painting the thorax and/or abdomen of the bees with

typewriter correcting fluid. For X-ray observations, the bees were marked

with a piece of wire glued to their notum. Nests were obtained from different

locations on Oahu and placed on the roof of the Entomology building of the

University of Hawaii at Manoa, where they were kept in the open, under a

rain-shelter. These nests were taken about twice a week to an X-ray machine

and radiographed. Additional observations were conducted in natural nest¬

ing sites the most noteworthy of which was the “Fashion Fabrics” store of

Kapiolani and Piikoi Ave. in Honolulu, where several hundred 2.5 x 5 cm

thick redwood poles of various lengths had been used for decorative pur¬

poses. These served as nesting substrates for numerous carpenter bees (Fig.

1). The experiment station of the University of Hawaii at Waimanalo, where

Dr. T. Nishida has conducted his studies in the past (Nishida, 1963), also

served as a place of observations. X-ray radiograms were taken with a “Hew¬

lett Packard” “Faxitron” at about 3 mAmp, 25-70 KVp, for 5-15 seconds

and a distance of 30 cm; Kodak AA+ or Agfa Industrex films were used.

VOLUME 58, NUMBER 4

337

Results

Nesting substrates and nest structure. — Hurd (1978) listed numerous nest¬

ing substrates used by this bee. In this study, I found nests in structural

timber, mainly introduced redwood timbers ( Sequoia sempervirens Endl.)

that are abundantly used for construction, panelling, decoration and fence

building. Nests were found in a variety of shapes and sizes of this wood,

provided its thickness exceeded about 2 cm. Dead parts of plants also served

as nesting sites. The ones observed in this study, branches of umbrella trees

(.Bassaia actinophylla ), Hibiscus spp. and baobab (. Adansonia digitata L.).

There were also reports of nests in dead Agave flowering stalks, dead branches

of Plumeria spp. and dead trunks of palm trees.

Nest digging was initiated by the bees even in unsuitable substrates, such

as 0.75 cm thick wooden stakes. A number of such cases were recorded

during February 1980 on the island of Molokai were the bees dug cavities,

and even holes through pine and redwood marking-stakes in pineapple fields

(Fig. 2). At times, when the bees encountered wooden stakes that had been

piled up, they dug tunnels through a number of them, acting as if they were

a solid wooden mass. A similar activity was seen in an old pile of redwood

boards in the Waimanolo experiment station on Oahu, where the bees not

only dug through 2, overlaying, 1 x h thick boards but even nested in this

compound.

It has been observed in other carpenter bee species that, although they

often dig extensive tunnels, they rarely penetrate the nest walls to the outside.

X. sonorina was the first carpenter bee species in which I observed several

cases in which the nest wall was broken often. This usually took the shape

of small holes or longitudinal clefts in the wood. Additionally, I found

secondary entrance holes to nests, that were obtained through the digging

action of the bees from the inside of the nest (Fig. 3). These were especially

common when nests have been dug in extensive wooden structures such as

the roof of the Molokai airport building, in which probably many generations

of bees have already developed.

Although the X. sonorina female starts her nest with 1 tunnel, the nests

usually become more complex and often have 4-5 tunnels each (Table 1

and Figs. 3-6). The number of bees developing simultaneously varied from

1-8, the high numbers including progeny of different ages, usually very young

bees as well as pupae. All of the eggs were laid by the same female, however

a number of bees often resided in the same nest. Tunnel length varied from

4 to 21.5 cm in the 4 nests studied, but tunnels in other nests were found

to reach over 30 cm.

Developmental history and daily activity. — Ovipositions took place the

year round. During the winter, from December to March, their frequency

seemed to recede with the decline in the bees’ activity, and the duration of

development took some 10-15 days longer than in the fall. However, pollen

v*-

338

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-7. Fig. 1. Decorative redwood sticks at the Fashion Fabrics store in Honolulu,

perforated by nest holes of X. sonorina. Fig. 2. Holes made by females of X. sonorina in marking

stakes made out of wood lathe used in the pineapple fields of Molokai. Fig. 3. X-ray radiogram

VOLUME 58, NUMBER 4

339

Table 1. Nesting data for 4 nests of X. sonorina.

Nest

no.

Date

No. of

tunnels/nest

With

Sum progeny

No. of

progeny

Nest tunnel lengths

(cm)

Per

nest

Per

tunnel

(max)

No. of

adult

bees

27.X

10, 10, 13, 20

26.XI

19.11

10.VI

9.2

15.V

25.IX

5, 9, 11, 11, 21.5

16.X

30.X

20.IX

4, 7, 10, 14

26.X

18, 19, 20

was collected and eggs were laid. Periodic visits to Waimanalo, on the wind¬

ward side of the Island indicated that nesting activity was discontinued there,

probably due to the much higher rainfall.

Duration of development out of doors from egg to adult was about 41-

50 days. The breakdown was as follows: Eggs 2-3; Larva I 3-4; Larva II 3-

5; Larva III 3-5; Non feeding larva (prepupa) 9-14; Pupa 21-27 days. The

time from beginning of pollen slant deposition to egg laying varied since,

like X. pubescens (Gerling, Hurd, and Hefetz, 1981) X. sonorina often re¬

moves the pollen slant after its formation. At times when it was not removed,

pollen slant build-up that preceded oviposition lasted about 4 days.

Tunnel digging is, apparently, done in increments. A bee was observed

digging in a redwood board for 10 consecutive hours, reaching 2-3 cm in

depth, and then stopping. We have no record on the continuation of this

excavation, but the X-ray records of nest No. 1, to which a 5th tunnel was

added (Table 1) showed that in that case, digging took 14 days from start

to the final length of 9 cm, and was done in 3 or more increments, with

several days’ cessation between each.

of nest No. 1, A—on Jan. 4, and B—on Apr. 22, 1980. V, original; X, secondary entrance holes.

Figs. 4-6. X-ray radiograms of nests No. 2-4 respectively. Fig. 7. Wings of A. young, B. active,

and C. old X. sonorina females.

340

PAN-PACIFIC ENTOMOLOGIST

Table 2. Incidence of regurgitation May 15, 1980.

Hour

Temp, in

shade

°C

No. of guard

bees at nest

entrances

No. of bees

seen

regurgitating

% of bees

regurgitating

715*

840

24.2

1020

35.3

1210

61.3

1525

32.5

55.9

* 1 st bee seen regurgitating.

The emerging females are black but their wings are milky-white. The wings

darken within 24-28 hours, the bees attain the ability to buzz during the

first week and to fly under laboratory conditions after two or more weeks.

Bees that developed in the field appeared to fly earlier.

The emerging adults clean their nest from fecal and nest-shaving debris,

but usually do not disturb younger progeny that might be in the same tunnels.

They are fed by their mother during the teneral stage, and in the lab. they

readily fed on honey from about the 4th day after emergence.

Different phases in the life cycle of carpenter bees from emergence to death

can be followed by observing the condition of various body parts and organs

that change with time (Daly, 1966; Gerling and Hermann, 1978). Accord¬

ingly, it is expected that the wings of bees will be more worn and their ends

frayed with increase of field activity (Fig. 7), the yellow gland will be de¬

veloped in actively nesting bees (Gerling, Orion, and Ovadia, 1979) and the

oocytes in the ovarioles will be larger as oviposition time draws near.

Table 3 depicts the conditions of 6 parameters that were examined in 28

bees. Two were overwintering bees, that emerged from their pupae in nest

No. 2 between Nov. 15 and 30 and were removed from that nest and

dissected on Feb. 25 and 26. Nine of the rest were collected while visiting

flowers, in particular Crotalaria mucronata Desr. at Waimanalo, and the

remaining 17 are from “Fashion Fabrics” and were caught with a net when

leaving or approaching their nests. Aside from the two bees that were re¬

moved from the nests, all were collected from the middle of March on, i.e.,

during nesting time.

Two of the 10 unmated bees, one from flowers at Waimanalo and the

other from the nests at “Fashion Fabrics,” had very worn wings that indi¬

cated extensive flight activity. The two bees that were removed from their

nests had completely new wings, were unmated, but had a crop that was full

of pollen and the ovaries were well developed. Ovarial development was

also relatively well advanced (at least one oocyte reaching x h or more of

VOLUME 58, NUMBER 4

341

Table 3. Results of dissections of X. sonorina Feb.-Apr. 1980.

Date

Place

Wing

wear

Pollen 1

Ovary

S 3

Glands 2

Weight

(g)

Stom¬

ach

Rec¬

tum

25.11

—

>/2- 2 /3

—

26.11

—

>/2- 2 /3

—

13.Ill

F.F.

+ + +

—

.668

21.Ill

F.F.

—

>/3

—

21.Ill

F.F.

—

+ +

21.Ill

F.F.

—

+ + + +

23.Ill

F.F.

+ + + +

‘/3

—

24.Ill

F.F.

—

y 2

—

24.Ill

F.F.

+ + +

28.Ill

F.F.

—

y 2

—

30.Ill

F.F.

+ + +

—

y 2 - 2 /3

—

.554

30.Ill

F.F.

—

y4-y 3

—

.638

30.Ill

F.F.

+ + + +

—

y 2

.690

30.Ill

F.F.

+ + + +

.597

30.Ill

F.F.

+ + +

—

>/2

30.Ill

F.F.

+ +

y»

—

.573

30.Ill

F.F.

>/2

.677

30.Ill

F.F.

+ + + +

—

y 2 -y3

—

.535

30.Ill

F.F.

—

>/3

.677

8.IV

+ + + +

y»

—

14.IV

+ + +

y 2

+ +

14.IV

+ +

—

‘/3

—

14.IV

—

‘/4

—

14.IV

+ + + +

—

.715

14.IV

+ + +

y 3

14.IV

+ + +

—

y 3

14.IV

+ + +

+ +

15.IV

+ + +

—

y 2

+ +

M—Manoa; F.F.—Fashion Fabrics; W—Waimanalo; Ovary: expressed as a fraction of max¬

imal size attainable by the oocyte; S—Spermatheca; D—Dufour; Y—Yellow gland; ND—No

data. 1 + = pollen present, 2 + = full, 3 + = well developed.

maximal size) in four more unmated bees indicating that mating was not a

prerequisite for such an occurrence. The pollen in the crop of the two young

bees might have come from one or more of a number of sources, like bee

bread or its remnants in the nest, new pollen from a pollen slant prepared

by the older bee residing in that nest, or trophallaxis with that female. Most

bees had many pollen remnants in their rectum indicating that they not only

carried this material in their crop for bee bread preparation (Schremmer,

1972) but also fed on it.

Yellow glands were well developed only in bees that showed ovarial de-

342

PAN-PACIFIC ENTOMOLOGIST

velopment, in one case even in an unmated bee. There were, however, several

cases of both mated and unmated bees in which these glands were white

and undeveloped.

The laboratory material was reared in glass tubes lined with absorbent

paper. Following adult emergence these were placed on one tray allowing

bees to enter and leave their breeding place. The bees, that first remained

inside the tubes, started showing up at the entrance on the 5th post-emergence

day and from the 9th day they left their nest occasionally, for varying lengths

of time and walked on the tray and in its vicinity. They usually found their

way back to the tubes but did not necessarily enter the same tube in which

they developed. Thus, marked bees from a number of tubes were found

concentrated in one tube, and showed no animosity towards each other.

Further confirmation of the lack of specific attraction to the place of emer¬

gence was furnished when the locations of the tubes were changed during

the absence of the bees, and when the absorbent paper linings of the tubes,

that bear many of the odors deposited by the bees during their development,

were interchanged. The re-entry of the bees into the tubes after their de¬

parture could not be correlated with either tube location or order.

In Manoa, nesting activity diminished during the winter months of De-

cember-March, no new diggings were observed, but the bees continued to

be active and some new progeny developed. The progeny that had emerged

during the fall remained as unmated individuals with their mother through

the short winter. Thereafter some females nested in the same tunnel complex

as their mothers, whereas others dug themselves new nests. Nest digging

was resumed during March and, once started, continued even during stormy

weather.

Due to the continuous breeding of X. sonorina throughout the year, there

is an extensive overlap of generations and it is possible to find old mothers

side by side with young females, both mated and unmated. It is also possible

to find females who dig and start new nests at the same time as others

establish a 2nd or 3rd generation in already established nests. From May

14 at 1230 until May 15 at 1230 the activity of the bees at the “Fashion

Fabrics” location in Honolulu was followed. For this purpose, all of the bees

in the air were counted at that location each Vi hour. The results show (Fig.

8) that most of X. sonorina, like X. pubescens (Ben Mordechai et al., 1978),

fly about the break of day (0530-0600 in our case, when the sun started

shining in that place at 0615). Moreover, as soon as the boards with the

Figs. 8, 9. Fig. 8. Activity of X. sonorina at the Fashion Fabrics location from April 14,

1980 at 1200 to April 15, 1980 at 1200. Fig. 9. Diagram showing the location of the nectar

droplet and the maxillary motions performed during nectar regurgitation.

VOLUME 58, NUMBER 4

343

14. iv .80 15. iv .80

Hour

344

PAN-PACIFIC ENTOMOLOGIST

nests become shady, at 1900 the bees stopped flying in spite of the presence

of light and sufficiently high temperatures.

Active bees were seen regurgitating nectar at their nest entrance during

the warm hours of the day. The incidence of this phenomenon, that has also

been observed by Nishida (1963) for X. sonorina and by Corbet and Willmer

(1980) for X. mordax Smith, was recorded during May 15. From Table 2 it

is evident that, during the limited observation, the total number of nests

being guarded at the same time changed only a little, but the number of

regurgitating bees, and their percentage of the guards rose with time and

temperature increase. The lack of replication to these observations precludes

the possibility of deriving quantitative conclusions about this phenomenon,

but strengthens the impression that regurgitation is most prevalent at high

ambient temperatures. Regurgitation occurs when the bee moves a drop of

liquid from her mouth, out among the galeae and then imbibes it again.

This is followed by nearly regular opening and closing of the mandibles. The

sequence during the regurgitation cycles included fast and slow mandibular

movements, during some of which the drop appeared (Fig. 9).

Relationship among the cohabitant females. — The X. sonorina female may

nest solitarily. Moreover, her progeny may reach adulthood, both in the field

and in the laboratory, even in the absence of their mother. However, very

often the bees nest gregariously, with several females being present contin¬

uously in the same nest. They share guard duties and the field-going female

feeds the other through trophallaxis. This facultative, gregarious condition

is created from the natural growth and development of the progeny within

the nest, or from the joining of a number of females to form a group within

the same nest. As far as could be observed, only one female, the mother,

lays eggs whereas the role of the other bees may be confined to guarding

duties as in X. pubescens (Gerling, Hurd, and Hefetz, 1981). As said, oc¬

casional unmated females were found to have very worn wings. This was

considered unusual since the normal amount of nectar collecting done before

mating is not large and would not be expected to result in worn wings.

The males left the nest once the teneral stage had elapsed, except during

the winter, when they remained in the nest until spring-nesting activities

start during March and April. Thereafter they were found in old, abandoned

carpenter bee nests in dead Hibiscus branches, and in redwood boards. They

were seen feeding on nectar of flowers such as Samanea saman (Jacq.) and

Bougainvillea sp. I was unable to find the location of their territorial flights

and have but one record of such an occurrence about a Bougainvillea hedge

near a fence in which X. sonorina nests existed on the Island of Kauai.

Flower relations.— Xylocopa species, especially the tropical ones, are re¬

puted as having a strong tendency to rob nectar (Barrows, 1980; Faegri and

van der Pijl, 1979), and X. sonorina is no exception. The extent of the

phenomena and the mechanisms involved have been discussed in the lit-

VOLUME 58, NUMBER 4

345

Table 4. Flower species on which nectar robbery was observed from September 1979

to June 1980.

Family

Flower species

Previously

recorded

Abun¬

dance

of cuts

Acanthaceae

Asystasia gangetica (L.)

T. Andres

Barrows, 1980

+ + +

Sanchesia nobilis Hook.

—

+ + +

Apocynaceae

Allamanda oenotherifolia Pohl

Nishida, 1963

Catharantus (Vinca) roseus (L.)

—

+ +

Ervatamia divaricata (L.)

—

Thevetia peruviana (Pers.) Schum.

Barrows, 1980

+ + +

Bignoniaceae

Doxantha unguis-cati (L.)

—

+ + +

Tabebuia argenta (Bur. & K.) Schum.

—

+ + +

Tabebuia pentaphylla (L.)

—

+ + +

Caprifoliaceae

Lonicera japonica Thunb.

—

+ +

Boraginaceae

Cordia subcordata Lam.

—

Malvaceae

Hibiscus rosa-sinensis L.

Barrows, 1980

Hibiscus rosa-sinensis x H.

schizopetalus

—

Hibiscus schizopetalus Hooker

—

+ + +

Solanaceae

Brunfelsia americana L.

—

+ +

Verbenaceae

Stachytarpheta mutabilis

(Jacq.)

Barrows, 1980

+ +

erature (van der Pijl, 1954; Schremmer, 1972; Barrows, 1980). A mechanism

that involves the activity of ants, which prevent the bees from robbing flowers

that have extra floral nectaries, has also been described and named “ant

guard” (van der Pijl, 1954).

