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6IPTITI ITi Ad!

Journal of the

New York Entomological Society

published by

The New York Entomological Society

Contents Volume 92, 1984, Numbers 1-4
Number 1

Directory of systematic entomologists and acarologists in the People’s Republic of
China Lloyd Knutson and W. L. Murphy 2-18

Vegetational complexity and parasitism of green peach aphids ( Myzus persicae (Sul-
zer) (Homoptera: Aphidae)) on collards David J. Horn 1 9-26

The Membracidae and other Homoptera described by Asa Fitch, 1851, and Ebenezer
Emmons, 1855: Historical perspective and analysis Jeffrey K. Barnes 27-34

Association of Paratrechina arenivaga (Hymenoptera: Formicidae), with nymphs of

Oecleus borealis (Homoptera: Cixiidae) C. R. Thompson 35-41

A new species of Sigara from western Oregon and Washington (Hemiptera: Corixidae)

Gary M. Stonedahl 42-47

Carvalhoma (Hemiptera: Miridae): Revised subfamily placement

Randall T. Schuh and Michael D. Schwartz 48-52

Life history and laboratory rearing of Euschistus ictericus (Hemiptera: Pentatomidae),
with descriptions of immature stages J. E. McPherson and S. M. Paskewitz 53-60

Laboratory rearing of Amaurochrous cinctipes (Hemiptera: Pentatomidae: Podopinae)
with descriptions of immature stages J. E. McPherson and S. M. Paskewitz 61-68

A conspectus of Pentatomini of the western hemisphere. Part 3. (Hemipera: Penta-
tomidae) L. H. Rolston and F. J. D. McDonald 69-86

Temporal and spatial variation in sex ratio and mating frequency in soldier beetles

Kim Holder Bailey, David E. McCauley, and W. David Sissom 87-92

Book Reviews

Bark beetles in North American conifers Stephen L. Wood 93-94

Biological diversification in the tropics: Proceedings of the Fifth International Sym-
posium for the Association for Tropical Biology David Grimaldi 94-96

Number 2

Revision of Arocera Spinola (Hemiptera: Pentatomidae) F. J. D. McDonald

Parathroscinus, a new genus of beetles from Southeast Asia (Limnichidae: Cephal-
obyrrhinae) Day id P. Wooldridge

Fleas (Siphonaptera) from nests of woodpeckers in Alaska

Glenn E. Haas and Nixon Wilson

Synopsis of Ommatius Wiedemann (Diptera: Asilidae) from Hispaniola

A. G. Scarbrough

Propleural transformations with respect to the disposition of propleural suture in
order Hymenoptera Malkiat S. Saini

Tool use by the ant, Novomessor albisetosus (Mayr) Philip McDonald

Foraging behavior of the bees Halictus ligatus (Hymenoptera: Halictidae) and Cer-
atina calcarata (Hymenoptera: Anthophoridae): Foraging speed on early-summer
composite flowers Howard S. Ginsberg

On the Pseudoscorpion-mimicking spider Cheliferoides (Araneae: Salticidae)

Norman I. Plat nick

Aufeius impressicollis (Hemiptera: Rhopalidae): Easternmost U.S. record, host plant
relationships, and laboratory rearing A. G. Wheeler, Jr.

Compsidolon ( Coniortodes ) salicellum (Hemiptera: Miridae): A predaceous plant bug,
new to the United States, found on Filbert

John D. Lattin and Russell H. Messing

Notes and Comments

Notes on the distribution of Dermaptera in some Costa Rican cacao plantations

Allen M. Young

Book Review

Naturalists’ Handbooks A. G. Wheeler, Jr.

97-120

121-124

125-130

131-149

150-155

156-161

162-168

169-173

174-178

179-183

184-187

188-192

Number 3

A Revision of the Black Grass Bug Genus Irbisia Reuter (Heteroptera: Miridae)

Michael D. Schwartz 193-306

Number 4

Ormenaria rufifascia (Homoptera: Fulgoroidea: Flatidae): descriptions of nymphal
instars and notes on field biology Stephen W. Wilson and James H. Tsai 307-3 1 5

A revision of the genus Horridipamera Malipatil (Hemiptera: Lygaeidae)

James A. Slater and Le-Yi Zheng 316-341

New synonymy and a new species in the genus Mormidea (Hemiptera: Pentatomidae)

L. H. Rolston 342-343

Pellaea santarosensis (Hemiptera: Pentatomidae), a new species from Costa Rica

L. H. Rolston 344-348

Karyotype of Conomyra flava (McCook) (Hymenoptera: Formicidae)

James C. Cokendolpher and Oscar F. Francke 349-351

Key to the males of the nominate subgenus of Euschistus in South America, with
descriptions of three new species (Hemiptera: Pentatomidae) L. H. Rolston 352-364

Chemical analysis of the male aedeagal bladder in the fire ant, Solenopsis invicta

Buren David E. Ball, H. J. Williams and S. B. Vinson 365-370

Revision of the harvestman genus Leptobunus and dismantlement of the Leptobun-

idae (Arachnida: Opiliones: Palpatores) James C. Cokendolpher 371-402

Notes and Comments

Oviposition behavior of Chrysops atlanticus (Diptera: Tabanidae)

Roy K. Sofield and Randy Gaugler 403-404

Carrion as an attractant to the American dog tick, Dermacentor variabilis (Say)

Kenneth A. Kneidel 405-406

A morphological distinction between the larvae of Cheumatopsyche campyla and C.
pettiti (Trichoptera: Hydropsychidae) Martin H. Smith 407-410

Book Review

A Field Guide to the Beetles of North America

Honorary, Life, and Sustaining Members
Reviewers for 1984

Quentin D. Wheeler

411-413

414

414

4

/:u& / *>

— — —

Vol. 92 JANUARY 1984 No. 1

Journal

of the

New York

Entomological Society

(ISSN 0028-7199)

Devoted to Entomology in General

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY

Editor: Randall T. Schuh, Department of Entomology, American Museum

of Natural History, Central Park West at 79th Street, New York, New
York 10024

Book Review Editor: Quentin D. Wheeler, Department of Entomology,

Cornell University, Ithaca, New York 14853
Publications Committee: Louis Trombetta, St. Johns University, Queens,

New York, Chairman; Alfred G. Wheeler, Jr., Pennsylvania State De-
partment of Agriculture, Harrisburg; Joseph M. Cerreta, St. Johns Uni-
versity, Queens, New York.

The New York Entomological Society
Incorporating The Brooklyn Entomological Society

President: Gerard Iwantsch, Department of Biological Sciences, Fordham

University, Bronx, New York 10458

Vice President: Henry M. Knizeski, Jr., Department of Biology, Mercy

College, Dobbs Ferry, New York 10522
Secretary: Irene E. Matejko, Science Department, The New Lincoln School,

New York, New York 10021

Assistant Secretary: Dennis J. Joslyn, Department of Biology, Rutgers

University, Camden, New Jersey 08102
Treasurer: Louis Sorkin, Department of Entomology, American Museum

of Natural History, New York, New York 10024
Trustees: Class of 1984— Joseph Cerreta, St. Johns University, Queens,

New York; Durland Fish, Fordham University, Bronx, New York;
Class of 1985— Peter Chabora, Queens College, New York; Charles
Porter, Fordham University, Bronx, New York.

Annual dues are $18.00 for established professionals with journal, $10.00 without journal,
$10.00 for students with journal, $5.00 without journal. Sustaining memberships are $48.00
per year, institutional memberships are $120.00 per year, and life memberships are $300.00.
Subscriptions are $27.00 per year domestic and $30.00 foreign. All payments should be made
to the Treasurer. Back issues of the Journal of the New York Entomological Society, the Bulletin
of the Brooklyn Entomological Society, Entomologica Americana, the Torre-Bueno Glossary of
Entomology and other Society publications can be purchased from Lubrecht and Cramer, RFD
1, Box 227 , Monticello, New York 12701.

Meetings of the Society are held on the third Tuesday of each month (except June through
September) at 8 p.m. in the American Museum of Natural History, Central Park West at 79th
Street, New York, New York.

Mailed June 19, 1984

The Journal of the New York Entomological Society (ISSN 0028-7199) is published quarterly (January, April, July,
October) for the Society by Allen Press, Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at
New Brunswick, New Jersey and at additional mailing office.

Known office of publication: American Museum of Natural History, New York, New York 10024.

Journal of the New York Entomological Society, total copies printed 600, paid circulation 443, mail subscription 443, free
distribution by mail 7, total distribution 450, 150 copies left over each quarter.

ANNOUNCEMENT

BEQUEST AND PUBLICATION FUND

Late in 1983 the Society was informed that it would be the recipient of
generous bequests from the estates of the late Dr. Charles Alexander and
Mrs. Patricia Vaurie.

Dr. Alexander was a world renowned expert on crane flies, which interest
he pursued during his career as a professor of entomology at the University
of Massachusetts in Amherst. Mrs. Vaurie took up entomology as an a vo-
cational interest, but soon established a strong working relationship with the
Department of Entomology of the American Museum of Natural History in
New York City, where as a Research Associate she pursued her interests in
beetle classification, particularly of the families Curculionidae, Languriidae,
and Scarabaeidae. Both Dr. Alexander and Mrs. Vaurie stipulated that their
gifts be used for the benefit of the Society’s publications.

The interests of C. P. Alexander and Patricia Vaurie were strongly oriented
towards monographic work on the groups of insects which they studied. The
same interests have long been associated with the publications of the New
York Entomological Society and the Brooklyn Entomological Society, which
were incorporated in 1968.

The Society has therefore established a “Journal Publication Endowment
Fund,” the proceeds of which will be devoted to the costs of publication of
the Journal of the New York Enntomological Society. The Society officers
intend the endowment to: 1) encourage the publication of monograph-length
papers; 2) increase the number of pages published annually in the Journal,
3) reduce the per-page cost to authors; and 4) maintain the costs of mem-
bership and subscription at reasonable and competitive rates.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 2-18

DIRECTORY OF SYSTEMATIC ENTOMOLOGISTS AND
ACAROLOGISTS IN THE PEOPLE’S REPUBLIC OF CHINA

Lloyd Knutson and William L. Murphy

Insect Identification and Beneficial Insect Introduction Institute,

U.S. Department of Agriculture, Beltsville, Maryland 20705

A bstract.— Listings of 188 systematic entomologists in the People’s Republic of China are
given by institution, speciality and name.

This directory is a revised and expanded version of a list that originally appeared
in the volume Biological Control of Pests in China (p. 249 ff.), which was published
in 1982 by the Office of International Cooperation and Development (OICD), United
States Department of Agriculture (USD A). The directory lists 188 persons who are
conducting or who have recently conducted taxonomic studies of insects and mites
at 63 institutions in the People’s Republic of China (PRC). The directory is derived
from articles published in Entomotaxonomia, Contributions of the Shanghai Institute
of Entomology, and Acta Entomologica Sinica; from lists prepared by J. R. Coulson
and D. R. Davis during their visits to the PRC in 1979; from information obtained
by P. M. Marsh and E. E. Grissell during their visit in 1980; and from information
obtained by L. Knutson and R. D. Gordon during their visit in 1981. Many sub-
sequent additions and corrections have been made by L. Knutson.

Names that appear in parentheses below the name of an institution indicate per-
sonal visits to the pertinent institution by the named individual (J. R. Coulson, E.
E. Grissell, R. D. Gordon, P. M. Marsh, and L. Knutson IIBIII, USDA; D. R. Davis,
Department of Entomology, Smithsonian Institution). These persons may be able to
provide additional information about these institutions to requestors.

This directory is arranged according to the historical arrangement of directories in
the PRC— by province, generally from the northeast to the southwest, then the center,
then the central and coastal southeast, then lastly the tropical southern provinces.
Within each province the institutions are arranged according to the following se-
quence: 1) institutions affiliated with Academia Sinica; 2) other research institutions
(universities, health stations, etc.); and 3) museums. Old names of provinces are
included in brackets at first mention. Names of persons are arranged alphabetically
within each institution, except for the large Institutes in Beijing and Shanghai.

Since 1949, formal titles and a hierarchical ranking system have generally not been
used in universities, colleges, and scientific units in the PRC. Exceptions to this apply
to persons who have received a doctoral degree in an overseas university and, in
some cases, persons who received a doctoral degree in China prior to 1949. Most
units apparently have someone who serves as “Chairman” or “Director” in the
Western sense of the term, although they may not be officially titled as such. Western
titles such as “Dr.”, “Mr.”, “Miss”, and “Mrs.” are appropriate in addressing persons
verbally. Note that married women retain their family name but may be addressed

verbally as “Madame .” In addressing someone in writing in China, titles are

usually not used, but a female character is placed after the name of a female. West-

1984

CHINESE SYSTEMATISTS

3

erners addressing persons in China in writing should use a title or not as seems
appropriate. Where known, we have included Western-style titles in this directory.

In this directory, the modern (pinyin) transliteration of the names of persons is
given first, followed by the old spelling (Wade-Giles), where both are known. In a
few instances, the spelling used here is that used by the scientist, particularly for
persons who have published under the old transliteration. In the Chinese system,
the family name is written first and the given name second. In the past few years,
the hyphen in the given name often has been dropped, so that now the given name
consists of one word. However, the use of the hyphen, and in fact the spelling of the
given name, may be inconsistent; different spellings have been used for two different
articles in even one issue of a journal. The pinyin system is used exclusively for place
names herein. Names or parts of names have been underlined where the spelling is
uncertain. Names preceded by a question mark and in brackets are neither pinyin
nor Wade-Giles and cannot be corrected without the appropriate Chinese characters.

We would like to thank A. C. F. Hung, Beneficial Insect Introduction Laboratory
(BIIL), IIBIII, for reviewing the directory and correcting the spelling of many names.
We also thank William L. Davis, formerly Agricultural Counselor, Beijing, for his
helpful comments on the formation of Chinese names; Prof. Yang Pinglan, Director,
Institute of Entomology, Academia Sinica, Shanghai, for his useful comments; and
Huo Shaotang, Academy of Agricultural Sciences, Yangling, Shaanxi Province, for
offering many important corrections and additions and for assisting in the organi-
zation of the information.

No words of appreciation would be complete without recognition of the support
received from the USDA’s Office of International Cooperation and Development,
which has coordinated the U.S.-China agricultural exchanges, sponsored visits to
the PRC by Agricultural Research Service entomologists, and encouraged publication
of results— an outgrowth of which is the present effort. We are most appreciative of
the various kinds of expert assistance and advice provided by the OICD staff and
especially by Thomas F. Kelly.

This directory, as well as several other directories of insect systematists and bio-
logical control workers, is being maintained and updated in the Insect Identification
and Beneficial Insect Introduction Institute as part of the Institute’s computerized
Biosystematic and Biocontrol Information System. Thus, it would be greatly appre-
ciated if any corrections or additions to this directory would be sent to Lloyd Knutson,
Chairman, Insect Identification and Beneficial Insect Introduction Institute, Room
1, Bldg. 003, Beltsville Agricultural Research Center— West, Beltsville, MD 20705,
to be incorporated into future revisions.

INSTITUTIONAL INDEX

1 . Institute of Zoology, Department of Insect Taxonomy and Faunology, Academia
Sinica, BEIJING [PEKING] (Coulson— 79, Davis— 79, Grissell and Marsh— 80,
Knutson and Gordon — 81)

Mr. Meng Xianling (Meng Hsiang-Ling) [Director for Research, Administrative
Section, Institute of Zoology].

Mr. Zhao Jianming (Chao Chien-ming) [Vice Director, Associate Research Sci-
entist]—Diptera: Tachinidae, Sarcophagidae.

4

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Prof. Dr. Zhu Hongfu (Chu Hung-fu) [Deputy Director] — Lepidoptera: Geometri-
dae, Sphingidae, Saturniidae, Bombycidae, Drepanidae, Hepialidae, Brahmaei-
dae, Uraniidae, Pterothysanidae, Callidulidae, Endromidae, Episopeiidae. Ho-
moptera: Aphidoidea.

Coleoptera

Mr. Chang Youwei [Assistant Research Scientist] — Scarabaeoidea.

Prof. Dr. Chen Shixiang (Chen Secien or Chen Shih-Hsiang) [Director] — Chrys-
omelidae.

Ms. Chen Yuanquing [Research Assistant]— Curculionoidea.

Mr. Huang Fusheng [Associate Research Scientist] — Scolytidae. Protura. Zorap-
tera.

Mr. Li Hongxing (Lee Hung-shing) [Assistant Research Scientist] — Bruchidae, Cur-
culionidae.

Ms. Pu Fuji (Pu F. J.) [Assistant Research Scientist]— Cerambycidae.

Ms. Tan Zhuanjie (T’an Chuanchieh) [Associate Research Scientist] — Meloidae,
Cicindelidae, Eumolpidae, fossil insects.

Ms. Yin Hui-fen (Hweifeng) [Associate Research Scientist] — Scolytidae, Platypod-
idae.

Ms. Yu Peiyu [Associate Research Scientist] — Carabidae, Criocerinae.

Lepidoptera

Mr. Cai Rongquan [Assistant Research Scientist]— Notodontidae, Limacodidae.

Ms. Chao Chungling [Assistant Research Scientist] — Lymantriidae, Psychidae,
Thyatridae.

Mr. Chen Yixin [Assistant Research Scientist]— Noctuidae, Agaristidae, Cossidae.

Ms. Fang Chenglai [Assistant Research Scientist]— Arctiidae, Amatidae, Hypsidae,
larvae and pupae of Noctuidae.

Mr. Hou Taogion [Assistant Research Scientist] — Lasiocampidae, Thaumeto-
poeidae. Coleoptera: Buprestidae.

Prof. Li Zhuanlong (Li Chuanlung)— Rhopalocera, Papilionidae.

Mr. Liu Yujiao (Liu Yu Chiao) [Associate Research Scientist]— Tortricidae, Co-
chylidae, Ethmiidae.

Mr. Wang Pingyuan [Associate Research Scientist] — Pyralidae, Zygaenidae.

Hymenoptera

Mr. Chen T&iXu—Tricho gramma.

Mr. Liao Dingxi (Liao Tingshi) [Associate Research Scientist] — Chalcidoidea.

Ms. Wang Jinyan [Research Assistant] — Braconidae.

Ms. Wang Sufang [Assistant Research Scientist]— Bombus, Ichneumonidae.

Ms. Wu Yanru (Wu Yenju) [Assistant Research Scientist]— Apoidea, Proctotru-
poidea.

Diptera

Mr. Li Tiesheng [Assistant Research Scientist]— Ceratopogonidae. Hymenoptera:
Vespoidea.

Ms. Ma Sufang [Assistant Research Scientist] — Culicidae.

Ms. Shi Yongshan [Assistant Research Scientist]— Tachinidae, Syrphidae. (Works
with Mr. Zhao.)

1984

CHINESE SYSTEMATISTS

5

Ms. Sun Caihong [Assistant Research Scientist] — Syrphidae, Anthomyiidae, Cal-
liphoridae, Muscidae.

Ms. Wang Zunming (Mrs. Li Tiesheng) [Research Assistant] — Tabanidae. (Visited
at Oregon State University, 1982.)

Homoptera

Mr. Wang Ziqing (Wang Tzeching) [Assistant Research Scientist]— Coccoidea.
Mr. Zhang Guangxue (Chang Guangshyue) [Assistant Research Scientist]— Aphi-
doidea. Thysanoptera.

Mr. Zhong Tiesen— Aphidoidea ( Entomotaxonomia ).

Orthoptera

Liu Jupeng— Acrididae.

Acari

Mr. Teng Kuofan [Associate Research Scientist]— Ixodoidea, Gamasoidea. Ano-
plura.

Ms. Wang Xiangfu [Assistant Research Scientist]— Tetranychidae.

2. Department of Plant Protection, Beijing Agricultural University, BEIJING (Coul-
son— 79)

Prof. Yang Ji-kun (Yang Chi-Kun)— Neuroptera: Chrysopidae. Other beneficial
insects. Diptera: Rhagionidae. Coleoptera: Stylopidae.

3. Forest Research Institute, Chinese Academy of Forestry, BEIJING (Knutson and
Gordon— 81)

Ms. Huang Xiaoyun— Hymenoptera: Tenthredinoidea.

Mr. Li Guan-wu— Hymenoptera: Tenthredinoidea.

Prof. Xiao Gangrou (Hsiao Kang-Jou) [Vice Director] — Hymenoptera: Tenthre-
dinoidea.

Dr. Zhou Shushi— Hymenoptera: Tenthredinoidea.

4. Institute of Microbiology and Epidemiology, Chinese Academy of Military Med-
ical Sciences, P.O. Box 130, BEIJING

Dou Guilan— Siphonaptera.

Ji Shuli— Siphonaptera.

Li Peisi— Diptera: Ceratopogonidae, Culicidae.

Liu Chi-ying— Siphonaptera: Ceratophyllidae.

Liu Quan — Siphonaptera: Ceratophyllidae.

Prof. Lu Baolin (Luh Paoling)— Diptera: Ceratopogonidae, Culicidae.

Ren Shiming— Siphonaptera.

Wang Tai-hua— Diptera: Ceratopogonidae.

Wu Houyong— Siphonaptera: Hystrichopsyllidae.

Mr. Yu Yixin— Diptera: Ceratopogonidae.

5. Museum of Natural History, BEIJING (Davis— 79)

Mr. Liu Sinan (Liu Szenang)— General entomology.

Ms. Wang Xicheng (Wang Sheehan)— General entomology.

(Both are involved mainly in exhibit work at present; no publications.)

6. Laboratory of Taxonomy and Ecology, Shanghai Institute of Entomology, Aca-

6

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92( 1 )

demia Sinica, Chungkin Road (S.), 225 SHANGHAI (Coulson— 79, Davis— 79, Gris-
sell and Marsh— 80, Knutson and Gordon— 81)

Prof. Xia Kai-ling (Hsia Kai-ling) [Laboratory Head] — Orthoptera: Acridoidea.
Isoptera: Rhinotermitidae, Termitidae. Diptera: Asilidae.

Protura

Prof Ms. Yin Wen-ying (Yin Wen-Ying) [Vice Director] — Eosentomidae.
Mr. Zhang Zhi-yuan (Zhang Zhi-yang)— Eosentomidae.

Isoptera

Mr. Fan Shu-de (Fan Shu-teh)— Rhinotermitidae.

Ms. Han Meizhen.

He Xiu-song— Termitidae.

Orthoptera

Ms. Bi Dao-ying— Acridoidea.

Ms. Jin Xing-bao— Acridoidea.

Homoptera

Mr. Hu Jing-lin (Hu Jin-lin)— Coccoidea: Diaspididae.

Prof. Yang Pinglan (Bain-ley Young) [Director of the Institute] — Coccoidea: Dias-
pididae. Aleyrodidae.

Diptera

Mr. Chen Zhi-zi (Chen Tzi-Tze) [Assistant Engineer]— Anthomyiidae, Calliphor-
idae, Muscidae, Sarcophagidae.

Prof. Fan Zi-de (Fan Tze-teh)— Anthomyiidae, Muscidae, Calliphoridae (especially
synanthropic calyptrate Diptera), Hypodermatidae, Syrphidae, Sarcophagidae
(not including Tachinidae).

Prof. Liu Wei-de (Liu Wi-teh) [Vice Director]— Tabanidae, Culicidae.

Yao Yun-mei— Tabanidae.

Lepidoptera

Mr. Li Chengzhang (Li Cheng-chang)— Pyralidae.

Hymenoptera

Mr. Shi Dasan [Deputy Head of the Laboratory]— Trichogramma, Aphidiidae.
Diptera: Syrphidae.

Acari

Mr. Wang Xiaozu (Wang Shiao-tsu)— Acaridae, Cheyletidae, Glycyphagidae.

Hemiptera

Ms. Jin Quinying.

7. Department of Biology, Fudan University, SHANGHAI (Knutson and Gor-
don—81)

Acari

Ms. Ding Tingzhong— Tarsonemidae (cyclamen mites), Eriophyidae.
Ms. Dong Huiqing— Eriophyidae.

Mrs. Ke Lisheng— Phytoseiidae.

1984

CHINESE SYSTEMATISTS

7

Mr. Liang Lairong— Phytoseiidae.

Prof. Xin Jieliu (Shin Kai-lou)— Especially Trombidiidae.

8. Shanghai College of Agriculture, SHANGHAI
Mr. Ma En-Pei— Acari: Tetranychidae.

9. Shanghai First Medical College, SHANGHAI
Prof. Hsu Yinchi— Acari: chiggers, Demodex.

Prof. Wen Tinghuan— Acari: Acariformes, Trombiculidae.

10. Second Military Medical College, Department of Parasitology, SHANGHAI
(Davis— 79)

Prof. Xu Feng-yi (Chu Fengi)— Diptera: Culicidae, Ceratopogonidae.

1 1. Museum of Natural History, SHANGHAI (Davis— 79)

Ms. Yuan Yilan— Acari: Tetranychidae.

12. Department of Biology, Nankai University, TIANJIN (TIENTSIN)

Mr. Zheng Leyi (Zheng Le-yi)— Hemiptera: Lygaeidae.

13. Tianjin Animal and Plant Quarantine Service, Pu-Cow Road, TIANJIN

Mr. Guan Lianghua— Taxonomy of stored-product pests, especially Coleoptera,
and quarantine.

14. Tianjin (Tientsin) Museum of Natural History, TIANJIN
Mr. Liu Shengli— Hemiptera: Acanthosomatidae.

15. First Control and Research Institute of Endemic Disease, JILIN [KIRIN] PROV-
INCE

Ma Liming— Acari: Parasitiformes: Rhodacaridae.

16. Northeastern Normal University, Changchun, JILIN PROVINCE
Zhang Fengling— Orthoptera: Acrididae.

17. Health and Anti-epidemic Station of Benxi City, LIAONING PROVINCE
Xue Wanqi— Diptera: Muscidae.

18. Sanitary and Epidemiological Station of Liaoning Province, LIAONING
PROVINCE

Ma Zhong-yu— Diptera: Anthomyiidae.

19. Shenyang Agricultural College, SHENYANG, LIAONING PROVINCE (Coul-
son— 79)

Ma Ning— Homoptera: Cicadellidae.

Wang Jinling— Hymenoptera: Trichogrammatidae.

Mr. Zhang Jing— Hymenoptera: Trichogrammatidae.

20. Nei-Monggol (Inner- Mongolian) College of Agriculture and Animal Husbandry,

HUHHOT, INNER MONGOLIA
Li Hong-Zhang (Lee Hung-Chang)— Coleoptera: Curculionidae.

21. North West Plateau Institute of Biology, Academia Sinica, YANGLING,
SHAANXI [SHENSI] PROVINCE

Yin Xiangchu— Orthoptera: Acrididae.

8

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

22. Academy of Agricultural Sciences, YANGLING, SHAANXI PROVINCE
Huo Shaotang— Hymenoptera: biotypes of Trichogramma.

23. Shaanxi Medical College, SHAANXI PROVINCE

Lu Yin-lian— Diptera: Muscidae, Calliphoridae, Sarcophagidae.

Wang Chang-jing— Diptera: Muscidae, Calliphoridae, Sarcophagidae.

24. Forest Research Institute, YANGLING, SHAANXI PROVINCE
Mr. Dang Xinde— Parasitic Hymenoptera.

Mr. Jin Buxian— Parasitic Hymenoptera.

25. Department of Biology, Shaanxi Normal University, XIAN, SHAANXI PROV-
INCE

Prof. Zheng Zhemin— Orthoptera: Acrididae, Tetrigidae.

26. Xian Medical College, XIAN, SHAANXI PROVINCE
Xiao Aixiang— Siphonaptera: Ischnopsyllidae.

27. Health and Anti-epidemic Station ofYanbei Region, SHAANXI PROVINCE
Wang Mingfu— Diptera: Muscidae.

28. Department of Plant Protection, Northwestern College of Agriculture, YANG-
LING, SHAANXI PROVINCE (Knutson and Gordon-81)

Mr. Chen Tong [Assistant]— Apterygota.

Prof. Dr. Chou Io (Zhou Yao)— Homoptera: Membracidae, Fulgoridae, Cercopi-
dae. Coccoidea: Diaspididae. Hymenoptera: Aphidiidae. Lepidoptera: Plusiinae,
Noctuidae. Apterygota. Mecoptera.

Mr. Liu Shaoyou [Lecturer] — Syrphidae (Syrphus and Sphaerophoria ).

Prof. Lu Jinsheng— Hemiptera: Nabidae.

Ms. Lu Zheng (Lu Tseng) (Mrs. Chou Io)— Homoptera: Cercopidae. Acari. Lepi-
doptera: Noctuidae.

Ms. Ran Ruibi [Lecturer] — Neuroptera: Chrysopidae.

Mr. Wei Jianhua [Lecturer]— Coleoptera: Coccinellidae.

Mr. Xiang Longcheng— Hymenoptera: Aphidiidae.

Mr. Yuan Feng [Lecturer] — Homoptera: Membracidae.

29. Lanzhou University, LANZHOU, GANSU [KANSU] PROVINCE
Wang Xin-e— Siphonaptera: Hystrichopsyllidae.

30. 201st Institute of Gansu Province, GANSU PROVINCE
Zhang Rongguang— Siphonaptera: Hystrichopsyllidae.

31. Institute of Forestry, Qinghai Academy of Agriculture & Forestry, QINGHAI
[CHINGHAI] PROVINCE

Xu Chenkuo (Hsu Chenkuo)— Lepidoptera: Lasiocampidae.

32. Qinghai Province (West) Institute of Biology, XINING, QINGHAI PROVINCE
Beng Guiying— Acari.

Chen Xiudao— Acari.

Su Tizhi— Acari.

1984

CHINESE S Y STEM ATISTS

9

33. Institute of Plant Protection, Sichuan Academy of Agriculture, CHENGDU, SI-
CHUAN [SZECHUAN] PROVINCE (Davis- 79)

Prof. Chen Fang-jie [Institute Director]— Homoptera: Coccidae.

Mr. Jiang Guang-Zao (K. T. Chiang)— Homoptera: Coccidae.

34. Southwestern Agricultural College, CHONGQING (=CHUNGKING), SI-
CHUAN PROVINCE (Coulson— 79)

Prof. Jiang Shunan (Chiang Shunan) [President of the College] — Coleoptera: Cer-
ambycidae.

Prof Li Longshu— Acari.

35. Citrus Research Institute, CHONGQING, SICHUAN PROVINCE

Yu Zhiren— Acari: Phytoseiidae. (Studied at University of California, Riverside,
1981-1982.)

36. Department of Parasitology, Chongqing Medical College, CHONGQING, SI-
CHUAN PROVINCE

Jeu Minghaw— Diptera: Ceratopogonidae.

Rong Yunlong— Diptera: Ceratopogonidae.

37. Division of Medical Zoology and Entomology, Health & Anti-epidemic Station
of Sichuan, SICHUAN PROVINCE

Chen Ningyu— Siphonaptera.

Wei Shufeng— Siphonaptera.

38. Department of Biology, Nanchung Teachers College, NANCHUNG, SICHUAN
PROVINCE

Mr. Zheng Fa-ke— Coleoptera: Staphylinidae.

39. Chengdu Institute of Termite Control, SICHUAN PROVINCE
Gong Anhu— Isoptera: Kalotermitidae.

40. Academy of Agricultural Sciences, SHANDONG [SHANTUNG] PROVINCE
Feng Jianguo— Hymenoptera: biotypes of Trichogramma.

41. Nanjing Agricultural College, JIANGSU [KIANGSU] PROVINCE
Ding Jin-Hua— Homoptera: Delphacidae.

Mr. Tian Li-Xin— Homoptera: Delphacidae.

42. Military Medical Research Institute, NANJING, JIANGSU PROVINCE
Wu Chenglin— Acari: Acariformes: Trombiculidae.

Wu Guanghua— Acari: Acariformes: Trombiculidae.

Yang Zhesheng— Acari: Acariformes: Trombiculidae.

43. Nanjing Institute of Termite Control, JIANGSU PROVINCE
Gao Daorong— Isoptera: Kalotermitidae.

Zhu Benzhong— Isoptera: Kalotermitidae.

44. Institute of Agriculture, NANTUNG, JIANGSU PROVINCE
You Qi-Ti— Hymenoptera: Ichneumonidae.

You Shi-Jin— Hymenoptera: Ichneumonidae.

10

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

45. Anhui Agricultural College, ANHUI [ANHWEI] PROVINCE
Prof. Ge Zhong-Lin (Ge Chung-Lin)— Homoptera: Delphacidae.

Meng Xuwu— Lepidoptera: Sphingidae.

46. Sanitarian and Anti-epidemic Station, HUBEI [HUPEH] PROVINCE
Zhang Di— Siphonaptera: Ceratophyllidae.

47. Hupei Medical College, Wuhan, HUBEI PROVINCE
Zhang Mengyu— Diptera: Sarcophagidae.

48. Kunming Institute of Zoology, Academia Sinica, KUNMING, YUNNAN
PROVINCE (Davis- 79)

Mr. Chao Wanyuan— Homoptera: Aphididae.

Mr. Gan Yunxing [Head of Division of Taxonomy] — Diptera: Calliphoridae. Lep-
idoptera: Noctuidae.

Mr. Kiong Tiang— Hemiptera: Coreidae.

49. Division of Forest Protection, Yunnan Forestry Institute, Hot Springs, KUN-
MING, YUNNAN PROVINCE

Wu Yi— Hymenoptera: Diprionidae.

50. Health and Epidemic Station of Jianchuan, YUNNAN PROVINCE
Yang Xue-Shi— Siphonaptera.

51. Control and Research Institute of Epidemic Diseases of Yunnan, YUNNAN
PROVINCE

Hu Gui— Siphonaptera.

Gong Zhengda— Siphonaptera: Hystrichopsyllidae, Ceratophyllidae.

Xie Pao-Ji (or Baoqi) (Hsieh, Pao-Chi) — Siphonaptera: Hystrichopsyllidae, Cer-
atophyllidae.

Yang Guang-Rong— Acari. Siphonaptera: Hystrichopsyllidae.

Yu Shi- Jin— Acari.

52. Guiyang (Kweiyang) Medical College, GUIZHOU [KWEICHOW] PROVINCE
Chen Hanbin— Diptera: Culicidae.

Prof. Jin Daxiong (Chin Ta-hsiung)— Anoplura: Enderleinellidae.

Prof. Li Guizhen (Li Kuei-chen)— Siphonaptera: Ceratophyllidae.

53. Guangdong Institute of Entomology, Insect Taxonomy Division, Academia Sin-
ica, GUANGZHOU [CANTON], GUANGDONG [KWANGTUNG] PROVINCE
(Coulson— 79, Davis— 79)

Mr. Chu Jie-lin— Isoptera.

Mr. Dai Zi-Rong— Isoptera.

Mr. Li Guixiang (Li Guei-shan)— Isoptera.

Ms. Li Liying (Mrs. Pang Xiongfei) [Director] — Hymenoptera: Trichogrammati-
dae.

Mr. Li Yan Chuyan [Curator of Collection] — General classification.

Mr. Lin Ping [Vice Director] — Coleoptera: Scarabaeidae: Rutellinae.

Lin Shanxiang— Collembola.

Mr. Ping Zhengming— Isoptera.

1984

CHINESE SYSTEMATISTS

11

Mr. Ren Huai [Research Assistant— Student] — Hymenoptera: Chalcidoidea, es-
pecially Chalcididae.

Mr. Wu Weinan— Acari: Phytoseiidae: Amblyseius.

54. Zhongshan University (=Chung San University or Sun Yat Sen University), De-
partment of Biology, Entomology Research Institute, GUANGZHOU, GUANG-
DONG PROVINCE (Coulson — 79, Davis- 79)

Chen Chunyao— Hemiptera: Scutelleridae, Pentatomidae.

Prof. Gjoan Yuwein (Djou Yu- Wen)— Hymenoptera: parasites of scale insects.
Hua Lizhong— Coleoptera: Cerambycidae.

Prof. Pu Zhelong (Pu Chih-Lung) [Department Chief]— Coleoptera: Hydrophilidae
and other aquatic Coleoptera.

55. Department of Plant Protection, South China Agricultural College, GUANG-
ZHOU, GUANGDONG PROVINCE (Coulson-79, Knutson and Gordon-81)

Mr. Chen Shoujian— Coleoptera: Carabidae (biological control of citrus insects).
Ms. Liu Chuanlu— Lepidoptera: Phyllocnistidae: Phyllocnista. Diptera: Muscidae.
Prof. Pang Xiongfei (Pang Hsui-fei) [Deputy Director] — Hymenoptera: Tricho-
grammatidae. Coleoptera: Coccinellidae. (Worked in Beltsville, Maryland in
1982 with A. C. F. Hung and R. D. Gordon, Insect Identification and Beneficial
Insect Introduction Institute, USDA.)

Mr. Zhang Weiqiu (Chang Wai-chiu)— Thysanoptera: Phlaeothripidae.

56. Jiangxi (=Kiangsi) Communist Labour University, JIANGXI [KIANGSI]
PROVINCE

Jin I-Shou— Hemiptera: Pentatomidae.

Prof. Zhang Shimei— Hemiptera: Pentatomidae.

57. Jiangxi University, Department of Biology, NANCHANG, JIANGXI (KIANG-
SI) PROVINCE

Dr. Chu Chimin— Acari: Phytoseiidae.

Dr. Jiang (Chiang) [President of the University]— Coleoptera: Cerambycidae.

Lin Yujian — Hemiptera: Pentatomidae.

58. Fujian Agricultural College, SHAXIAN, FUJIAN [FUKIEN] PROVINCE
Mr. Chen Jiahua— Hymenoptera: Aphidiidae (biological control of vegetable in-
sects).

Prof. Huang Bangkan— Coleoptera: Coccinellidae (biological control of fruit in-
sects).

Dr. Zhao Xiufu (Chao Hsiu-fu)— Hymenoptera: Ichneumonoidea (biological con-
trol of rice insects).

59. Zhejiang Agricultural University, Department of Plant Protection, HANG-
ZHOU, ZHEJIANG [CHEKIANG] PROVINCE (Coulson-79, Knutson and Gor-
don-81)

Mr. He Junhua (He Chuen-hua)— Hymenoptera: Ichneumonoidea: Braconidae
(biological control of rice insects).

Prof Li Xueliu (Lee H. L.)— Hymenoptera: Chalcidoidea: Encyrtidae, Aphelinidae
(biological control of citrus insects).

Prof. Tang Jiao— Hymenoptera: Formicidae.

12

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

60. Zhejiang Provincial Museum, HANGZHOU, ZHEJIANG PROVINCE
Mr. Cai Chanmo.

Mr. Zhou Wenbao— Odonata. Coleoptera: Cerambycidae.

Complete addresses for the following individuals have not been located. The in-
formation provided here, however, may be of use to some researchers.

61. Sanitary and Epidemiological Station of Henan Province, ZHENGZHOU,
HENAN [HONAN] PROVINCE

Ge Feng-Xiang— Diptera: Anthomyiidae.

62. Team of Epidemic Prevention, Logistics Department of Xizang Military Region,
PLA, LASA, XIZANG AUTONOMOUS REGION

Wu Fu-lin— Diptera: Calyptrata.

Zhong Yin-Hong— Diptera: Calyptrata.

63. Institute of Tropical Forestry, Chinese Academy of Forest Science (city and prov-
ince unknown)

Chen Zhi-qing— Orthoptera: Acrididae.

TAXONOMIC INDEX

Numbers in parentheses following researchers’ names correspond to numbers as-
signed in previous section to institutions. Hyphens have been omitted from the second
(given) name.

Acari

Acaridae: Wang Xiaozu (6)

Cheyletidae: Wang Xiaozu (6)

Eriophyidae: Ding Tingzhong (7); Dong Huiqing (7)

Gamasoidea: Teng Kuofan (1)

Glycyphagidae: Wang Xiaozu (6)

Ixodoidea: Teng Kuofan (1)

Phytoseiidae: Chu Chimin (57); Ke Lisheng (7); Liang Lairong (7); Wu Weinan
(53) ( Amblyseius ); Yu Zhiren (35)

Rhodacaridae: Ma Liming (15)

Tarsonemidae: Ding Tingzhong (7)

Tetranychidae: Ma Enpei (8); Wang Xiangfu (1); Yuan Yilan (1 1)

Trombiculidae: Hsu Yinchi (9) ( Demodex ); Wen Tinghuan (9); Wu Chenglin (42);

Wu Guanghua (42); Xin Jieliu (7) (Trombidiidae); Yang Zhesheng (42)

Acari, General: Beng Guiying (32); Chen Xiudao (32); Li Longshu (34); Lu Zheng
(28); Su Tizhi (32); Yang Guangrong (51); Yu Shijin (51); Zhang Fengling (16)
Apterygota

Apterygota, General: Chen Tong (28); Chou Io (28)

Protura

Eosentomidae: Yin Wenying (6); Zhang Zhiyuan (6)

Protura, General: Huang Fusheng (1)

Collembola

Collembola, General: Lin Shanxiang (53)

1984

CHINESE SYSTEMATISTS

13

Odonata

Odonata, General: Zhou Wenbao (60)

Orthoptera

Acrididae: Chen Zhiqing (63); Liu Jupeng (1); Yin Xiangchu (21); Zhang Fengling
(16); Zheng Zhemin (25)

Acridoidea: Bi Daoying (6); Jin Xingbao (6); Xia Kailing (6)

Tetrigidae: Zheng Zhemin (25)

Isoptera

Kalotermitidae: Gao Daorong (43); Gong Anhu (39); Zhu Benzhong (43)
Rhinotermitidae: Fan Shude (6); Xia Kailing (6)

Termitidae: He Xiusong (6); Xia Kailing (6)

Isoptera, General: Chu Jielin (53); Dai Zirong (53); Han Meizhen (6); Li Guixiang
(53); Peng Zengming (53); Ping Zhengming (53)

Zoraptera

Zoraptera, General: Huang Fusheng (1)

Anoplura

Enderleinellidae: Jin Daxiong (52)

Anoplura, General: Teng Kuofan (1)

Thysanoptera

Thysanoptera, General: Zhang Guangxue (1)

Phlaeothripidae: Zhang Weiqiu (55)

Hemiptera

Acanthosomatidae: Liu Shengli (14)

Aphidoidea: Zhu Hongfu (1)

Coreidae: Kiong Tiang (48)

Lygaeidae: Zheng Leyi (12)

Nabidae: Lu Jinsheng (28)

Pentatomidae: Chen Chunyao (54); Jin Ishou (56); Lin Yujian (57); Zhang Shimei
(56)

Scutelleridae: Chen Chunyao (54)

Hemiptera, General: Jin Qinying (6)

Homoptera

Aleyrodidae: Yang Pinglan (6)

Aphididae: Chao Wanyuan (48)

Aphidoidea: Zhang Guangxue (1); Zhong Tiesen (1)

Cercopidae: Chou Io (28); Lu Zheng (28)

Cicadellidae: Ma Ning (19)

Coccidae: Chen Fangjie (33); Jiang Guangzao (33); Wang Ziqing (1)

Delphacidae: Ding Jinhua (41); Ge Zhonglin (45); Tian Lixin (41)

Diaspididae: Chou Io (28); Hu Jinglin (6); Yang Pinglan (6)

Fulgoridae: Chou Io (28)

Membracidae: Chou Io (28); Yuan Feng (28)

Neuroptera

Chrysopidae: Ran Ruibi (28); Yang Jikun (2) (and other beneficial insects)
Coleoptera

Bruchidae: Li Hongxing (1)

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Buprestidae: Hou Taogion (1)

Carabidae: Chen Shoujian (55) (biological control of citrus pests); Yu Peiyu (1)
Cerambycidae: Hua Lizhong (53); Jiang (57); Jiang Shunan (34); Pu Fuji (1); Zhou
Wenbao (60)

Chrysomelidae: Chen Shixiang (1); Yu Peiyu (1) (Criocerinae)

Cicindelidae: Tan Zhuanjie (1)

Coccinellidae: Huang Bangkan (58) (biological control of fruit insects); Pang Xiong-
fei (55); Wei Jianhua (28)

Curculionoidea: Chen Yuanquing (1); Li Hongxing (1); Li Hongzhang (20)
Eumolpidae: Tan Zhuanjie (1)

Hydrophilidae: Pu Zhelong (54) (and other aquatic Coleoptera)

Meloidae: Tan Zhuanjie (1)

Platypodidae: Yin Huifen (1)

Scarabaeidae: Lin Ping (53)

Scarabaeoidea: Chang Youwei (1); Lin Ping (53) (Rutellinae)

Scolytidae: Huang Fusheng (1); Yin Huifen (1)

Staphylinidae: Zheng Fake (38)

Stylopidae: Yang Jikun (2)

Mecoptera

Mecoptera, General: Chou Io (28)

Lepidoptera

Agaristidae: Chen Yixin (4)

Amatidae: Fang Chenglai (1)

Arctiidae: Fang Chenglai (1)

Bombycidae: Zhu Hongfu (1)

Brahmaeidae: Zhu Hongfu (1)

Callidulidae: Zhu Hongfu (1)

Cochylidae: Liu Yujiao (1)

Cossidae: Chen Yixin (4)

Drepanidae: Zhu Hongfu (1)

Endromidae: Zhu Hongfu (1)

Episopeiidae: Zhu Hongfu (1)

Ethmiidae: Liu Yujiao (1)

Geometridae: Zhu Hongfu (1)

Hepialidae: Zhu Hongfu (1)

Hypsidae: Fang Chenglai (1)

Lasiocampidae: Hou Taogion (1); Xu Chenkuo (31)

Limacodidae: Cai Rongquan (1)

Lymantriidae: Chao Chungling (1)

Noctuidae: Chen Yixin (4); Chou Io (28); Fang Chenglai (1) (larvae and pupae);

Gan Yunxing (48); Lu Zheng (28)

Notodontidae: Cai Rongquan (1)

Papilionidae: Li Chuanlung (1)

Phyllocnistidae: Liu Chuanlu (55) ( Phyllocnista )

Plusiinae: Chou Io (28)

Psychidae: Chao Chungling (1)

Pterothysanidae: Zhu Hongfu (1)

1984

CHINESE SYSTEMATISTS

15

Pyralidae: Li Chengzhang (6); Wang Pingyuan (1)

Rhopalocera: Li Chuanlung (1)

Satumiidae: Zhu Hongfu (1)

Sphingidae: Meng Xuwu (45); Zhu Hongfu (1)

Thaumetopoeidae: Hou Taogion (1)

Thyatridae: Chao Chungling (1)

Tortricidae: Liu Yujiao (I)

Uraniidae: Zhu Hongfu (1)

Zygaenidae: Wang Pingyuan (1)

Diptera

Anthomyiidae: Chen Zhizi (6); Ge Fengxiang (61); Ma Zhongyu (18); Sun Cai-
hong (1)

Asilidae: Xia Kailing (6)

Calliphoridae: Chen Zhizi (6); Fan Zide (6) (especially synanthropic calyptrate
Diptera); Gan Yunxing (48); Lu Yinlian (23); Sun Caihong (1); Wang Changjing
(23)

Calyptrata: Wu Fulin (62); Zhong Yinhong (61)

Ceratopogonidae: Jeu Minghaw (36); Li Tiesheng (1); Li Peisi (6); Rong Yunlong
(36); Wang Taihua (4); Xu Fengyi (10); Yu Yixin (4); Yu Yixin (4); Zhang Zhi-
yuan (6)

Culicidae: Chen Hanbin (52); Li Peisi (4); Liu Weide (6); Lu Baolin (4); Ma Sufang
(1); Xu Fengyi (10)

Muscidae: Chen Zhizi (6); Fan Zide (6); Liu Chuanlu (55); Lu Yinlian (23); Wang
Mingfu (27); Xue Wanqi (17); Sun Caihong (1); Wang Changjing (23)
Rhagionidae: Yang Jikun (2)

Sarcophagidae: Chen Zhizi (6); Fan Zide (6) (not including Tachinidae); Lu Yinlian
(23); Wang Changjing (23); Zhang Mengyu (47); Zhao Jianming (1)

Syrphidae: Fan Zide (6); Liu Shaoyou (28) ( Syrphus and Sphaerophoria ); Shi Dasan
(6); Shi Yongshan (1); Sun Caihong (1)

Tabanidae: Liu Weide (6); Wang Zunming (1); Yao Yunmei (6)

Tachinidae: Shi Yongshan (1); Zhao Jianming (1); Yang Xueshi (50)
Siphonaptera

Ceratophyllidae: Gong Zhengda (51); Li Guizhen (52); Liu Chiying (4); Liu Quang
(4); Xie Paoji (51); Zhang Di (46)

Hystrichopsyllidae: Gong Zhengda (51); Wang Xine (20); Wu Houyong (4); Xie
Paoji (51); Yang Guangrong (51); Zhang Rongguang (30)

Ischnopsyllidae: Xiao Aixiang (26)

Siphonaptera, General: Chen Ningyu (37); Dou Guilan (4); Hu Gui (51); Ji Shuli
(4); Ren Shiming (4); Wei Shufeng (37)

Hymenoptera

Aphidiidae: Chen Jiahua (58) (biological control of vegetable insects); Chou Io
(28); Shi Dasan (6); Xiang Longcheng (28)

Apoidea: Wang Sufang (1) ( Bombus ); Wu Yanru (1)

Braconidae: He Junhua (59) (biological control of rice insects); Wang Jinyan (1)
Chalcidoidea: Li Xueliu (59) (Encyrtidae, Aphelinidae; biological control of citrus
insects); Liao Dingxi (1); Ren Huai (53) (especially Chalcididae)

Diprionidae: Wu Yi (49)

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Formicidae: Tang Jiao (59)

Ichneumonidae: Wang Sufang (1); You Shijin (44); You Qiti (44)

Ichneumonoidea: He Junhua (59); Zhao Xiufu (58) (biological control of rice pests)

Proctotrupoidea: Wu Yanru (1)

Tenthredinoidea: Huang Xiaoyun (3); Li Guanwu (3); Xiao Gangrou (3); Zhou
Shushi (3)

Trichogrammatidae: Chen Tailu (1) ( Trichogramma)', Feng Jianguo (40) (biotypes
of Trichogramma ); Huo Shaotang (22) (biotypes of Trichogramma)', Li Liying
(53); Pang Xiongfei (55); Shi Dasan (6) ( Trichogramma ); Wang Jinling (19);
Zhang Jing (19)

Vespoidea: Li Tiesheng (1)

Parasitic Hymenoptera: Dang Xinde (24); Gjoan Yuwein (54) (parasites of scale
insects); Jin Buxian (24)

Miscellaneous

Classification, General: Li Yan Chuyan (53)

Entomology, General: Liu Sinan (5); Wang Xicheng (5)

Fossil Insects: Tan Zhuanjie (1)

Taxonomy of Stored Product Pests (especially Coleoptera) and Quarantine: Guan
Lianghua ( 1 3)

Unknown Speciality: Cai Chanmo (60)

SPECIALIST INDEX

Chen Yuanquing (1)

Chu Chimin (57)
Chu Jielin (53)

Chou Io (28)

Chen Zhiqing (63)
Chen Zhizi (6)

Chen Tong (28)
Chen Xiudao (32)
Chen Yixin (4)

Cai Chanmo (60)
Cai Rongquan (1)
Chang Youwei (1)
Chao Chungling ( 1 )

Chen Chunyao (54)
Chen Fangjie (33)
Chen Hanbin (52)
Chen Jiahua (58)
Chen Ningyu (37)
Chen Shixiang (1)
Chen Shoujian (55)
Chen Tailu (1)

Chao Wanyuan (48)

Beng Guiying (32)
Bi Daoying (6)