The present observations confirmed the knowledge about the flower species

robbed by X. sonorina and added several species to the list (Table 4). More¬

over, noticing the behavior of X. sonorina while visiting flowers, one can

expect numerous additional flowers of the Sympetalae to be added to this

list as the number of observations grows. Visits made to the flowers of the

genus Hibiscus proved to be of special interest. Hawaii abounds in Hibiscus

belonging to numerous varieties, most of which are hybrids or cultivars of

H. rosa sinensis L. Possibly, the most common hybrid is the cross H. rosa

sinensis x H. schizopetalus Hooker that produced red, pink or white flowers

and is reared in large numbers as a hedge plant. The flowers of this variety

have a long, slender, and thin walled calyx as in H. schizopetalus, the bracts

are short, and the petals are somewhat deflexed and a little serrate at the

edge. They differ in these characteristics from many cultivars of H. rosa

346

PAN-PACIFIC ENTOMOLOGIST

sinensis that have long bracts hugging the calyx that is goblet-shaped and

thick walled, and that have smooth edged petals. Both variety groups have

the typical nectaries located at the base of the corolla, hidden by the calyx,

and both produce nectar, that is then visited by ants.

Observations were conducted in order to understand the relationships of

the bees to the above mentioned Hibiscus species. These were carried out

in a number of ways: 1. Watching the behavior of the visiting bees; 2.

Following the blooming cycle of marked flowers from the bud to the wilting

stage; 3. Sampling flowers and registering their condition as to the presence

of nectar, ants, and incisions made by the visiting bees.

The behavior of the bees consisted of approaching the flower, flying near

it for a few seconds and either departing, or landing on the petals. In the

former cases, ants {Pheidole megacephala (F.)) were sometimes, but not

always, found in the calyx. In the latter case, the bees always turned with

their head to the petals and walked down to the calyx, where they robbed

the nectar of the flower. This was done by cutting the calyx either from the

top margin down by slitting it along its side or, rarely, by biting it. The

mouthparts are inserted in the slit and nectar is imbibed. At times, the bees

utilized present slits.

The blooming cycle of Hibiscus plants lasts one day. They open in the

early morning hours, and close in the afternoon or evening. However, the

opening time of the two varieties under observation differed. H. rosa sinensis

flowers opened at sunrise, whereas the flowers of the crosses with H. schi-

zopetalus usually opened later. This was true especially of the white and pink

flowered varieties, that opened between 0730 and 0830, and in shady spots

even later.

The behavior of the bees and the conditions of the flowers of H. rosa

sinensis (r), H. schizopetalus (s) and the cross (rxs) were noted at various

times and in several locations. Following notation that differences existed

between flower visits on (r) and (r x s), counts were made of flowers on two

hedges, one of (r) and one of (rxs) on the Manoa campus. In January,

February and March, the flowers were counted and sorted once a day, late

in the afternoon. In April and May several daily counts were made of the

same flowers. The results are depicted in Table 2, whereas Figures 10 and

11 show the dynamics of the flower visits. Figure 10 compares the visit from

sunrise to 1300 in (r) and (rxs) whereas Figure 11 depicts a whole day’s

visiting sequence on (rxs).

Although the degree to which (r) was visited varied greatly, it was always

less than that of (r x s), cuts in calyces of which occurred usually on 100%

of the flowers (Table 5). Visits to (r) began early in the morning and by 0800

bees have visited already over half of the flowers that were going to be visited

on that day. Most visits to (r x s) started, in nice weather, about 0800, when

Period of day

Figs. 10, 11. Fig. 10. Percent flower-calyx incisions made by X. sonorina during May 2, 1980 on 100 flowers of the hybrid H. rosa

sinensis x H. schizopetalus on the Manoa campus. Fig. 11. Percent flower-clayx incisions made by X. sonorina during April 15, 1980 on 100

flowers of 2 hibiscus varieties (H. rosa sinensis = r and H. rosa sinensis x H. schizopetalus = rxs) on the Manoa campus.

VOLUME 58, NUMBER 4 347

348

PAN-PACIFIC ENTOMOLOGIST

Table 5. Afternoon counts of total flower numbers visited on hedges of H. rosa sinensis (r)

and H. rosa sinensis x schizopetalus (r x s). The two records of May 5 on H. rosa sinensis were

taken from the same hedge.

Variety

Place

Date

Uncut

flowers

Top cut

Side cut

Top and

side cut

No.

rxs

Hawaii Kai

27.1

Manoa

29.1

37.5

62.5

Manoa

12.11

16.6

33.3

5.3

Manoa

10.III

100 +

Manoa

11.Ill

0.6

150

Manoa

2.V

100

Hawaii Kai

27.1

Manoa

29.1

Manoa

12.Ill

Manoa

10.III

8.5

87.1

4.2

Manoa

11.Ill

1.6

Manoa

2.V

100

Manoa

2.V

100

the flowers opened and later under cloudy and rainy weather. Once started,

the bees visited most of the flowers within two hours (Fig. 10).

More than one cut per flower was made in (r x s) and, to a lesser degree,

in (r). Multiple cuts started to show up when uncut flowers were still available

(Figs. 10, 11), at the same time bees were seen feeding through cuts made

by others, an activity that occasionally caused an enlargement of the slit in

the calyx.

Definite preference existed to “top cuts” from the margin of the calyx

down, in (r) and to side slits in (r x s). Top cuts appear in (r x s) mainly after

side slits have been made in all of the flowers. These are sometimes enlarged

through the pushing of the bees’ head, and may join a side slit to make a

substantial tear in the calyx.

Van der Pijl’s (1954) observation that ants visiting the nectaries preclude

robbery by bees prompted an examination of the subject in the case of

Hibiscus and X. sonorina. In the evening of January 27, 25 flowers of (r x s)

and 50 flowers of (r) were examined for both visits by bees as evidenced by

cuts and slits in the calyces, and the presence of ants. All of the calyces of

(r x s) and 20 calyces of (r) showed visits by bees (Table 5), two of the former

and 17 of the latter had also ants in them. In addition, 15 (r) flowers that

have not been cut by bees had ants in them. The possibility that the ants

have reached the flowers after the bees have finished visiting them was

examined by direct observation and was found to be incorrect.

VOLUME 58, NUMBER 4

349

Additional observations of behavior of Hibiscus- visiting bees revealed

that bees may approach flowers, hover above them and at times land on

them, and leave without robbing nectar. In all cases examination of these

flowers showed that ants of the species P. megacephala were inside. No such

behavior was observed when other ant species, especially Tetramorium si-

milis (Fr. Smith) were in the flower. Moreover, only a few of the flower-

visiting ants found in the count taken on January 27 belonged to P. me¬

gacephala.

Discussion

In general, the biology and ecology of X. sonorina conforms with that of

other Xylocopa species. Some points in its biology may be noteworthy either

because they are different than those of other species studied, or because

they may shed some light upon the social relationships of the bees. X.

sonorina starts to develop a nest as a single female, and later may be joined

by additional females. The latter may be her progeny, or other bees that

have come to reside with her. This phenomenon has also been observed

with X. pubescens (Ben Mordechai et al., 1978). Similarly, each nesting X.

virginica (L.) female is often joined by a non nesting female who remains

with her for the duration of the nesting period. However, in the latter case,

we do not know if there is familial relationship between the two bees (Gerling

and Hermann, 1978). Yet, in all known cases, only one female did the work

of collecting, cell preparing, and ovipositing; whereas the other bees per¬

formed mainly guard, and nest cleaning duties.

Hierarchy of the bees within the nest, as studied in some Xylocopa species

(Bonelli, 1976; Gerling, Hurd, and Hefetz, 1981) is such that the mother

does all the outside work whereas one daughter usually guards the entrance

and the others stay inside until they are ready to leave, mate, and establish

their own nest. The latter is often near, or a continuation of, the mother’s

nest.

The finding of females that were unfertilized but had worn wings indicates

that they did more than their normal share of outdoor activity. It is possible

that they were active for an unusually long time out of doors, perhaps until

providing food for other nest inhabitants. This extended period of activity

while being unmated may have resulted from inability to find mates due to

seasonal or topographical problems. It also might have been the consequence

of pheromonal suppression of the mating capacity of these bees by others,

possibly their sisters. The possibility of such pheromonal disruption of ac¬

tivity may be indicated by the fact that mating activity in some carpenter

bee species was shown to be regulated by pheromones (Velthuis and Gerling,

1980).

The floral relationships of X. sonorina differ from those of several other

studied species. Van der Pijl (1954) recorded the frequent visits of Xylocopa

350

PAN-PACIFIC ENTOMOLOGIST

species on Calotropis gigantea R. Br. Likewise strong association between

X. pubescens and X. sulcatipes Maa and C. procera (Ait.) Ait. f. exists (Ger-

ling, personal observations). Yet X. sonorina was never seen visiting the

abundant C. gigantea growing on Oahu, even when it was flowering near

the bees’ nests. Such an association was also never mentioned by Nishida

(1963) in his discussion of pollination and flower visits by X. sonorina.

The range of plant species that are robbed is also interesting, since several

of these occur as introduced plants, both in Hawaii and in Israel. Indeed,

some of the plant species, most notoriously Lonicera japonica Thunb., are

robbed in both countries. However, neither Hibiscus species, nor Thevetia

peruviana were found to be robbed in Israel in spite of their abundance in

the ranges of several Xylocopa species.

When the range of plant species that are robbed is compared with the

bees’ natural distribution, it becomes apparent that X. pubescens robs mainly

(or perhaps only) plants that originate in that range (e.g., Lonicera italiana

is wild in Israel, and Clerodendron inerme Gaertn. is from India). X. sono¬

rina, on the other hand, robs both Thevetia peruviana that is a New World

species, and Hibiscus that is of Asiatic origin.

Apparently, an element of experimenting and learning exists in the adop¬

tion of the new flower species and robbing them of their nectar, and X.

sonorina has been able to expand her flower range much more than X.

pubescens in that respect.

Acknowledgments

This work was done during my stay at the University of Hawaii at Manoa,

to the faculty and staff of which I am indebted for their assistance in many

ways. Special thanks are due to Drs. F. Haramoto, J. W. Beardsley, Jr., T.

Nishida, and W. Mitchell, to Mr. Lee Yudin and to Mrs. M. Uegawatchi all

of the Department of Entomology; to Drs. S. Margolin and P. Kroopnick

of the Hawaii Institute of Geophysics for allowing the use of X-ray facilities;

and to Dr. H. Nakasone of the Department of Botany for aid in identifying

the Hibiscus varieties.

Thanks are also due to the American Israeli Binational Science Foundation

that financed some of the observations made, under grant No. BSF 1562;

to the artist Mrs. R. Suzin and the photographers Ms. L. Maman and M.

Weinberg and to the typist Mrs. C. Meyer.

Literature Cited

Anzenberger, G. 1977. Ethological study of African carpenter bees of the genus Xylocopa

(Hymenoptera, Anthophoridae). Z. Tierpsychol., 44:337-374.

Barrows, E. M. 1980. Robbing of exotic plants by introduced carpenter and honey bees in

Hawaii with comparative notes. Biotropica, 12:23-29.

Ben Mordechai, Y., R. Cohen, D. Gerling, and E. Moscovitz. 1978. The biology of Xylocopa

VOLUME 58, NUMBER 4

351

pubescens Spinola (Hymenoptera Anthophoridae) in Israel. Isr. J. Entomol., 12:107-

121 .

Bonelli, B. 1976. Osservazioni eto-ecologiche sugli Imonotteri aculeati dell’Etiopa. Xylocopa

(Mesotrichia) combusta Smith (Hymenoptera-Anthophoridae). Boll. 1st. Entomol. Univ.

Studi Bologna, 32:1-31.

Corbet, S. A., and P. G. Willmer. 1980. Pollination of the yellow passion fruit: Nectar, pollen

and carpenter bees. J. Agric. Sci., 95:655-666.

Daly, H. V. 1966. Biological studies on Ceratina dallatorreana, an alien bee in California

which reproduces by parthenogenesis (Hymenoptera: Apoidea). Ann. Entomol. Soc. Am.,

59:1138-1154.

Faegri, K., and L. van der Pijl. 1979. Principles of pollination ecology. Pergamon Press,

Oxford, 244 pp.

Gerling, D., and H. R. Hermann. 1978. Biology and mating behavior of Xylocopa virginica

L. Behav. Ecol. Sociobiol., 3:99-111.

-, P. D. Hurd, Jr., and A. Hefetz. 1981. In-nest behavior of the carpenter bee Xylocopa

pubescens Spinola (Hymenoptera: Anthophoridae). J. Kans. Entomol. Soc., 54:209-218.

-, T. Orion, and M. Ovadia. 1979. Morphology, histochemistry and ultrastructure of

the yellow glands of Xylocopa pubescens Spinola (Hymenoptera, Anthophoridae). Int. J.

Insect Morphol. Embryol., 8:123-134.

Hurd, P. D., Jr. 1978. An annotated catalog of the carpenter bees (genus Xylocopa Latreille)

of the Western Hemisphere (Hymenoptera: Anthophoridae). Smithsonian Institution

Press, Washington, D.C, 106 pp.

Nishida, T. 1963. Ecology of the pollination of the passion fruit. Hawaii. Agric. Exp. Stn.

Tech. Bull., 55:21-31.

Pijl, L. van der. 1954a. Xylocopa and flowers in the tropics I. The bees as pollinators. Lists

of the flowers visited. Proc. K. Ned. Akad. Wet., ser. C, Biol. Med. Sci., 57:413-423.

-. 1954b. Xylocopa and flowers in the tropics II. Observations on Thunbergia, Ipomoea,

Costus, Centrosema, and Canavallia. Proc. K. Ned. Akad. Wet., ser. C, Biol. Med. Sci.,

57:541-551.

-. 1954c. Xylocopa and flowers in the tropics III. Observations on some Papilionaceae,

Melastoma, Calotropis, Cassia and some orchids, with general considerations. Proc. K.

Ned. Akad. Wet., ser. C, Biol. Med. Sci., 57:552-562.

Schremmer, F. 1972. Der Stechsaugruessel, der Nektarraub, das Pollensammeln und der

Bluetenbesuch der Holzbienen ( Xylocopa) (Hymenoptera: Apidae). Z. Morphol. Tiere,

72:263-294.

Velthuis, H. H. W., and D. Gerling. 1980. Observations on territoriality and mating behaviour

of the carpenter bee Xylocopa sulcatipes. Entomol. Exp. Appl., 28:82-91.

Footnote

1 Submitted for publication March 12, 1981.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 352-364

Published 30 December 1983

FIVE NEW SPECIES AND NEW SYNONYMIES FOR THE GENUS

DERAEOCORIS (HETEROPTERA: MIRIDAE) FROM

WESTERN NORTH AMERICA

Vincent Razafimahatratra 1 and John D. Lattin

Systematic Entomology Laboratory, Department of Entomology,

Oregon State University, Corvallis 97331

The mirid genus Deraeocoris Kirschbaum contains approximately 200

species distributed throughout much of the world. Most species are believed

to be predacious on small arthropods and some, like Deraeocoris brevis

(Uhler), are important predators of economically important insects. The new

species described here and the proposed synonymies are the result of a recent

study of the Deraeocoris of western North America completed as partial

fulfillment of the requirements for the Ph.D. degree at Oregon State Uni¬

versity by the senior author. Approximately 10,000 specimens were ex¬

amined, including the relevant type specimens. All measurements are given

in millimeters. Institutional abbreviations are given under Acknowledg¬

ments.

Deraeocoris fraserensis Razafimahatratra and Lattin, new species

Diagnosis. — Similar in general aspect to D. validus (Reuter) but cuneus

distinctly red; scutellum dark red; calli not punctate as in D. poecilus McAtee;

differs from D. ornatus Knight in having antennal segment II greater in

length than pronotum; male and female genitalia distinctive but showing

relationship to D. validus.

Male. Length 4.00. Head: Length 0.56, width 0.88, vertex 0.40, dark brown

to black, spots on middle of front, along eye margin, on median line and at

each side of tylus pale white to yellow; carina of vertex slightly convex,

white, trapezoidal shaped; rostrum reaching upon middle coxae. Antennae:

Segment I, 0.32, piceous, sparsely pubescent; segment II, 1.04, constricted

at base, nearly cylindrical but slightly thicker toward apex, piceous to black,

covered with short, dense pubescence and with several erect setae; segment

III, 0.40; segment IV, 0.40; last two segments paler, densely pubescent and

with erect setae. Pronotum: Length 0.96; anterior angles 0.80; width at base

1.76; uniformly yellow to testaceous, slightly darker on each side of median

line, punctures coarse and black, lateral margins of pronotum carinate; calli

separated by one deep puncture, piceous to black, latero-anterior angles

invaded with pale; area before calli pale white, a few punctures may be

present at antero-lateral angles of calli; scutellum reddish brown, lateral

VOLUME 58, NUMBER 4

353

angles, apex and apical half of median line ivory-white, punctures black.

Hemelytra: Width 1.92; yellowish testaceous, more or less translucent in

one specimen, corium marked with brown at apex and middle; punctures

black; cuneus red, darker at apex; membrane clear, apical half and areoles

fumate. Legs: Red to piceous; femora with two pale bands near apex, with

a row of darker spots on anterior surface; tibiae triannulated with pale; basal

segments of tarsi pale; claws, piceous, deeply cleft. Genitalia parameres as

in Figure 1; vesica as in Figure 2; both vesica and parameres showing close

relationship to D. validus.

Female. Similar to male but darker in color and larger in size. Length

4.48; maximum width 2.24. Second antennal segment, 0.96, black, slightly

paler at middle, with short pubescence on apical third and with several erect

setae, gradually thickening toward apex. Posterior wall of bursa copulatrix

as in Figure 3; sclerotized rings as in Figure 4. The female genitalia also

show a close relationship to D. validus.

Holotype. — Male, Wells, British Columbia, Wendell Park, 8-4-1949, G.

J. Spencer, deposited in the collection of the University of British Columbia.

Paratypes: One topotypic male; one topotypic female; one male and three

females from Quesnel, British Columbia, 6-18-1949, G. J. Spencer (UBC).

The species name, fraserensis, has been derived from the name of the type

locality of the type series, the Fraser Plateau.

Distribution.— At present, this species is only known from the above lo¬

calities.

Biology. — One of the specimens from Quesnel bears the label ‘on Alder.’

Deraeocoris fraserensis is structurally related to D. validus. Moreover, Ques¬

nel and Wells are located along the Fraser and the Willow Rivers respectively.

It is probable that D. fraserensis lives on vegetation bordering the Fraser

River and its tributaries. It is noted that D. validus is found mostly on

vegetation bordering streams and lakes, usually on Salix sp.

Deraeocoris picipes Knight, new status

Deraeocoris incertus var. picipes Knight, 1921:111, 116; Knight, 1927:37.

Deraeocoris incertus var. carneolus Knight, 1921:111, 116, NEW SYN¬

ONYMY.

Diagnosis. — Very similar to D. incertus but larger in size; femora piceous,

only pale at very apex; tibiae usually with only one pale band on apical half;

genitalia, although related to D. incertus, show differences in the left and

right claspers and in the sclerotized rings of the female.

Male. Length 5.28-7.02. Head: Length 0.64, width 1.04, vertex 0.32;

markings similar to D. incertus ; dorsal width of an eye greater than vertex.

Antennae: Segment I, length 0.40; segment II, 1.76-1.92, piceous to black,

nearly cylindrical, covered with dense, short pubescence and with fewer

354

PAN-PACIFIC ENTOMOLOGIST

Figs. 1-8. Figs. 1-4, Deraeocoris fraserensis. Fig. 1, claspers. Fig. 2, vesica. Fig. 3, posterior

wall of bursa copulatrix. Fig. 4, sclerotized rings. Figs. 5-8, Deraeocoris cochise. Fig. 5, claspers.

Fig. 6, vesica. Fig. 7, posterior wall of bursa copulatrix. Fig. 8, sclerotized rings.

exserted longer setae; segment III, 0.48; segment IV, 0.40; last two segments

piceous, cylindrical, with short pubescence and with a few longer setae.

Pronotum: Length 0.96-1.04, anterior angles 0.72, width at base, 1.76; disk

of pronotum uniformly piceous, lighter-colored specimens having pale spots

behind calli; lateral and basal margins of pronotum narrowly ivory-white,

lateral margins more carinate and more sinuate than in D. incertus\ calli

piceous to black, convex, area before calli usually pale; scutellum, punctate,

dark brown to piceous black, angles narrowly pale. Hemelytra: Width 2.24;

uniformly reddish brown to piceous, translucent in light-colored specimens;

VOLUME 58, NUMBER 4

355

basal area of corium, embolium and clavus pale; cuneus dark brown to

piceous, basal one-third pale in light-colored specimens; membrane infus-

cated except narrowly behind cuneus and areoles, clear. Legs: Reddish brown

to piceous; femora narrowly pale at apices; tibiae banded with one pale

annulus on apical half, row of spinose setae on anterior face of tibiae; tarsi

and claws piceous. Venter: Piceous, covered with golden pubescence. Gen¬

italia: Parameres as in Figure 20, right clasper distinct from that of D.

incertus, vesica as in Figure 21, closely related to that of D. incertus.

Female. Length 4.96-5.76. Maximum width 2.24-2.48. Second antennal

segment, length 1.60, slender, thickening toward apex. More broadly pale,

more robust and more ovate than male. Dorsum often tinged with red.

Latero-anterior angles of calli invaded with pale. Very similar to D. incertus

but larger in size, tibiae usually with only one pale annulus; femora narrowly

pale at apices. Posterior wall of bursa copulatrix as in Figure 22; sclerotized

rings as in Figure 23; genitalia showing close relationship to D. incertus but

distinct.

Holotype.— Female, Webber’s Camp, 7800', Sta. Catalina Mts., Arizona,

July 26, H. H. Knight. This specimen is in the U.S. National Museum

Collection.

Material examined. —Arizona: Cochise Co.: Deer Park, Chiricahua Mts.,

8000', 7-7-1927, J. A. Kusche (CAS); S.W.R.S., 5 mi. W Portal, 5400', 7-18-

1957, M. Statham (AMNH); Rustler Park, Chiricahua Mts., 8200', 6-22-

1955, A. Anderson and M. Statham (AMNH); Huachuca Mts., 8-2-?, H. G.

Barber (USNM); Chiricahua Mts., 8-9000', on Pinus ponderosa, 8-31-1976,

J. D. Pinto (UCR): 7-8-1932, R. H. Beamer (UK). Coconino Co.: Flagstaff,

7-27-1936, R. H. Beamer (UK); Williams, 8-4-?, H. H. Knight (USNM).

Graham Co.: Graham Mts., 7-6-1955, Ordway and Statham (AMNH). Col¬

orado: Archuleta Co.: Pagosa Spgs., 6-24-1964, H. R. Burk (TAM). Larimer

Co.: Pingree Park, 8-8-1925, Beamer and Lawson (UK). Las Animas Co.:

Trinidad, 9000', 8-8-1925, H. H. Knight (TAM). Saguache Co.: 1 mi. E

North Pass, 8-19-1969, J. C. Schaffner (TAM). New Mexico: Catron Co.:

Bursum Camp, 18 mi. E Alma, 9000', 9-11 to 12-1961, R., P. and J. Rindge

(AMNH); 2 mi. W La Cueva, 7900', 7-30-1961, F., P. and J. Rindge (AMNH).

Colfax Co.: 5 mi. E Eagle’s Nest, 8-8-1969, J. C. Schaffner (TAM). Otero

Co.: Cloudcroft, 6-27-1940, R. H. Beamer (UK); 2-3 mi. E Cloudcroft, 1947

(AMNH). Santa Fe Co.: 8 mi. E Santa Fe, 8700', Hyde State Park, 7-29-

64, F., P. and J. Rindge (AMNH). Taos Co.: Sipapu, 9-3-1965, R. G. Jones

(USU); Tres Ritos, 9-25-1968, J. C. Schaffner (TAM). Torrence Co.;Tajique,

6-28-1947, R. H. Beamer (UK). Utah: Beaver Co.: Beaver, 6-23-1966, G.

F. Knowlton (USU).

Material illustrated. — Male, Cloudcroft, New Mexico, 6-27-1940, R. H.

Beamer (UK); female, Flagstaff, Arizona, 7-27-1936, R. H. Beamer (UK).

Distribution.— Deraeocoris picipes is only found in the southern parts of

356

PAN-PACIFIC ENTOMOLOGIST

Figs. 9-15. Figs. 9-11, Deraeocoris gilensis. Fig. 9, claspers. Fig. 10, posterior wall of bursa

copulatrix. Fig. 11, sclerotized rings. Figs. 12-15, Deraeocoris knightonius. Fig. 12, claspers.

Fig. 13, vesica. Fig. 14, posterior wall of bursa copulatrix. Fig. 15, sclerotized rings.

VOLUME 58, NUMBER 4

357

Figs. 16-23. Figs. 16-19, Deraeocoris schuhi. Fig. 16, claspers. Fig. 17, sclerotized rings.

Fig. 18, posterior wall of bursa copulatrix. Fig. 19, vesica. Figs. 20-23, Deraeocoris picipes. Fig.

20, claspers. Fig. 21, vesica. Fig. 22, posterior wall of bursa copulatrix. Fig. 23, sclerotized

rings.

358

PAN-PACIFIC ENTOMOLOGIST

the Rocky Mountains, in Utah, Colorado, Arizona, New Mexico and prob¬

ably in Mexico.

Biology.— The habits of this species are not known. Collection records

indicate that it may have only one generation per year and that it overwinters

in the egg state. D. picipes is known to breed on conifers: Pinus ponderosa

Dougl., Picea sp. and Pseudotsuga menziesii var. glauca (Beissn.) Franco.

Deraeocoris picipes was described by Knight (1921) as a variety of D.

incertus. It is elevated to the species level because it differs from D. incertus

chiefly in the genital structure and in size. The variety carneolus Knight of

D. incertus does not differ structurally from D. picipes. It appears to be only

the light color form of the latter and should not be given taxonomic rank.

Deraeocoris cochise Razafimahatratra and Lattin, new species

Diagnosis.— Dorsum pale yellow, nearly glabrous; hemelytra translucent;

scutellum impunctate; legs uniformly pale yellow; claws deeply cleft near

bases; genitalia distinctive for the species.

Male. Length 5.92. Head: Length 0.80, width 1.04, vertex 0.48, pale yel¬

low, median line of front, carina and spot near margin of eye ivory-white;

narrow spot between anterior margin of eye and antennal base, dark brown;

carina slightly carinate; collum pale yellow, brown on sides; rostrum reaching

upon middle coxae. Antennae: Segment I, length 0.48, pale yellowish with

erect setae; segment II, 1.36, nearly cylindrical but more slender base, pale

yellow, black at apex, covered with short pubescence and a longer erect setae;

segment III, 0.54; segment IV, 0.40; last two segments pale yellow covered

with short pubescence and longer erect setae. Pronotum: Length 1.28, an¬

terior angles 0.36, width at base 2.16; pale yellowish only slightly darker on

each side of median line and behind calli, coarsely punctuate, with short

and sparse pubescence near lateral margins; lateral margins nearly straight,

slightly carinate; calli slightly convex, confluent, smooth, largely pale yel¬

lowish, anterior and posterior margins irregularly delimited by heavy black

lines, transverse spot inside each callus and spots at their latero-anterior

angles, dark brown or black; two deep punctures present at middle of pos¬

terior margin of calli; scutellum, impunctate, convex, pale to ivory-white,

spots on each side of median line, dark brown. Hemelytra: Width 2.48,

uniformly pale yellowish, translucent, punctures fine, yellowish brown, cu-

neus pale, translucent; membrane pale, translucent. Legs: Pale yellow; femora

with long erect setae on ventral side; tibiae only slightly brownish at apex

covered, beset with long setae; claws cleft near bases. Venter: Brownish

yellow and tinged with reddish, covered with semi-erect setae. Genitalia:

Parameres as in Figure 5; vesica as in Figure 6.

Female. Length 6.22, width 2.88. Very similar in general appearance to

male but more robust and more ovate. Second antennal segment, length

VOLUME 58, NUMBER 4

359

1.28, pale yellowish and slender, apical one-fifth thicker and brown. Genitalia

as in Figures 7 and 8.

Holotype. — Male, Cochise Co., Arizona, vicinity of Portal, 1500-1700

meters, May 20, 1978, on Quercus oblongifolia Torr., R. T. Schuh; deposited

in American Museum of Natural History, New York. Allotype: Female,

same data as holotype. Paratypes: 120 males and females, collected on the

same date and at the same location as the types by R. T. Schuh (AMNH).

Other specimens examined: Molino Basin, Mt. Lemon, Arizona, V-19-1953,

A. and H. Dietrich (CU); Arizona, Cornell University Lot 34' (CU).

This species is closely related to D. hyalinus, described by Carvalho and

Schaffner (1973) from Mexico. It differs from the latter by the following

characters: vertex of front devoid of black spots; length of first antennal

segment equal to width of vertex; scutellum having two black spots on each

side of median line; embolium distinctly punctate, although punctures are

often concolorous; genital claspers very distinct from that of D. hyalinus.

According to R. T. Schuh, this species is very abundant on oak at the type

locality. However, at present, its biology is not known.

The sensory lobe of the left clasper on some of the paratypes is shorter

than that of the holotype. This species resembles closely D. knightonius

which occurs in California and Oregon but they are easily separated by the

genital structures.

Deraeocoris schuhi Razafimahatratra and Lattin, new species

Diagnosis. — Dorsum largely pale yellowish brown; calli with dark brown

and red impression; scutellum dark brown, transversally rugose, lateral an¬

gles, apex and apical half of median line pale yellow; wings of male long;

membrane of female barely surpassing tip of abdomen; sensory lobe of left

clasper very elongate; related to D. piceicola Knight and D. navajo Knight.

Male. Length 6.24. Head: Length 0.80, width 1.12, vertex 0.48; pale yel¬

lowish, longitudinal spots along margin of eye, on each side of tylus, on juga,

on lorae and bucculae, and row of transversal spots on each side of median

line of front, red; front moderately convex; rostrum reaching middle coxae.

Antennae: Segment I, length 0.48, pale yellow; segment II, 1.60, nearly

cylindrical, basal one-fourth narrower, densely pubescent, beset with several

erect setae, dark brown, paler at base; segment III, 0.56; segment IV, 0.48;

last two brown, pale pubescent and beset with long erect setae. Pronotum:

Length 1.36, anterior width 0.96, posterior width 2.40; disk, densely and

coarsely punctate, pale yellowish brown, darker along posterior margin, pu¬

bescence short; pronotal collar, pale, pruinose; calli moderately convex, dark

brown, outlined with red, antero-lateral angles invaded by pale, ray from

posterior margin of each callus curving posteriorly and anteriorly to anterior

margin of pronotum, each callus separated by two deep punctures; area

between and before calli pale; posterior half of pronotal margin distinctly

360

PAN-PACIFIC ENTOMOLOGIST

carinate; ostiolar peritreme white, tinged with red; scutellum dark brown,

lateral angles, apex and apical half of median line pale, transversely rugose,

shortly and sparsely pubescent. Hemelytra: Width 2.72; embolar margin

nearly parallel; short pubescence, pale yellowish, translucent apex of clavus,

apex and middle of corium and along claval suture, dark brown; cuneus

concolorus with corium, apical half, dark reddish brown; membrane infus-

cated, veins reddish brown. Legs: Yellowish brown; femora biannulate with

dark red on apical half; tibiae triannulate with dark red, beset with long

prominent setae; tarsi darker at apices; claws slender, not cleft at base. Venter:

Piceous. Genitalia: Claspers as in Figure 16, sensory lobe of left clasper very

elongate; vesica as in Figure 17, related to D. piceicola.

Female. Length 5.60, width 2.76. Similar to male but paler in coloration

and more ovate. Second antennal segment, length 1.60, slender, apical one-

fourth thicker, pale yellow, brown near apex; calli and scutellum more broad¬

ly pale; membrane barely surpassing tip of abdomen. Posterior wall as in

Figure 18, distinctive for the species; sclerotized rings as in Figure 19.

Holotype. — Male, 9 mi. E Shasta City, California, on Abies magifica var.

shastensis Lemmon, August 8, 1956, J. Schuh; deposited in the California

Academy of Sciences. Paratypes: Two males and eight females taken with

the types (OSU); one male, Plaskett Meadows, Glenn Co., California, 6000',

7-27-1960, H. B. Leech (CAS); one female, Martin Springs, Lassen Co.,

California, 7-31-1922, J. O. Martin (CAS); one male, Haskell Creek, Sierra

Co., at light, 8-18-1971, S. Frommer (UCR); one male, Biledo Meadow,

Madera Co., 7-27-1946, R. L. Usinger (UCB).

This species is named after the late Mr. J. Schuh, Klamath Falls, Oregon,

who collected the type series of this species.

Deraeocoris schuhi is one of the most difficult species of the genus because

of its coloration and form. This species is distributed widely in northern

California. It may occur also in southern Oregon.

Deraeocoris gilensis Razafimahatratra and Lattin, new species

Diagnosis.— Closely related to D. vanduzeei Knight, but setae on dorsum,

antennae and legs denser and distinctly longer; punctures on dorsum sparser,

except for cluster of punctures behind calli; male genitalia similar to but

distinct from that of D. vanduzeei, female genitalia distinctive for the species.

Male. Length 4.64. Head: Length 0.64, width 0.96, vertex 0.48; pale yel¬

lowish; front more convex than in D. vanduzeei; pubescence longer; posterior

margin of vertex more sinuate; rostrum reaching hind margins of middle

coxae. Antennae: Segment I, length 0.40, pale yellow; segment II, 1.12,

gradually thickening toward apex, pale yellowish, slightly darker at apex,

covered with long erect setae, length reaching three times greatest thickness

of segment; segment III, 0.48, pale yellow; segment IV, 0.40, pale brown.

Pronotum: Length 1.12, anterior width 0.96, posterior width 2.00, very

VOLUME 58, NUMBER 4

361

similar to D. vanduzeei, pale yellow, darker on posterior half of disk; punc¬

tures deeper than in D. vanduzeei ; cluster of deep punctures behind calli;

calli as in D. vanduzeei, pair of punctures separating calli more widely sep¬

arated; setae of pronotal disk distinctly longer; scutellum pale yellow, spots

on each side of median line black, with dense and long setae. Hemelytra:

Width 2.30, pale yellow, black punctate, punctures deeper and more sparse,

setae longer than in D. vanduzeei ; spots at middle of corium, at apices of

embolium clavus and corium brown, translucent; cuneus, pale, apical one-

third and basal margin black; membrane infuscated, veins darker. Legs: Pale

yellow, densely clothed with long erect setae, distinctly longer than in D.

vanduzeei ; femora with two red bands near apices; tibiae with three reddish

bands, apical band paler, setae on tibiae reaching in length twice the thickness

of article; claws deeply cleft. Venter: Reddish, pale near sides. Genitalia

claspers as in Figure 9, sensory lobe of left clasper distinct from that of D.

vanduzeei ; vesica not illustrated, related to that of D. vanduzeei but different

in the shape of the spicules (the vesica of the male did not inflate).