Dai Zirong (53)
Dang Xinde (24)
Ding Jinhua (41)
Ding Tingzhong (7)
Dong Huiqing (7)
Dou Guilan (4)

Fan Shude (6)

Fan Zide (6)

Fang Chenglai (1)
Feng Jianguo (40)
Gan Yunxing (48)
Gao Daorong (43)
Ge Fengxiang (61)
Ge Zhonglin (45)
Gjoan Yuwein (54)
Gong Anhu (39)
Gong Zhengda (51)
Guan Lianghua (13)
Han Meizhen (6)

He Junhua (59)

He Xiusong (6)

Hou Taogion (1)
Hsu Yinchi (9)

Hu Gui (51)

1984

CHINESE SYSTEMATISTS

17

Hu Jinglin (6)

Hua Lizhong (53)
Huang Bangkan (58)
Huang Fusheng (1)
Huang Xiaoyun (3)
Huo Shaotang (22)
Jeu Minghaw (36)

Ji Shuli (4)

Jiang (57)

Jiang Guangzao (33)
Jiang Shunan (34)
Jin Buxian (24)

Jin Daxiong (52)

Jin Ishou (56)

Jin Qinying (6)

Jin Xingbao (6)

Ke Lisheng (7)
Kiong Tiang (48)

Li Chengzhang (6)

Li Chuanlung ( 1 )

Li Guanwu (3)

Li Guixiang (53)

Li Guizhen (52)

Li Hongxing (1)

Li Hongzhang (20)
Li Liying (53)

Li Longshu (34)

Li Peisi (4)

Li Tiesheng (1)

Li Xueliu (59)

Li Yan Chuyan (53)
Liang Lairong (7)
Liao Dingxi (1)

Lin Ping (53)

Lin Shanxiang (53)
Lin Yujian (57)

Liu Chiying (4)

Liu Chuanlu (55)

Liu Jupeng (1)

Liu Quang (4)

Liu Shaoyou (28)

Liu Shengli (14)

Liu Sinan (5)

Liu Weide (6)

Liu Yujiao (1)

Lu Baolin (4)

Lu Jinsheng (28)

Lu Yinlian (23)

Lu Zheng (28)

Ma Enpei (8)

Ma Liming (15)

Ma Ning (19)

Ma Sufang (1)

Ma Zhongyu ( 1 8)
Meng Xianling (1)
Meng Xuwu (45)
Pang Xiongfei (55)
Ping Zhengming (53)
Pu Fuji (1)

Pu Zhelong (54)

Ran Ruibi (28)

Ren Huai (53)

Ren Shiming (4)
Rong Yunlong (36)
Shi Dasan (6)

Shi Yongshan (1)

Su Tizhi (32)

Sun Caihong (1)

Tan Zhuanjie (1)
Tang Jiao (59)

Teng Kuofan (1)

Tian Lixin (41)

Wang Changjing (23)
Wang Jinling (19)
Wang Jinyan (1)
Wang Mingfu (27)
Wang Pingyuan (1)
Wang Sufang (1)
Wang Taihua (4)
Wang Xiangfu (1)
Wang Xiaozu (6)
Wang Xicheng (5)
Wang Xine (29)
Wang Ziqing (1)
Wang Zunming (1)
Wei Jianhua (28)

Wei Shufeng (37)
Wen Tinghuan (9)
Wu Chenglin (42)

Wu Fulin (62)

Wu Guanghua (42)
Wu Houyong (4)

Wu Weinan (53)

Wu Yanru (1)

18

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Wu Yi (49)

Xia Railing (6)

Xiang Longcheng (28)
Xiao Aixiang (26)
Xiao Gangrou (3)

Xie Paoji (51)

Xin Jieliu (7)

Xu Chenkuo (31)

Xu Fengyi (10)

Xue Wanqi (17)

Yang Guangrong (51)
Yang Jikun (2)

Yang Pinglan (6)

Yang Xueshi (50)
Yang Zhesheng (42)
Yao Yunmei (6)

Yin Huifen (1)

Yin Wenying (6)

Yin Xiangchu (21)
You Qiti (44)

You Shijin (44)

Yu Peiyu (1)

Yu Shijin (51)

Yu Yixin (4)

Yu Zhiren (35)

Yuan Feng (28)

Yuan Yilan (11)

Zhang Di (46)

Zhang Fengling ( 1 6)
Zhang Guangxue (1)
Zhang Jing (19)

Zhang Mengyu (47)
Zhang Rongguang (30)
Zhang Shimei (56)
Zhang Weiqiu (55)
Zhang Zhiyuan (6)
Zhao Jianming (1)
Zhao Xiufu (58)

Zheng Fake (38)

Zheng Leyi ( 1 2)

Zheng Zhemin (25)
Zhong Tiesen (1)
Zhong Yinhong (61)
Zhou Shushi (3)

Zhou Wenbao (60)
Zhu Hongfu (1)

Received September 15, 1983; accepted October 27, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 19-26

VEGETATIONAL COMPLEXITY AND
PARASITISM OF GREEN PEACH APHIDS
(MYZUS PERSICAE (SULZER) (HOMOPTERA: APHIDAE))
ON COLLARDS

David J. Horn

Department of Entomology, The Ohio State University,
Columbus, Ohio 43210

Abstract. — In 1 978-1979, parasitized green peach aphids (GPA) were recovered from collards
in adjacent weedy and weedless habitats. During each season, 22 percent of GPA were para-
sitized, primarily in weedless plots wherein Diaeretiella rapae comprised 94 percent of primary
parasitism. Parasitism by Aphidius spp. was concentrated on GPA from collards in weedy plots.
Secondary parasitism occurred much more frequently in weedless plots, wherein 59 percent of
primary parasitoids were parasitized. Secondary parasitoids were: Asaphes lucens (53 percent
of all parasitoids recovered), Aphidencyrtus aphidivorus (34 percent), Alloxysta nr. megourae
and nr. discreta (7 percent), Phaenoglyphis ambrosiae and n. sp. (3 percent), Dendrocerus niger
(3 percent) and D. incompletus (< 1 percent); A. lucens was more abundant in 1978 than 1979
while the reverse was noted in Alloxysta spp. Secondary parasitoids were evidently attracted
to higher densities of primary hosts on collards located in weedless plots.

Influence of vegetational complexity on interactions between parasitic Hymenop-
tera and their hosts is a subject of ongoing interest. A widely-held tenet is that added
environmental complexity may impart stability to otherwise disequilibrial systems.
Such increased complexity may be spatial heterogeneity (Huffaker, 1958), addition
of predators (Paine, 1966), vegetational diversity (Pimentel, 1961) or some combi-
nation of these. In agricultural production there is increasing interest in development
of minimum-tillage systems that increase complexity by encouraging the growth of
ground cover. Little information exists on the effects of reduced tillage on insect
host-parasitoid interactions. The present study reports initial results on the effects
of weed cultivation on parasitism of green peach aphids (hereinafter GPA) by Aphi-
diidae (Hymenoptera) on collards ( Brassica oleracea var.) in central Ohio. Weeds
were expected to provide greater availability of alternate hosts and nectar sources
for adult parasitoids. Van Emden (1963) found densities of Ichneumonidae to be
higher in weedy than in tilled plots, and Copland (1 979) noted reduction in secondary
parasitism of hop aphids when weeds were eliminated. I (Horn, 1981) found that
densities of aphidophagous Coccinellidae, Chrysopidae, and Syrphidae were higher
on collards surrounded by unmanaged weeds despite lower densities of GPA there.
When weeds were regularly trimmed to 10 cm maximum height, predators moved
to more dense populations of GPA on adjoining weedless collards. In the present
study I therefore expected that primary and secondary parasitism on GPA might
increase when weeds were uncontrolled.

20

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

METHODS

Fieldwork was conducted on the Horticultural Farm, Ohio State University. Col-
lards (var. “Georgia”, Ferry-Morse Seed Co.) were started in a greenhouse, and during
early May were transplanted into a 9 m x 28 m area outdoors. Transplants were 50
cm apart (later thinned to 1 m) in 1 m rows. The experimental area was subdivided
into 8 4 m x 7 m plots. Plots were tilled weekly by machine and hand. The other 4
plots were untilled; the ground cover 1 July was Amaranthus retroflexus L. (65
percent) and Chenopodium album L. (20 percent) with scattered Xanthium stru-
marium L., Portulaca oleracea, Polygonum spp., and grasses.

Four randomly-selected collard plants per plot were sampled weekly from 1 5 May
through 20 July 1978. 1 counted all GPA and collected all parasitized mummies into
gelatin capsules (#000) which were held at 25°C and 1 5 hr photophase for emergence
of parasitoids. Mummies from which had emerged Chalcidoidea or Cynipoidea were
dissected to verify evidence of secondary parasitism. Voucher specimens of each
species were deposited in the Ohio State University Collection of Insects.

Overhead sprinklers irrigated the area when necessary. Surrounding areas were
planted to onions, squash, melons and tomatoes, which were treated heavily and
repeatedly with carbaryl and malathion. This may have limited immigration of GPA
and parasitoids from adjacent crops.

In 1979 I followed similar procedures except as follows: Weeds were trimmed
weekly to a maximum height of 10 cm, and weedless plots were kept free of weeds
by black plastic placed over the soil surface between rows. Plots were 4 m x 8 m,
and sampling continued through August. Major weeds in 1979 were grasses (60
percent), Chenopodium album, Cirsium spp., Ambrosia artemisiifolia L., and Polyg-
onum spp. The experiment was located in a community garden with local and in-
termittent insecticide application, probably allowing immigration of GPA and para-
sitoids from nearer sources than in 1978.

RESULTS

Figure 1 indicates abundance of living aphids on collards in weedy and weedless
plots. All aphids were GPA except late in 1978 when there was a small influx of
cabbage aphids ( Brevicoryne brassicae L.). Cultivated collards, contrasting with either
bare soil or black plastic, were more readily colonized by alate GPA (Horn, 1981)
and their higher intensities on weedless collards reflect this. Subsequent production
of apterous GPA resulted in significantly higher total intensity of functioning aphids
on weedless collards especially in 1979.

Eleven species of parasitic Hymenoptera were recovered from mummified GPA
during this study. Diaeretiella rapae McIntosh accounted for 94 percent of all primary
parasitism. It is a solitary, cosmopolitan endoparasitoid of many species of crucifer-
infesting aphids (Read et al., 1970). Figure 2 shows seasonal trends in primary
parasitism on GPA within weedy and weedless subplots. In 1978, there appeared no
significant difference in percent parasitism between habitats, though total parasitism
was higher on weedless collards, where more hosts were present (Fig. 1). Aphidius
spp. were recovered almost exclusively from weedy collards. In 1979, these trends
were more pronounced; parasitism was very low on GPA from weedy collards.

Figure 3 depicts secondary parasitism on D. rapae. Table 1 lists the complex of

Total aphids, 16 plants Total aphids, 16 plants

1984

VEGETATIONAL COMPLEXITY

21

JUNE JULY

Fig. 1 . Total living Myzus persicae on weedy and weedless collards, Columbus, Ohio. 1978:
weeds not trimmed. 1979: weeds trimmed to 10 cm tall.

22

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Fig. 2. Percent parasitism by Aphidiidae on Myzus persicae. Proportion of parasitism due
to Diaeretiella rapae in weedy, 84 percent in 1978, 98 percent in 1979; in weedless, 100 percent
each year.

% secondary parasitism % secondary parasitism

1984

VEGETATIONAL COMPLEXITY

23

JUNE JULY

JUNE JULY AUG

Fig. 3. Percentage of Aphidiidae secondarily parasitized on weedy and weedless collards.

24

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Table 1. Secondary parasitoids recovered from parasitized mummies of green peach aphid,
Columbus, Ohio, 1978-1979.

1978

1979

Parasitoid

Weedy

Weedless

Weedy

Weedless

Pteromalidae

Asaphes lucens Provancher
Encyrtidae

17

209

12

209

Aphidencyrtus aphidivorus Mayr
Alloxystidae

2

58

0

226

Alloxysta nr. megourae (Ashmead)
Alloxysta nr. discreta (Foerster)

0

16

0

39

Phaenoglyphis spp.
Megaspilidae

0

2

0

20

Dendrocerus incompletus Muesebeck

0

0

0

4

D. niger (Howard)

J_

6

_0

23

Totals

20

291

12

521

secondary parasitoids. The secondary parasitoids occurred primarily in weedless
habitats. There was little seasonal succession in the complex, except that A. aphid-
ivorus was earliest to arrive (mid- June) and Alloxystidae and Dendrocerus spp. were
limited to when GPA was most abundant (July).

DISCUSSION

The greater intensity of GPA on weedless collards apparently reflects the ease with
which the plants are located. Their contrast with backgrounds of bare soil or black
plastic enhances their attractiveness to colonizing GPA. The weedy plots harbor a
variety of aphidophagous predators whose activity serves to further depress GPA
populations on collards in weedy plots (Horn, 1981).

The complex of parasitoids reported here is similar to that found elsewhere on
Brassica (Chua, 1977; Takada, 1976a, b) and other crops regardless of vegetational
complexity (Copland, 1979; Jones, 1979; Sullivan and van den Bosch, 1971; van
den Bosch et al., 1979). Usually there are a few species of Aphidiidae, one or two
dominant secondary parasitoids (often Asaphes or Alloxysta), and several minor
secondary parasitoids.

In my experiments parasitism of GPA by D. rapae was concentrated on weedless
collards, especially when weeds in adjacent plots were trimmed (1979). Proximity
of weedy and weedless plots may have permitted these widely-ranging parasitoids to
quickly locate, and concentrate oviposition on, GPA populations of higher intensity
on weedless collards. Parasitism by Aphidius spp., by contrast, occurred mostly in
weedy plots, where alternate hosts were available, especially on Chenopodium. De-
spite a greater diversity of primary parasitoids, a lower percentage parasitism resulted,
at least in 1979, on GPA in the vegetationally more diverse habitat.

Secondary parasitoids occasionally parasitize one another (Bennett and Sullivan,
1978; Griswold, 1923), so that estimates based on rearing may not accurately reflect

1984

VEGET ATIONAL COMPLEXITY

25

the relative abundance of searching parasitoids in the field. However, excepting A.
lucens, there was very little secondary parasitism of Aphidiidae on collards in weedy
plots (Table 1). Higher GPA densities on collards in weedless plots led to increased
primary parasitism which in turn supported higher secondary parasitism and a more
diverse array of parasitoids. The proximity of plots may have contributed, as rela-
tively mobile insects could cross the 1 m gap rather easily. Higher densities of par-
asitized GPA concentrated an attractive resource for secondary parasitoids. Root
(1973) found diversity of all Hymenoptera to be higher in collard monoculture than
on collards planted within an old-field habitat. Vegetational complexity, therefore,
might dilute the apparency of primary parasitoids such as D. rapae rather as it hides
collards from colonizing GPA.

ACKNOWLEDGMENTS

The Department of Horticulture, Ohio State University, provided field space and the Graduate
School, OSU, provided funding for parts of the study. I was assisted afield by D. Dix, D. L.
Faustini, C. A. and R. B. Horn, and S. Thrall. Hymenoptera were identified by: E. Grissell
(Encyrtidae, Pteromalidae), P. Marsh (Aphidiidae, Megaspilidae), F. Andrews and A. Menke
(Alloxystidae). J. Briggs’ collection of obscure reprints was valuable, as was the hospitality of
L. Ehler and the University of California, Davis. The manuscript has been improved by com-
mentaries of R. Dowell, L. Ehler, R. Hall, R. S. Horn, D. Simonet, G. Stairs, and G. Walker.

LITERATURE CITED

Bennett, A. W. and D. J. Sullivan, S.J. 1978. Defensive behavior against tertiary parasitism
by the larvae of Dendrocerus carpinteri, an aphid hyperparasitoid. J. New York Ent. Soc.
86:153-160.

Chua, T. H. 1977. Population studies of Brevicoryne brassicae (L.), its parasites and hyper-
parasites in England. Res. Pop. Ecol. 19:125-140.

Clausen, C. P. 1940. Entomophagous Insects. McGraw-Hill Book Co., New York.

Copland, M. J. W. 1 979. Hymenoptera in hop gardens, with particular reference to parasitoids
associated with the damson-hop aphid, Phorodon humuli. Entomologia Experimentalis
et Applicata 25:146-152.

Emden, H. F. van. 1963. Observations on the effects of flowers on the activity of parasitic
Hymenoptera. Ent. Mon. Mag. 98:265-270.

Force, D. C. 1974. Ecology of insect host-parasitoid communities. Science 184:624-632.

Griswold, G. H. 1929. On the bionomics of a primary parasite and of two hyperparasites of
the geranium aphid. Ann. Ent. Soc. Amer. 22:438-457.

Gutierrez, A. P. and R. van den Bosch. 1 970. Studies on the host selection and host specificity
of the aphid hyperparasite Charips victrix (Hymenoptera: Cynipidae) 1 . Review of hy-
perparasitism and the field ecology of Charips victrix. Ann. Ent. Soc. Amer. 63:1345-
1354.

Horn, D. J. 1981. Effect of weedy backgrounds on colonization of collards by green peach
aphid, Myzus persicae, and its major predators. Environ. Ent. 10:285-289.

Huffaker, C. B. 1958. Experimental studies on predation: dispersion factors and predator-
prey oscillations. Hilgardia 27:343-383.

Jones, M. G. 1979. Abundance of aphids on cereals from before 1973 to 1977. J. Appl. Ecol.
16:1-22.

Paine, R. T. 1966. Food web complexity and species diversity. Amer. Nat. 100:65-75.

26

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Pimentel, D. 1961. Natural control of aphid populations on cole crops. Ann. Ent. Soc. Amer.
54:885-888.

Read, D. R., P. P. Feeny and R. B. Root. 1970. Habitat selection by the aphid parasite
Diaeretiella rapae (Hymenoptera:Braconidae) and hyperparasite Charips brassicae (Hy-
menoptera:Cynipidae). Can. Ent. 102:1567-1578.

Root, R. B. 1973. Organization of a plant-arthropod association in simple and diverse habitats:
the fauna of collards. Ecol. Monogr. 43:95-124.

Sedlag, U. 1958. Untersuchungen iiber Bionomie und Massenweschel von Diaeretiella rapae
(Curtis) (Hymenoptera: Aphidiidae). Transactions of the First Conference on Insect
Pathology and Biological Control, Prague, pp. 367-373.

Sekhar, P. 1958. Studies on Asaphes Jletcheri (Crawford), a hyperparasite of Aphidius testa-
ceipes (Cresson) and Praon aguti (Smith), primary parasites of aphids. Ann. Ent. Soc.
Amer. 51:1-7.

Sullivan, D. J. and R. van den Bosch. 1971. Field ecology of the primary parasites and
hyperparasites of the potato aphid, Macrosiphon euphorbiae, in the east San Francisco
Bay area. Ann. Ent. Soc. Amer. 64:389-394.

Takada, H. 1976a. Studies of aphids and their parasites on cruciferous crops and potatoes.

I. Parasite complex of aphids. Kontyu 44:234-253.

Takada, H. 1976b. Studies of aphids and their parasites on cruciferous crops and potatoes.

II. Life cycle. Kontyu 44:366-384.

van den Bosch, R. et al. 1979. Biological control of the walnut aphid in California: impact
of the parasite, Trioxys pallidus. Hilgardia 47:1-13.

Received March 2, 1983; accepted December 9, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 27-34

THE MEMBRACIDAE AND OTHER HOMOPTERA DESCRIBED
BY ASA FITCH, 1851, AND EBENEZER EMMONS, 1855:
HISTORICAL PERSPECTIVE AND ANALYSIS1

Jeffrey K. Barnes

Biological Survey, New York State Museum,

The State Education Department, Albany, New York 12230

Abstract.— The historical background to Fitch’s and Emmons’ publications on Homoptera
is discussed. Correct publication dates and authorships of new taxa described in Fitch’s Ho-
moptera catalogue and Emmons’ report on injurious insects are established using information
derived from recently discovered Fitch manuscript notes and correspondence and from obscure
government documents. Collecting data and specimen numbers are provided for types of new
species of Membracidae described in Emmons’ report. Types of Homoptera described in Fitch’s
catalogue are discussed briefly.

Since the death of Dr. Asa Fitch in 1879, several papers dealing with his type
specimens of Membracidae and other Homoptera have appeared (Sanborn and Lint-
ner, 1879; Goding, 1893; Lintner, 1893; Osborn, 1905; Felt, 1910; Sanders, 1910;
Funkhouser, 1915; Woodruff, 1924; McCabe and Johnson, 1980). Inaccuracies in
this literature have raised questions about the true dates of publication of the species
names, the correct authorship of new species described in Emmons’ Insects of New
York, and the identity of type specimens of Fitch’s and Emmons’ species. Archival
research has helped to resolve this confused situation.

HISTORICAL BACKGROUND

On April 15, 1836, the New York State Legislature directed the Governor to employ
competent persons to make a geological survey of the State, and to “furnish a full
and scientific description of its rocks, soils and minerals, and of its botanical and
zoological productions” (Laws of New York, Chapter 142). The reports of this early
survey were published in the voluminous series entitled “Natural History of New
York.” James E. DeKay was placed in charge of the zoological department of the
survey, but he chose not to report on the insects (Assembly Document 137, 1855),
despite Asa Fitch’s desire for that assignment.2 Ebenezer Emmons was given charge
of the agricultural portion of the survey, and with the assistance of Dr. Fitch he
reported on the insects of interest to farmers. Dr. Fitch also responded to a request
from the Regents of the University of the State of New York that he build a collection
of insects for the State Cabinet of Natural History. He authored two catalogues based
on specimens in that collection (Fitch, 1849, 1851).

Many interesting details concerning the history of Emmons’ report and Fitch’s

1 Published as New York State Museum Journal Series Number 416.

2 Fitch to William S. Robertson, letter dated March 26, 1840; in Fitch’s Diary 8, Yale
University Library.

28

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

work on Homoptera can be gleaned from Fitch’s letters to Thaddeus William Harris.3
Dr. Harris was librarian of Harvard University and an eminent entomologist. Mallis
(1971) stated that Fitch and Harris apparently had no dealings with each other, but
this series of correspondence attests differently.

“At present I am engaged in preparing some account of our noxious insects for
Dr. Emmons’ forthcoming volume on the Agricultural Geology of the state, which
is to form one of the ‘by authority’ quartos of the State Natural History. The season
for making personal observations was nearly past when Dr. E. first intimated a desire
for a paper of this kind; consequently I am obliged to borrow much from books, and
particularly from your invaluable ‘Report.’ I trust, however, that no one on this or
any other occasion, will have grounds for accusing me of borrowing without giving
due credit. In this paper I intend describing several new species, mostly of the sub-
order Homoptera: —the time allowed for completing the article will scarcely enable
me to work out much more than this that is new.” [Letter of December 30, 1846.]

“. . . I will, as I now think revise the Homoptera, and devote next year assiduously
to further collecting, studying and describing our species, and have scarcely a doubt,
that, with the facilities which you tender me, I can by the close of the year be prepared
to present this sub-order in a very accurate and perfect shape.” [Letter of June 22,
1848.]

“The Homopterous Hemiptera, which have already been adverted to in our cor-
respondence are again before me. I have been collecting further, and am now preparing
a descriptive catalogue of all the species I have been able to find, their synonyms,
the plants on which they occur, etc. — which catalogue will be published in the course
of the coming winter. Dr. Emmons is also having a number of them very nicely
figured for his final volume ( Deo gratiasl ) of the State Nat. Hist, series .... I am
quite anxious to work these Homopterous insects out, in the best manner I am capable
of doing.” [Letter of November 13, 1850.]

“Last evening’s mail brought me also the proof-sheets of what I had prepared upon
our Homoptera. Those species that I had been able to make out to my satisfaction
had been ticketed and placed in the State Cabinet at Albany a few weeks ago— and
the paper containing my descriptions of them (28 pages octavo) the printers are now
waiting to strike off. In this emergency I have concluded to go down to Albany
tomorrow morning, to see if I can get them to delay a day or two, to enable me to
compare your specimens with mine and change my names for yours, in those instances
where we both have had the same species.” [Letter of February 20, 1851.]

“I reached home from Albany last evening, and hasten to inform you of the result
of my visit. The printers reluctantly consented to defer their work for one day, to
enable me to re-examine the specimens in the State Cabinet and make alterations
in my copy .... Herewith are forwarded copies of my article upon these insects. My
final revisal of the proof I perceive was scarcely looked at by the printers; hence there
are some typographical errors, but I see none of moment.” [Letter of March 1-4,
1851.]

“I have not yet obtained Emmons’ ‘big book.’ Indeed, I dread looking into it,
knowing what a hotch-potch thing it must be, and not knowing what use has been

3 Fitch to Harris, 1 5 letters dated December 30, 1846, to February 26, 1855; file bMu 998. 10.2,
Museum of Comparative Zoology, Harvard University.

1984

HOMOPTERA: FITCH AND EMMONS

29

made of my name in it. Last September, as I was packing up to start next day on a
journey, the Dr. unexpectedly dropped in on me, with a bundle of the engravings,
for me to name the figures therein. The specimens from which the drawings were
made he did not think it was necessary to bring, as the figures were ‘perfectly accurate’
showing everything which the specimens showed! On hastily glancing over the plates
(for I had not time to examine deliberately) some of the figures I recognized, others
I could only guess at, & others still I could form no conception what they were. He
also looked over my cabinet and took down some names from thence, although
informed they were of no value, some of the families having been ticketed many
years ago, when I had scarcely any authorities at hand to aid me in the work. As
mineralogy is the Doctor’s speciality, and I have some minerals which I should like
to have named, I think it will be but fair for me to ask the Dr. to reciprocate the
favor he asked of me, and send him ‘accurate drawings’ of these minerals, that he
may name them for me. Like his volume on Fruits, this on Insects, I think, must
fall still bom from the press.” [Letter of February 26, 1855.]