Female. Length 4.88; maximum width 2.56; segment II of antennae, length

1.20; very similar to male but slightly large in size. Posterior wall of bursa

copulatrix as in Figure 10, sclerotized rings as in Figure 11, distinct from

those of D. vanduzeei.

Holotype: Male, Gila Co., 2 mi. W Miami, Arizona, 3800' elev., on oak,

5-4-1979, J. D. Pinto and E. M. Fisher; deposited in the collection of the

University of California, Riverside. Paratypes: One female and three males

with same data as the type (UCR); three males from Molino Basin, Mt.

Lemmon, Arizona, 5-19-1953, A. and H. Dietrich (CU).

Like D. vanduzeei, this species lives on oak. The above specimens have

been compared to the holotype of D. vanduzeei which was in the possession

of the authors at the time of the writing of this description.

The head of some of the paratypes are marked with brownish as in D.

comanche.

The species name gilensis is derived from the name of Gila County, the

type locality.

Deraeocoris knightonius Razafimahatratra and Lattin, new species

Deraeocoris cerachates, sensu Knight, 1921:187, nec Uhler, 1894:265.

Diagnosis. — More elongate in form than D. cerachates Uhler; dorsum,

especially the pronotum, very coarsely punctate; with short and sparse pu¬

bescence, except near anterior margin of pronotum where a few longer setae

are present; calli more convex than in D. cerachates.

Male. Length 5.76-6.72. Head: Length 0.80, width 0.96-1.12, vertex 0.48-

0.56; pale yellowish, marked brown or dark brown on sides and base of

tylus, on each side of median line on front, along eye margin and on each

362

PAN-PACIFIC ENTOMOLOGIST

side of median line of vertex; front moderately convex; lorae and genae

marked with reddish to blackish brown; rostrum reaching upon middle

coxae. Antennae: Segment I, length 0.48-0.56, pale brownish; segment II,

1.44-1.76, pale yellowish, apical one-fourth infuscated, slender, thickening

toward apex, clothed with pale pubescence and longer setae; segment III,

0.64; segment IV, 0.48. Pronotum: Length 1.12-1.52, anterior width 0.96-

1.12, posterior width 1.92-2.40; disk deeply and more coarsely punctate

than D. cerachates, with sparse and short pubescence, few long setae often

present on anterior angles and lateral margins; pale yellowish to rich brown¬

ish, posterior half-brown to dark brown, median line pale; lateral margins

slightly concave when viewed from above, posterior half carinate; calli dis¬

tinctly convex, heavily outlined with brown or black as in D. cerachates’,

scutellum moderately convex, pale to ivory-white, often brownish to black

on each side of median line. Hemelytra: Width 2.40-3.04; pale yellow,

corium, clavus, and embolium irregularly brown, dark coloration spreading

from punctures, pubescence sparse and shorter than in D. cerachates ; cuneus

pale, often narrowly at apex and along basal margin brown; membrane

slightly infuscated, sometimes darker on apical half; veins dark brown. Legs:

Pale yellow, covered with long setae as in D. cerachates ; femora slightly

brownish on apical half; tibiae slightly brownish at apex, not very distinctly

banded with brown at middle, claws deeply cleft. Genitalia: Claspers as in

Figure 12, vesica as in Figure 13, showing close relationship to D. cerachates.

Venter: Brownish and marked with piceous, pale pubescent.

Female. Length 5.44-6.72, maximum width 2.40-3.20; segment II of

antennae 1.44-1.76, very similar to the male in size and coloration. Geni¬

talia: Posterior wall as in Figure 14, sclerotized rings as in Figure 15.

Holotype.— Male, June 9, Santa Cruz County, California, 600 feet, W. M.

Gifford; deposited in the California Academy of Sciences.

Paratypes, 40 males and females from the following localities.—California:

Alameda Co.: two females, Oakland Hills, 5-26-1928, R. L. Usinger (UCB);

two males, one female, Leona Heights, 5-15-1933, R. H. Beamer (UK).

Fresno Co.: one male, Fresno, 5-6-1979, E. L. Paddock (CDFA). Inyo Co.:

three males, Lone Pine, on alder, 6-8-1929, E. P. Van Duzee (CAS); one

female, Bishop, 6-20-1929, R. L. Usinger (CAS). Calaveras Co.: one male,

Angels Camp, 5-22-1930, E. P. Van Duzee. Eldorado Co.: one specimen,

Alpine Creek, near Tahoe, 7-30-1925, E. P. Van Duzee (CAS). Mariposa

Co.: one specimen, Yosemite, 3880-4000', 5-22-1931, R. L. Usinger (UCB).

Riverside Co.: five males, Soboda Spgs., 5-30-1917, E. P. Van Duzee (CAS).

San Bernadino Co.: five females, Mill Creek, 9-22-1923, E. P. Van Duzee

(CAS); three females, Mill Creek, 6000' on Alnus rhombifolia Nutt., 7-26-

1947, Timberlake (UCR). Santa Barbara Co.: one specimen, 3 mi. N Refugio

Beach, 6-21-1965, G. A. Gorelick (UCB). San Mateo Co.: one female, Pes-

cadero, 6-15-1917, C. L. Fox (CAS). Tuolumne Co.: two males, two females,

VOLUME 58, NUMBER 4

363

Bear River, 5-17-1930, E. P. Van Duzee (CAS); one male, one female,

Tuolumne Co., on Artemesia, 7-22-1975, Bingham (CDFA). Oregon: Benton

Co.: one male, Lobster Valley, on Alnus, 7-23-1977, J. D. Lattin (OSU).

Jackson Co.: one male, Carberry Creek, 7-8-1962, J. D. Vertrees (AMNH).

Klamath Co.: one female, 11 mi. NE Bly, 5000', 8-14-1975, Schuh and

Denning (AMNH); three males, one female, Sand Creek, on Alnus, 7-31-

1956, J. Schuh (OSU).

Distribution.— This species is known from California and Oregon. It has

a more northern distribution than D. cerachates.

Biology.— The adults of this species are collected from May to September.

It is known to breed on Alnus sp., and A. rhombifolia Nutt.

This species was thought by H. H. Knight (1921) to be Uhler’s D. cera¬

chates because he had not seen Uhler’s type himself. He asked E. P. Van

Duzee to compare a male specimen with Uhler’s type. The latter declared

them identical. If fact, they are different in the structure of pronotum, hair¬

iness and in general appearance. Deraeocoris cerachates is ovate in form and

distinctly pilose while the specimens described by Knight as “ cerachates ”

are rather elongate and not conspicuously hairy.

Deraeocoris incertus Knight

Deraeocoris incertus Knight, 1921:114.

Deraeocoris rufusculus Knight, 1921:116, NEW SYNONYMY.

Deraeocoris incertus and D. rufusculus were described by H. H. Knight

from specimens collected by A. A. Nichol in Portland, Oregon. The authors

have examined the types of these taxa and find no differences in the para-

meres and that both have exserted setae on the second antennal segment.

Deraeocoris rufusculus is only the dark color form of D. incertus. These color

forms are often found in the same local populations.

Deraeocoris fulvescens (Reuter)

Camptobrochis ( Euarmosus ) fulvescens Reuter, 1909:53, 56.

Deraeocoris fulvescens, Knight, 1921:167.

Deraeocoris pilosus Knight, 1921:169, NEW SYNONYMY.

The specimens described by H. H. Knight as D. pilosus do not show any

structural difference from D. fulvescens. The genitalia of these two taxa are

identical in structure. Deraeocoris pilosus is believed to be only the light

color form of D. fulvescens.

Deraeocoris cerachates Uhler

Deraeocoris cerachates Uhler, 1894:265.

Deraeocoris californicus Knight, 1921:185, NEW SYNONYMY.

364

PAN-PACIFIC ENTOMOLOGIST

The authors have compared the types of these taxa and find them to be

identical. The specimen which is believed to be Uhler’s type is in the col¬

lection of the California Academy of Sciences. This specimen bears the

following labels: San Jose del Cabo, female, Deraeocoris cerachates Uhler.

Acknowledgments

The authors would like to thank the following curators and institutions

for allowing us to examine or borrow the specimens for this paper: P. H.

Arnaud, Jr., California Academy of Sciences (CAS); P. D. Ashlock, Uni¬

versity of Kansas (UK); R. C. Froeschner, U.S. National Museum of Natural

History (USNM); W. J. Hanson, Utah State University (USU); A. R. Hardy,

California Department of Food and Agriculture (CDFA); Oregon State Uni¬

versity (OSU); L. L. Pechuman, Cornell University (CU); J. D. Pinto, Uni¬

versity of California, Riverside (UCR); J. A. Powell, University of California,

Berkeley (UCB); J. C. Schaffner, Texas A and M University (TAM); G. G.

E. Scudder, University of British Columbia (UBC).

Literature Cited

Carvalho, J. C. M., and J. C. Schaffner. 1973. Neotropical Miridae, CLX: Descriptions of

two new species of Deraeocorini (Hemiptera). Rev. Bras. Biol., 33(suppl.):61-64.

Knight, H. H. 1921. Monograph of the North American species of Deraeocoris (Heteroptera:

Miridae). Rep. Minn. State Entomol., 18:76-210.

-. 1927. Notes on the distribution and the host plants of some North American Miridae.

Can. Entomol., 59:34-44.

Reuter, O. M. 1909. Bemerkungen liber nearktische Capsiden nebst Beschreibung neuer Arten.

Acta Soc. Sci. Fenn., 36:1-86.

Uhler, P. R. 1894. Observations upon the heteropterous Hemiptera of Lower California, with

descriptions of new species. Proc. Calif. Acad. Sci., (2)4:223-295.

Footnote

1 Present address: Boite Postale 4329, Tananarive, Madagascar.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 365-366

Published 30 December 1983

SCIENTIFIC NOTE 1

TABANIDAE (DIPTERA) ATTRACTED TO ARTIFICIAL

LIGHTS IN CALIFORNIA

During July and August last summer, three specimens (two males, one

female) of the rather primitive horse fly, Pilimas abaureus (Philip), and a

single male of Hybomitra californica (Marten) were collected in a black light

trap by Williams 2.5 miles south of West Point in Calaveras County. These

records constitute the third and fourth tabanid species from California (out

of the 7 3 species) that have been collected at artificial lights (the first in black

light flight traps), and the first records of these species from Calaveras County.

Pilimas californica (Bigot) and Apatolestes parked (Philip) have also been

taken while attracted to lights in the state (Middlekauff and Lane, 1980,

Bull. Calif. Insect Surv., 22:1-99). The significance of this phenomenon

among habitually day-flying tabanids is not understood, but either attraction

of these flies while resting on nearby vegetation or that their flight patterns

may include crepuscular or nocturnal habits is suggested.

None of the aforementioned tabanids has been captured from, or observed

feeding on, vertebrates, but H. californica has been taken in C0 2 -baited

insect flight traps in Mendocino County (Middlekauff and Lane, 1980). The

latter finding suggests that H. californica is hematophagous, whereas P.

abaureus and P. californica, both of which have been swept from flowers,

as well as A. parked, may be autogenous, i.e., capable of producing eggs

without an ovigenous blood meal.

Larvae and pupae of H. californica were described by Lane (1979, J. Med.

Entomol., 16:142-149), who collected larvae from soil and mosses bordering

a woodland vernal pool. The immatures of none of the three known Pilimas

species has been described; therefore it is hoped that the black light trap site

may furnish a clue for discovering the larval habitat of P. abaureus next

season. Very little is known about the biologies of immature (or adult)

Pilimas or other indigenous members of the generalized subfamily Pango-

niinae (. Apatolestes, Brennania, Stonemyia). For instance, larvae and pupae

of only two of 16 pangoniines from California have been found, namely,

Brennania hera (Osten Sacken) and Apatolestes actites Philip and Steffan

(Middlekauff and Lane, 1980). Detailed descriptions of the immatures of

these two species will be presented in a forthcoming report (Lane and Philip,

in prep.). Both species were found in terricolous habitats along the coast

where availability of potential vertebrate host blood is minimal.

The two males and lone female of P. abaureus are passed among the

audience for comparison with two males of the related P. californica, which

was the subject of an earlier meeting note by Philip (1978, Pan-Pac. Entomol.,

54:74). The latter males were taken (and photographed) while hovering

366

PAN-PACIFIC ENTOMOLOGIST

(probable mating postures!) at the tops of 300-foot high giant sequoia trees.

The identities of both sexes of P. abaureus were confirmed by Dr. Robert

S. Lane of the University of California, Berkeley, who also determined the

male of H. californica. The hind tibial fringes of the last are entirely black

in contrast to the usual reddish fringes in the females.

Cornelius B. Philip, California Academy of Sciences, Golden Gate Park,

San Francisco 94118, and Stanley C. Williams, Department of Biological

Sciences, San Francisco State University, San Francisco 94132.

1 This note was submitted for publication in the Proceedings of the 406th meeting of the

Pacific Coast Entomological Society, 20 February 1981.—Ed.

CORRECTION, page 8, line 13

Before the turn of the century T. D. A. Cockerell was on the staff of the

New Mexico College of Agriculture and Mechanic Arts, Las Cruces (as

second Professor of Entomology and Entomologist of the Agricultural Ex¬

periment Station). Professor Cockerell was never employed by the University

of New Mexico. We thank Professor J. G. Watts of New Mexico State

University, Las Cruces, for calling this correction to our attention.—Editor.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 367-368

Published 30 December 1983

SCIENTIFIC NOTE

THE LARVA OF MELOE FRANCISCANUS VAN DYKE AND

MACSWAIN’S MELOE SP. A (COLEOPTERA: MELOIDAE)

In his important study of meloid first instar larvae, MacSwain (1956,

Univ. Calif. Publ. Entomol., 12:1-182) described the highly distinctive larva

of a species of Meloe which he referred to as Meloe sp. a. He noted that

although the adults were unknown, the larvae of this species were the most

commonly encountered Meloe on bees in California. He considered the

possibility that the adults were subterranean and unknown. Considering the

collecting locales of Meloe sp. a, Pinto and Selander (1970, Univ. Ill. Biol.

Monogr., 42:1-222) believed this larva almost certainly belonged to M.

franciscanus Van Dyke, M. californicus Van Dyke, or M. vandykei Pinto

and Selander.

It now appears that MacSwain’s Meloe sp. a is assignable to Meloe fran¬

ciscanus. I collected several adults of this species feeding on Astragalus

lentiginosus Douglas on 2 February 1973 on the sand dunes at the north

end of Panamint Valley in Inyo County, California. The females oviposited

and first instar larvae were obtained.

The first instar larva of M. franciscanus fits MacSwain’s description of

Meloe sp. a in most details. The four long subequal caudal setae, black

coloration, large size (2.0-2.5 mm), longitudinally divided sterna on ab¬

dominal segments I-IV, lack of obvious sclerotization on the thoracic sterna,

and short robust legs are characteristic of Panamint Valley M. franciscanus

and Meloe sp. a. No other known North American Meloe has this combi¬

nation of characteristics. The four elongate caudal setae (subequal to the

abdomen in length) are unique. In other species only two of the caudal setae

are elongate.

MacSwain’s material differs from Panamint Valley specimens in two ways.

He described the terminal seta of the antenna as being slightly shorter than

the antenna itself. In the Panamint material the terminal seta is ca. ‘A longer

than the antenna. Also, MacSwain’s specimens that I have examined (col¬

lected in Berkeley, California) have substantially larger spiracles. For ex¬

ample, the first abdominal spiracle is V 2 as long as tergum I. In the Panamint

material this spiracle is only % the length of tergum I.

Although the possibility that Meloe sp. a represents M. californicus or M.

vandykei cannot be totally discounted, either association is unlikely. First

of all, adults of M. californicus and M. vandykei are much less common in

California than are those of M. franciscanus. More significantly, such an

association would be inconsistent with the current classification of the sub¬

genus Meloe. Minor anatomical differences notwithstanding, the unique fea-

368

PAN-PACIFIC ENTOMOLOGIST

tures of the larva of M. franciscanus from Panamint Valley and of Meloe

sp. a are consistent with the unique traits of M. franciscanus adults. These

adult characteristics led Pinto and Selander (1970) to place M. franciscanus

in its own species group. Meloe californicus and M. vandykei were assigned

to the Angusticollis Group along with the majority of North American Meloe.

The first instar larvae of four species of the Angusticollis Group are known

(Pinto and Selander, 1970). All are relatively similar to one another and, as

a group, differ distinctly from Meloe sp. a and the confirmed larva of M.

franciscanus.

Meloe franciscanus is known from several localities in central and southern

California. It also occurs in northern Arizona, western Utah, and western

Nevada (Pinto and Selander, 1970). In California it appears to be primarily

cismontane, although it has now been taken on three dune systems (Eureka

Valley, Kelso, and Panamint) in the high desert. So far, adults have been

collected in numbers and during daylight hours only on sand dunes. Most

of the other scattered records are based on single specimens. This is un¬

doubtedly the reason MacSwain did not consider M. franciscanus as the

identity of the larva of Meloe sp. a.

The reason for the small number of collections of adults of M. franciscanus

is unknown. However, I suspect that it is due to nocturnal behavior of the

adults of cismontane populations, as well as to the adults occurring primarily

during winter when collectors are less likely to find them. Although data are

lacking for M. franciscanus in areas other than desert dunes, the adults of

two other cismontane California species, Meloe strigulosus Mannerheim and

M. occultus Pinto and Selander, are active at night and in winter. I have

observed numerous M. strigulosus adults near La Grange, California, in

Stanislaus County in early January feeding on grass as late as midnight when

air temperatures adjacent to the beetles were as low as 0°C. During the day

they were inactive and found under logs and boards (Pinto, unpublished).

Similarly, I have taken a few specimens of M. occultus at night in January

in Menifee Valley in SW Riverside County. Mr. P. A. Sullivan (Eaton Canyon

Nature Center, Pasadena, California; pers. comm.) has collected this species

near Devil’s Punchbowl County Park, on the north side of the San Gabriel

Mountains in Los Angeles County at night in January and, in this case, with

snow on the ground. Three other uncommon species of California Meloe

(M. californicus, M. vandykei, and M. barbarus) may also be nocturnal.

John D. Pinto, Department of Entomology, University of California, Riv¬

erside 92521.

PAN-PACIFIC ENTOMOLOGIST

October 1982, Vol. 58, No. 4, pp. 369-377

Published 30 December 1983

PACIFIC COAST ENTOMOLOGICAL SOCIETY

D. H. Kavanaugh

President

J. E. Hafemik, Jr.

President-elect

R. E. Somerby

Secretary

P. H. Amaud, Jr.

Treasurer

PROCEEDINGS

FOUR HUNDRED AND EIGHTH MEETING

The 408th meeting was held on Friday, 22 May 1981, at 8:00 P.M., in the Morrison Audi¬

torium, California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr.

David H. Kavanaugh presiding. A total of twenty-nine persons was present, of which 14 signed

as regular members and 14 as guests.

Minutes of the meeting held on Friday, 24 April 1981, were summarized. Dr. Wojciech J.

Pulawski was elected as a regular member of the society.

Dr. Edward L. Smith gave a note on fossil insect wing venation. The development of venation

patterns needs re-evaluation due to the complex manner of development.

The Executive Board approved the following dues and subscription increases to begin on 1

January 1982: Regular membership—$15.00; Subscriptions—$20.00; Student members—$7.50;

Retired members—$7.50. The Executive Board also approved an increase in page charges of

$36.00 to non-members who publish in the Pan-Pacific Entomologist. If, however, an author

decides to become a PCES member, he may do so and then be charged the $24.00 page charge.

Dr. Kavanaugh introduced Dr. Arthur M. Shapiro, University of California at Davis, who

presented “Butterfly Faunas of the Trinity Alps and Mount Eddy and Their Relationships.”

The talk was illustrated, and Dr. Shapiro discussed the local geology (serpentines, granites) and

flora as they related to the butterfly fauna.

A question and answer period followed at. which time the following comments entitled

Precinctive is more precise zoogeographic term than etymologically ambiguous “endemic” were

contributed by Dr. C. B. Philip: Without implying any criticism of the excellent address we

have just heard by Professor Shapiro on butterfly communities in the Trinity Alps of northern

California, I should like to take this pertinent opportunity to again call attention of students

and others to the increasingly ambiguous use of the term “endemic” by zoo- (and phyto-)

geographers whereas it has been a standard, longstanding term used in a different sense by

epidemiologists and other medical disciplines in the context of the correlative terms, endemic

and epidemic as adjectives in studies of infectious, mostly human diseases, e.g., endemic typhus

and epidemic typhus, two long recognized, specific human diseases without geographic restric¬

tion (other than related to arthropod vectors). The equivalent accepted terms which refer to

diseases of lower animals are enzootic and epizootic as, I presume, most of you know. In

contrast, if we speak of an endemic insect, what then is an epidemic one? Etymologically, the

derivation is wrong as now used by zoogeographers. According to lexicons, the stem is trans¬

literated from the Greek “demas, bodily frame, usually of man, Homer, rarely of other animals,”

or “demos, population as referred to people.” The prefixes refer to “in” or “on.” But there is

even less reason for biologists to perpetuate use of the ambiguous term endemic, when the

precise terms, precinctive and/or indigenous, or sometimes native, are available where they

apply. I hope younger biologists can be weaned away from the use of the ambiguous term of

endemic, and they should not be subtly disdained by using this more perceptive scholarly

terminology. When I discussed this subject in the zoogeographic section of the Montreal En¬

tomological Congress in 1956, the vigorous response by the listeners that medical people had

usurped the term is not substantiated etymologically, as discussed above.

Refreshments were served in the entomology conference room following the meeting.—

R. E. Somerby, Secretary.

370

PAN-PACIFIC ENTOMOLOGIST

FOUR HUNDRED AND NINTH MEETING

The 409th meeting was held on Friday, 16 October 1981, at 8:15 P.M., in the Morrison

Auditorium, California Academy of Sciences, Golden Gate Park, San Francisco, with President

Dr. David H. Kavanaugh presiding. Approximately thirty-five persons were present, of which

24 signed as regular members and 7 as guests.

Minutes of the meeting held on Friday, 22 May 1981 were summarized.

Six new regular members, S. W. Wilson, D. Ubick, M. Garcia-Vidal, C. Parisek, A. M. Pollen,

and W. E. Steiner, Jr., and five student members, D. A. Polhemus, D. Nakamoto, S. Stein, M.

A. Ivie, and D. M. Gordon, were elected as society members.

Dr. Edward L. Smith gave a note on fossil records of lithobiid centipedes and Pedipalpida

for the mid-Devonian. Dr. C. B. Philip called to our attention the usefulness of the 1943 Post

Office Guide for finding old rare localities.

Dr. Kavanaugh introduced Dr. Richard W. Rust, who presented “Great Basin Sand Dune

Arthropods.” He gave an interesting overview of the arthropods taken on the dunes.

Refreshments were served in the entomology conference room following the meeting.—

R. E. Somerby, Secretary.

FOUR HUNDRED AND TENTH MEETING

The 410th meeting was held on Friday, 20 November 1981, at 8:00 P.M., in the Morrison

Auditorium, California Academy of Sciences, Golden Gate Park, San Francisco, with President

Dr. David H. Kavanaugh presiding. Approximately fifty-four persons were present, of which

32 signed as regular members and 22 as guests. Minutes of the meeting held on Friday, 16

October 1981 were summarized.

The following note was presented:

Comparative flower constancy o/Steniolia tibialis Handlirsch and Bombus vosnesenskii Ra-

doszkowski.— Many of the published studies on hymenopteran flower visitation are concerned

with the bees, especially Apis and Bombus, or with specialized wasps such as the Agaonidae

which are the only pollinators of the genus Ficus. Few studies of the flower visitation patterns

of other wasps exist. The general dogma among pollination biologists is that wasps, particularly

sphecid wasps, are casual flower visitors of little importance in pollination of flowers. Bum¬

blebees are, on the other hand, considered to be highly competitive, efficient pollinators.

To test these ideas comparative observations on flower visitation patterns of bumblebees

(.Bombus vosnesenskii Radoszkowski) and sphecid wasps ( Steniolia tibialis Handlirsch) were

made in the north central Sierra Nevada from July 6 to August 17, 1981. Steniolia spp. all

have an unusually long proboscis, ranging from 10 to 19 mm. The adults of these wasps are

only known to feed on floral nectar, while bumblebees feed on nectar and pollen. Results

indicate that B. vosnesenskii and S. tibialis adults are similar in their high frequency of visits

to a mint Monardella odoratissima Benth. in areas where other more abundant sympatric plants

flower simultaneously, but dissimilar in that Bombus chooses a much wider variety of flowers

(10 spp.) than does Steniolia (4 spp.).

This study suggests that the general belief that wasps are casual flower visitors requires

reexamination.— Timothy L. Tyler, Department of Biological Sciences, San Francisco State

University, San Francisco, California 94132.

Dr. Edward L. Smith gave a review of a note he gave at the 409th meeting relative to mid-

Devonian arthropods including insects. He also brought to our attention two books, a new atlas

and a book on insect morphology, published by Haden and Sons, Philadelphia.

Dr. Kavanaugh read a letter from Hugh B. Leech to the Pacific Coast Entomological Society,

thanking the Society for honoring him in the first issue of the 57th volume of the Pan-Pacific

Entomologist.

Dr. Kavanaugh introduced Dr. Lawrence W. Swan, Professor, Department of Biology, San

VOLUME 58, NUMBER 4

371

Francisco State University, who presented “Tibet and High Altitude Life.” His presentation

was well illustrated with slides and anecdotal information.

Refreshments were served in the entomology conference room following the meeting.—

R. E. Somerby, Secretary.

FOUR HUNDRED AND ELEVENTH MEETING

The 411th meeting was held on Friday, 18 December 1981, at 8:00 P.M., in the Morrison

Auditorium, California Academy of Sciences, with President Dr. David H. Kavanaugh intro¬

ducing our new president. Dr. John E. Hafernik, Jr. The following names were brought forth

from the nominating committee for office in 1982: President-elect Edward L. Smith, Treasurer

Wojciech J. Pulawski, and Secretary Vincent F. Lee. The slate of candidates was duly elected

to office.

Approximately 51 persons were present, of which 33 signed as members and 18 as guests.

Minutes of the meeting held 20 November 1981 were summarized. The following reports were

given. New members presented and elected were Roger W. Daggs, Karl A. Giljum, Richard E.

Hill,.and William B. Warner as student members, and Alec M. Balmy, Herb C. Field, Magdy

M. A. Sayed, and Richard J. Skalski as regular members.

Dr. Paul H. Amaud, Jr., Treasurer, reported that there was a total income of $18,708

(including $8,162 from dues and subscriptions). The income from dues and subscriptions was

$16 less than that received in 1980. The statement of income, expenditures and changes in

fund balances will be published and available to all members through its publication in the

Proceedings of the Society.

The treasurer’s office is indebted to Mrs. Vashti Hawley Getten (as volunteer) and Mrs. Gail

Freihofer (Entomology Secretary) for their handling of the Society’s accounts, billings, and

mailing of publications, and to our member Mr. H. Vannoy Davis of Walnut Creek, California,

for his annual audit of the Treasurer’s records and completion of the Society’s tax forms.

Helen K. Court gave the auditing committee report, and Dr. Kavanaugh gave the historical

committee report. As a point of interest, he also read a letter from Horn to Rivers in reference

to Casey. The publication committee’s two new replacement nominees were Dr. Arnaud and

Dr. Marius S. Wasbauer.

The first note was given by Dr. Edward S. Ross, who gave a brief overview of his six months

in North Africa studying eucerine bees. Next, David M. Katz presented slides on some select

local insects. This was followed by Dr. Edward L. Smith who presented information on a new

class of Crustacea, Remipedia, and mentioned the report of terrestrial mayfly nymphs 10 cm

long. As a final report, Dr. Cornelius B. Philip presented a motion-picture film on an unusual

number of mutillid wasps that occurred near the Rocky Mountain Laboratory.

Dr. Hafernik introduced Dr. Kavanaugh who presented a talk entitled “Entomological Ex¬

ploration of the Queen Charlotte Islands, British Columbia.” The talk was well presented with

slides and interesting anecdotes; a fascinating report on summer held work in 1981.

Refreshments were served in the entomology conference room following the meeting.—

R. E. Somerby, Secretary.

PACIFIC COAST ENTOMOLOGICAL SOCIETY

STATEMENT OF INCOME, EXPENDITURES AND

CHANGES IN FUND BALANCES

Years Ended September 30, 1981 and 1980

1981

Income

Dues and subscriptions . $ 8,162

Reprints and miscellaneous . 5,611

Sales of Memoirs .

1980

$ 8,178

10,646

191

372

PAN-PACIFIC ENTOMOLOGIST

Interest . 3,909 2,484

Dividends, American Telephone & Telegraph Company. 416 492

Increase (decrease) in value of capital stock

of American Telephone & Telegraph Company. 610 (250 )

$18,708 $21,741

Expenditures

Publication costs—Pan-Pacific Entomologist . $17,925 $14,293

Reprints, postage and miscellaneous. 4,049 3,545

$21,974 $17,838

Increase (decrease) in fund balances . (3,266) 3,903

Fund balances October 1, 1980 and 1979 . 38,219 34,316

Fund balances September 30, 1981 and 1980 . $34,953 $38,219

STATEMENT OF ASSETS, September 30, 1981 and 1980

1981 1980

Cash in bank

Commercial account. $ 3,345 $ 1,548

Savings accounts & certificates of deposit

General fund . 4,777 13,304

Memoir—Fall fund. 17,617 15,431

Life membership fund . 4,454 3,786

Total cash in bank . $30,193 $34,069

Investment in 80 shares of American Telephone

& Telegraph Co. common stock (life member¬

ship and Fall funds), at market value. 4,760 4,150

$34,953 $38,219

Notes to the Financial Statements Year Ended September 30, 1981

Summary of significant accounting policies.

Accounting Method: Income and expenses are recorded by using the cash basis of accounting.

Marketable Securities: American Telephone & Telegraph Co. common stock is carried at market

value. Increases and decreases in value are reflected in income. Income Tax: The Society is

exempt from Federal income and California franchise tax. Accounts Receivable: As of September

30, 1981 accounts receivable aggregated $3,229 as follows:

September, 1981 billings. $3,049

Prior billings . 180

$3,229

Accounts Payable: As of September 30, 1981 there were no unpaid bills.

As Chairman of the Auditing Committee, and in accordance with its bylaws, I have reviewed

the financial records of the Society. During the course of this review nothing was noted which

indicated any material inaccuracy in the foregoing statements.

/s/ H. Vannoy Davis

H. Vannoy Davis

Chairman of the Auditing Committee

VOLUME 58, NUMBER 4

373

PACIFIC COAST ENTOMOLOGICAL SOCIETY

J. E. Hafemik, Jr. E. L. Smith V. F. Lee W. J. Pulawski

President President-elect Secretary Treasurer

PROCEEDINGS

FOUR HUNDRED AND TWELFTH MEETING

The 412th meeting was held Friday, 22 January 1982, 8:00 p.m., in the Morrison Auditorium,

California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. John E.

Hafernik, Jr. presiding.

The minutes of the meeting held 18 December 1981 were read, corrected, and accepted.

Three new members were elected to the Society: Mr. Phillip L. Anderson as regular member,

and Ms. Linda A. Wilcox and Mr. Keve Ribardo as student members.

Dr. Edward L. Smith announced three recent publications— Fresh-water invertebrates of the

United States, second edition, by R. W. Pennak; Insect phylogeny, by W. Hennig (English

translation by A. C. Pont); and “ Triarthrus eatoni (Trilobita): Anatomy of its exoskeletal,

skeletomuscular, and digestive systems,” by J. L. Cisne, in Palaeontographica Americana, 9(53).

Mr. Jerry Scribner, Director of the Medfly Project, lectured on “The Medfly Problem: Pros¬

pects for the Future.” The slide-illustrated lecture gave an account of the history of medfly

introductions and control of the medfly in the United States, with particular emphasis on

California’s attempt to eradicate the pest in 1980 through the present.

The social hour was held in the entomology conference room following the meeting.

A total of 47 persons was present, of which 29 signed as members and 15 signed as guests.—

V. F. Lee, Secretary.

FOUR HUNDRED AND THIRTEENTH MEETING

The 413th meeting was held Friday, 19 February 1982, at 8:05 p.m., in the Morrison Au¬

ditorium, California Academy of Sciences, Golden Gate Park, San Francisco, with President

Dr. John E. Hafernik, Jr. presiding.

The minutes of the meeting held 22 January 1982 were read and accepted. Two new regular

members were elected to the Society: Mr. Norman L. Markowitz and Dr. Patricia A. Malone.

Dr. Cornelius B. Philip presented a note with a photographic slide of a horse fly Tabanus

vittiger Thomson that takes its blood meal from the marine iguana of the Galapagos Islands.

Mr. Evan A. Sugden presented slides illustrating the mating and burrowing behavior of the rain

beetle Pleocoma rubiginosa transsierrae Hovore. Dr. Edward L. Smith reported that Dr. Jarmila

Kukalova-Peck wrote to him about the discovery of a remarkable fossil immature mayfly from

the Upper Carboniferous of Bohemia. The nymph and associated adult possess unusual struc¬

tures and occur in habitats quite different from recent mayflies.

Dr. Edward S. Ross, Curator Emeritus of the Department of Entomology, California Academy

of Sciences, presented “An Entomologist’s Tour of North Africa and Southern Europe.” The

well-illustrated slide lecture recounted in travelogue style a six-month trip to this area. Em-

bioptera collecting, the Ophrys orchids which visually and chemically mimic several insects,

and the site of the Cro-Magnon race were highlighted.

The social hour was held in the Goethe Room following adjournment of the meeting.

A total of 86 persons was present, of which 33 signed as members and 30 as guests. —V. F.

Lee, Secretary.

FOUR HUNDRED AND FOURTEENTH MEETING

The 414th meeting was held Friday, 19 March 1982, at 8:05 p.m., in the Morrison Audi¬

torium, California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr.

John E. Hafemik, Jr. presiding.