FITCH’S HOMOPTERA CATALOGUE

The full title of Fitch’s Homoptera catalogue is “Catalogue with References and
Descriptions of the Insects Collected and Arranged for the State Cabinet of Natural
History,” and it appeared in the Fourth Annual Report of the Regents of the Uni-
versity, on the Condition of the State Cabinet of Natural History, and the Historical
and Antiquarian Collections Annexed Thereto, Made to the Senate, January 14,
1851. That report is also known as Senate Document 30. Fitch dated his introduction
to the catalogue February 22, 1851. In his letter of March 1-4, 1851, he informed
Harris that he was forwarding copies of his “article upon these insects,” and he
mentioned the typographical errors that it contained. That article was unmistakably
his Homoptera catalogue. Evidently, it was printed and ready for distribution some-
time between February 22 and March 1, so February 28, 1851, is the earliest date
of publication demonstrated by evidence, in the meaning of the International Code
of Zoological Nomenclature, Article 21. J. A. Lintner had the catalogue reprinted,
with corrections and notes by E. P. Van Duzee and C. V. Riley, in 1893 in his Ninth
Report on the Injurious and Other Insects of the State of New York for the Year 1892
[From the Forty- sixth Report of the New York State Museum].

EMMONS’ REPORT

The correct title of Emmons’ report is Insects of New York, although this title
appears only on the first page of text, above the title of the first chapter. The first
“title page” reads “Natural History of New York, By Authority,” and it is followed
by a page that reads “Agriculture of New York: Comprising an account of the Clas-
sification, Composition and Distribution of the Soils and Rocks, and of the Climate
and Agricultural Productions of the State; Together with Descriptions of the More
Common and Injurious Species of Insects, Volume V.” The latter page also bears
the date 1854, and the Preface is dated July 25, 1854. In New York State Assembly
Document 53, dated January 30, 1855, E. W. Leavenworth and T. Romeyn Beck,
commissioners appointed to superintend the completion of the publication of the

30

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

“Natural History of New York,” reported that . . another volume has been com-
pleted, viz: the treatise of Dr. Emmons on Insects Injurious to Agriculture. It will
probably be ready for general distribution during the month of January.” In his letter
of February 26, 1855, Fitch told Harris that he had not yet seen Emmons’ report,
but he seemed to imply that it was available. The publication date, in the meaning
of the code, therefore, seems to be January 31, 1855.

EMMONS’ MEMBRACIDAE

Thirty-five species of Membracidae are described in Emmons’ report, two of them
simply as named figures. Emmons credited Fitch for providing specimens and names
of the genera and species. In his letter of November 13, 1850, Fitch told Harris that
he was preparing a descriptive catalogue of all the species of Homoptera he was able
to find, and that Emmons was having some of them figured for the “Natural History
of New York.” It is evident that Fitch presented Emmons with specimens, some of
which he had identified and others of which he had recognized as belonging to new
species that he named. According to Article 50 of the Code, the author of a species
name is the person who first publishes it in a way that satisfies the criteria of avail-
ability, “unless it is clear from the contents of the publication that . . . some other
person is alone responsible both for the name and the conditions that make it avail-
able.” Because Emmons credited Fitch with the species names, but not the descrip-
tions or illustrations, Emmons must be considered the author of any new species
named in his report. Goding’s (1893) conclusion that “there is no valid reason for
crediting such species to Emmons because the descriptions “were written by Dr.
Fitch, as is stated by Emmons” is misleading and supported by false evidence.

Emmons attributed eight of the species described in his report to previous authors.
By comparing Fitch’s manuscript species notes— most of which were recently found
in the New York State Museum— with Emmons’ report, it becomes apparent that
eleven or twelve more species should have been attributed to previous authors. These
include Thelia lutipes, which is apparently a misspelling of latipes Say. The corre-
sponding figure in Emmons’ report is labelled Enchophyllum latipes, and in Fitch’s
notes it is given as Enchenopa latipes Say, with the generic names Enchophyllum
and Membracis crossed out. The status of Gargara inermis is discussed below. The
remaining fifteen or sixteen membracid species names in Emmons’ report must be
attributed to Emmons. Some of them appear in Fitch’s notes, but others do not.

fitch’s specimen registers

The types of Emmons’ species came from Fitch’s personal collection. Fitch never
designated a type for any of his species, but he did list individual specimen code
numbers in his notes under each species name. The listed specimens that were
collected before the publication date of each of his species names must be considered
the specimens of his type series. With each code number he gave the collecting locality,
date, and frequently other valuable information. The code numbers also correspond
with four registers that list the numbers and give the collecting conditions of nearly
all of the specimens in his personal collection. In one of the registers, now in the
New York State Museum, Fitch wrote:

1984

HOMOPTERA: FITCH AND EMMONS

31

“All (nearly) of the specimens in my collection have a number upon the pin.
Numbers written on white paper with black ink, are all from the State of New York;
and are registered in this book; or, those with one or more red-ink lines across the
number, are registered in another book similar to this. Specimens from other parts
of the U. States, & from Canada are numbered with red ink on white paper, &
recorded in a separate book. Specimens numbered with black ink upon colored paper,
are from other parts of the world, & are registered in a fourth book.”

In another register, Fitch wrote:

“List of Insects with numbers attached crossed with a single red line up to No.
10,000— two red lines, 10,000 to 20,000 .... These are all taken in the state of New
York.”

In a third register, also in the possession of the New York State Museum, he wrote:

“List of Insects numbered with red ink. These are all North American, but collected
out of the state of New York, but in the other United States or Canada or north of
there.”

In the fourth register, now in the library of the Museum of Science, Boston, Fitch
recorded his specimens from places other than the United States or north of there.

It is evident that most of Fitch’s specimens, if recognized as such, can be matched
with the proper date and locality given in his manuscript notes or registers. It is also
evident, as Funkhouser (1915) found, that Fitch’s specimens that carry a number in
red ink and those that are marked with a red line across the label are not necessarily
the true types. The only possible types are those specimens that are listed in Fitch’s
manuscript notes and registers and that were collected before the species name was
published.4

TYPES OF EMMONS’ MEMBRACIDAE

Emmons’ membracid species, together with excerpts from the existing notes on
type series in Fitch’s manuscripts, are listed below. Any number preceded by an
asterisk is a specimen number that Fitch crossed with a single red line.

1 . Telamona acclivata Emmons, 1855:155 [=Heliria cristata (Fairmaire)] . 11,782,
Stillwater, NY, 16.vii.1846, on oaks.

2. Cyrtosia arcuata Emmons, 1855:154 [ =Cyrtolobus arcuatus (Emmons)].

3. Gargara cinereum Emmons, 1855:156 [ =Cyrtolobus cinereus (Emmons)].
*4536, Salem, NY, 26.vi.1846, on oaks. 1 1,760, Stillwater, NY, 16.vi.1846, on oaks.

1 1,' 766-777, Stillwater, NY, on plants.

Goding (1893) stated that a specimen numbered 4536 was in the U.S.N.M., but
I have been unable to locate it.

4. Gargara discoidalis Emmons, 1855:157 [=Cyrtolobus discoidalis (Emmons)].
3038, Salem, NY, 24.vi.1846, on white oak trees, beat from leaves. 4837, Salem,
NY, 5.vii.l847, on white oak tree.

4 Fitch’s notes on Psyllidae and Coccidae are in the Smithsonian Institution Archives, those
on Aphididae are with the United States National Museum aphid collection in Beltsville, those
on Collembola are in the library of the Museum of Science in Boston, and those on all other
arthropod groups, constituting the bulk of his manuscript species notes, are in the New York
State Museum in Albany.

32

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Goding (1 893) stated that specimens numbered 3038 and 4837 were in the U.S.N.M.,
but I have been unable to locate them.

5. Cyrtosia fuliginosa Emmons, 1855:154 [=Cyrtolobus fuliginosus (Emmons)].
1 1,756, 1 1,764, Stillwater, NY, 16.vii.1846, on oak in big swamp.

Goding (1893) found specimen number 1 1,764 in the U.S.N.M., but I have been
unable to locate it.

6. Gargara inermis Emmons, 1855:157 [=Cyrtolobus inermis (Emmons)]. *2550-
*2555, Salem, NY, 20. v. 1846, beat from trees (chiefly chestnut and oak). 1787, 1788,
3047, Tullehassie, AR [presumably Tullahassee, Wagoner Co., OK], July 1851, from
Wm. S. Robertson.

Although Fitch’s notes indirectly suggest that this name should have been attributed
to Fabricius, the species Emmons described and illustrated was apparently new. The
name is usually attributed to Emmons (Funkhouser, 1927; Metcalf and Wade, 1966;
Kopp and Yonke, 1973).

7. Cyrtosia intermedia Emmons, 1855: plate 13, figure 16 [=Xantholobus inter-
medius (Emmons)]. *4635, Salem, NY, 28.vi.1847, on oaks.

8. Gargara maculifrontis Emmons, 1855:156 [=Cyrtolobus maculifrontis (Em-
mons)]. *3039, Salem, NY, 24. vi. 1846, beat from white oaks. 11,762, Stillwater,
NY, 16.vii.1846, on oaks in big swamp.

9. Gargara majus Emmons, 1855:156 [=Carynota mera (Say)].

10. Gargara nigricephala Emmons, 1855:1 57 [=Atymna castaneae (Fitch)]. *3598,
Salem, NY, 24.vii. 1846, on chestnut. *5181, Stillwater, NY, 15.vii.1847, beat from
chestnut bushes.

11. Telamona ornata Emmons, 1855:155 [=Telamona concava (Fitch)]. 5419,
Stillwater, NY, 1834, a female.

12. Gargara pallidifrontis Emmons, 1855: plate 13, figure 7 [=Cyrtolobus palli-
difrontis (Emmons)]. *3034, Salem, NY, 24.vi, 1 846, beat from white oak trees. 1 1 ,763,
Stillwater, NY, 16.vii. 1846, on oaks in the big swamp. *4897, Salem, NY, 5.vii.l847,
on black oak, bank of the kill. *5107, Stillwater, NY, 15.vii.1847, on oak bushes.

Goding (1893) found specimen number 11,763 in the U.S.N.M., and Dr. James
P. Kramer has confirmed its existence (pers. comm.). It is labelled “1 1,763/pallidi-
frontis/ Lectotype, Cyrtolobus pallidifrontis (Emmons), L. B. Woodruff.”

13. Gargara pectoralis Emmons, 1855:157 [=Stictocephala lutea (Walker)].

14. Gargara pubescens Emmons, 1855:157 [=Ophiderma pubescens (Emmons)].

15. Telamona turriculata Emmons, 1855:155 [=Glossonotus turriculatus (Em-
mons)].

16. Smilia viridis Emmons, 1855:154 [=Cyrtolobus viridis (Emmons)]. 11,767,
Stillwater, NY, 16.vii.1846, on plants. *5178-79, Stillwater, NY, 15.vii. 1847, on
walnut bushes. *5364, Salem, NY, 23.vii.1847, on white oak.

fitch’s homoptera

The Homoptera collection that Fitch described in his 1851 catalogue is in the New
York State Museum, successor to the State Cabinet of Natural History. The specimens
in it were taken from his personal collection and renumbered. Fitch enumerated
consecutively all specimens in his two catalogues and in the State Cabinet. Specimens
referred to in his 1849 catalogue were numbered 1-608, and those in his 1851

1984

HOMOPTERA: FITCH AND EMMONS

33

catalogue were numbered 609-874. These numbers bear no relation to Fitch’s species
notes or specimen registers.

In the Homoptera catalogue, Fitch described 6 new genera, 85 new species, and 5
new subspecies, including 3 genera, 1 5 species, and 2 subspecies of Membracidae.
The State Museum’s collection still contains original specimens of 60 of the species
and subspecies. The specimens of Psyllidae, many of the Aphididae, and a few
specimens from other families were destroyed by museum pests (Lintner, 1 893). The
remaining specimens, as well as specimens from Fitch’s personal collection that were
collected prior to February 28, 1851, must be considered types. Fitch’s notes indicate
that for most species he had more than one specimen prior to the publication of his
catalogue. In these cases, a lectotype should be designated if that has not already
been done. Funkhouser (1915) designated specimens from the State Museum col-
lection as lectotypes for Fitch’s species and subspecies of Membracidae that were
described in his catalogue. Specimens of Homoptera from Fitch’s personal collection
are in the U.S.N.M. and in the T. W. Harris Collection, Museum of Comparative
Zoology, Harvard University.

A catalogue to the taxa and known existing specimens described by Fitch, a bib-
liography of his entomological papers, and a biography of Fitch are in preparation.

ACKNOWLEDGMENTS

I thank Drs. L. L. Deitz and J. P. Kramer for supplying information and for critically reviewing
the manuscript, and I gratefully acknowledge permission to publish excerpts from the T. W.
Harris correspondence held by the Museum of Comparative Zoology, Harvard University.

LITERATURE CITED

Emmons, E. [1855]. Insects of New York. Agriculture of New York, volume 5. Natural history
of New York. Albany, 272 pp.

Fitch, A. 1849. Catalogue of the insects in the State Cabinet of Natural History, December
1, 1848. Pages 27-39 in: Second annual report of the Regents of the University, on the
condition of the State Cabinet of Natural History, with catalogues of the same. Made to
the Senate January 12, 1849. Albany, 103 pp.

Fitch, A. 1851. Catalogue with references and descriptions of the insects collected and arranged
for the State Cabinet of Natural History. Pages 43-69 in: Fourth annual report of the
Regents of the University, on the condition of the State Cabinet of Natural History, and
the historical and antiquarian collection, annexed thereto. Made to the Senate, January
14, 1851. Albany, 146 pp.

Felt, E. P. 1910. Insect types in the New York State Museum. New York State Museum
Bulletin 141:119-122.

Funkhouser, W. D. 1915. Types of Fitch’s species of Membracidae. Bull. Brooklyn Ent. Soc.
10:45-50.

Funkhouser, W. D. 1927. General catalogue of the Hemiptera, Fascicle I. Membracidae.
Smith College, Northampton, Massachusetts, 581 pp.

Goding, F. W. 1893. Fitch’s types of N. A. Membracidae. Can. Ent. 25:171-196.

Kopp, D. D. and T. R. Yonke. 1973. The treehoppers ofMissouri: Part 3. Subfamily Smiliinae;
tribe Smiliini. J. Kansas Ent. Soc. 46:375-421.

Lintner, J. A. 1893. Catalog of the known Homoptera of the State of New York in 1851.
Annual Report of the New York State Museum 46( 1 892):38 1—4 1 3.

34

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Mallis, A. 1971. American entomologists. Rutgers University Press, New Brunswick, New
Jersey, 549 pp.

McCabe, T. L. and L. M. Johnson. 1980. Catalogue of the types in the New York State
Museum insect collection. New York State Museum Bulletin 434:1-38.

Metcalf, Z. P. and V. Wade. 1965. General catalogue of the Homoptera. A supplement to
Fascicle I— Membracidae, of the general catalogue of the Hemiptera. Membracoidea.
North Carolina State University, Raleigh, 1552 pp.

Osborn, H. 1905. Jassidae of New York State. New York State Museum Bulletin 97:498—
545.

Sanborn, F. G. and J. A. Lintner. 1879. An account of the collections which illustrate the
labors of Dr. Asa Fitch. Psyche 2:273-276.

Sanders, J. G. 1910. A review of the Coccidae described by Dr. Asa Fitch. Proc. Ent. Soc.
Washington 12:56-61.

Woodruff, L. B. 1924. Critical observations in the membracid genus Cyrtolobus Goding.
(Hemip.-Homop.). J. New York Ent. Soc. 32:1-62.

Received May 24, 1983; accepted November 22, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 35-41

ASSOCIATION OF PARATRECHINA ARENIVAGA
(HYMENOPTERA: FORMICIDAE), WITH NYMPHS OF
OECLEUS BOREALIS (HOMOPTERA: CIXIIDAE)1 2

C. R. Thompson1 2 3

Department of Entomology and Nematology, University of Florida,
Gainesville, Florida 32611

Abstract.— The previously unknown nymphs of the planthopper Oecleus borealis Van Duzee
were found in the subterranean nests of the ant Paratrechina arenivaga (Wheeler). The nymphs
appear to associate only with this ant species. Food plants of the O. borealis nymphs are
suspected to be the roots of sand pine Pinus clausa (Engelm.) Sarg., saw palmetto Serenoa
repens (Bartr.) Small, or turkey oak Quercus laevis Walter, although radioisotope experiments
to establish this were inconclusive.

The family Cixiidae is a member of the superfamily Fulgoroidea, or planthoppers.
This family, to which Oecleus belongs, has an extensive distribution in the temperate
regions of the world (Van Duzee, 1908). J. P. Kramer (1977) listed 40 species of
Oecleus in the U.S. The distribution of the genus is mostly western, with only 3
species occurring east of the Mississippi River. O. borealis occurs from the Gulf
states north to New York and west and north to Texas and Kansas (Kramer, 1977).
Literature on cixiid biology is extremely sparse and is contained for the most part,
in small notes following taxonomic descriptions. Dozier (1926) took both adult sexes
of O. borealis by sweeping grasses in pinewoods. He speculated that one of these
plants was probably the food plant. Ball and Klingenberg (1935) collected adults
“throughout the season” and suspected this was the hibernating stage. They found
adults on many trees and shrubs and believed the species was a general feeder, or
that the nymphal hosts were low, widespreading plants and adults flew to higher
vegetation.

Although one species was reported to oviposit in Yucca (Townsend, 1892), Myers
(1929) was the first to report the habitat of cixiid nymphs: Mnemosyne cubana Stal
and Bothriocera signoreti Stal from Cuba. Instars of the Mnemosyne were found in
a rotted stump in March. They had filamentous wax tails and were feeding on roots
covered with this wax. Myers found the nymphs in apparent association with the
ponerine ant, Odontomachus haematoda insularis var. pallens Wheeler. He stated
that the ant was tending the nymphs, although he did not actually observe this activity.
In addition, the ponerines are primitive ants; none are known to attend homopterans
(Nixon, 1951). Late instar nymphs of Bothriocera were found under stones feeding
on rootlets in March. The nymphs were covered with a white flocculent wax. In
neither case, apparently, did Myers determine the identity of the food plants.

1 This research was a portion of a M.S. thesis submitted to the Graduate School of the
University of Florida, Gainesville, Florida.

2 Fla. Agric. Exp. Stn. Journal Series No. 4870.

3 Present address: 3205 S.W. College Avenue, Ft. Lauderdale, Florida 33314.

36

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92( 1 )

The nymphs of the cixiid Oliarus vicarius (Walker) have been found feeding on
roots beneath rotted stumps and logs in the same ecosystem as O. borealis (Thompson
et al., 1979). Two species of ants seemed to be associated with these nymphs. The
ant-Homoptera relationship reported in this paper is distinct, and appears to be
obligatory for the Oecleus nymphs.

MATERIALS AND METHODS

The nest of P. arenivaga is marked by a distinctive crater. This ant is the predom-
inant ant species in the sand pine habitats in which it is found. Excavations were
undertaken in sand pine woods, Pinus clausa (Engelm.) Sarg. ca. 27 km east of Silver
Springs, Florida (Marion County) in Ocala National Forest, on the south side of
Highway 40. The vegetation in this area was composed of saw palmetto, Serenoa
repens (Bartr.) Small, and turkey oak, Quercus laevis Walter.

After experimentation with various collecting methods, I found two effective in
excavating maximum numbers of nymphs. One method was to shovel 4 vertical 20
cm thrusts to form a square around the crater of a Paratrechina nest. The entire
clump was lifted and transported to a flat surface. It was broken apart with the hands
while having an aspirator ready. The clump often broke, particularly when wet, along
the tunnels of the ants. A second method, utilizing the element of surprise, was to
thrust a shovel downward 3 to 4 cm from a crater and then carefully flip the shovel
upwards. The nymphs were usually found among the greatest concentrations of ants.
In this quick-attack method, the nymphs could be collected before they had time to
escape. They were placed in separate vials from the ants.

Difficulties were encountered during warm weather in keeping the nymphs, ant
queens and even ant workers alive until they were brought to the laboratory. It was
found that moisture must be available at all times. To maintain collected live spec-
imens in good condition, shell vials containing the insects were buried in a shaded
place. Upon departure, tubs were filled with cool, moist sand, into which the vials
were reburied. This method allowed successful transportation of live specimens dur-
ing hot weather.

Captured Oecleus nymphs were brought to the laboratory and placed in Wilson
cells (Wilson, 1962) with moist cotton floors. To approximate the Oecleus natural
habitat, roots and sand from their original chambers were placed with the nymphs.
The nymphs were kept under observation and disturbed as little as possible.

Radioisotype tracer studies and field observation were used to study food plant
preferences of Oecleus nymphs. Prior to the isotope experiment, roots of three dif-
ferent plants were repeatedly found in the Paratrechina nests. Two of these root types
were identified as pine and saw palmetto. Subsequently, young pine and saw palmetto
plants were dug up and brought to the laboratory. Their roots were sandwiched
between small pieces of non-absorbent cotton (to stop possible isotope leaks in the
external root surfaces), then between small, back-to-back petri dish lids. The root
space was taped shut around the sides of the petri dishes with water-proof tape, then
packed tightly full of sand. The protruding roots were placed in water containing 100
juCi P32 per plant. After 24 hr, the nymphs were placed in cells through which passed
roots in direct contact with the tracer solution. After contact with the plants for 36
hr, the nymphs were tested for radioactivity by both Geiger-Mueller and liquid-
scintillation counters.

1984

PARATRECHINA AND OECLEUS

37

In a preliminary experiment, nymphs exposed to radioactive pine roots died after
becoming trapped in water film which had collected on the inside chamber wall. To
reduce film formation, the facing plastic of each chamber, sealed with clay against
the petri dish, was perforated with a hot needle.

RESULTS AND DISCUSSION

Nymphs of O. borealis were discovered while excavating nests of the ant P. ar-
enivaga in Ocala National Forest, Marion County, Florida. Oecleus borealis nymphs
were also found in P. arenivaga nests in Putnam County, ca. 48 km north of the first
discovery site, and later, in Broward and Orange Counties. These areas had similar
habitats and were characterized by areas of white sand.

Many hundreds of nests were dug up during the course of this study. Oecleus
nymphs were found in the Paratrechina nests from December through August (Table
1). Subsequent collections have been made in September and November (Wilson et
al., 1983). The largest number of nymphs found in a single nest was five, but the
actual numbers present could not be determined because of the methods used to
excavate the nests. The ratios of nests with, to those without, Oecleus varied, but
there was a pattern. During most months the ratio was 1:5, 6 or 7. In December-
February the ratio was often 1:20 or more. At the other extreme, during July the
ratio was reduced to 1:2.

Paratrechina nests were not found in heavily shaded areas. The preferred habitats
of the ants were clearings, pathways within the woods, or disturbed areas including
roadways. Nests were found only where the sand was clear of thick debris. A cover
of sand pine needles was sufficient to exclude the ants from an area. Even when the
nests were in the middle of a wide road, the roots of sand pine, saw palmetto, and
what was probably turkey oak were found thickly matted beneath the soil surface.
In some cases, measurement showed the nearest plant to be 3-4 m away.

Areas with similar vegetation, but without Paratrechina nests, were also excavated
in attempts to locate Oecleus nymphs. These efforts were not successful. Oecleus
nymphs were not found except in the company of the ants.

The nymphs collected varied in length from 0.8 mm to 3.7 mm. The smallest
nymphs were faint yellow, darkening in later instars to bright yellow with orange
bands on the abdominal segments. This coloration made the nymphs highly visible
against the sand of the ant nests. Late instars had wing pads and dark reddish orange
markings on the abdomen. The posterior white wax filaments were rarely found on
the nymphs in the field, but grew out on nymphs in the laboratory within 1 2 hr. Full
descriptions and a key to the nymphs are in Wilson et al., 1983.

Oecleus nymphs were repeatedly observed directly in the chambers of the ants. In
at least 6 cases (2 in the laboratory, 4 in the field) ants were observed carrying first
or second instar nymphs in their mandibles. This appeared to be a reaction to colony
disturbance. When nests were dug between 9 or 10 A.M. and 4 P.M., few if any,
nymphs were found. Temperatures are highest during this time and the sand becomes
dry and hot in the upper layers. The nymphs and ants may avoid the heat by retreating
down into the sand. During this time I was unable to find the ants or nymphs by
digging downward ca. 30 cm. One nest was excavated to a depth of 80 cm and
searched laterally, layer by layer, but no queens, brood or nymphs were found. The
nymphs were also less frequently found if no rain had recently fallen. This was

38 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Table 1. Collections of Oecleus borealis nymphs in

Paratrechina arenivaga nests.

Date

Numbers
of nymphs

Location

Notes

1/11/75

5

Ocala

1/25/75

4

Ocala

—

3/22/75

3

Ocala

2-L nymphs

4/5/75

4

Ocala

—

4/13/75

5

Putnam

1-S, 2-M, 2-L nymphs

4/21/75

1

Putnam

1-S nymph in worker’s mandibles

6/18/75

9

Ocala

8-S, 2-M, 2-L nymphs

6/19/75

13

Ocala

—

6/29/75

2

Ocala

—

7/16/75

18

Ocala

—

7/25/75

1

Ocala

—

8/2/75

10

Ocala

—

8/4/75

3

Gainesville

1-M nymph

8/12/75

0

Ocala

—

12/-/75

1

Ocala

1-M nymph

1/22/76

0

Ocala

—

2/19/76

1

Ocala

1-M nymph

3/22/76

1

Ocala

1-S nymph

3/31/76

8

Ocala

5-S, 3-M nymphs

4/1/76

2

Ocala

2-L nymphs

4/5/76

2

Ocala

3-L nymphs

4/6/76

1

Ocala

—

4/15/76

1

Ocala

1-L nymph

4/27/76

8

Ocala

2-S, 6-M nymphs

5/10/76

6

Ocala

2-L nymphs

7/2/76

3

Putnam

1-M, 2-L nymphs

7/15/76

24

Putnam

24-S nymphs

Explanation of letters: S, M, L = estimates of Oecleus nymph size (Small, Medium, Large).

probably due in part to the tendency of the dry sand blocks to fall apart during
excavation.