374

PAN-PACIFIC ENTOMOLOGIST

The minutes of the meeting held 19 February 1982 were read and accepted. Three new

members were elected to the Society: Mr. Wesley A. Malfei as student member, and Mr. Russ

B. Parman and Mr. Michael A. Wolf as regular members.

Dr. Edward L. Smith presented his new interpretations of arachnid segmentation and tag-

mosis.

Dr. Ward Watt, Stanford University, lectured on “Energy Balance and Adaptation in But¬

terflies.” His presentation showed the results of his research on thermoregulation of Colias in

the field, tied in with laboratory studies of life history and biochemical genetics.

The social hour was held in the entomology conference room following adjournment of the

meeting.

A total of 43 persons was present, of which 30 signed as members and 13 signed as guests.—

V. F. Lee, Secretary.

FOUR HUNDRED AND FIFTEENTH MEETING

The 415th meeting was held Friday, 16 April 1982, at 8:10 p.m., in the Morrison Auditorium,

California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. John E.

Hafemik, Jr. presiding.

Several guests were introduced, including Dr. Charles A. Triplehom of Ohio State University

and Dr. Robert S. Lane of University of California, Berkeley.

The minutes of the meeting held 19 March 1982 were read and accepted. Four new members

were elected to the Society: Mr. Herbert H. Daubner, Dr. William P. MacKay, and Dr. John

D. Stein as regular members, and Mr. Larry Haimowitz as student member.

Mr. Charles E. Griswold, University of California, Berkeley, lectured on “Enemies and

Defenses of Spiders, with Examples from a Neotropical Rain Forest.” He presented a review

of parasitoids and predators of spiders with some examples from La Selva Biological Station

in Costa Rica where he did field work last year with Dr. Rollin E. Coville. They collaborated

in studying Trypoxylon wasps and the wasps’ provisioned spiders.

The social hour was held in the entomology conference room following adjournment of the

meeting.

A total of 70 persons was present, of which 30 signed as members and 24 as guests. —V. F.

Lee, Secretary.

FOUR HUNDRED AND SIXTEENTH MEETING

The 416th meeting was held Friday, 21 May 1982, at 8:10 p.m., in the Morrison Auditorium,

California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. John E.

Hafemik, Jr. presiding.

The minutes of the meeting held 16 April 1982 were read and accepted. Two new student

members were elected to the Society: Ms. Mary Ann Tenorio and Mr. Martin Galindo-Ramirez.

Dr. Paul H. Amaud, Jr. introduced Mrs. Zun-ming Wang of Academia Sinica, Beijing, and

Dr. Kenneth S. Hagen introduced Mr. Yu-hua Yan of the Beijing Agricultural University.

President Hafemik announced the death of Dr. J. Linsley Gressitt, a life member of the Society.

Dr. Edward L. Smith noted the deaths of Dr. Paul D. Hurd, Jr., also a member, and Dr. Robert

E. Silberglied.

Ms. Linda A. Wilcox presented a note on an analysis of the fecal shield of the larva of Lema

nigrovittata Guerin-Meneville which proved to have alkaloids similar to its host plant Datura

meteloides A. DC. Dr. Smith discussed additional interpretations of arachnid segmentation.

Mr. Larry G. Bezark announced the recent publication by the California Department of Food

and Agriculture of color photo keys to California whiteflies and scale insects.

The featured speaker Dr. Herbert Baker, University of California, Berkeley, presented “Nectar

and Pollen Rewards to Flower Visitors.” His slide-illustrated lecture summarized the current

knowledge of the biochemical aspects of nectar and pollen and their relationships to insect

flower visitors.

VOLUME 58, NUMBER 4

375

The social hour was held in the entomology conference room following adjournment of the

meeting.

A total of 61 persons was present, of which 32 signed as members and 25 as guests.—V. F.

Lee, Secretary.

FOUR HUNDRED AND SEVENTEENTH MEETING

The 417th meeting was held Friday, 22 October 1982, at 8:10 p.m., in the Morrison Audi¬

torium, California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr.

John E. Hafemik, Jr. presiding.

The minutes of the meeting held 21 May 1982 were read and accepted. Seventeen persons

were elected as members of the Society: Mr. Paul E. Blom, Mr. Clinton L. Campbell, Mr. Jeffrey

A. Halstead, Mr. Paul Hanson, Mr. Dennis M. Kubly, Mr. Matthew A. Owen, and Mr. John

A. Skinner as student members; and Dr. Richard A. Arnold, Dr. Alan B. Bond, Mr. Stanley

E. Brecht, Mr. Daniel Heffem, Ms. R. Deedee Kathman, Mr. Timothy N. King, Dr. Joseph G.

Morse, Mr. Wayne J. Steele, Mr. Ildy Szabo, and Dr. Robert H. Turnbow, Jr. as regular members.

Dr. Edward L. Smith announced the deaths of Dr. Charles P. Alexander, an honored member

of the Society, and Dr. Clarence E. Mickel. Mr. James E. Tobler I, Dr. Ronald E. Stecker, and

Dr. Robbin W. Thorp introduced some new visitors. Dr. J. Gordon Edwards introduced Major

and Mrs. Alan R. Gillogly, members of the Society, and family, who recently returned from

overseas.

Mr. Tobler announced that specimens from his Bali-Sarawak trip in 1969-1970 will be on

display during the social hour. Dr. Harvey I. Scudder reported the discovery of the largest

North American thrips from Miocene fossil beds in Nevada. Dr. Smith noted his involvement

with the Pomona insect collection when he was a graduate student there. A portion of the

collection was recently transferred to the Department of Entomology of the California Academy

of Sciences. He also reported on the recently proposed geological period, the Ediacarian Period.

The featured speaker, Dr. Richard A. Arnold, University of California, Berkeley, presented

“Ecological Studies and Current Status of Six Species of Endangered Butterflies.” His slide-

illustrated lecture reported on the endangered butterflies of California, with emphasis on his

research of the Lange’s Metalmark at the Antioch dunes.

The social hour was held in the entomology conference room following adjournment of the

meeting.

A total of 61 persons was present, of which 38 signed as members and 13 as guests.—V. F.

Lee, Secretary.

FOUR HUNDRED AND EIGHTEENTH MEETING

The 418th meeting was held Friday, 19 November 1982, at 8:15 p.m., in the Morrison

Auditorium, California Academy of Sciences, Golden Gate Park, San Francisco, with President

Dr. John E. Hafemik, Jr. presiding.

The minutes of the meeting held 22 October 1982 were read, corrected, and accepted. Four

persons were elected as members of the Society: Ms. Beverly A. Friesen, Mr. Daniel F. Gross,

and Ms. Laurie M. Swiadon as student members, and Dr. Sara S. Rosenthal as regular member.

President Hafemik announced the formation of two committees: the auditing committee,

consisting of Mr. H. Vannoy Davis and Dr. Paul H. Amaud, Jr., and the nominating committee,

consisting of Dr. Stanley C. Williams (chairperson), Dr. Jerry A. Powell, and Dr. David H.

Kavanaugh. Dr. Williams presented a preview of the slate of officers for 1983: Dr. Edward L.

Smith as president, Dr. Harvey I. Scudder as president-elect, Dr. Wojciech J. Pulawski as

treasurer, and Mr. Vincent F. Lee as secretary. Dr. Smith asked for volunteers for the 1983

program committee.

Dr. Smith noted an article in a recent issue of Science in which researchers at Stonybrook

376

PAN-PACIFIC ENTOMOLOGIST

demonstrated the presence of hematite below the stomach of honeybees. The hematite appar¬

ently acts as an “autocompass.” Dr. Williams added that Dr. Jack Tomlinson will be talking

about this subject to the San Francisco Bee Club in February 1983.

The featured speaker Dr. Rollin E. Coville, University of California, Berkeley, presented

“Natural History of Costa Rican Anthophorid Bees in the Genus Centris .” His slide-illustrated

lecture reported on the pollination activity, nesting habit, territorial behavior of males, and

mating behavior of these bees in Guanacaste Province.

The social hour was held in the entomology conference room following adjournment of the

meeting.

A total of 53 persons was present, of which 39 signed as members and 10 as guests.—V. F.

Lee, Secretary.

FOUR HUNDRED AND NINETEENTH MEETING

The 419th meeting was held Friday, 17 December 1982, at 8:10 p.m., in the Morrison

Auditorium, California Academy of Sciences, Golden Gate Park, San Francisco, with President

Dr. John E. Hafemik, Jr. presiding.

The minutes of the meeting held 19 November 1982 were read, corrected, and accepted.

Four persons were elected as members of the Society: Ms. Marciana J. Rinaldi and Ms. Teena

K. Stader as student members, and Mr. Gary A. Freed and Ms. Leslie Saul as regular members.

Dr. Paul H. Amaud, Jr. gave a report from the auditing committee. He also reported for the

historical committee on the receipt of historical materials from Mr. Hugh B. Leech and Dr.

Cornelius B. Philip and of scientific notes from the late Mr. Philip H. Timberlake, articles on

Timberlake from Dr. E. Gorton Linsley, and the loaning of an original illustration from Kellogg’s

American Insects to the Oakland Museum. Dr. Amaud read the report of the Treasurer, who

was ill. Special mention was made of the indebtedness of the Society to Mrs. Vashti Getten, a

volunteer for 39 years, who maintained the Society’s accounts. She retired this past summer,

and Mrs. Roberta L. Brett kindly assumed her post. The Society also is grateful to Mr. H.

Vannoy Davis, who reviewed the Treasurer’s records and completed the annual federal and

state income tax forms. Dr. Stanley C. Williams, chairperson of the nominating committee,

presented the slate of candidates for officers of the Society for 1983: Dr. Edward L. Smith as

president, Dr. Harvey I. Scudder as president-elect, Dr. Wojciech J. Pulawski as treasurer, and

Mr. Vincent F. Lee as secretary. The members who were present unanimously elected these

persons to office. Dr. Hafemik thanked a number of persons who helped the Society during his

tenure as president.

Dr. Smith, president for 1983, announced that the program committee consists of Dr. G.

Andrew H. McClelland and Dr. Marius S. Wasbauer.

Dr. Hafemik presented a note on the presence of an aggregation of the monarch butterflies

in Golden Gate Park at 41st Avenue and South Drive. Dr. Smith noted the longevity of

entomologists and especially that of the late Dr. Charles P. Alexander. Dr. Alexander had a

remarkably large library, and he was the most prolific taxonomist—he described over 10,800

crane flies. Dr. Smith also announced the recent publication of Rainer F. Foelix’s Biology of

Spiders, published by Harvard University Press.

The featured speaker, Dr. John E. Hafemik, Jr., San Francisco State University, presented

the presidential address “Population Structure and Distribution of the Rare Damselfly Ischnura

gemina, a Bay Area Endemic.” His slide-illustrated lecture reported on the habitats, population

structure, and natural history of the most restricted Odonata of western North America.

The social hour was held in the entomology conference room following adjournment of the

meeting.

A total of 40 persons was present, of which 31 signed as members and 7 as guests.— V. F.

Lee, Secretary.

VOLUME 58, NUMBER 4

377

PACIFIC COAST ENTOMOLOGICAL SOCIETY

STATEMENT OF INCOME, EXPENDITURES AND

CHANGES IN FUND BALANCES

Years Ended September 30, 1982 and 1981

1982 1981

Income

Dues and subscriptions . $10,078 $ 8,162

Reprints and miscellaneous . 6,046 5,482

Sales of Memoirs . 256 129

Interest . 5,727 3,909

Dividends, American Telephone & Telegraph Company. 432 416

Increase (decrease) in value of capital stock

of American Telephone & Telegraph Company. (150 ) 610

$22,389 $18,708

Expenditures

Publication costs—Pan-Pacific Entomologist . $ 6,418 $17,925

Reprints, postage and miscellaneous. 5,039 4,049

$11,457 $21,974

Increase (decrease) in fund balances . 10,932 (3,266)

Fund balances October 1, 1982 and 1981 . 34,953 38,219

Fund balances September 30, 1982 and 1981 . $45,885 $34,953

STATEMENT OF ASSETS, September 30, 1982 and 1981

1982 1981

Cash in bank

Commercial account. $12,023 $ 3,345

Savings accounts & certificates of deposit

General fund . 5,049 4,777

Memoir—Fall fund . 19,229 17,617

Life membership fund . 4,860 4,454

Total cash in bank . 41,161 30,193

Undeposited receipts . 114 0

Investment in 80 shares of American Telephone

& Telegraph Co. common stock (life member¬

ship and Fall funds), at market value. 4,610 4,760

$45,885 $34,953

See accompanying notes to the financial statements.

Notes to the Financial Statements Year Ended September 30, 1982

Summary of significant accounting policies.

Accounting Method: Income and expenses are recorded by using the cash basis of accounting.

Marketable Securities: American Telephone & Telegraph Co. common stock is carried at market

value. Increases and decreases in value are reilected in income. Income Tax: The Society is

exempt from Federal income and California franchise tax. Accounts Receivable: As of September

378

PAN-PACIFIC ENTOMOLOGIST

30, 1982, accounts receivable aggregated $4,091. Accounts Payable: As of September 30, 1982,

there were no material amount of unpaid bills.

As Chairman of the Auditing Committee, and in accordance with its bylaws, I have reviewed

the financial records of the Society. During the course of this review nothing was noted which

indicated any material inaccuracy in the foregoing statements.

/s/ H. Vannoy Davis

H. Vannoy Davis

Chairman of the Auditing Committee

HONORED MEMBERS OF THE

PACIFIC COAST ENTOMOLOGICAL SOCIETY

Melville H. Hatch

Hugh B. Leech

E. Gorton Linsley

Abraham E. Michelbacher

Robert C. Miller

THE PAN-PACIFIC ENTOMOLOGIST

INDEX TO VOLUME 58

(New taxa indicated by boldface type)

Acanthoscelides fraterculus, 38

Acari: Argasidae, 96; Bdellidae, 216; Cera-

neidae, 180; Ixodidae, 96; Pygmephoridae,

216; Trombidiidae, 180

Acrididae, 130

Acromyrmex landolti, 258

Actia sp., 42

Adelonia costipennis, 87; filiformis, 87; insu-

laris, 85; quadricollis, 87; sulcatula, 87

Adistemia watsoni, 79

Aedes aegypti, 129

Aegialia blanchardi, 319; conferta, 319; cras-

sa, 319; crescenta, 319; hardyi, 319; lacus-

tris, 319; latispina, 319; magnified, 319;

punctata, 319

Aegialites fuchsi, 163 \ subopacus, 163

Ageniella arcuata, 224; fuscipennis, 224

Ageniella ( Priophanes) arizonica concolor, 224

Agenioideus ( Agenioideus ) humilis, 225

Agroeca sp., 223

Agromyza ( Liriomyza) fiaveola, 303; pusilla,

304; subpusilla, 304

Aleurocanthus woglumi, 312

Alopecosa gertschi, 227; kochi, 227

Amauris, 281

Amaurobius ferox, 216

Ambrysus drakei\ plautus, 326

Amiocentrus aspilus, 212

Amobia floridensis, 290

Amphicerus cornutus, 180

Ampulex, 129

A nap he moloneyi, 181

Ancistrocerus waldenii flavidulus, 259

Andrena kristina, 309; melanochroa, 309; pi-

peri, 309; primulifrons, 311; trapezoidea, 311

Andrena ( Callandrena ) accepta, 111, 287;

haynesi, 284; helianthi, 111, 286

Anoplius ( Anoplius ) dreisbachi, 227; imbellis,

227; ithaca, 227; toluca, 227

Anoplius ( Pompilinus ) bequaerti, 226; inso¬

le ns, 226; marginatus complex, 226; splen-

dens, 226; townesi 226

Anoplosiagum cubensis, 330; flavicolle, 330;

oteroi, 332; pallidum, 330; peltatum, 330;

rufum, 330; rutilus, 330; scabrosum, 330;

scaramuzzai, 330; simplicipes, 330; swe-

zeyi, 333; turquinensis, 334; variabile, 330;

zayasi, 335

Anthidium, 179

Anthophora, 179

Anthrax tigrinus, 180

Anuroctonus phaiodactylus, 164

Anyphaena pacifica, 227

Apatolestes, 365; actites, 365; parkeri, 365

Apiocera clavator, 296; haruspex, 296; ma-

ritima, 296; painteri, 296

Apis mellifera, 178

Aplomyiopsis xylota, 149

Aporinellus completus, 229

Aporus hirsutus, 224; luxus, 224

Aptostichus stanfordianus, 224

Arachnis picta, 180

Araneae: Agelenidae, 228; Antrodiaetidae, 216;

Anyphaenidae, 227, Araneidae, 225, 289;