During collecting trips in December and January, the gasters of Paratrechina work-
ers dug up were extremely distended with a clear liquid. This coincided with the
presence of relatively large Oecleus nymphs in the ant nests. It is known that 1 aphid
can produce copious amounts of honey dew relative to its size and weight and that
the honey dew of some aphids is high in amino acids, protein and other nutritional
substances (Way, 1963). If the Paratrechina are obtaining honey dew from the Oe-
cleus, a relatively small number of nymphs might be able to support an entire colony—
or at least to provide a substantial amount of its food supply.

Observations of Paratrechina behavior support this hypothesis. Paratrechina ar-
enivaga is a crepuscular and mostly nocturnal ant and excavates or repairs its nests
at that time. Many nights I have observed the ants, at the moment the head lamp
light hit them, doing no foraging; carrying only sand grains. The light caused them
to retreat: the workers quickly dropped the sand on the crater and returned down

1984

PARATRECHINA AND OECLEUS

39

the entrance. When offered baits of honey, sugar water, com syrup, egg or raw
hamburger, the ants were most attracted to the sweet solutions, but consumed all
the baits. Aside from the bait experiment, however, Paratrechina workers were not
observed returning to a nest with anything in their mandibles or with distended
gasters. They were rarely found more than 20 cm from their nest entrances. It is
possible that these ants obtain their food from an underground source, perhaps the
cixiid nymphs, at least during the winter months.

Attempts to rear Oecleus nymphs were not successful until several large nymphs
were collected in March 1975. These were placed in Wilson cells with Paratrechina
workers. An adult emerged several days later, but was almost entirely eaten by the
ants. The ants were removed and a second adult emerged. It was found in an arm
of the Wilson cell. The exuvia was clinging by the tarsi to the inside cell wall. This
adult was determined to be Oecleus borealis Van Duzee.

In the pine and saw palmetto experiment, the radiation levels of roots removed
only from the nymphal chambers varied from 113 to 2,585 cpm on the Geiger-
Mueller counter and from 637 to 21,450 cpm on the liquid scintillation counter.
Every set of roots had a count well above background on even the less sensitive
Geiger-Mueller counter. None of the 5 nymphs recovered were radioactive.

A number of factors may have caused the non-radioactivity of the nymphs. Some
of these are: 1) the plants were too young, possessing roots too small, or lacking
sufficient sap; 2) sap flow in the plants was affected by the methods used to handle
the plants or roots; 3) the nymphs were not able to feed due to handling methods or
unnatural surroundings; or 4) neither of these plants is the natural food plant of O.
borealis nymphs.

The difficulty involved in ascertaining the food plant of the Oecleus nymphs in
the field lies with tearing apart the soil of the nest without disturbing these sensitive
insects. Many times a nest was dug to find the nymphs already disturbed and jumping
about (some can jump at least 8 cm). Other subterranean Homoptera known to
associate with ants, such as coccids and aphids, are relatively sessile organisms. Even
when alarmed, these insects are not capable of jumping and running, while this is
the usual reaction of an Oecleus nymph. Even if the nymphs feed on ant nest roots
and the ants are obtaining honey dew, it is doubtful that the ants could tend large
nymphs directly. This is because the nymphs appear to avoid the ants (see below)
and can move much more quickly than the ants themselves.

A number of interesting behaviors were noted in laboratory Wilson cell groups of
nymphs and ants. The nymphs would not remain with the ants. If antennated by the
ants, the nymphs moved quickly away. If antennated a number of times within a
short period, the nymphs jumped away from the ants. The nymphs rested on roots
in the cell, but did not seem to select one kind of root over another to rest upon.
One nymph on a root was observed to move around the root and, in a squirrel-like
motion, kept the root between itself and a nearby ant. The nymphs were extremely
sensitive to any jarring of the cell, which caused them to move around or to jump.

During a 24 hr observation period, the wax filaments of a nymph’s tail slowly
lengthened ca. 2 mm. At one point, the nymph jumped, and when it landed the tail
filaments had been flicked off. The most unusual behavior noted for a nymph occurred
at 7:52 A.M., 21 hr after observation had begun. A medium-sized nymph was ob-
served arching its head and abdomen downward, bending itself into a C shape. These

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

movements continued periodically until 9:37 A.M., when the abdominal interseg-
mental membranes began to protrude from between the original abdominal plates.
By 10:1 1 A.M., three new abdominal bands as wide as the 4 abdominal plates were
present. They were a slightly lighter yellow-orange and glistening. The nymph arched,
often while clinging to a root, and the haemolymph was observed flowing up and
down in the proboscis and head. A change occurred in the nymph’s behavior. Al-
though it still avoided the ants, the nymph became extremely active. It had jumped
only 3 times prior to 10:30 A.M. (twice in response to ant antennation), but between
10:30 and 1 1:00 A.M., when observation was terminated, it jumped 15 times. The
nymph also continued to arch periodically during this time.

CONCLUSIONS

The nymphs of the planthopper O. borealis were found in the subterranean nests
of the ant, P. arenivaga. These nymphs, members of the homopteran family Cixiidae,
were previously unknown. Adding to Wilson’s (1971) catalog of associations, Cixiidae
is the ninth homopteran family known to be symbiotic with ants. Nymphs were
present in the ant nests throughout the year.

It is suggested that the ant-nymph relationship is obligatory for Oecleus, involving
the need for Oecleus to reach its host plant roots, which it does by penetrating a
Paratrechina nest where roots are exposed by excavations of the ants. In all known
ant-homopteran relationships, the ant obtains honey dew from the homopteran. In
the ant -Oecleus relationship, this is also presumed to occur, but was not observed
in field or laboratory colonies.

Although attempts to determine the Oecleus food plant with P32 were not successful,
the author believes the technique was valid with the exception of nymph handling.
Better handling techniques are needed since many nymphs succumbed to or did not
feed under experimental conditions.

ACKNOWLEDGMENTS

I wish to thank the late Dr. William F. Buren for identification of the ant and Dr. James P.
Kramer for identification of the planthopper.

LITERATURE CITED

Ball, E. D. and P. Klingenberg. 1935. The genus Oecleus in the United States (Homoptera:
Fulgoridae). Ann. Ent. Soc. Am. 28:193-213.

Dozier, H. L. 1 926. The Fulgoridae or plant-hoppers of Mississippi, including those of possible
occurrence. Tech. Bull. Miss. Agri. Exp. Stn. 14:1-152.

Kramer, J. P. 1977. Taxonomic study of the planthopper genus Oecleus in the United States
(Homoptera: Fulgoroidea: Cixiidae). Trans. Am. Ent. Soc. 103:379-449.

Myers, J. G. 1929. Observations of the biology of two remarkable cixiid plant-hoppers (Ho-
moptera) from Cuba. Psyche 36:283-292.

Nixon, G. E. J. 1951. The Association of Ants with Aphids and Coccids. Commonw. Inst.
Ent., London, 36 pp.

Thompson, C. R., J. C. Nickerson and F. W. Mead. 1 979. Nymphal habitat of Oliarus vicarius
(Homoptera: Cixiidae), and possible association with Aphaenogaster and Paratrechina
(Hymenoptera: Formicidae). Psyche 86:321-325.

1984

PARATRECHINA AND OECLEUS

41

Townsend, C. H. T. 1892. Oviposition of a homopterous insect in yucca. Psyche 6:353-354.

Van Duzee, E. P. 1908. Studies in North American Fulgoridae. Proc. Acad. Nat. Sci. Phila-
delphia 1907:467-498.

Way, M. J. 1963. Mutualism between ants and honey-dew producing Homoptera. Annu. Rev.
Ent. 8:307-344.

Wilson, E. O. 1962. Chemical communication among workers of the fire ant, Solenopsis
saevissima (Fr. Smith). 1. The organization of mass foraging. Anim. Behav. 10:134-137.

Wilson, E. O. 1971. The Insect Societies. Belknap Press of Harvard Univ. Press, Cambridge,
Massachusetts, 548 pp.

Wilson, S. W., J. H. Tsai and C. R. Thompson. 1983. Descriptions of the nymphal instar of
Oecleus borealis (Homoptera: Fulgoroidea: Cixiidae). J. New York Ent. Soc. 91:41 8—
423.

Received July 21, 1983; accepted October 31, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 42—47

A NEW SPECIES OF SIGARA FROM WESTERN OREGON
AND WASHINGTON (HEMIPTERA: CORIXIDAE)

Gary M. Stoned ahl

Systematic Entomology Laboratory, Department of Entomology,
Oregon State University, Corvallis, Oregon 97331

Abstract.— -A new corixid, Sigara krafti, is described from specimens collected in western
Oregon and Washington. This species seems to prefer shallow, slow-moving water in streams
and small rivers, and is most abundant over silty or muddy substrates. Illustrations of the male
abdominal terga, right pala and right paramere of the male, left hemelytron of the female, and
the mesepimeron are provided for Sigara krafti and three related species ( S . mckinstryi Hun-
gerford, S. vandykei Hungerford, S. washingtonensis Hungerford) found in Oregon and Wash-
ington.

Hungerford (1948) reported the following five species of the genus Sigara from
Oregon and Washington: S. alternata (Say), S. omani (Hungerford), S. ornata (Ab-
bott), S. vandykei Hungerford, S. washingtonensis Hungerford. All of these species
are common in the Pacific Northwest except for S. ornata, which appears to have
been erroneously recorded from this region. Hungerford reported S. ornata from an
unspecified locality in Oregon, basing his identification on three specimens from the
Uhler collection (USNM). I have examined a male and two female specimens from
the Uhler collection that I believe are the ones Hungerford said were from Oregon.
The writing on the labels is barely discernible, but it appears to read “Oro, 4-21-
00.” Hungerford apparently thought the label read “Ore” as in Oregon. However, it
is more probable that “Oro” pertains to a specific locality or its abbreviation (e.g.,
Orono, Maine). In any case, all other records of S. ornata are from the northeastern
United States and Canada, and it is doubtful this species occurs outside of that region.

Besides the species reported by Hungerford (1948), four other Sigara species occur
in Oregon and/or Washington. These are S. decoratella (Hungerford), S. grossolineata
Hungerford, S. mckinstryi Hungerford, and a new species, S. krafti, described below.
Sigara krafti is distinguished from other Sigara species of the Pacific Northwest by
the following combination of characters: body length 5.3-6. 1 mm; ostiole located
about midway between lateral bend and apex of mesepimeron; mesepimeral process
broadest just anterad of ostiole, then slightly narrower to lateral bend (Fig. 13);
anterior one-third to one-half of metaepistemum fuscous; posteromedial projection
of male seventh abdominal tergum triangular (Fig. 9); right paramere of male thick-
ened preapically (Fig. 5); and male pala with 29-34 pegs (Fig. 17).

The following abbreviations are used for specimen depositories cited in the locality
data: American Museum of Natural History (AMNH); California Academy of Sci-
ences (CAS); John T. Polhemus, personal collection (JTP); Oregon State University
(OSU); National Museum of Natural History, Smithsonian Institution (USNM).

1984

NEW WESTERN CORIXID

43

vandykei

krafti washingtonensis mckinstryi

Figs. 1-8. 1-4. Hemelytron of female. 5-8. Right

vandykei

Sigara krafti, new species
(Figs. 1, 5, 9, 13, 17, 18)

Description. SIZE: Length: males, 5. 3-5. 8 mm; females, 5.4-6. 1 mm. Width of
head across eyes: males, 1.6-1. 8 mm; females, 1.7-1. 9 mm. Synthilipsis: males, 0.5-
0.6 mm; females, 0.6-0. 7 mm. COLOR: Pale yellow to yellowish brown ground color
with brown to fuscous markings; pronotal disk and hemelytra grayish yellow with
fuscous bands, sometimes lightly tinged with red. Pronotum with eight or nine trans-
verse bands, dark bands equal in width or slightly narrower than pale intervening
regions; middle one or two bands often broken, sometimes extending only across
central portion of disk. Clavus with regular transverse bands, sometimes forked at

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

mckinstryi

vandykei

Figs. 9-17. 9-12. Abdominal terga of male. 13-16. Mesepimeron. 17. Male pala.

outer margin; bands tending to coalesce along commissure of hemelytra. Corial
pattern distinctly transverse, bands coalescing to form longitudinal stripe along em-
bolium and short longitudinal band across from apex of clavus. Membranal pattern
irregular, sometimes fading along inner margin; membrane separated from corium
by pale line. Embolium smoky, often fading to yellowish brown beyond nodal furrow.
Head pale yellow, sometimes brownish yellow between eyes. Sternum and venter
mostly darkened; sternum sometimes lighter in females. Mesepimeron paid, mes-

1984

NEW WESTERN CORIXID

45

epimeral process moderately to heavily infuscated; metaepisternum pale, anterior
one-third to one-half fuscous. Legs pale, coxae lightly to moderately infuscated.
STRUCTURAL CHARACTERISTICS: Head half as long as pronotal disk; synthi-
lipsis about equal to width of eye; postocular space narrow; vertex slightly produced;
facial hairs sparse; fovea of male broad and shallow. Pronotum, clavus, and corium
strongly rastrate. Pronotum 1 .5 x -1.7 x as wide as long; lateral angles acute; posterior
margin angulate; median carina small but distinct. Mesepimeron broadest just an-
terad of ostiole, slightly narrower and parallel-sided between ostiole and lateral bend;
ostiole about two-fifths the distance from the tip to the lateral bend. Metaxyphus
distinctly broader than long. Pala of male with 29-34 pegs arranged in single, curved
row (Fig. 17). Male abdomen dorsally as in Fig. 9; abdominal strigil small, sub-
spherical, composed of four or five combs; posteromedial projection of seventh
abdominal tergum triangular. Right paramere of genital capsule as in Fig. 5.

Etymology. This species is named for Dr. Gerald F. Kraft (Biology Department,
Western Washington University, Bellingham, Washington); while under his super-
vision I developed a strong interest in the family Corixidae, as well as other aquatic
and semiaquatic Hemiptera.

Comparative notes. Sigara krafti is very similar to three other species of Sigara
found in Oregon and Washington. In Hungerford (1948) and Lauck (1979), S. krafti
keys to a couplet containing two of these species, S. mckinstryi and S. washing-
tonensis. Sigara krafti differs from these species by the more extensively darkened
embolium, fuscous anterior region of the metaepisternum, weakly produced vertex,
and by the shape of the right paramere of the male (compare Figs. 5-7). Sigara krafti
is further differentiated from S. washingtonensis by the more regular transverse
banding of the corium (compare Figs. 1 and 2), smaller abdominal strigil, and by
the shape of the posteromedial projection of the seventh abdominal tergum of the
male (compare Figs. 9 and 10). The pronotal disk of S. krafti is 1.5 x -1.7 x as broad
as long, whereas in S. washingtonensis the disk is nearly twice (1.8 x -1.9 x) as broad
as long. These two species also differ in the width of the postocular space. When
measured at the inner posterior angle of the eye, the postocular space of S. krafti is
distinctly narrower than the width of the middle tarsus. In S. washingtonensis, the
postocular space is equal to or greater than the tarsal width. Finally, males of S.
krafti usually have more palar pegs (29-34) than males of S. washingtonensis (26-
30), but some overlap occurs in the range of peg numbers of these two species.

The hemelytral patterns of S. krafti and S. mckinstryi are similar, but the dark
bands are usually narrower in S. krafti (compare Figs. 1 and 3). Sigara krafti also
differs from S. mckinstryi by the shape of the mesepimeral process. In the latter
species, the process is parallel-sided or weakly divergent from the ostiole to the lateral
bend (Fig. 15), whereas in S. krafti the process is broadest just anterad of the ostiole,
then slightly narrower to the lateral bend (Fig. 1 3).

Sigara krafti is superficially similar to a third Sigara species in Oregon and Wash-
ington, S. vandykei. The latter species is easily distinguished from S. krafti by the
subapical position of the ostiole on the mesepimeral process (compare Figs. 1 3 and
16), the explanate anterolateral margin of the clavus that projects laterally over the
clavoprunia, the protuberance on the posterolateral margin of the female hemelytron
(compare Figs. 1 and 4), and by the right paramere of the male (compare Figs. 5 and
8).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

SCALE IN MILES
50

0 50 100 150 O

SCALE IN KILOMETERS \

Fig. 18. Distribution of Sigara krafti Stonedahl, new species.

Distribution (Fig. 18). Sigara krafti is distributed west of the Cascade Range in
Oregon and Washington. The majority of records are from streams and small rivers,
but several specimens have been collected from small ponds (near rivers) in west-
central Oregon. This species appears to be most abundant in shallow, static or slow-
moving water over silty or muddy substrates. Three other species of Sigara are found

1984

NEW WESTERN CORIXID

47

in similar habitats in western Oregon and Washington, and are sometimes collected
with S. krafti. Of these, S. washingtonensis is encountered most often; the other
species are S. vandykei and S. omani.

Holotype. Male: Oregon, Benton Co., Mary’s R., 1 mi. W jet. St. Hwy, 34 on US
20, 2 Aug. 1983, G. M. Stonedahl (CAS).

Paratypes. OREGON. Benton Co.: 19 males and 19 females, same data as holotype
(AMNH, CAS, JTP, OSU, USNM); 4 males, 4 mi. S Blodgett, 26 March 1959 (OSU);
1 male, Log pond, W end Philomath, 17 April 1962, B. Courtright (OSU); 1 male,
Mary’s R., 3 mi. W Philomath, 26 April 1956, J. D. Lattin (OSU). Clatsop Co.: 1
male, East Humbug Crk., 23 April 1966, S. G. Jewett, Jr. (OSU). Josephine Co.: 1
male and 8 females, 10 mi. W Selma, 13 May 1972, Wu Shuang Lee (OSU). Lincoln
Co.: 1 female, Kasner Pond, 2 mi. E Burnt Woods, 22 April 1962, D. Hipply (OSU).
Multnomah Co.: 1 male, Rooster Rock, Columbia R. Gorge, 15 April 1972 (OSU).
WASHINGTON. Lewis Co.: 8 females, Chehalis R. at Alexander Park, 1 1 Sept.
1977, G. M. Stonedahl (author’s collection); 1 male, T13N, R3W, Sec. 18, 12 Sept.
1977, G. M. Stonedahl (author’s collection). Pacific Co.: 1 male, T14N, R8W, Sec.
19, 12 Sept. 1977, G. M. Stonedahl (author’s collection); 2 males and 2 females,
Willapa R., 9 Sept. 1930 (OSU); 19 males and 20 females, Willapa R., T13N, R8W,
Sec. 25, 12 Sept. 1977, G. M. Stonedahl (author’s collection). Thurston Co.: 1 female,
Tumwater, 3 Sept. 1929 (OSU).

ACKNOWLEDGMENTS

I thank John D. Lattin and Gerasimos Cassis (OSU), and Thomas J. Henry (SEL, ARS,
USDA, % USNM) for comments on the manuscript. Special thanks to Thomas J. Henry for
lending specimens of S. ornata from the National Museum of Natural History, and to Julie A.
Stonedahl for typing of the manuscript. Bonnie B. Hall provided the illustration of the hem-
elytron of S. krafti. The University of California Press, Berkeley granted permission to use the
illustrations of the hemelytra and male abdomens of S. mckinstryi, S. vandykei, and S. wash-
ingtonensis from Lauck (1979).

LITERATURE CITED

Hungerford, H. B. 1948. The Corixidae of the Western Hemisphere (Hemiptera). University
of Kansas Science Bulletin 32:1-827.

Lauck, D. R. 1979. Corixidae. Pages 87-123 in: A. S. Menke (ed.), The semiaquatic and
aquatic Hemiptera of California. Bulletin of the California Insect Survey 21:1-166.

Received October 31, 1983; accepted December 13, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 48-52

CARVALHOMA (HEMIPTERA: MIRIDAE):

REVISED SUBFAMILY PLACEMENT

Randall T. Schuh and Michael D. Schwartz

Department of Entomology, American Museum of Natural History,

New York, New York 10024

Abstract. — Carvalhoma Slater and Gross is transferred from the Phylinae to the Cylapinae.
The nymphal and adult male habitus, male genitalia, and claws of Carvalhoma malcolmae
Slater and Gross are figured. Characters used to determine subfamily placement are discussed,
and it is suggested that the Cylapinae, Isometopinae, and Psallops Usinger form a monophyletic
group.

Slater and Gross (1977) described from Australia two species of Carvalhoma on
the basis of female specimens. They placed the genus in the mirid subfamily Phylinae
on the basis of its hairlike (setiform) parempodia, simple sclerotized rings, and the
narrow straplike interramal sclerites in the female. We question the correctness of
this subfamilial assignment for the following reasons: the habitus of Carvalhoma is
unusual in the Phylinae; setiform parempodia occur in cylapine, deraeocorine, and
bryocorine mirids in addition to the Phylinae and are therefore not diagnostic of any
one of these groups; and the simple structure of the female genitalia illustrated by
Slater and Gross is not necessarily diagnostic for the Phylinae. We have examined
a male specimen of Carvalhoma malcolmae collected in forest litter at Talbingo
Mountain, 47.8 km S of Tumut, New South Wales, Australia, on April 3, 1976. The
specimen is illustrated in Figure 1. The nymph is illustrated in Figure 2. The male
of C. malcolmae is similar in appearance to the female as illustrated by Slater and
Gross (1977) but has a slightly more elongate and swollen pronotum and somewhat
differently shaped hemelytra which cover less of the abdomen. We have studied the
pretarsal structures (Figs. 3, 4) and male genitalia in detail (Figs. 5-9). We believe
our examination supports placement of Carvalhoma in the subfamily Cylapinae (we
have tentatively placed it in the tribe Cylapini based on the structure of the head
and the condition of the ostiolar peritreme which is without an ocelloid shining
tubercle). Our reasons are as follows.

1 . The claws are elongate and slender and lack pulvilli, attributes which, in con-
junction with the setiform parempodia, have traditionally been used to diagnose the
Cylapinae. None of these pretarsal characteristics is unique to the Cylapinae, however,
and therefore do not provide suitable defining characters for the group. The one
unique structural detail of the pretarsus of Carvalhoma and all other cylapines of
which we are aware is the subapical tooth, seen in Figures 3 and 4. This feature has
been illustrated by Knight (1918), Carvalho (1955), and Schuh (1976), but was not
observed in Carvalhoma by Slater and Gross. A similar claw structure occurs in the
Isometopinae as shown by McAtee and Malloch (1924) and Schuh (1976) and in
Psallops Usinger as shown by Schuh (1976), strongly suggesting that these three groups
together form a monophyletic unit in the Miridae. The strength of this argument lies

1984

CARVALHOMA : A CYLAPINE

49

Fig. 1 . Carvalhoma malcolmae. Adult male, dorsal view.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Figs. 2-4. Carvalhoma malcolmae. 2. Nymph, dorsal view. 3. Pretarsus. 4. Detail of claw.

in the fact that the subapical tooth occurs only in the Miridae and does not occur in
those subfamilies and/or tribes of Miridae which have been recognized as mono-
phyletic on the basis of other characters.

2. The Phylinae are recognized by the unique structure of the male genitalia, with
the straplike vesica, the phallotheca attached to the posterior wall of the genital
capsule (rather than the phallobase which is the condition in all other Miridae), and
the characteristic shape of the left clasper, which supports the phallotheca in repose.
The male genitalia of Carvalhoma malcolmae are clearly unlike those of any Phylinae
and appear not to have been derived from a phyline type. The parameres (Figs. 5,
6) are of a form similar to that found in some Cylapinae (see Carvalho and Lorenzato,
1978, and Carvalho, 1982) and in some Isometopinae (see Slater and Schuh, 1969).
In C. malcolmae the right and left parameres are almost identical in shape and size,
whereas in some cylapines (e.g., Fulvius spp.) the right paramere is extremely small.
The membranous vesica (Fig. 7) is of a type found in many cylapines (see Carvalho
and Lorenzato, 1978) and is similar in structure to that of some other non-phyline
Miridae. The small membranous vesica of malcolmae has a large secondary gonopore
(as also occurs in species of Cylapocerus and Peltidocylapus [Carvalho and Fontes,
1968]) with an additional heavily sclerotized ringlike structure basad of it which
surrounds the ductus seminis (Fig. 8). The phallotheca surrounds the vesica and is
attached to the phallobase; the dorsal surface is sclerotized and tapering distally; the
ventral surface is membranous (Fig. 9).

3. The structure of the head, with its protruding eyes and depressed vertex, and

1984

CARVALHO MA\ A CYLAPINE

51

Figs. 5-9. Carvalhoma malcolmae. 5. Left paramere, lateral view. 6. Right paramere, lateral
view. 7. Vesica, dorsal view. 8. Ring-like structure of vesica, ventral view. 9. Phallotheca,
ventral view.

the very long appendages, appear to be a synapomorphy for Carvalhoma with some
mirid taxa placed in the Cylapinae by Carvalho (1957) (e.g., Cylapus Say and Cor-
I covadocola Carvalho, the latter also known to dwell in the litter [Carvalho, 1948]).

The characters used by Slater and Gross to assign Carvalhoma to the Phylinae are
not unique to that group of Miridae. The claws and male genitalia appear to offer a
character unique to a grouping of mirid taxa which has not been formally recognized
by previous authors. Further study of other characters in the Cylapinae, Isometopinae,
and Psallops, will help to determine the validity of the theory that these three taxa
form a monophyletic group.

Our justification for the subfamilial placement of Carvalhoma is based on an
interpretation of characters which contradicts the hypothesis of relationships pre-
sented by Schuh (1976). In that paper the absence of ocelli was interpreted as a
character uniting all Miridae except Isometopinae and the subapical claw tooth was
considered to be plesiomorphic within the Miridae and to have been lost in the
phyline-orthotyline and mirine-deraeocorine-bryocorine clades. Furthermore, Schuh
gave no unequivocal defining characters for Psallops or the Cylapinae as groups. A
more parsimonious interpretation of character distributions is to lose the ocelli twice
and to define a group including Isometopinae, Cylapinae, and Psallops on its pos-
session of the subapical claw tooth.

The idea that Psallops and Isometopinae are related is not new; it was first suggested
by Carvalho (1956) and reassessed by Schuh (1974, 1976). In fact there seems to be

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

little justification for considering the pronotal collar of Cylapinae as homologous
with that found in the Mirinae-Deraeocorinae-Bryocorinae as was done by Schuh
(1976); the Bothriomirini lack a pronotal collar, a point overlooked by Schuh, and
in addition several orthotyline genera have well developed rounded collars.

ACKNOWLEDGMENTS

We thank Alex Slater for the gift of the male specimen of Carvalhoma malcomae, J. C. M.
Carvalho for the loan of type specimens of Corcovadocola, J. A. Slater for discussion, and
Kathleen Schmidt for preparation of the figures.

LITERATURE CITED

Carvalho, J. C. M. 1948. Mirideos Neotropicais. XXXV. Generos “Corcovadocola” n.g.,
“Guanabarea” n.g., e “Caulotops” Bergroth (Hemiptera). Rev. Brasil. Biol. 8:525-533.
Carvalho, J. C. M. 1955. Key to the genera of Miridae of the World. Bol. Paraense Emilio
Goeldi 11:1-151.

Carvalho, J. C. M. 1957. Catalogue of the Miridae of the World. Part I. Arq. Mus. Nac., Rio
de Janeiro 44:158 pp.

Carvalho, J. C. M. 1965. Insects of Micronesia. Heteroptera: Miridae: Miridae. Bernice P.
Bishop Museum, Honolulu 7:1-100.

Carvalho, J. C. M. 1968. Mirideos Neotropicais. CI. Revisao do complexo “ Cylapus" Say
com descricoes de generos e especies novos (Hemiptera). Rev. Brasil. Biol. 28:273-282.
Carvalho, J. C. M. 1982. Mirideos Neotropicais. CCXXXIX. Descricoes de algumas especies
de Cylapinae do Amazonas (Hemiptera). Acta Amazonica 12:81 1-815.

Carvalho, J. C. M. and L. M. Lorenzato. 1978. The Cylapinae of Papua New Guinea (He-
miptera: Miridae). Rev. Brasil. Biol. 38:121-149.

Knight, H. H. 1918. Synoptic key to the subfamilies of Miridae (Hemiptera-Heteroptera). J.
New York Ent. Soc. 26:40-44.

McAtee, W. L. and J. R. Malloch. 1924. Some annectant bugs of the superfamily Cimicoidea
(Heteroptera). Bull. Brooklyn Ent. Soc. 19:69-83.

Schuh, R. T. 1974. The Orthotylinae and Phylinae (Hemiptera: Miridae) of South Africa with
a phylogenetic analysis of the anti-mimetic tribes of the two subfamilies for the World.
Ent. Amer. 47:332 pp.

Schuh, R. T. 1976. Pre tarsal structure in the Miridae (Hemiptera) with a cladistic analysis of
relationships within the family. Amer. Mus. Novitates 2601:39 pp.

Slater, J. A. and G. F. Gross. 1977. A remarkable new genus of coleopteroid Miridae from
southern Australia (Hemipera: Heteroptera). J. Austr. Ent. Soc. 16:135-140.

Slater, J. A. and T. Schuh. 1 969. New species of Isometopinae from South Africa (Hemiptera:
Miridae). J. Ent. Soc. Southern Africa 32:351-366.

Received December 10, 1983; accepted January 6, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 53-60

LIFE HISTORY AND LABORATORY REARING OF
EUSCHISTUS ICTERICUS (HEMIPTERA: PENTATOMIDAE),
WITH DESCRIPTIONS OF IMMATURE STAGES

J. E. McPherson and S. M. Paskewitz1

Department of Zoology, Southern Illinois University,

Carbondale, Illinois 62901

Abstract.— The life history of Euschistus ictericus was studied in southern Illinois, and the
immature stages were described. The bug was also reared from egg to adult in the laboratory.
Adults emerged from overwintering sites in early May, began feeding on Carex comosa Boott,
and reproduced shortly thereafter. Eggs were generally laid on C. comosa and nymphs were
found only on this plant. No individuals were found after mid-August. E. ictericus was reared
on green beans under a 16L:8D photoperiod and constant temperature of 23.9 ± 1.1°C. The
incubation period averaged 7.33 days. Durations of the 5 subsequent stadia averaged 3.99,
7.72, 7.27, 9.03, and 12.62 days, respectively. The external anatomy of the egg and each of the
5 nymphal instars is described.

Euschistus ictericus (L.) occurs from eastern Canada south to Florida, and west
and southwest to Utah, Oklahoma, and Texas (McPherson, 1982). It has been col-
lected on sedges, Iris versicolor L., water lilies, Saururus cernuus L., Salix, yellow
thistle, cow-parsnip, Cuscuta, soybeans, and common mullein (McPherson, 1982).

Little has been published on the field life history of this stink bug. It overwinters
as adults (Blatchley, 1895; Parshley, 1 923). Torre-Bueno and Engelhardt (1910) found
mating specimens on 24 July on water lilies in North Carolina. This paper presents
information on the life history and laboratory rearing of E. ictericus and includes
descriptions of the immature stages.

MATERIALS AND METHODS

Life history. On 12 June 1981, several adults, including copulating pairs, were
observed on the heads, leaves, and stems of Carex comosa Boott growing in water
near the shore of La Rue Swamp, Union County, Illinois. The presence of copulating
adults suggested a life history study was possible and, therefore, a study was conducted
from June 1981 to July 1982. Collection of data during 1981 ended in August after
the insects left the host plants. During 1982, data were collected from April to July,
before the bugs appeared on, and after they left the plants.

Data were collected weekly during the study and consisted primarily of counts of
all stages present. Few animals were actually collected because the population size
appeared small. The only exceptions were a few egg clusters and young nymphs that
were reared to adults to confirm their identities and, with nymphs, to confirm their
instars when collected. Data gathered during the 2 years of the study were combined
to gain a better understanding of the annual life cycle.

1 Present address: Department of Entomology, University of Georgia, Athens, Georgia 30602.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Yol. 92(1)

Laboratory rearing. Fifteen adults (9 66, 699) were collected from C. comosa at La
Rue Swamp on 12 June 1981, taken to the laboratory, and placed in a 1-qt. (ca. 0.95-
liter) Mason jar covered on the bottom with filter paper. Green snap beans ( Phaseolus
vulgaris L.) served as food and were placed on end in the bottom of the jar. A strip
of paper toweling was added which, together with the filter paper, increased absorption
of excrement and provided a good walking surface for the nymphs. The jar was closed
with wire screen and paper toweling and secured with the band of the 2-piece Mason
jar lid. A cheesecloth strip (ca. 6 x 20 cm), which served as an oviposition site, was
placed inside the jar with 1 end over the lip and held in place by the band.

The cage was examined daily for eggs. Cheesecloth, with attached egg clusters, was
removed and placed on moist filter paper in the bottoms of petri dishes (ca. 9 cm
diam., 2 cm depth) and covered with the lids. Water was added daily to keep the
filter paper moist.

The first instars (an apparently nonfeeding stage) were also kept in the petri dishes.
The second through fifth instars were kept in Mason jars prepared similarly to the
oviposition cage except for the absence of cheesecloth.

Food, filter paper, and strips were changed every 3-4 days. The dishes and jars
were kept in an incubator maintained at 23.9 ± 1.1°C and a 16L:8D photoperiod.

Descriptions of immature stages. The description of each stage is based on 10
individuals that were collected from the laboratory culture and preserved in 95
percent ETOH. The first instar is described in detail, but only major changes from
previous instars are described for subsequent instars. Comparative statements refer
to previous instars (e.g., “more numerous”). Length is measured from tip of tylus to
tip of abdomen; width is measured across the mesonotum. Dimensions are expressed
in mm as x ± SE. Drawings were made with the aid of a camera lucida, measurements
with an ocular micrometer.

RESULTS AND DISCUSSION

Life history. Fifty-nine adults, 70 eggs, and 35 nymphs were found during this
study. Adults emerged from overwintering sites in early May and began feeding on
the heads of C. comosa. Shortly thereafter, they were observed copulating on the
heads, leaves, and stems of the host plant; as in many other pentatomoids (Mc-
Pherson, 1982), the copulating position was end-to-end.

The 70 eggs were laid in 6 clusters, an average of 1 1.7 eggs/cluster (range, 6-14).
These clusters and those deposited in the laboratory usually consisted of 2 to 4 rows
of eggs. The first cluster was collected on 14 May and had been deposited on a head
of C. comosa. The 5 remaining clusters were collected on 18 June; 4 had been laid
on C. comosa (2 on heads, 2 on bracts) and 1 on Sagittaria latifolia Willdenow (on
a leaf) that was growing in a stand adjacent to C. comosa.

No first or second instars were collected. Five third instars were collected between
4 June and 3 July, 12 fourths between 12 June and 30 July, and 18 fifths between
1 8 June and 6 August; all were collected on C. comosa. Adults occurred on C. comosa
until mid- August; most (55.2 percent) were observed between 12 and 18 June.

From these limited data, it is difficult to decide whether E. ictericus was uni- or
bivoltine at Pine Hills. Both interpretations are reasonable.

Laboratory rearing. The 230 eggs deposited (Table 1) were laid on the cheesecloth

1984

BIOLOGY OF EUSCHISTUS ICTERICUS

55

Table 1.

Duration (in days) of each immature stage of E. ictericus.

No.

Days

Stage

completing

stadium

Range

x ± SE

Cumulative
mean age

Egg

227a

6-8

7.33 ± 0.04

7.33

Nymph
1 st instar

226

3-5

3.99 ± 0.05

11.32

2nd instar

205

6-15

7.72 ± 0.09

19.04

3rd instar

184

5-14

7.27 ± 0.10

26.31

4th instar

171

6-18

9.03 ± 0.13

35.34

5th instar

165

9-21

12.62 ± 0.15

47.96

a 230 eggs were laid.

in 14 clusters, an average of 16.4 eggs/cluster (range, 8-31). The incubation period
averaged 7.33 days. The eye spots and mouth parts were visible in 3-4 days.

The first instars were gregarious and remained atop the egg shells unless disturbed.
They apparently did not feed. The duration of this stadium averaged 3.99 days.

The durations of the second through fifth stadia averaged 7.72, 7.27, 9.03, and
12.62 days, respectively. Total developmental time from egg to adult averaged 47.96
days. Of the 226 individuals that began the second stadium, 73.0 percent (165
individuals) reached adults.

Descriptions of immature stages. EGG (Fig. 1). Length, 1.13 ± 0.01; width, 1.00 ±
0.0 1 . Generally laid in clusters of 1 4; each egg kettle-shaped, yellowish white. Chorion
with irregular triangular and quadrangular reticulations; spine at apex of each angle.
Operculum present, surrounded by 34-38 micropylar processes, each process ca. 0.06
mm long and slightly dilated at tip.

FIRST INSTAR (Figs. 2, 3). Length, 1.67 ± 0.02; width, 1.23 ±0.01. Body oval,
greatest width at abdominal segments 2-3. Punctures present dorsally and ventrally,
all punctures minute.

Head declivent, anterolateral margins sinuate; yellowish brown to light brown
dorsally with vertex yellow medially and tylus reddish to yellowish brown, often with
red U-shaped marking between eye and midline of head; tylus exceeding juga; line
extending from each eye posteromedially and disappearing beneath pronotum. Eyes
red. Antennae 4-segmented; segments 1-3 brownish to red; segment 4 largest, fusi-
form, reddish brown to brown; incisures albidus; distinct constrictions at junctures
of 2-3 and 3-4; ratio of antennal segment lengths ca. 10:11:12:25. Ventral surface
of head whitish to yellowish brown. Beak 4-segmented, whitish to brown, segments
3-4 darker.

Thoracic nota reddish brown to brown, yellow medially, yellow mediolongitudinal
line extending from anterior margin of pronotum nearly to or reaching posterior
margin of metanotum; lateral margins entire; pro- and mesonota sclerotized, posterior
margins arcuate; metanotum sclerotized except posteriorly, posterior margin of me-
tanotal plate slightly arcuate medially, bending cephalad laterally; faint calli occa-
sionally present on all segments; intersegmental line between metanotum and first

56

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

5

Figs. 1-5. Immature stages of E. ictericus. 1. Egg. 2, 3. First instar. 4. Second instar. 5.
Third instar.

1984

BIOLOGY OF EUSCHISTUS ICTERIC US

57

abdominal tergum often sclerotized either side of midline ca. one-half way between
lateral margin and midline of body. Pleura light brown; pro- and mesopleura fused
to respective nota; metapleura separated from metanotal plate by membranous area.
Spiracles located on posterior margins of pro- and mesopleura. Sterna concolorous
with ventral surface of abdomen. Coxae whitish, each with central brown mark on
lateral surface; trochanters pale brown; femora brownish with apices reddish; tibiae
reddish with apices yellowish, front tibiae each with bifurcate spine on inner posterior
margin of distal one-third; tarsi 2-segmented, yellowish, apex of segment 2 darker;
tarsal claws and pul villi brown to yellowish brown.

Dorsum of abdomen white, or yellow, with red markings; sparsely punctate light
brown to brown medial and lateral plates present. Faint pseudointersegmental lines
on all but first and last segments, each originating at inner margin of lateral plate.
Eight medial plates present; plates 1-2 small, plate 2 may be weakly sclerotized; plate
3 narrowed medially; plate 4 subrectangular with markedly irregular margins, slightly
narrower than, and 4 to 5 times medial length of, plate 3; plate 5 subtrapezoidal with
markedly irregular margins, slightly narrower than, and 5 to 6 times medial length
of, plate 3; plate 6 often consisting of anterior transverse strip and paired posterior
pieces; plate 7 variable in shape, often split or partially split medially; plate 8 fused
to laterals; paired ostioles of scent glands located on plates 3-5. Nine lateral plates
present, subelliptical, extending dorsally and ventrally from margin of abdomen;
plate 1 small; plates 2-6 largest; remainder generally decreasing in size posteriorly,
i Sterna generally concolorous with dorsum, occasionally redder; segments 5-9 oc-
casionally with weakly sclerotized subrectangular medial plates, that of sternum 9
fused with laterals. Spiracles located on segments 2-8, those of segment 8 reduced,
each with brown peritreme. A single trichobothrium located posteromesad to each
spiracle on segments 3-7.

SECOND INSTAR (Fig. 4). Length, 2.15 ± 0.04; width, 1.77 ± 0.02. Body broadly
j pyriform; dorsum of head and thorax with numerous punctate brown spots, punctures
large, brown; ventral punctures still minute.

Head less declivent, anterolateral margins more sinuate; white to yellow dorsally
with posterior margin dark brown, oval brown spot slightly medial to each eye, red
U-shaped markings of first instar absent, yellow area on vertex replaced by faint
I yellow V-shaped marking, tylus white to reddish. Antennal segments 1 and 4 brown;
segments 2-3 reddish brown to brown, each red distally; incisures albidus to red;
ratio of antennal segment lengths ca. 1 2:2 1 :20:36. Ventral surface of head dark brown
except for white area either side of beak and white to red strip beneath beak.

Thoracic nota white to yellow, lateral margins explanate, dentate, edged with brown
(pro- and mesonota), posterior margins brown; mesonotum with medial area ex-
tended posteriorly; intersegmental line between metanotum and first abdominal ter-
gum sclerotized either side of midline ca. one-half way between lateral margin and
midline of body; pro- and metanota each with 1 pair, and mesonotum with 2 pairs,
of brown calli. Pleura white, each edged with brown and with brown spots and brown
longitudinal stripes. Sterna red. Coxae white, each with lateral edge and central spot
brown; trochanters white; femora each with proximal one-half white, distal one-half
brown, apex often red; tibiae reddish brown to brown, red basally, carinate; tarsi
brown.

58

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Dorsum of abdomen white to yellow with numerous, short, longitudinal, red mark-
ings. Medial and lateral plates with minute punctures. Medial plates 1-2 absent;
plates 3-8 brown; plate 3 with transverse white stripe; plate 4 subtrapezoidal without
marked irregular margins, ca. 3 times medial length of plate 3, transverse white
marking present that is often anchor-shaped; plate 5 without marked irregular mar-
gins, ca. 4 times medial length of plate 3, white markings present and consisting of
transverse anchor-shaped mark and short mediolongitudinal mark posterior to an-
chor; plate 6 small, linear, not divided into anterior strip and posterior pieces; plate
7 often undivided, white marking present medially; plate 8 with white marking present
medially. Lateral plates white, margins brown, generally 1-4 brown spots in white
areas. Ventrally, linear sclerite present posterior to each metacoxa. Sterna concolorous
with dorsum in lateral one-half, whitish medially with sparse red markings; medial
plates brown with minute punctures, now often on segments 4-9. Two trichobothria
posterior to each spiracle on segments 3-7.

THIRD INSTAR (Fig. 5). Length, 4.04 ± 0.06; width, 2.40 ± 0.02. Dorsum of
head and thorax with punctate brown spots limited primarily to humeri, remaining
punctures more numerous and relatively smaller.

Head with anterolateral margins more sinuate; dorsally, yellow to yellow with
lateral one-half of juga reddish, oval brown spot of second instar near each eye now
yellow to yellowish brown; 2 red ocelli present posteromedially but often obscured
by brown along posterior margin of head. Antennal segment 1 yellow to brown, with
red to brown markings; segments 2-3 red; segment 4 reddish brown to brown; ratio
of antennal segment lengths ca. 13:30:25:40. Ventral surface of head brown except
for white areas either side of beak and around bases of antennae, yellow area which
is often present between beak and eye, and white to red strip beneath beak.

Thoracic nota with paired calli yellow to brown; pronotum with medial area slightly
extended posteriorly. Femora each with proximal two-thirds white, distal one-third 1
mottled.

Dorsum of abdomen yellow with numerous short, longitudinal, yellow to red
markings; terga 1-7 each with red to brown spot midway between lateral margin and i
midline of abdomen, spot may be obscure. Medial plate 3 yellow with brown anterior
and posterior margins; plates 4-5 with markings similar to third instar but yellow
instead of white, plates 3-5 often with brown spots on yellow areas; plates 7-8 yellow
with brown spots. Lateral plates with brown spots small and more numerous. Sterna
with medial plates on segment 4-9 varying from almost transparent with brown spots
to brown; intersegmental lines between segments 2-3, 3-4, and 4-5 often sclerotized
either side of midline.

FOURTH INSTAR (Fig. 6). Length, 6.20 ± 0.09; width, 3.62 ± 0.04. Dorsum
of head and thorax with punctures more numerous in some areas and generally
relatively smaller.

Head yellow dorsally with lateral one-third of juga reddish; ocelli now always
visible. Antennal segment 1 yellow, apex brown, with red to brown spots; segment
2 yellow to reddish yellow; segment 3 yellow to red; segment 4 red to yellow proxi-
mally, brown distally; ratio of antennal segment lengths ca. 5:13:10:14. Ventral sur-
face of head yellow except for lateral dark brown stripe extending from each eye at
least to base of antennal segment 1, dark brown mark behind eye that may be
continuous with dark brown mark on either side of midline along posterior margin

1984

BIOLOGY OF EUSCHISTUS ICTERICUS

59

Figs. 6, 7. Immature stages of E. ictericus. 6. Fourth instar. 7. Fifth instar.

of head, and occasional pink spot beneath beak. Beak with segment 1 yellow to yellow
with apex brown; segment 2 yellow to brown; segments 3-4 yellowish brown to
brown.

Thoracic nota with brown borders along posterior margins reduced or absent;
humeri with brown spots more numerous; calli yellow, those of meso- and metanota
often with brown central spot. Meso- and metanotal wing pads ca. the same length,
extending onto first abdominal segment. Pleura with brown spots and brown lon-
gitudinal stripes limited primarily to lateral one-half except for 1 brown spot at base
\ of each coxa. Sterna yellow. Coxae white to yellowish white, each with light brown
band on lateral edge absent and central brown spot occasionally absent; trochanters
whitish yellow; femora yellow to reddish yellow proximally, yellow with reddish to
brown spots distally; tibiae yellow to reddish brown, with red to brown markings;
tarsal segment 1 yellow, apex often darker, segment 2 yellow basally, brown distally.

Dorsum of abdomen yellow with red spots medially. Medial plates 3-5 with brown
spots more numerous; plates 4-5 yellow to brown and, if yellow, anchor-shaped
markings of third instar obscured; plates 7-8 with brown spots more numerous.
Lateral plates transparent, each with brown margins broken and sometimes reduced
to large brown spot at base of pseudointersegmental line on plates 2-7; small brown
I spots often more numerous but may be absent. Ventrally, linear sclerite posterior to
jeach metacoxa reduced to 1-4 brown spots. Sterna yellow; medial plates reduced or
transparent, to transparent with a few small brown spots; sclerotized stripes on
1 intersegmental lines between segments 2-3, 3-4, and 4-5 not evident.

FIFTH INSTAR (Fig. 7). Length, 9.30 ± 0.09; width, 5.38 ± 0.12. Dorsum of
head and thorax with punctures more numerous in some areas and generally relatively
' smaller.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Head with posterior margin yellow to yellowish brown, yellow to yellowish brown
oval spot near eye of third and fourth instars now broadly crescent-shaped. Antennal
segment 1 yellow to reddish, with brown spots; segment 2 yellow to red, with brown
spots; segments 3-4 red to yellowish red, apex of segment 4 darker; ratio of antennal
segment lengths ca. 7:22:15:19. Ventral surface of head without pink spot beneath
beak, brown mark either side of midline along posterior margin of head often reduced
or absent.

Humeri with brown spots more numerous and extended anteriorly in band along
each lateral margin. Meso- and metanotal wing pads ca. same length, extending onto
third abdominal segment. Pleura yellow, brown spot at base of each coxa, propleura
each with brown mark present laterally, other brown spots and brown stripes of
fourth instar absent. Coxae white, without central brown spot on lateral surface;
femora and tibiae yellowish, with red to brown spots.

Medial plates 3-5 with brown spots more numerous; plate 6 reduced or absent.
Lateral plates with brown margins much reduced, large brown spot at base of pseu-
dointersegmental lines still present but limited to 3-7, other small brown spots of
fourth instar absent or much reduced. Ventrally, linear sclerite posterior to each
metacoxa not evident; medial plates generally obscure or absent, brown spots absent.

ACKNOWLEDGMENTS

We thank Karen A. Schmitt, Scientific Photography and Illustration Facility, SIU-C, for the
final illustrations of the immature stages. Cost of the illustrations were met by the Office of
Research Development and Administration, SIU-C, and the Department of Zoology.

LITERATURE CITED

Blatchley, W. S. 1895. Notes on the winter insect fauna of Vigo County, Indiana— II. Psyche
7:267-270.

McPherson, J. E. 1 982. The Pentatomoidea (Hemiptera) of Northeastern North America with
Emphasis on the Fauna of Illinois. Southern Illinois University Press, Carbondale and
Edwardsville, 240 pp.

Parshley, H. M. 1923. Families Pentatomidae, Cydnidae, Scutelleridae. In W. E. Britton,
Guide to the insects of Connecticut. Part IV. The Hemiptera or sucking insects of
Connecticut. Conn. State Geol. Natur. Hist. Surv. Bull. 34:1-807.

Torre-Bueno, J. R. de la and G. P. Engelhardt. 1910. Some Heteroptera from Virginia and
North Carolina. Can. Entomol. 42:147-151.

Received March 2, 1983; accepted July 27, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 61-68

LABORATORY REARING OF AMAUROCHROUS CINCTIPES
(HEMIPTERA: PENTATOMIDAE: PODOPINAE) WITH
DESCRIPTIONS OF IMMATURE STAGES

J. E. McPherson and S. M. Paskewitz1

Department of Zoology, Southern Illinois University,

Carbondale, Illinois 62901

Abstract.— Amaurochrous cinctipes (Say) was reared in the laboratory under a 16L:8D pho-
itoperiod and constant temperature. Egg and first instar development averaged 6.13 and 4.98
days, respectively. The second through fifth stadia for nymphs reared on Sagittaria latifolia
Willdenow averaged 15.91, 16.80, 15.33, and 20.00 days, respectively; and on green beans
averaged 14.20, 18.67, 15.67, and 17.00 days, respectively. The external anatomy of the egg
and each of the 5 nymphal instars is described.

Amaurochrous cinctipes (Say) occurs from Quebec and New England south to the
Carolinas, and west and southwest to Minnesota, Nebraska, Kansas, Missouri, Lou-
isiana, and Texas (Barber and Sailer, 1953). It generally seems to occur on vegetation
in low marshy places or near bodies of water (McPherson, 1982) and to overwinter
i beneath cover near water (Blatchley, 1926). It has been collected on Typha, Scirpus,
Carex, Juncus gerardi Loiseleur-Deslongchamps, and at Spartina consocies (Mc-
Pherson, 1982); nymphs have been found in high numbers on Typha (Davis, 1925)
and J. gerardi (Parshley, 1923). The egg, first, third(?), and fifth instars have been
briefly described (Parshley, 1923). This paper presents information on the laboratory
rearing of A. cinctipes and descriptions of the immature stages.

1

j MATERIALS AND METHODS

Laboratory rearing. Between 12 June and 3 July 1981, 23 adults, including cop-
ulating pairs, were observed on the leaves and petioles of duck-potato ( Sagittaria
latifolia Willdenow) growing in water near the shore of La Rue Swamp, Union County,
; Illinois. Ten specimens (3<$<3, 799) were collected, taken to the laboratory, and placed
in an oviposition cage.