Clubionidae, 223; Ctenidae, 225; Ctenizi-

dae, 224; Gnaphosidae, 180, 228; Lycosi-

dae, 223; Oxyopidae, 224; Philodromidae,

226; Pisauridae, 225; Salticidae, 224; Ther-

idiidae, 180; Thomisidae, 180, 225

Araneus bispinosus, 225; cornutus, 225

Archips, 259

Arctosa, 227; sp., 226

Argiope trifasciata, 225

Armadillium vulgare, 180

Amaud, aggregation Coelopa, 245; publica¬

tions received, 201, 205, 215, 222, 230

Atrichopogon epicautae, 71; farri, 1 1; fusculus,

64; meloesugans, 71; oedemararum, 71;

pavidus, 71; pollinivorus, 71

Atta sexdens, 258

Attagenus megatoma, 79

Atypoides river si, 2 16

Auplopus, 228; architectus metallicus, 224; ni-

grellus, 224

Axinopalpus biplagiatus, 180

Aysha gracilis, 228

Bakerdania damboldti, 221; longisetus, 221

379

380

PAN-PACIFIC ENTOMOLOGIST

Barnard, biting midges flight periodicity, 64

Bathythrix sp., 42

Battus philenor, 278; polydamus, 280

Bembix brullei, 129; oculata, 129; rostrata, 129

Benbow and Foster, biology Eutreta diana, 19

Bezzia pulverea, 64; setulosa, 64

Blapstinus, 180

Blepharicera capitata, 117; micheneri, 117;

tenuipes, 117

Bombus vosnesenskii, 370

Bombylius androgynus, 237; armeniacus, 237;

cinerarius, 237; quadrifarius, 237; testacei-

ventris, 237; vlasovi, 237

Bombylius ( Zephyrectes ) anthophoroides,

236; cruciatus, 236; incanus, 236; montan-

us, 236; nicholsonae, 236; rams', 236

Brachycentrus americanus, 212; occidentalis,

214

Brachymeria ovata, 42

Brachymyrmex depilis, 258

Brennania, 365; /zcra, 365

Brontes dubius truncatus, 180

Bruchophagus mexicanus, 41; sp. (prob. raex-

icanus ), 38

Byers, a new Neopanorpa, 92

Callandrena, 284

Callosamia promethia, 278

Camponotus sp., 180; spp., 196

Caryobruchus buscki, 244; gleditsiae, 244; v<?-

seyi, 240

Chaetocoelopa sydneyensis, 248

Chaetophora, 36

Charitographa, 166

Chelostoma minutum, 235

Chilocorus stigma, 314

Chilometopon abnorme, 84; brachystomum,

81; castaneum, 84; cribricolle, 85; ensifer,

84; pallidum, 84; rugiceps, 85

Chionodes psiloptera, 77; sp., 259

Chiracanthium sp., 224

Chlorion aerarium, 130

Chlorochroa ligata, 4 1; uhleri, 4 1

Chlorocytus sp., 40

Chrysis coerulans, 260

Chrysopa spp., 314

Chrysoteuchia topiaria, 77

Cinara balachowskyi, 200; piniradicis, 200;

puerca, 200; radicivora, 196; saskensis, 200;

sorini, 200

Clark, Frohlich, and Comanor, shrike preda¬

tion on scorpion and solpugid, 164

Clavipalpus, 330

Clement and Miller, insect seed predation, 38

Clisodon, 179

Cnephasia sp., 259

Coelopa frigida, 248; pilipes, 248

Coelopa ( Neocoelopa) vanduzeei, 245

Coleoptera: Anthicidae, 180; Bostrichidae, 180;

Bruchidae, 38, 240; Buprestidae, 180; Byr-

rhidae, 31; Carabidae, 180; Cerambycidae,

180; Chrysomelidae, 139; Coccinellidae,

314; Cucujidae, 180; Curculionidae, 180;

Dermestidae, 79, 180; Elateridae, 180, 202;

Endomychidae, 180; Hydraenidae, 163;

Lathridiidae, 79; Meloidae, 184, 367; Me-

lyridae, 163; Nosodendridae, 53; Ostomi-

dae, 180; Rhizophagidae, 180; Salpingidae,

163; Scarabaeidae, 159, 319, 330; Staphy-

linidae, 163; Tenebrionidae, 79, 81, 177,

180

Collembola: Entomobryidae, 180

Comperilla, undet. genus near, 23

Condylostylus chrysoprasi, 314

Conoderus amplicollis, 202; exsul, 202; falli,

202

Cotinis mutabilis, 159; nitida, 159; texana, 159

Coville, biology of Ancistrocerus, 259

Cry modes devastator, 7 3

Cryptocheilus severini, 223

Cupiennius salei, 225

Curimopsis, 36

Cybaeus sp., 228

Cycloneda sp., 314

Cyrtacanthacridinae, 130

Danaus, 281; gilippus berenice, 281; gilippus

strigosus, 281; plexippus, 281

Davis, financial statement 1981,371; financial

statement 1982, 377

Dasyhelea grisea, 64; mutabilis, 64

Delphastus pusillus, 314

Denning, new western Trichoptera, 206

Deraeocoris, 352; brevis, 352; californicus, 363;

cerachates, 361; cochise, 358; fraserensis,

352) fulvescens, 363; gilensis, 360; hyalinus,

359; incertus, 353; incertus carneolus, 353;

knightonius, 361; navajo, 359; ornatus, 352;

picipes, 353; piceicola, 359; pilosus, 363;

poecilus, 352; rufusculus, 363; schuhi, 359;

validus, 352; vanduzeei, 360

VOLUME 58, NUMBER 4

381

Diabrotica longicornis barbari, 141; virgifera

virgifera, 141

Dialeurodes citrifolii, 313

Diaulota densissima, 163; vandykei, 163

Dibrachys cavus, 42

Dicerca sp. (prob. horni ), 180

Dioptopsis dismalea, 117; sequoiarum, 117

Diptera: Agromyzidae, 302; Apioceridae, 296;

Asilidae, 250, Blephariceridae, 117; Bom-

byliidae, 180, 236; Coelopidae, 245; Doli-

chopodidae, 314; Sarcophagidae, 289; Ta-

banidae, 365; Tephritidae, 19, 25; Tipulidae,

153

Dolomedes, 225

Dorr and Neff, nesting of Pseudomasaris, 124

Dowell, biology of Tetraleurodes, 312

Doyen, new western Tenebrionidae, 81

Ectemnius, 180

Ection burchelli, 258

Efferia, 298

Ehler, Trialeurodes in garden, 291

Eleodes sp., 164

Embolostoma, 166

Encarsia sp., 316

Endeodes collaris, 163

Ephialtes Ontario, 42

Episyron quiquenotatus hurdi, 225

Eretmocerus IportoricensLs, 314

Eschatomoxys, 90

Eucallipterus tiliae, 63

Eupelmus, 23

Euphoria, 159

Eupompha decolorata, 184; edmundsi, 184;

elegans, 184; imperialis, 184; viridus, 184;

vizcaina, 184

Eurytoma sp., 23

Eutreta diana, 19

Evenhuis, notes on Bombylius ( Zephyrectes ),

236

Farula, 210; davisi, 210; honeyi, 210; jewetti,

210; malkini, 210; petersoni, 210; rainieri,

210; reapiri, 210; wigginsi, 210

Financial statement, 371, 377

Forcipomyia bipunctata, 64; brevipennis, 64;

inornatipennis, 71

Formica aquilonia, 258; montana, 258; opa-

civentris, 258; rufa, 258; subintegra, 258

Garcia-Vidal, new Cuban Anoplosiagum, 330

Garrison, new Mexican dragonfly, 135

Gastrancistrus sp., 23

Gelis tenellus, 42

Geolycosa raphaelana, 229

Gerling, biology Hawaiian Xylocopa, 336

Giuliani, intertidal beetles California, 163

Glaucopsyche lygdamus, 41

Gnaphosa muscorum, 228

Gryllotalpa hexadactyla, 130

Habronattus, 226

Haemaphysalis leporispalustris, 98

Hafemik, mimetic selection Papilio, 278

Halticoptera sp., 40

Harpyllus sp., 180

Hartman and Hynes, immature stages Tipula,

153

Hemiptera: Miridae, 180, 352; Naucoridae,

326

Heppner, revision Thaumatographa, 165

Heteroptera: Miridae, 352

Hilarographa, 165

Homoptera: Aleyrodidae, 291, 312; Aphidi-

dae, 196; Delphacidae, 105

Honored members, 378

Hurd, Linsley, and Hall, In Memoriam — Phil¬

ip Hunter Timberlake, 2; correction, 366

Hybomitra californica, 365

Hydropsyche andersoni, 206; dorata, 208; tana,

208

Hylemya anane, 39

Hymenoptera: Andrenidae, 111, 284; Antho-

phoridae, 336; Aphelinidae, 316; Apidae,

180; Apoidea, 309; Chrysididae, 180; Dry-

inidae, 107; Encyrtidae, 23; Eulophidae, 23;

Eumenidae, 259; Eupelmidae, 23; Eurytom-

idae, 23, 38; Formicidae, 180, 258; Masar-

idae, 124; Megachilidae, 180, 231; Pompili-

dae, 109, 223; Pteromalidae, 23, 40;

Signiphoridae, 316; Sphecidae, 129, 180,288

Idiothauma, 166

Incisitermes minor, 180

In Memoriam: Paul David Hurd, Jr., 262;

Philip Hunter Timberlake, 2

Irianassa, 166

Iridomyrmex nitidus, 182

Ischnura gemina, 376

Isopoda: Armadillididae, 180; Oniscidae, 180

Isoptera: Kalotermitidae, 180; Hodotermiti-

dae, 180

382

PAN-PACIFIC ENTOMOLOGIST

Itoplectis behrensii, 42

Ixodes pacificus, 96; peromysci, 96; rugosus,

98; signatus, 96

Johnson, new genus Byrrhidae, 31

Kamm, Protagrotis a grass pest, 73

Kimsey, female Pompilus wasbaueri, 109

Kimsey and Backus, biology Dioptopsis se-

quoiarum, 117

Krombein and Linsley, In Memoriam—Paul

David Hurd, Jr., 262

Kulhavy, Schwandt, and Hobbs, indicators of

host trees of wounded tree beetle, 53

Lane, Miller, and Collins, ticks from Califor¬

nia Channel Islands, 96

Lanham, new Colorado Andrena, 309

Larra analis, 130

Lebia cyanipennis, 149

Lee, Proceedings 412th-419th meetings, 373

Lema nigrovittata, 374

Lepidoptera: Arctiidae, 180; Dioptidae, 42;

Gelechiidae, 77; Hesperiidae, 260; Noctui-

dae, 73; Papilionidae, 278; Pieridae, 162;

Pyralidae, 77; Tortricidae, 165

Lepidostoma roemhildi, 214; spicata, 214

Limenitis archippus, 281; arthemis arthemis,

279; arthemis astyanax, 278

Liometopum occidentale, 180

Liparocephalus cordicollis, 163

Liriomyza brassicae, 302; dianthi, 303; fla-

veola, 303; huidobrensis, 302; langei, 303;

munda, 304; orbona, 303; pictella, 304; pu-

silla, 304; sativae, 302; subpusilla, 304; tn-

foliearum, 302; trifolii, 302

Liris, 130

Lissonota clypeator montana, 77

Lycosa coloradensis, 223; frondicola, 229; gu-

losa, 228; helluo, 226; sp., 227, 228

Maevia vittata, 226

Marpissa, 226

Mastrus aciculatus, 42

Mecoptera: Panorpidae, 92

Megachile, 181; pacifica, 178; rotundata, 182

Meloe barbarus, 368; californicus, ?>61\fran-

ciscanus, 367; occultus, 368; sp. a, 367; stri-

gulosus, 368; vandykei, 367

Mesidiopsis sp., 62

Metaphidippus aeneolus, 224

Metepeira arizonica, 225; grandiosa, 289

Metoponium convexicolle, 180

Micrandrena, 309

Micrasema alexanderi, 211; bactro, 211; di-

teris, 211; etra, 211; onisca, 211; oregoni,

211

Microbembex monodonta, 129

Microchaetes, 36

Microgramme arga, 79

Micromes, 90

Mictocommosis, 166

Mictopsichia, 165

miltogrammine flies, 289

Misumenops sp., 225

Mycocepurus goeldii, 258

My das, 299

Neopanorpa burmana, 95; byersi, 95; globu-

lifer, 92; parvula, 94

Neuroptera: Chrysopidae, 314

Nexosa, 166

Nomada ( Gnathias ) opacella, 114

Nomada ( Pachynomada ) sp., 112, 286

Nosodendron californicum, 53

O’Brien, Trypargilum in trap nests, 288

O’Brien and Atsatt, biology of Trirhabda, 139

Ochthebius vandykei, 163

Odonata: Libellulidae, 135

Oedipodinae, 130

Olkowski, Olkowski, van den Bosch, Horn,

Zuparko, and Klitz, Trioxys established on

elm aphid, 59

Ornithodoros talaje, 96

Orodrassus, 228

Osmia, 179

Osmia ( Nothosmia) albiventris, 234; cordata,

234; marginata, 231; pumila, 234

Otobius megnini, 98

Oxybelus, 132; emarginatus, 129

Oxygonodera, 90

Oxyopes salticus, 224; sp. near tridens, 224

Page, polyandry in Brachymyrmex, 258

Palpomyia tibialis, 64

Paltothemis cyanosoma, 135; lineatipes, 135

Papilio androgeus, 281; dardanus, 281; glau-

cus, 278; lycophron, 281; polyxenes, 278;

troilus ilioneus, 279; troilus troilus, 278

Parandra marginicollis marginicollis, 180

VOLUME 58, NUMBER 4

383

Pardosa lowriei, 227; ramulosa, 227; sp., 180,

223, 227

Parker and Bohart, biology Andrena helianthi,

111

Parker and Griswold, biological notes An¬

drena, 284

Parker and Tepedino, Osmia nesting behav¬

ior, 231

Parrella, taxonomy Liriomyza spp., 302

Pedomoecus sierra, 211

Pellenes, 226; oregonensis, 229; sp., 229; vi-

ridipes, 229

Pemphredon, 129

Perillus splendidus, 149

Pfaffenberger, final larval instar Caryobru-

chus, 240

Pheidole megacephala, 346

Phidippus cardinalis, 226

Philip, precinctive more precise than “endem¬

ic”, 369

Philip and Williams, Tabanidae at artificial

lights, 365

Philorus, 117; yosemite, 117

Phryganidia californica, 42

Pieris occidentalis, 162; protodice, 162

Pilimas abaureus, 365; californica, 365

Pinto, larva of Meloe, 367; new Eupompha,

184

Pinto, Oatman, and Platner, redescription Tri-

chogramma australicum, 48

Pleocoma rubiginosa transsierrae, 373

Podalonia, 132

Pogonomyrmex barbatus, 258; desertorum,

258; maricopa, 258; occidentalis, 258; ru¬

ga sus, 258

Polhemus and Polhemus, notes on Naucori-

dae, 326

Pompilus (Ammosphex) luctuosus luctuosus,

227; wasbaueri, 109

Pompilus ( Archnospila ) arctus, 228; fumipen-

nis eureka, 228; scelestus, 228

Porcellio sp., 180

Potamocoris, 328; beckeri, 328; robustus, 328;

parvus, 328

Prenolepis imparis, 258

Priocnemis, 228

Priocnemis ( Priocnemis) cornica, 223

Prionyx albisectus, 131; atratus, 132; par ken,

130; subfuscatus, 131

Proceedings of PCES, 369-377

Proctacanthella leucopogon, 250, 299

Proctacanthus, 300

Prometopion, 85

Protagrotis obscura, 73

Pseudogonatopus arizonicus, 107

Pseudomasaris coquilletti, 124; edwardsii, 124;

macneilli, 127; maculifrons, 124; margi-

nalis, 124; occidentalis, 124; phaceliae, 124;

texanus, 124; vespoides, 124; zonalis, 124

Pseudopygmephorus atypoides, 218

Pseudoscorpionida, 180

Pseudotephritis sp., 41

Psoloessa delicatula, 132

Razafimahatratra and Lattin, new species De-

raeocoris, 352

Reimer and Goeden, life history Stobaera, 105

Rhagoletis cornivora, 25; mendax, 25; po-

monella, 25; zephyria, 25

Rhaphiomydas, 299

Rhizophagus sp., 180

Rhyncolus langulans, 180

Rust and Hanks, biology of Aegialia, 319

Scaphandrena, 311

Scaphinotus, 248

Sceliphron, 288

Schizocosa mccooki, 227; pacifica, 228

Scolopendra sp., 180

Scorpionida: Vaejovidae, 164

Sericopompilis, 228; apicalis, 225; neotropi-

calis, 225

Shapiro, elevational record Pieris, 162

Shelly and Pearson, predatory behavior Proc¬

tacanthella, 250

Shepard, records of Thymelicus, 260

Sierraclava, 31; cooperi, 32

Signiphora sp., 316

Smith and Olson, Coleoptera in stored mari¬

juana, 79

Sokoloff, Wilson, Mulder, and Faustini, Tri-

bolium brevicornis populations, 177

solpugid, 164

Solpugida, 164

Somerby —Proceedings 408th-411th meet¬

ings, 369

Speyeria diana, 278

Steiner, use of proboscis in Prionyx, 129

Steniola spp., 370; tibialis, 370

Stenopelmatus, 164

Stibia, 90

Stictia signata, 129

384

PAN-PACIFIC ENTOMOLOGIST

Stobaera tricarinata, 105

Stone, peach beetle in California, 159

Stone and Wilcox, elaterid light trap collec¬

tions, 202

Stonemyia, 365

Strymon melinus, 41

Symbiotes montanus, 180

Tabanus vittiger, 373

Tachypompilus ferrugineus burrus, 225; fer-

rugineus ferrugineus, 226

Tachysphex, 130

Tapinoma sessile, 180

Tenebroides crassicornis, 180

Tetraleurodes acaciae, 312

Tetramorium similis, 349

Tetrastichus cecidophagus 23; sp., 23, 42

Thaumatographa aurosa, 174; cubensis, 170;

decoris, 166; eremnotorna, 165 \jonesi, 172;

leucopyrga, 166; olympica, 172; regalis, 16.6,

170; youngiella, 172; zapyra, 166

Thereva-group, 300

Thinobatis, 85

Thymelicus lineola, 260

Tibellus gertschi, 226

Tinocallis platani, 59

Tinodes, 208; belisa, 208; cascadia, 208; con-

suetus, 208; gabriella, 208; parvula, 208;

powelli, 208; provo, 208; schusteri, 209; si-

godana, 208; siskiyou, 208; twila, 208; usil-

la, 208

Tipula acuta, 153; graminivora, 153; quaylii,

153; silvestra, 153; simplex, 153

Tongamya miranda, 296

Trialeurodes packardi, 291; vaporariorum, 291

Tribolium anaphe, 181; antennatum, 182;

apiculum, 181; audax, 182; brevicornis, 177;

castaneum, 111', confusum, 79, 177; de¬

structor, 179; madens, 181; myrmecophi-

lum, 182

Trichiotes, 90

Trichogramma australicum, 48; californicum,

51; chilonis, 48; retorridum, 51

Trichoptera: Brachycentridae, 211; Hydro-

psychidae, 206; Lepidostomatidae, 214;

Limnephilidae, 210; Psychomyiidae, 208

Trigona, 182; carbonaria, 181

Trimytis, 85

Triorophus, 90

Trioxys curvicaudus, 63; hartorum, 59; te-

nuicaudus, 59

Triphalopsis californicus, 87; impressicollis, 90;

minor, 90; partida, 90

Triphalus, 90

Trirhabda canadensis, 139; confusa, 139; fla-

volimbata, 139; geminata, 139; luteocincta,

139; nitidicollis, 139; pilosa, 139; sericotra-

chyla, 139

Trogoderma orbatum, 180; variabile, 19

Trypargilum tridentatum, 288

Trypoxylon, 374

Tuberculatus sp., 62

Tychius soltaui, 41

Tyler, comparative flower constancy of Ste-

niolia and Bombus, 370

Vacusus confinus, 180

Vincent and Rack, a new Pseudopygmephorus,

216

Voegtlin, a new Cinara, 196

Wasbauer, pompilid prey records, 223

Westcott, differentiating Rhagoletis pomonel-

la, 25

Wharton, biology of Apiocera, 296

Xylocelia, 129

Xylocopa, 179; mordax, 342; pubescens, 339;

sonorina, 336; sp., 344; sulcatipes, 350; ta-

baniformes orpifex, 180; virginica 349

Young, pupal parasites California oakworm,

Zenillia virilis, 42

Zootermopsis angusticollis, 180

Published by the

Pacific Coast Entomological Society

in cooperation with

The California Academy of Sciences

VOLUME FIFTY-EIGHT

1982 .

EDITORIAL BOARD

G. A. MARSH, Editor

C. PLEINES, Editorial Assistant

R. M. BOHART

H. B. LEECH

E. S. ROSS

J. A. CHEMSAK

W. J. PULAWSKI, Treasurer

PUBLICATION COMMITTEE

1982 1983

G. A. H. McClelland, Chairman K. W. Cooper

F. G. Andrews J. E. Hafemik, Jr.

1984

R. M. Bohart

J. G. Edwards

1985

P. H. Amaud, Jr.

M. S. Wasbauer

San Francisco, California

1982

CONTENTS FOR VOLUME 58

Arnaud, P. H., Jr. —Aggregation of Coelopa ( Neocoelopa ) vanduzeei Cresson

on the Monterey Peninsula Coast, California, and notes on the family (Diptera:

Coelopidae) . 245

Arnaud, P. H., Jr.—P ublications received. 201, 205, 215, 222, 230

Atsatt, P. R. —See O’Brien, P. Y.

Backus, E. A.—See Kimsey, R. B.

Barnard, D. R.—Flight periodicity in Colorado biting midges (Diptera: Cer-

atopogonidae) . 64

Benbow, S. M., and D. E. Foster—B iology of Eutreta diana Osten Sacken on

sand sagebrush Artemisia filifolia Torr. (Diptera: Tephritidae). 19

Bohart, G. E. —See Parker, F. D.

van den Bosch, R. —See Olkowski, W.

Byers, G. W.—A new Neopanorpa (Mecoptera: Panorpidae) from Laos ..... 92

Clark, W. EL, D. R. Frohlich, and P. L. Comanor—S hrike predation on the

scorpion Anuroctonus phaiodactylus (Wood) and on a solpugid (Scorpion-

ida: Vaejovidae; Solpugida) . 164

Clement, S. L., and D. H. Miller—I nsect seed predation on Astragalus bi-

sulcatus (Hook.) Gray (Leguminosae). 38

Collins, P. W. —See Lane, R. S.

Comanor, P. L. —See Clark, W. H.

Coville, R. E.—A note on the biology of Ancistrocerus waldenii flavidulus

Bequaert (Hymenoptera: Eumenidae). 259

Davis, H. V.—Financial statements . 371, 377

Denning, D. G.—New and interesting Trichoptera from the western United

States . 206

Dorr, L. J., and J. L. Neff —Pseudomasaris marginalis nesting in logs in

Colorado (Hymenoptera: Masaridae). 124

Dowell, R. V. —Biology of Tetraleurodes acaciae (Quaintance) (Homoptera:

Aleyrodidae) . 312

Doyen, J. T.—New species of Tenebrionidae from western North America

(Coleoptera) . 81

Ehler, L. E. —Observations on Tria/eurodes packardi (Morrill) in a community

garden (Homoptera: Aleyrodidae). 291

Evenhuis, N. L.—Notes on the subgenus Bombylius ( Zephyrectes ) (Diptera:

Bombyliidae) . 236

Faustini, D. —See Sokoloff, A.

Foster, D. E. —See Benbow, S. M.

Frohlich, D. R. —See Clark, W. H.

Garcia-Vidal, M.—The genus Anoplosiagum Blanchard (Coleoptera: Scara-

baeidae) in Cuba. Part I. Descriptions of five new species. 330

Garrison, R. W.—Paltothemis cyanosoma, a new species of dragonfly from

Mexico (Odonata: Libellulidae). 135

Gerling, D.—Nesting biology and flower relationships of Xylocopa sonorina

Smith in Hawaii (Hymenoptera: Anthophoridae). 336

Giuliani, D.—Notes on a collection of intertidal beetles from the Farallon

Islands, California .. 163

Goeden, R. D. —See Reimer, N. J.

Griswold, T. —See Parker, F. D.

Hafernik, J. E., Jr.—M imetic selection and subspecific variation in the spice-

bush swallowtail, Papilio troilus Linnaeus (Lepidoptera: Papilionidae). 278

Hall, J. C. — See Hurd, P. D., Jr.

Hanks, L. M. — See Rust, R. W.

Hartman, M. J., and C. D. Hynes—T he immature stages of Tipula simplex

Doane and T. acuta Doane (Diptera: Tipulidae). 153

Heppner, J. B.—Revision of American Thaumatographa, with a new species

from Cuba (Lepidoptera: Tortricidae: Chlidanotinae). 165

Hobbs, S. D.—See Kulhavy, D. L.

Hom, R.—See Olkowski, W.

Hurd, P. D., Jr., E. G. Linsley, and J. C. Hall—P hilip Hunter Timberlake,

1883-1981 . 2

Hynes, C. D. —See Hartman, M. J.

Johnson, P. J.—A new genus and species of California pill beetle (Coleoptera:

Byrrhidae). 31

Kamm, J. A. —Protagrotis obscura Barnes and McDunnough (Lepidoptera: Noc-

tuidae): A pest of grasses grown for seed in the Pacific northwest. 73

Kimsey, L. S.—The female of Pompilus (Ammosphex ) wasbaueri Evans (Hy-

menoptera: Pompilidae) . 109

Kimsey, R. B., and E. A. Backus—O bservations on the biology and life history

of the net-winged midge Dioptopsis sequoiarum (Alexander) (Diptera:

Blephariceridae) . 117

Klitz, W. —See Olkowski, W.

Krombein, K. V., and E. G. Linsley—P aul David Hurd, Jr., 1921-1982 . . . 262

Kulhavy, D. L., J. W. Schwandt, and S. D. Hobbs—U nderstory plants as

indicators of host trees of the wounded tree beetle, Nosodendron californicum,

in northern Idaho (Coleoptera: Nosodendridae). 53

Lane, R. S., S. E. Miller, and P. W. Collins—T icks (Acari: Argasidae and

Ixodidae) from the California Channel Islands. 96

Lanham, U. N.—A new species of Andrena (Micrandrena ) from Colorado, with

other notes on the group (Hymenoptera: Apoidea). 309

Lattin, J. D. —See Razafimahatratra, V.

Lee, V. L.—Proceedings . 373

Linsley, E. G. —See Hurd, P. D., Jr.

Linsley, E. G. —See Krombein, K. Y.

Miller, D. H.— See Clement, S. L.

Miller, S. E. —See Lane, R. S.

Mulder, G. —See Sokoloff, A.

Neff, J. L.—See Dorr, L. J.

Oatman, E. R. —See Pinto, J. D.

O’Brien, M. F . — Trypargilum tridentatum (Packard) in trap nests in Oregon

(Hymenoptera: Sphecidae: Trypoxylinae) . 288

O’Brien, P. Y., and P. R. Atsatt— Life history and general bionomics of

Trirhabda sericotrachyla Blake (Coleoptera: Chrysomelidae) in southern

California. 139

Olkowski, H. —See Olkowski, W.

Olkowski, W., H. Olkowski, R. van den Bosch, R. Hom, R. Zuparko, and

W. Klitz— The parasitoid Trioxys tenuicaudus Stary (Hymenoptera: Aphi-

diidae) established on the elm aphid Tinocallis platani Kaltenbach (Homop-

tera: Aphididae) in Berkeley, California. 59

Olson, C. A.—See Smith, R. L.

Page, R. E., Jr. —Polyandry in Brachymyrmex depilis Emery (Hymenoptera:

Formicidae). 258

Parker, F. D., and G. E. Bohart—N otes on the biology of Andrena ( Callan -

drena ) helianthi Robertson (Hymenoptera: Andrenidae). Ill

Parker, F. D., and T. Griswold—B iological notes on Andrena ( Callandrena )

haynesi Viereck and Cockerell (Hymenoptera: Andrenidae). 284

Parker, F. D., and V. J. Tepedino—B ehavior of Osmia ( Nothosmia ) marginata

Michener in the nest (Hymenoptera: Megachilidae). 231

Parrella, M. P.— A review of the history and taxonomy of economically im¬

portant serpentine leafminers (Liriomyza spp.) in California (Diptera: Agro-

myzidae) . 302

Pearson, D. L.—See Shelly, T. E.

Pfaffenberger, G. S. —Description of and phylogenetic comments on the final

larval instar of Caryobruchus veseyi (Horn) (Coleoptera: Bruchidae). 240

Philip, C. B.—Precinctive is more precise zoogeographic term than etymolog¬

ically ambiguous “endemic”. 369

Philip, C. B., and S. C. Williams—T abanidae (Diptera) attracted to artificial

lights in California. 365

Pinto, J. D.—A new Eupompha from Baja California with additional infor¬

mation on E. decolorata (Horn) (Coleoptera: Meloidae) . 184

Pinto, J. D.—The larva of MeloefranciscanusVan Dyke and MacSwain’s Meloe

sp. a (Coleoptera: Meloidae). 367

Pinto, J. D., E. R. Oatman, and G. R. Platner— Trichogramma australicum

Girault (Hymenoptera: Trichogrammatidae): Redescription and lectotype

designation . 48

Platner, G. R. —See Pinto, J. D.

Polhemus, D. A.—See Polhemus, J. T.

Polhemus, J. T., and D. A. Polhemus—N otes on Neotropical Naucoridae IE

A new species of Ambrysus and review of the genus Potamocoris (Hemip-

tera). 326

Rack, G. —See Vincent, L. S.

Razafimahatratra, V., and J. D. Lattin—F ive new species and new syn¬

onymies for the genus Deraeocoris (Heteroptera: Miridae) from western North

America. 352

Reimer, N. J., and R. D. Goeden—L ife history of the delphacid planthopper

Stobaera tricarinata (Say) on western ragweed, Ambrosia psilostachya De-

candolle, in southern California (Hemiptera-Homoptera: Delphacidae) .... 105

Rust, R. W., and L. M. Hanks—N otes on the biology of Aegialia hardyi Gordon

and Cartwright (Coleoptera: Scarabaeidae). 319

Schwandt, J. S. —See Kulhavy, D. L.

Shapiro, A. M. —A new elevational record for Pieris protodice in California

(Lepidoptera: Pieridae). 162

Shelly, T. E., and D. L. Pearson—D iurnal variation in the predatory behavior

of the grassland robber fly, Proctacanthella leucopogon (Williston) (Diptera:

Asilidae). 250

Shepard, J. H. —Additional records of Thymelicus lineola (Ochsenheimer) in

British Columbia (Lepidoptera: Hesperiidae). 260

Smith, R. L., and C. A. Olson—C onfused flour beetle and other Coleoptera

in stored marijuana . 79

Sokoloff, A., R. Wilson, G. Mulder, and D. Faustini—O bservations on

populations of Tribolium brevicornis LeConte (Coleoptera: Tenebrionidae).

II. The habitat niche of a local population in southern California. 177

Somerby, R. E. —Proceedings. 369

Steiner, A. L.—Use of the proboscis for prey-piercing and sucking by sphecid

wasps of the genus Prionyx (Hymenoptera: Sphecidae), a case of convergent

evolution. 129

Stone, M. W.—The peach beetle, Cotinis mutabilis (Gory and Percheron), in

California (Coleoptera: Scarabaeidae). 159

Stone, M. W., and J. Wilcox— Light trap collections of three introduced Co-

noderus species (Coleoptera: Elateridae) in southern California. 202

Tepedino, V. J. —See Parker, F. D.

Tyler, T. L. — Comparative flower constancy of Steniolia tibialis Handlirsch

and Bombus vosnesenskii Radoszkowski. 370

Vincent, L. S., and G. Rack— Pseudopygmephorus atypoides Rack, new species

(Acari: Pygmephoridae) associated with the fossorial mygalomorph spider

Atypoides riversi O. P.-Cambridge (Araneae: Antrodiaetidae) in Cali¬

fornia . 216

Voegtlin, D.—A new aphid species, Cinara radicivora (Homoptera: Aphididae)

living on white fir. 196

Wasbauer, M. S.—Prey records for some North American spider wasps (Hy¬

menoptera: Pompilidae) . 223

Westcott, R. L.—Differentiating adults of apple maggot, Rhagoletis pomonella

(Walsh) from snowberry maggot, R. zephyria Snow (Diptera: Tephritidae) in

Oregon. 25

Wharton, R. A.—The biology of Apiocera haruspex Osten Sacken (Diptera:

Apioceridae) in central California, and comparison with other Asiloidea . . 296

Wilcox, J. —See Stone, M. W.

Williams, S. C. — See Philip, C. B.

Wilson, R. —See Sokoloff, A.

Young, L. C. —Key to the pupal parasites of California oakworm, Phryganidia

californica (Lepidoptera: Dioptidae), based on larval exuviae. 42

Zuparko, R. —See Olkowski, W.

THE PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

Papers on the systematic and biological phases of entomology are favored, including short notes or articles up to ten printed pages,

on insect taxonomy, morphology, ecology, behavior, life history, and distribution. Excess pagination must be approved and will be

charged to the author. Papers are published after acceptance in approximately the order that they are received. Papers of less than a

printed page will be published as space is available, in Scientific Notes.

All manuscripts will be reviewed before acceptance.

Manuscripts for publication, proofs, and all editorial matters should be addressed to the editor.

General. — The metric system is to be used exclusively in manuscripts, except when citing label data on type material, or in direct

quotations when cited as such. Equivalents in other systems may be placed in parentheses following the metric, i.e. “1370 m (4500

ft) elevation”.

Typing. — Two copies of each manuscript must be submitted (original and one xerox copy or two xerox copies are suitable). All

manuscripts must be typewritten, double-spaced throughout, with ample margins, and be on bond paper or an equivalent weight.

Carbon copies or copies on paper larger than 8 'A X 11 inches are not acceptable.

Underscore only where italics are intended in the body of the text. Number all pages consecutively and put authors name on each

sheet. References to footnotes in text should be numbered consecutively. Footnotes must be typed on a separate sheet.

Manuscripts with extensive corrections or revisions will be returned to the author for retyping.

First Page. — The page preceding the text of the manuscript must include (1) the complete title, (2) the order and family in parentheses,

(3) the author’s name or names, (4) the institution with city and state or the author’s home city and state if not affiliated (5) the

complete name and address to which proof is to be sent.

Names and descriptions of organisms. — The first mention of a plant or animal should include the full scientific name with the author

of a zoological name not abbreviated. Do not abbreviate generic names. Descriptions of taxa should be in telegraphic style. The

International Code of Zoological Nomenclature must be followed.

Tables. — Tables are expensive and should be kept to a minimum. Each table should be prepared as a line drawing or typed on a

separate page with heading at top and footnotes below. Number tables with Arabic numerals. Number footnotes consecutively for

each table. Use only horizontal rules. Extensive use of tabular material requiring typesetting may result in increased charges to the

author.

Illustrations. — No extra charge is made for line drawings or halftones. Submit only photographs on glossy paper and original drawings.

Authors must plan their illustrations for reduction to the dimension of the printed page (117 X 181 mm; 4 5 /s X 7'/s inches). If possible,