The oviposition cage consisted of a round, glass battery jar (ca. 16 cm diam., 20
| cm depth) with filter paper covering the bottom. A freshly cut petiole of duck-potato,
and attached leaf, served as food and was placed in a water-filled vial stoppered with
cotton. A fresh cutting was provided every 3-4 days, and the vial was refilled as
needed. The jar was closed with 2 layers of cheesecloth held in place with an elastic
band.

The cage was examined daily for eggs. Plant sections, with attached eggs, were
removed, placed on moist filter paper in the bottoms of petri dishes (ca. 9 cm diam.,

2 cm depth), and covered with the lids. Water was added daily to keep the filter
paper moist.

1 Present address: Department of Entomology, University of Georgia, Athens, Georgia 30602.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

The first instars (an apparently nonfeeding stage) were also kept in the petri dishes.
The second instars were transferred to one of two types of containers and reared to
adults. The first type consisted of a lantern globe (ca. 16.8 cm high) set on a clay
flower pot (ca. 10.5 cm ID at top) filled with white silica sand and placed in a
fingerbowl; water was added as needed to the sand to compact it and provide the
insects with better footing. A freshly cut petiole of duck-potato, with attached leaf,
served as food and was placed in a water-filled vial stoppered with cotton; the vial
was pushed into the sand up to the lip to provide the nymphs easy access to the
plant. A fresh cutting was provided every 3-4 days and the vial was refilled as needed.
The globe was closed with 2 layers of cheesecloth held in place with an elastic band.

The second type of rearing container consisted of a 1-qt. (ca. 0.95-liter) Mason jar
covered on the bottom with filter paper. Green snap beans ( Phaseolus vulgaris L.)
served as food and were placed on end in the bottom of the jar. A strip of paper
toweling was added and, together with the filter paper, increased absorption of ex-
crement and provided a good walking surface for the nymphs. The jar was closed
with wire screen and paper toweling and secured with the band of the 2-piece mason
jar lid. Food, filter paper, and paper toweling were changed every 3-4 days.

The oviposition cage and rearing containers were kept in an incubator maintained
at 23.9 ± 1.1°C and a 16L:8D photoperiod.

Descriptions of immature stages. The description of each stage is based on 10 !
individuals, unless stated otherwise, that were collected from the laboratory culture
and preserved in 95 percent ETOH. The first instar is described in detail, but only
major changes from previous instars are described for subsequent instars. Compar-
ative statements refer to previous instars (e.g., “more numerous”). Length is measured
from tip of tylus to tip of abdomen; width is measured across the mesonotum.
Dimensions are expressed in mm as x ± SE. Drawings were made with the aid of a
camera lucida, measurements with an ocular micrometer.

RESULTS AND DISCUSSION

Laboratory rearing. Eggs were deposited in clusters on the petioles and leaves of I
duck-potato. Each cluster consisted of an alternating double row of 9 or 10 eggs (N = -j
7, x = 9.71). Each egg was yellowish brown with a brown operculum. A triangular
egg burster was readily visible after hatching. The incubation period averaged 6.13
days (Table 1).

The first instars were gregarious and remained atop the egg shells unless disturbed.
They apparently did not feed. The duration of this stadium averaged 4.98 days.

The duration of the second through fifth stadia of nymphs reared on duck-potato
averaged 15.91, 16.80, 15.33, and 20.00 days, respectively; and on green beans ,
averaged 14.20, 18.67, 15.67, and 17.00 days, respectively. Total developmental
time from egg to adult on the 2 plants averaged 79.53 and 76.13 days, respectively.

Heaviest mortality on both duck-potato and green beans (an unnatural host) oc-
curred during the second stadium, the earliest stadium in which nymphs apparently
feed. This and the fact that no eggs or nymphs were ever found on duck-potato at
La Rue Swamp, and no adults after early July, suggest that this plant is not a natural
host.

Descriptions of immature stages. EGG (Fig. 1). Length, 0.93 ± 0.01; width, 0.63 ±
0.01. Generally laid in clusters of 10; each egg cylindrically shaped, yellowish brown

1984

BIOLOGY OF AMAUROCHROUS CINCTIPES

63

Table 1 . Duration (in days) of each immature stage of A. cinctipes.

No

Days

Stage

completing

stadium

Food

source

Range

Jc ± SE

Cumulative
mean age

Egg

37a

6-7

6.24 ±

0.07

6.24

27b

—

5-7

5.96 ±

0.16

5.96

Total

64

5-7

6.13 ±

0.08

-

Nymph

1st instar0

36

—

5-6

5.25 ±

0.07

11.49

27

—

4-5

4.63 ±

0.09

10.59

Total

63

4-6

4.98 ±

0.07

-

2nd instar

11

duck-potato

12-19

15.91 ±

0.58

27.40

5

green beans

13-16

14.20 ±

0.58

24.79

3rd instar

5

duck-potato

14-23

16.80 ±

1.59

44.20

3

green beans

16-22

18.67 ±

1.76

43.46

4th instar

3

duck-potato

15-16

15.33 ±

0.33

59.53

3

green beans

13-18

15.67 ±

1.45

59.13

5th instar

1

duck-potato

—

20.00

79.53

3

green beans

16-19

17.00 ±

1.00

76.13

a 38 eggs were laid (nymphs subsequently reared on duck-potato).
b 30 eggs were laid (nymphs subsequently reared on green beans).
c Nonfeeding stage.

with brown operculum at oviposition but fading to yellow after preservation. Chorion
with small, irregular pentagonal and hexagonal reticulations. Operculum surrounded
by 18-23 micropylar processes, each process ca. 0.03 mm long.

FIRST INSTAR (Figs. 2, 3). Length, 1.05 ± 0.02; width, 0.82 ± 0.01. Body
elliptical-ovoid, greatest width at abdominal segments 1-2. Large brown punctures
present dorsally but not ventrally.

Head declivent, anterolateral margins sinuate; brown dorsally; tylus exceeding juga.
Eyes red. Antennae 4-segmented; segments 1-3 brown to light brown; segment 4
largest, fusiform, yellowish brown to brown, apex darker; incisures albidus; distinct
constrictions at junctures of 2-3 and 3-4; ratio of antennal segment lengths ca. 21:
22:20:57. Ventral surface of head light brown. Beak 4-segmented, whitish to brown.

Thoracic nota brown, mediolongitudinal line extending from anterior margin of
pronotum nearly to or reaching posterior margin of metanotum; pro- and mesonota
sclerotized, posterior margins moderately arcuate; metanotum sclerotized except pos-
teriorly, posterior margin of plate straight medially, bending cephalad laterally. Pleura
concolorous with and reaching nota. Spiracles located on posterior margins of pro-
and mesopleura. Sterna concolorous with ventral surface of head and abdomen. Coxae
light brown to brown, each with central brown spot on lateral surface; trochanters
light brown to brown; femora and tibiae reddish brown to brown, tibiae slightly
carinate, dilated distally, front tibiae each with bifurcate spine or inner posterior
margin of distal one-third; tarsi 2-segmented, yellowish, apex of segment 2 darker;
tarsal claws and pulvilli yellow, translucent.

64 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Figs. 1-5. Immature stages of A. cinctipes. 1. Egg. 2, 3. First instar. 4, 5. Second instar.

1984

BIOLOGY OF AMAUROCHROUS CINCTIPES

65

Dorsum of abdomen whitish to reddish with punctate brown medial and lateral
plates. Faint pseudointersegmental lines on all but first and last segments, each
originating at inner margin of lateral plate. Eight medial plates present; plates 1-2
linear; plate 3 subrectangular, slightly constricted medially, partially fused to plate
4; plate 4 subtrapezoidal with posteromedial notch, subequal in width and ca. 3-4
times medial length of plate 3; plate 5 subtrapezoidal with posteromedial notch, ca.
three-fourths width and ca. 4-5 times medial length of plate 3; plate 6 often with
posteromedial notch that almost bisects the plate; plates 7-8 small, paired; paired
ostioles of scent glands located on plates 3-5. Nine lateral plates present, subelliptical,
extending dorsally and ventrally from margin of abdomen, plates 1-8 (occasionally
9) punctate; plate 1 small; plates 2-5 largest; remainder generally decreasing in size
posteriorly. Sterna concolorous with dorsum, segments 4-9 with faint medial plates.
Spiracles readily apparent only on segments 2-7. A single trichobothrium located
posterior to each spiracle only on segments 4-5.

SECOND INSTAR (Figs. 4, 5). Length, 1.59 ± 0.03; width, 1.14 ± 0.02. Body
elliptical, greatest width at metanotum and abdominal segments 1-2. Punctures more
numerous dorsally, now present ventrally.

Head less declivent, anterolateral margins more sinuate; yellow to brown dorsally
with 2 brown markings near base of tylus that extend and converge ca. one-half way
to posterior margin of head, and brown mark adjacent to inner margin of each eye
that occasionally extends back to and then medially along posterior margin of head;
2 small red ocelli present posteromedially. Eyes protruding. Antennal segments brown,
incisures albidus to red; ratio of antennal segment lengths ca. 10:13:12:29. Ventral
surface of head concolorous with dorsum except for white to yellow strip beneath
beak and along posterior margin of head. Beak whitish to yellow with apex of last
segment darker.

Thoracic nota yellow to brown, lateral margins yellowish, explanate, and dentate;
pro- and metanota each with 1 pair, mesonotum with 2 pairs, of brown calli; pro-
and mesonota with medial area extended posteriorly. Meso- and metapleura of ex-
panded specimens separated from respective nota by membranous area. Trochanters
white to brown; femora and tibiae brown, each lighter at base, tibiae more carinate.

Dorsum of abdomen with medial plates 3-5 and often 8 yellowish brown to brown;
lateral plates yellowish. Medial plates 1-2 poorly defined, often appearing as a trans-
verse row of punctures; plate 3 more constricted medially, not fused to plate 4; plates
4-5 subequal in size, slightly wider than, and ca. 4 times medial length of, plate 3,
each plate with posteromedial notch greatly reduced; plate 6 small, linear, without
posteromedial notch; plate 7 greatly reduced or absent; plate 8 fused to laterals, not
paired. Lateral plates slightly crenate along margin of abdomen. Sterna 5-9 (and
often 4) with sclerotized subrectangular medial plates, plates impunctate or nearly
so; nonsclerotized portions of sterna impunctate medially. Spiracles now apparent
on segments 2-8, those of 8 much reduced.

THIRD INSTAR (Figs. 6, 7). Length, 2.37 ± 0.06; width, 1.69 ± 0.02. Body with
greatest width at abdominal segments 1-3. Punctures more numerous dorsally and
ventrally.

Head less declivent, 2 brown markings near base of tylus converging two-thirds
way to posterior margin of head, brown marking along posterior margin of second
instar generally absent. Eyes dark reddish brown. Ratio of antennal segment lengths

66

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

0.5 mm

9

Figs. 6-9. Immature stages of A. cinctipes. 6, 7. Third instar. 8. Fourth instar. 9. Fifth instar.

1984

BIOLOGY OF AMAUROCHROUS CINCTIPES

67

ca. 7:9:8:19. Ventral surface of head yellowish brown to brown except for white to
yellow strip beneath beak and along posterior margin of head, and occasional yellow
area on either side of beak that is continuous with yellowish brown stripe between
it and base of antenna.

Thoracic nota yellow to brownish yellow; pro- and mesonota often with white to
yellow markings posterior to calli. Pleura brown. Sterna white to pink. Coxae pinkish
to light brown, each with central brown spot on lateral surface; trochanters white;
femora whitish basally, brown distally.

Dorsum of abdomen whitish to yellow, with red markings; medial plate 7 absent;
plate 8 yellow. Sterna 4-9 with sclerotized subrectangular medial plates, posterior
plates with a few large punctures; nonsclerotized portions of posterior sterna punctate
medially. A single trichobothrium located posterior to each spiracle on segments
3-7.

FOURTH INSTAR (Fig. 8). Length, 3.84; width, 2.24; 1 specimen examined.
Body with greatest width at abdominal segment 3. Punctures more numerous dorsally
and ventrally.

Head less declivent, yellowish brown, 2 brown markings near base of tylus con-
I verging near posterior margin of head; tylus and juga subequal in length. Ratio of
antennal segment lengths ca. 9:13:11:24. Ventral surface of head brown, markings
; similar to third instar except that yellow area on either side of beak is larger and
j yellowish brown stripe between it and base of antenna is now red.

Thoracic nota yellowish brown; lateral margins of metanotum not explanate or
dentate; pro- and mesonota with markings posterior to calli now whitish yellow,
i Meso- and metanotal wing pads ca. same length, extending onto first abdominal
segment. Pleura reddish brown. Sterna whitish. Coxae whitish with central brown
spot on lateral surface; trochanters whitish.

Dorsum of abdomen whitish yellow with red markings; medial plates 3-5 yellowish
brown.

FIFTH INSTAR (Fig. 9). Length, 5.00; width, 3.22; 1 specimen examined. Body
with greatest width at abdominal segments 2-3. Punctures more numerous dorsally
and ventrally.

Head brownish yellow. Antennal segment 1 yellowish brown; segment 2 yellowish
brown with apex red; segment 3 brown; segment 4 black; ratio of antennal segment
lengths ca. 12:21:14:34. Ventral surface of head dark brown, markings similar to
fourth instar except that red stripe extending from beak to base of antenna is now
j yellow.

Thoracic nota brownish yellow; pro- and mesonota with markings posterior to
i calli now white. Meso- and metanotal wing pads ca. same length, extending onto
third abdominal segment. Pleura brown, yellowish at bases of coxae. Coxae whitish,

! central brown spot faint; femora whitish basally, whitish with brownish markings
distally; tibiae yellowish brown.

Dorsum of abdomen with medial plates 3-5 yellow with brown margins.

ACKNOWLEDGMENTS

We thank Dr. Carl W. Schaefer, University of Connecticut, Storrs, for determining the
trichobothrial patterns of the first-third instars. We are also grateful to Karen A. Schmitt,

| Scientific Photography and Illustration Facility, SIU-C, for the final illustrations of the immature

68

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

stages. Costs of these illustrations were met by the Office of Research Development and Admin-
istration, SIU-C, and the Department of Zoology.

LITERATURE CITED

Barber, H. G. and R. I. Sailer. 1953. A revision of the turtle bugs of North America (Hemiptera:
Pentatomidae). J. Wash. Acad. Sci. 43:150-162.

Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America with Especial
Reference to the Faunas of Indiana and Florida. Nature Pub. Co., Indianapolis, 1,116

pp.

Davis, W. T. 1925. Note on Podops cinctipes and Solubea pugnax. Bull. Brooklyn Entomol.
Soc. 20:147.

McPherson, J. E. 1 982. The Pentatomoidea (Hemiptera) of Northeastern North America with
Emphasis on the Fauna of Illinois. Southern Illinois Univ. Press, Carbondale and Ed-
wardsville, 240 pp.

Parshley, H. M. 1923. On the ecology of Podops cinctipes Say and Rhytidolomia saucia Say,
(Hemiptera, Pentatomidae). Can. Ent. 55:69-71.

Received March 2, 1983; accepted July 27, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 69-86

A CONSPECTUS OF PENTATOMINI
OF THE WESTERN HEMISPHERE. PART 3
(HEMIPTERA: PENTATOMIDAE)

L. H. Rolston and F. J. D. McDonald

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University Agricultural Center, Louisiana State University,
Baton Rouge, Louisiana 70803, and
Department of Plant Pathology and Agricultural Entomology,

University of Sydney, Sydney, New South Wales 2006, Australia

Abstract.— A key is provided for those North American, Central American and West Indian
genera of Pentatomini that do not have a basal tubercle or spine on the abdominal venter.
Pentatoma antiguensis Westwood, Thyanta bimini Ruckes, T. elegans Malloch and T. elegan-
tula Jensen-Haarup are transferred to Cyptocephala Berg from Thyanta St&l. Tepa Rolston and
McDonald, new genus, is proposed for 6 species transferred from Thyanta: Pentatoma rugulosa
Say, type species, Thyanta brevis Van Duzee, T. jugosa Van Duzee, T. panda Van Duzee, T.
panctiventris Van Duzee and T. yerma Rolston. Caribo Rolston, new genus, is proposed for C.
subgibbus Rolston, new species from the Bahamas, and C. fasciatus Rolston, new species from
Jamaica and St. John, Virgin Islands. Neurohalys Bliven, 1960, is placed in the synonymy of
Prionosoma Uhler, 1863, and N. bucculatus Bliven, 1960, in the synonymy of P. podopioides
Uhler, 1863. Thyanta subgenus Parathyanta Jensen-Haarup, 1928, is a junior synonym of
Cyptocephala Berg, 1883. Phacidium euchlorum Breddin, 1912, whose type locality was er-
roneously reported as Costa Rica, is a junior synonym of Thyanta aeruginosa Berg, 1878, a
species known only from Argentina and Uruguay. A lectotype is designated for Phacidium
euchlorum Breddin.

We have previously provided keys for those genera of Pentatomini of the Western
Hemisphere which possess a spine or tubercle at the base of the abdominal venter
(Rolston et al., 1980; Rolston and McDonald, 1981). This paper concerns those
genera of Pentatomini that do not have such a spine or tubercle, but it is geographically
restricted to exclude South America.

KEY TO GENERA

1 . Margins of propleura before coxae produced, anterior margin covering base of

head from eyes mesad nearly to rostrum (Fig. 1) 48

Propleura not produced 2

2(1). Sulcus and/or ruga of various lengths extending laterad from metathoracic

ostiole (Figs. 2, 3) 3

Metathoracic ostiole unattended by sulcus or ruga 46

3(2). Superior surface of femora produced distally as stout spine (Fig. 4) 49

Superior surface of femora unarmed distally, at most obtusely produced 4

4(3). Ostiolar ruga extending less than % distance from mesial margin of ostiole to

lateral margin of metapleuron, usually auriculate in form (Fig. 3) 5

- Ostiolar ruga extending % or more of distance from mesial margin of ostiole

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Figs. 1-12. 1. Neottiglossa undata. Propleural production and reflexion. 2. Thyanta palli-

dovirens. Ostiolar sulcus. 3. Euschistus servus. Auriculate ostiolar ruga. 4. Loxaflavicollis. Apical
spine on superior surface of femur. 5. Sibaria englemani. Subapical spines on inferior surface
of femur. 6. Moromorpha tetra. Spine on posterior angle of pronotum. 7. Proxys punctulatus.
Eyes in relation to anterior pronotal margin. 8. Hymenarcys nervosa. Lobed type of bucculae.
9. Hymenarcys aequalis. Truncate type of bucculae. 10. Agroecus griseus. Buccula and first
rostral segment, lateral view. 1 1 . Tibraca limbativentris. Buccula and first rostral segment, lateral
view. 12. Galedanta myops. Evanescent type of bucculae. Abbreviations: buccula (b); femur (fe);
ostiolar sulcus (o.s.); propleural reflexion (pr.); pronotal spine (s); spine (sp.); tibia (ti.).

1984

CONSPECTUS OF PENTATOMINI: 3

71

5(4).

6(5).

7(6).

8(7).

9(8).

10(9).

11(10).

12(9).

13(12).

14(13).

15(14).

16(12).

17(8).

18(17).

19(18).

to lateral margin of metapleuron, usually bordering an elongate sulcus (Fig. 2)

34

At least anterior femora armed on inferior surface with one or more stout

subapical spines (Fig. 5) Sibaria

Femora unarmed or bearing low setose tubercles 6

Posterolateral angles of pronotum armed with small flat spine extending over

base of each corium (Fig. 6) Moromorpha

Posterolateral angles of pronotum unarmed 7

Eyes separated from pronotum by about lh their diameter, their base striated

between reticulation of eye and pronotum (Fig. 7) Proxys

Eyes usually contiguous with pronotum, their base smooth 8

Width of scutellum at distal end of frena more than Vi of basal width, usually

V5 or more 9

Width of scutellum at distal end of frena V2 or less of basal width 17

Bucculae lobed or truncate posteriorly (Figs. 8, 9) 12

Bucculae evanescent or arcuate posteriorly (Figs. 10, 11) 10

First rostral segment projecting beyond bucculae (Fig. 10); 2 rows of small

tubercles present on inferior surface of femora Agroecus

First rostral segment lying entirely between bucculae (Fig. 1 1); inferior surface

of femora lacking tubercles 11

Distance between ocelli less than % width of head across eyes ... (in part)

Tibraca

Distance between ocelli more than % width of head across eyes ... (in part)

Hymenarcys

Superior surface of tibiae sulcate 13

Superior surface of tibiae rounded 16

Anterior width of pronotum more than V2 width at humeri; coria scarcely or

not surpassing apex of scutellum Coenus

Anterior width of pronotum less than V2 width at humeri; coria usually ex-
tending well past apex of scutellum 14

Mesial margin of metathoracic ostiole Y-shaped; head and usually pronotum

with vesture of numerous long setae ... (in part) Trichopepla

Mesial margin of metathoracic ostiole rounded; head and pronotum glabrous

or with short, scattered setae 15

Metathoracic ostiole at right angle to plane of metapleuron, invisible from view
perpendicular to metapleuron; ventral border of anterolateral pronotal margins

punctate ... (in part) Hymenarcys

Metathoracic ostiole inclined to plane of metapleuron but entirely visible from
view perpendicular to metapleuron; anterolateral pronotal margins with ob-
scurely punctate ventral border of uneven width Codophila

Body hirsute; first rostral segment lying entirely between bucculae . . Prionosoma
Body glabrous; apex of first rostral segment projecting beyond bucculae ...

Cosmopepla

Bucculae lobed or abruptly truncated posteriorly (Figs. 8, 9) 18

Bucculae evanescent or arcuate posteriorly (Fig. 12) 20

Juga projecting beyond tylus by at least width of tylus at apex, acute apically

(Fig. 14) ... (in part) Dichelops

Juga projecting little if at all beyond tylus, if angulate the angle a right or obtuse

angle 19

Post-frenal portion of scutellum % or more of scutellar length; green or stra-
mineous dorsally; length 7.5 mm or less excluding hemelytral membranes . . .

(in part) Tepa, new genus

72

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Figs. 13-22. 13. Berecynthus delirator. Head. 14. Dichelops sp. Head. 15. Menecles insertus.

Head and anterior pronotal margin. 16. Runibia sp. Connexivum and hemelytron. 17. Eu-
schistus biformis. Macules on thoracic pleura. 18. Tepa rugulosa. Paramere. 19. Cyptocephala
antiguensis. Paramere. 20. Thyanta perditor. Paramere. 2 1 . Cyptocephala bimini. Genital cup.
22. Cyptocephala bimini. Paramere. Abbreviations: connexivum (c); paramere (cl); dorsal margin
(d.m.); pleural macule (p.m.).

- Post-frenal portion of scutellum less than % of scutellar length; black or brown

dorsally; length without hemelytral membranes 8.5 mm or more Padaeus

20(17). Superior surface of tibiae sulcate for most of their length 23

- Superior surface of tibiae rounded for most of their length 21

21(20). Large species, about 18 mm long 23

- Moderate sized species, about 1 2 mm or less in length 22

22(21). Apex of first rostral segment projecting past bucculae Mormidea

1984

CONSPECTUS OF PENTATOMINI: 3

73

23(20,21).

24(23).

25(24).

26(24).

27(26).

28(26).

29(28).

30(29).

31(30).

32(31).

33(32).

34(4).

35(34).

36(35).

First rostral segment lying entirely between bucculae Oebalus

Both juga and tylus angulate apically, a small incision separating each jugum
from tylus at apex of head; tylus usually spinose and projecting well past juga

(Fig. 13) Berecynthus

Either juga or tylus or both juga and tylus obtuse apically 24

Juga separated and acute apically, projecting beyond obtuse tylus; incision in

apex of head expanding toward apex 25

Juga no longer than tylus, or longer than tylus but either contiguous apically

or leaving a quadrate to rectangular incision in apex of head 26

Connexiva marked with black spot or line at transverse sutures ... (in part)

Chlorocoris

Connexiva immaculate ... (in part) Dichelops

Juga longer than tylus, contiguous apically or leaving quadrate to rectangular

incision in apex of head 27

Juga little or no longer than tylus 28

Black tubercle present in each basal angle of scutellum; humeral angles obtusely

rounded Galedanta

Basal angles of scutellum lacking tubercle; humeral angles acute to spinose . . .

(in part) Chlorocoris

Head deeply inserted into pronotum, imaginary line drawn across anterior

limits of pronotum bisecting eyes near middle (Fig. 15) Menecles

Imaginary line drawn across anterior limits of pronotum passing near posterior

margin of eyes 29

Lateral jugal margins strongly reflexed; antennae 4-segmented ... (in part)

Boea

Lateral jugal margins not reflexed; antennae 5-segmented 30

Costal angle of coria produced; lateral connexival margins strongly serrate (Fig.

16); bicolored species, orange with black markings ... (in part) Runibia

Costal angle of coria acute but not produced, lateral connexival margins entire

or weakly serrate; not colored as above 31

Metastemum moderately produced, sloping downward from anterior margin

to hind coxae; rostrum not attaining metacoxae Hypatropis

Metastemum nearly flat or shallowly sulcate, not produced; rostrum reaching

between or beyond metacoxae 32

Thoracic pleura with small black spot at base of each subcoxa (usually with
additional spot at anterolateral angle of propleuron and at distal end of supra-
coxal cleft on mesopleuron) and/or anterolateral margins of pronotum dentic-
ulate (Fig. 17) Euschistus

Thoracic pleura immaculate or otherwise marked; anterolateral pronotal mar-
gins entire 33

First and second antennal segments subequal in length; fovea in basal angles

of scutellum deep, black ... (in part) Tibraca

First antennal segment shorter than second segment; fovea in basal angles of

scutellum shallow, not black ... (in part) Chlorochroa

Post-frenal portion of scutellum comprising Vi or more of scutellar length ... 35

Post-frenal portion of scutellum less than V2 of scutellar length 37

Juga neither contiguous nor markedly convergent apically ... (in part)

' Trichopepla

Juga contiguous or convergent before tylus 36

Anterolateral margins of pronotum strongly concave Dendrocoris

Anterolateral margins of pronotum weakly concave at most ... (in part) . . .

Holcostethus

74

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

37(34).

38(37).

39(38).

40(39).

41(38).

42(41).

43(42).

44(43).

45(44).

46(2).

47(46).

48(1).

49(3).

Bucculae lobed or right-angularly truncate posteriorly; length of body 7.5 mm
or less; parameres bilobed, without serrations between lobes (Fig. 1 8) ... (in

part) Tepa, new genus

Bucculae evanescent or arcuate posteriorly, or if lobed length of body more

than 10 mm; parameres not as above 38

Ostiolar ruga reaching Vi or less of distance from inner margin of ostiole to

lateral margin of metapleuron 39

Ostiolar ruga reaching more than Vi distance from inner margin of ostiole to

lateral margin of metapleuron 41

Lateral jugal margins strongly reflexed; antennae 4-segmented; rostrum ending

at metacoxae ... (in part) Boea

Lateral jugal margins not reflexed; antennae 5 -segmented; rostrum reaching

onto abdomen 40

Costal angle of coria produced; lateral connexival margins strongly serrate (Fig.

16) ... (in part) Runibia

Costal angle of coria not produced; lateral connexival margins entire or weakly

serrate ... (in part) Chlorochroa

Anterolateral margins of pronotum reflexed in part and/or rimmed Arocera

Anterolateral margins of pronotum neither reflexed nor rimmed 42

Width of scutellum at distal end of frena Vi or more basal width; hemelytral

membranes fumose ... (in part) Holcostethus

Width of scutellum at distal end of frena % or less width at base; hemelytral

membranes vitreous 43

Juga contiguous or convergent before tylus, or projecting beyond tylus by
distance equal to or greater than apical width of tylus ... (in part) . . Chloropepla ,

Juga little or no longer than tylus 44

Anterolateral margins of pronotum vertically rounded; costal angle of coria

reaching last connexival segment Caribo, new genus

Anterolateral margins of pronotum carinate at least near humeri; costal angle

of coria reaching penultimate connexival segment 44

Parameres bilobed with fine denticulations between two of lobes (Fig. 1 9);
length including membrane usually less than 8 mm, rarely as much as 9 mm

Cyptocephala

Parameres simple, acute apically (Fig. 20); more than 8.5 mm in length

Thy ant a

Distal end of first rostral segment surpassing bucculae by distance subequal to
rostral width at posterior limit of bucculae; brightly colored species . . Murgantia
First rostral segment not or scarcely projecting past bucculae; dull colored

yellowish brown species 46

First segment of labium scarcely visible below bucculae from lateral view;
width of scutellum at distal end of frena about % width at base; last stemite

of male much shorter than preceding stemites combined Capivaccius

Distal portion of first labial segment largely visible below bucculae from lateral
view; width of scutellum at distal end of frena about % width at base; last
stemite of male more than twice length at meson of preceding stemites com-
bined Pylophora

Antennifer hidden from lateral view by anterior production of propleuron

Aelia

Antennifer not covered by anterior production of propleuron Neottiglossa

Last rostral segment lying entirely caudad of metacoxae and abdominal seg-
ments 3-5 shallowly sulcate mesially Fecelia

1984

CONSPECTUS OF PENTATOMINI: 3

75

Last rostral segment lying partially between metacoxae, or if lying entirely
caudad of metacoxae abdominal sulcus not extending onto sternite 5 49

50(49). Ostiolar ruga extending more than % of distance from mesial margin of ostiole

to lateral margin of metapleuron ... (in part) Chloropepla

Ostiolar ruga extending xh or less of distance from mesial margin of ostiole to
lateral margin of metapleuron 51

51(50). Ostiolar ruga extending less than % of distance from mesial margin of ostiole

to lateral margin of metapleuron Loxa

Ostiolar ruga extending lA to lh of distance from mesial margin of ostiole to
lateral margin of metapleuron Mayrinia

ANNOTATED LIST OF GENERA

1. Aelia Fabricius, 1803. Holarctic. One species in Western Hemisphere, A. amer-
icana Dallas, 1851.

2. Agroecus Dallas, 1851. Generic revision by Buckup (1957). One species in region
covered, A. griseus Dallas, 1851.

3. Arocera Spinola, 1837. Revision by McDonald (1984).

4. Berecynthus Stal, 1862. One species in region covered, B. delirator (Fabricius,
1787).

5. Boea Walker, 1867. Redescribed by Kormilev( 1950) as Willinerinia. One species
in region covered, B. costaricensis Distant, 1890.

6. Capivaccius Distant, 1893. Monotypic, based on C. bufo Distant, 1893.

7. Caribo Rolston and McDonald, new genus. See following pages.

8. Chlorochroa Stal, 1872 (as subgenus of Lioderma Uhler). Holarctic. Generic
synonymy, subgenus Rhytidilomia, and Opuntiae species group of nominate
subgenus treated by Thomas (1982). Species of Sayi species group treated by
Buxton et al. (1984).

9. Chlorocoris Spinola, 1837. Needs revision.

10. Chloropepla StM, 1867. One species in Panama. Grazia (1980a) gives a key to
species.

1 1 . Codophila Mulsant, 1 866 (as subgenus of Carpocoris Kolenati). Holarctic. Thom-
as (1974) gives diagnosis for genus, treats the 2 species in Western Hemisphere.

12. Coenus Dallas, 1851. Two species, C. delius (Say, 1832) and C. inermis Harris
and Johnson, 1936.

13. Cosmopepla Stal, 1867. Generic review by McDonald (1984).

14. Cyptocephala Berg, 1883. See following pages.

15. Dendrocoris Bergroth, 1891 (as new name for Liotropis Uhler, 1877). Generic
revision by Nelson (1955). This genus actually belongs in Section 2 (Rolston and
McDonald, 1981). However, males of D. humeralis (Uhler, 1877) and D. var-
iegatus Nelson, 1955, have no spine at base of abdominal venter.

16. Dichelops Spinola, 1837. Generic revision by Grazia (1978). One species in
region covered, D. bicolor Distant, 1890.

17. Euschistus Dallas, 1851. Revision of Middle American species by Rolston (1974).
Species in northeastern North America treated by McPherson (1982).

18. Fecelia Stal, 1872. Generic revision by Grazia (1976), corrections and one species
added by Grazia (1980b) another by Eger (1980). The 4 species of Fecelia occur
in the Greater Antilles.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

1 9. Galedanta Amyot and Serville, 1843. Generic revision by Grazia (1 967) modified
by Grazia (1981). One species in region covered, G. myops (Fabricius, 1803).

20. Holcostethus Fieber, 1860 ( =Peribalus Mulsant and Rey, 1866). Holarctic. Re-
vision of Western Hemisphere components by McDonald (1975), revised key
by McDonald (1982).

21. Hymenarcys Amyot and Serville, 1843. Generic review by Rolston (1973c).

22. Hypatropis Bergroth, 1 89 1 (as new name for Melpia Stal, 1 867). Needs revision.
One unidentified species in region covered.

23. Loxa Amyot and Serville, 1843. Revision by Eger (1978).

24. Mayrinia Horvath, 1925. Revised by Grazia-Vieira (1972). One species occurs
in the region covered, M. variegata (Distant, 1880).

25. Menecles Stfil, 1867. Revision by Rolston (1973b).

26. Mormidea Amyot and Serville, 1843. Revision by Rolston (1978).

27. Moromorpha Rolston, 1978. Monotypic, based on Moromorpha tetra (Walker,
1868).

28. Murgantia Stal, 1862. Needs review.

29. Neottiglossa Kirby, 1837. Holarctic. McPherson (1982) keys 4 of the 7 nominal
American species. Needs revision.

30. Oebalus Stal, 1862. Revision by Sailer (1944), synonymy and checklist by Sailer
(1957).

31. Padaeus Stal, 1862. Needs revision, of 6 nominal species apparently only 4
belong in this genus. These 4 are in region covered.

32. Prionosoma Uhler, 1863. Monotypic, based on P. podopioides Uhler, 1863.
McPherson (1982) summarizes literature concerning this species. See following (j
pages.

33. Proxys Spinola, 1837. Needs review.

34. Pylophora Van Duzee, 1923. Van Duzee (1923) described the 2 species. From
Gulf of California region.

35. Runibia Stal, 1861. Needs review.

36. Sibaria Stal, 1872. Reviewed by Rolston (1976).

37. Tepa Rolston and McDonald, new genus. See following pages.

38. Thyanta Stal, 1862. Needs revision. Thyanta antiguensis (Westwood), T. bimini
Ruckes, T. elegans Malloch, and T. elegantula Jensen-Haarup are transferred
to Cyptocephala Berg, and Thyanta rugulosa (Say), T. brevis Van Duzee, T.
jugosa Van Duzee, T. panda Van Duzee, T. punctiventris Van Duzee and T.
yerma Rolston are transferred to Tepa Rolston and McDonald, new genus. See
following pages.

39. Tibraca Stal, 1860. Needs revision.

40. Trichopepla Stal, 1867. Revised by McDonald (1976).

NEW GENERA, SPECIES, AND COMBINATIONS

Cyptocephala Berg, 1883

Cyptocephala Berg, 1883:209-210 (reprinted, 1884:25-26).

Crato Distant, 1893:457.

Thyanta subgenus Parathyanta Jensen-Haarup, 1928:186. New Synonymy.

CONSPECTUS OF PENTATOMINI: 3

77

1984

Type species. Cyptocephala cogitabunda Berg, 1883.

Diagnosis. Metathoracic ostiole a fissure in metapleuron, not on notable elevation;
ostiolar ruga sulcate proximally, reaching 7/i0-8/io distance from mesial margin of
ostiole to lateral margin of metapleuron. Post-frenal portion of scutellum comprising
about % of scutellar length; scutellar width at distal end of frena about % of basal
width. Bucculae arcuately truncate at posterior termination; basal segment of rostrum
not projecting beyond bucculae. Anterolateral margins of pronotum carinate, entire,
neither reflexed nor rimmed.

Parameres bilobed or trilobed apically, minutely denticulate between anterior and
lateral lobes (Figs. 21, 22).

! Species. Species here transferred from Thyanta to Cyptocephala are: Pentatoma
lantiguensis Westwood, 1837, Thyanta bimini Ruckes, 1952, Thyanta elegans Mal-
loch, 1919, and Thyanta ( Parathyanta ) elegantula Jensen-Haarup, 1928.

Comments. Cyptocephala differs from Thyanta and Tepa primarily in the lobed,
denticulate parameres (Figs. 19, 22). In Thyanta the parameres are apically spinose
or acute (Fig. 20); in Tepa the parameres have a thumb-like lateral lobe (Fig. 18);
denticles are absent from the parameres of Thyanta and Tepa. The aedeagus is quite
similar in these three genera, simple, and apparently of little taxonomic use.
i The spermathecae of Cyptocephala species are not entirely uniform. In C. anti-
guensis, C. bimini and C. elegans there is an enlargement at the base of the proximal
Iflange, but this structure is absent in C. cogitabunda and C. elegantula (Figs. 36-41).
This enlargement is present and variously modified in the Thyanta species examined
j!(Figs. 23-35); it is absent in Tepa species (Figs. 42-48). The membranous sac of the
Ispermathecal duct does not dilate the full length of the enclosed sclerotized rod in
Cyptocephala (Figs. 36-39), as it does in Tepa (Figs. 42, 43) and as it usually, but
not always, does in Thyanta (Figs. 23-26).

: New records for C. bimini are Cuba and Jamaica.

New records for C. elegantula are Catamarca, Cordoba and Tucuman provinces
in Argentina.

i The holotypes of Thyanta bimini and T. elegantula, and a male and female homo-
jtype of Cyptocephala cogitabunda were examined. Type material of Thyanta anti-
\guensis and T. elegans was not seen.

I

Tepa Rolston and McDonald, new genus

* Type species. Pentatoma rugulosa Say, 1832.

Diagnosis. Metathoracic ostiole a fissure in metapleuron, not on notable elevation;
i ostiolar ruga sulcate proximally, reaching %-3/5 distance from mesial margin of ostiole
to lateral margin of metapleuron. Post-frenal portion of scutellum % -V2 of scutellar
j length; width of scutellum at distal end of frena % -V2 of basal width. Bucculae lobed
or right-angularly truncate posteriorly. Parameres bilobed; dorsal lobe narrowly
rounded; lateral lobe smaller, thumb-like, variously bent dorsad; denticles absent
j (Fig. 1 8). Spermathecal bulb hemispherical, base of proximal flange unmodified (Figs.
42-48). Length less than 7.5 mm excluding hemelytral membranes.

Species. Species transferred from Thyanta to Tepa are: Pentatoma rugulosa Say,
1832, Thyanta brevis Van Duzee, 1904, Thyanta jugosa Van Duzee, 1923, Thyanta

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Figs. 23-32. 23-26. Spermatheca. 23. Thyanta calceata. 24. T. custator. 25. T. patruellis.
26. T. pseudocasta. 27-32. Spermathecal bulb and pump. 27. Thyanta calceata. 28. T. custator.
29. T. maculata. 30. T. nitidula. 31. T. pallidovirens. 32. T. patruellis.

panda Van Duzee, 1923, Thyanta punctiventris Van Duzee, 1904, and Thyanta
yerma Rolston, 1972.

Comments. The species of Tepa are included in a key to the small North American
species then considered to be in Thyanta, and the male genitalia are figured (Rolston,
1972).

Tepa differs from Thyanta and Cyptocephala primarily in the form of the para-

0-5

1984

CONSPECTUS OF PENTATOMINI: 3

79

Figs. 33-41. 33-35. Spermathecal bulb and pump. 33. Thyanta planifrons. 34. T. pseu-
docasta. 35. T. signoreti. 36-39. Spermatheca. 36. Cyptocephala antiguensis. 37. C. bimini. 38.
C. cogitabunda. 39. C. elegans. 40, 41. Spermathecal bulb and pump. 40. Cyptocephala bimini.
41. C. elegans.

meres. The parameres in Thyanta are acute or spinose apically (Fig. 22); in Cypto-
cephala they are bilobed with minute denticles between two of the lobes (Fig. 24).
Tepa also differs from the Thyanta species known to us with the exception of T.
aeruginosa Berg, and from Cyptocephala in having the posterior termination of the
bucculae lobed or right-angularly truncate. The spermatheca in all Tepa species is
simple without the enlargement that seems to be characteristic of Thyanta and
Cyptocephala species other than C. cogitabunda and C. elegantula.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

0 -5

0-5 0-5

Figs. 42-48. 42, 43. Spermatheca. 42. Tepa rugulosa. 43. T. yerma. 44-48. Spermathecal
bulb and pump. 44. Tepa brevis. 45. T. jugosa. 46. T. panda. 47. T. punctiventris. 48. T. yerma.

Tepa is confined to the western U.S.A. and northern Mexico with the exception
of T. panda which is recorded from California, Curasao and Florida.

Caribo Rolston, new genus
Type species. Caribo subgibbus Rolston, new species.

Diagnosis. Metathoracic ostiole inclined to plane of metapleuron; ostiolar ruga
sulcate proximally, extending from mesial margin of ostiole slightly more than V2-%
of distance to lateral margin of metapleuron. Juga no longer than tylus, their lateral
margins not reflexed. First antennal segment not reaching apex of head. Bucculae
evanescent posteriorly; basal segment of rostrum projecting beyond bucculae by less

1984

CONSPECTUS OF PENTATOMINI: 3

81

Figs. 49-54. 49-53. Caribo subgibbus. 49. Pygophore, ventral view. 50. Pygophore, lateral
view. 51. Aedeagus, lateral view. 52. Aedeagus, ventral view. 53. Genital plates, caudo ventral
view. 54. Caribo fasciatus. Pygophore, ventral view. Abbreviations: cavity (ca.); dorsal margin
(d.m.); basal plates (1 gx); lateral conjunctival appendage (l.c.ap.); median conjunctival appen-
dage (m.c.ap.); median penial lobes (m.p.l.); orifice (po.); ventral conjunctival appendage (v.c.ap.).

than thickness of segment at apex; second rostral segment reaching mesocoxae. An-
terolateral margins of pronotum rounded vertically, lacking distinct carinae except
at humeri, entire. Post-frenal part of scutellum about % of scutellar length; scutellar
width at distal end of frena nearly Vi of basal width; length and width of scutellum
subequal. Costal angle of each corium extending to last connexival segment. Pro-
sternum longitudinally sulcate; mesostemum feebly carinate mesially; metastemum
weakly concave. Femora unarmed; superior surface of tibiae flattened, obscurely
sulcate. Length less than 7.5 mm without hemelytral membranes.

Small orifice in lateral surface of pygophore on each side exposed only when
pygophore partially extracted (Fig. 50), opening into large internal cavity (Fig. 49).

Comments. The pair of cavities opening on the lateral pygophoral surfaces have
not been reported previously in pentatomids.

Caribo subgibbus Rolston, new species
Figs. 49-53

Description. Pale stramineous above and below; punctation irregular, fuscous to
black. Markings on head consisting of irregular fuscous to black longitudinal bands
on lateral margins of tylus basally, lateral margins of slightly elevated vertex from

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92( 1 )

base of head to eye (enclosing ocellus) on each side. Cicatrices outlined in black,
thickly so except anteriorly. Patches on subgibbous base of scutellum densely punc-
tate, black, irregular in form. Punctures on ventral disk of abdomen black, sparse,
variable in size, a few quite large; punctation along lateral borders small, rather
uniform, many concolorous with background. Spots on legs variable in size, coalescing
into irregular incomplete preapical band on femora; distal end of tibiae and tarsi
infuscate.

Jugal margins briefly parallel between anteocular concavity and rounded apex of
head; juga not convergent; tylus broadened at apex. Width of head across eyes 1.5
mm, length 1.3-1. 5 mm. Antennal segments 0. 3-0.4, 0.6-0. 7, 0.5-0. 6, 0.7, 0.7-0. 8
mm long; distal 3A of segment 5, distal Vi of segment 4, sometimes distal xh of segment
3 dark. Rostrum reaching onto stemite 4 (3rd visible); segments 2-4 about 1.0, 0.7-
0.8, 0.6-0. 7 mm long.

Anterior submargin of pronotum depressed; cicatrices elevated; additional irreg-
ularities in anterior pronotal disk. Thin impunctate line divides pronotum at meson.
Humeral angles somewhat produced laterad, rounded. Pronotal width across humeri
3. 6-3. 8 mm; mesial length 1.4-1. 5 mm.

Basal % of scutellum subgibbous; fovea in basal angles triangular, shallow, black.
Membranes of hemelytra projecting well beyond body, vitreous, each of 5 veins in
fumose band.

Abdominal stemites 3-4 and 5 basally with shallow longitudinal sulcus mesially.
Spiracles nearly concolorous with surrounding area of sternite. Posterolateral and
anterolateral angles of sternites with small black macule. Length of body 5. 2-5. 5 mm
excluding hemelytral membranes.

Posterior margin of each basal plate convexly arcuate (Fig. 53). Gonocoxae 2
diamond shaped, broader than long, carinate mesially. Surface of paratergites 9
concave, especially so adjacent to gonocoxae 2; their posterior margin broadly convex.
Stemite 10 small, triangular. Stemite 7 of female bearing large fuscous macule me-
sially at base.

Pygophore with U-shaped mesial emargination in posterior margin, black mesial
macule at anterior margin; posterior border bearing fringe of long setae (Fig. 49).
Ventral conjunctival appendages sclerotized apically, forming plate (Figs. 51, 52);
lateral appendages sclerotized, medial appendages membranous.

Holotype. 6, labeled “Bahamas:Mayaguana Isl. (3 words inked out) 28-VIII-63. C.
Murvosh. Blacklight trap.” Deposited in the Florida State Collection of Arthropods.

Paratypes. 2, labeled as holotype, deposited with holotype; <3 labeled as holotype,
deposited in senior author’s collection.

Caribo fasciatus Rolston, new species
Fig. 54

Description. Light stramineous above and below, appearing brown dorsally from
dense fuscous to black punctation. Pronotum traversed dorsally between cicatrices
and humeri by stramineous fascia with sparse fuscous punctation. Cicatrices mostly
black. Base of head, lateral margins of vertex and base of tylus densely black punctate.
Ivory callus present on mesopleura about xh distance from lateral margin; another
irregularly shaped and sparsely dark punctate callus in posterolateral corner of meta-

1984

CONSPECTUS OF PENTATOMINI: 3

83

pleura. Punctures on ventral disk of abdomen of various sizes, some quite large;
punctures along lateral borders small, more nearly uniform in size. Each abdominal
stemite bearing black dot at anterolateral and posterolateral corners. Legs sparsely
dotted with black; distal end of tibiae and tarsi infuscate.

Jugal margins subparallel between anteocular concavity and rounded apex of head.
Width of head across eyes 1 .8-2.0 mm, length 1 .5-1 .7 mm. Antennal segments about
0.4, 0.7, 0.7-0. 8, 1.0, 1.1 mm long; distal 3A of last antennal segment dark, remainder
of antennae pale. Rostrum reaching sternite 4 (3rd visible); length of segments 2-4
about 1.2, 0.7-0. 9, 0.7-0. 9 mm.

Anterolateral margins of pronotum concave. Humeri moderately produced laterad,
narrowly rounded. Cicatrices somewhat elevated; pronotal disk otherwise smooth
excepting shallow depression anterior to cicatrices. Width across humeri 4. 7-5. 4 mm;
mesial length 1. 7-2.0 mm.

Scutellum 2. 8-3. 3 mm wide at base, 2. 7-3. 2 mm long. Fovea in basal angles
shallow, small, triangular. Four ill-defined clusters of punctures evenly spaced along
base, similar in size to foveae. Scutellar disk evenly convex. Hemelytral membranes
vitreous with fumose mottling along 7-8 veins, projecting well past abdominal apex.

Abdominal sternite 3 and base of 4 shallowly sulcate mesially. Spiracles black.

Length of body 6. 5-7.4 mm excluding membranes.

Genital plates similar to those of C. subgibbus. Sternite 7 bearing large black macule
mesially at base.

Posterior margin of pygophore truncate with small U-shaped mesial notch; black
mesial macule present at base (Fig. 54).

Holotype. 6, labeled “Antilles:Jamaica. Falmouth. July 19, 1960. C. & P. Vaurie.”
Deposited in the American Museum of Natural History.

Paratype. 9 labeled (a) “Virgin Is:St. John, Est. Carolina, NW of Coral Bay, 31
May 1982, 250 ft” (b) “at ultraviolet light, W. B. Muchmore.” Deposited in senior
author’s collection.

ADDITIONAL NEW SYNONYMY

Prionosoma Uhler, 1863

Prionosoma Uhler, 1863:363-364.

Neurohalys Bliven, 1960:34. New Synonymy.

Prionosoma podopioides Uhler, 1863
Prionosoma podopioides Uhler, 1863:364-365.

Neurohalys bucculatus Bliven, 1960:34-36, PI. 5, figs. 1,1a. New Synonymy.

The type of N. bucculatus, type species of Neurohalys, in the California Academy
of Sciences, was examined. It is an unremarkable example of Prionosoma podopioides.

Thyanta aeruginosa Berg, 1878
Thyanta aeruginosa Berg, 1878:24.

Phacidium euchlorum Breddin, 1912:92-93. New Synonymy.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

The type series of Phacidium euchlorum, in the Universite Louis Pasteur, Stras-
bourg, France, was compared with specimens determined by H. Ruckes, one of them
designated a homotype, as Thyanta aeruginosa Berg. Without doubt Phacidium
euchlorum and Thyanta aeruginosa are synonyms of this quite distinctive species
from Argentina and Uruguay. The purported provenance of the type series of P.
euchlorum, Costa Rica, is obviously erroneous.

The type series of P. euchlorum consists of 2 males and 1 female, all labeled “Costa-
Rica, San-Jose” (in error) above a red label bearing “Type.” A male, here designated
lectotype, bears an additional label, presumably inscribed by Breddin with “Phaci-
dium euchlorum Bredd. n. gen. n. sp., typus” (the “t” in typus is not crossed). Pronotal
width of this specimen is 3.45 mm. The smaller male, pronotal width 3.25 mm, and
female are paralectotypes.

This species appears to be somewhat aberrant among other Thyanta species known
to us, and once Thyanta is revised Phacidium may stand as a valid generic or
subgeneric name. The species synonymy is mentioned now to resolve the previously
vexing question as to the identity of P. euchlorum.

ACKNOWLEDGMENTS

We are indebted to Drs. N. Moller Andersen (Universitetets Zoologiske Museum), P. H.
Amaud (California Academy of Sciences), H. Dodge Engleman, Richard C. Froeschner (U.S.
National Museum), Michael A. Ivie (Ohio State University), J. Matter (Universite Louis Pas-
teur), Frank W. Mead (Florida State Collection of Arthropods), Luis de Santis (Universidad
Nacional de LaPlata), Randall T. Schuh (American Museum of Natural History), and Donald
B. Thomas for the loan of specimens relevant to this work. We are especially grateful to Drs.
Joseph E. Eger, H. Dodge Engleman, J. E. McPherson and Donald B. Thomas for critically
reviewing the manuscript. Any errors or obfuscations that may have crept into the revision are
entirely the authors. Dr. Jocelia Grazia compared the female specimen of Cyptocephala cogi-
tabunda Berg with the holotype.

LITERATURE CITED

Berg, C. 1878. Hemiptera Argentina: ensayo de una monografia de los Hemipteros Heter-
opteros y Homopteros de la Republica Argentina. Anal. Soc. Cient. Arg. 6( 1 ):23— 36.
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193-217.

Berg, C. 1884. Addenda et emendanda ad Hemiptera Argentina. Bonariae.

Berg, C. 1891. Nova Hemiptera faunarum Argentina et Uruguayensis. Anal. Soc. Cient. Arg.
32:277-287.

Bliven, B. P. 1960. Studies on insects of the Redwood Empire III: new Hemiptera with notes
on others. The Occidental Entomologist 1:35-42.

Breddin, G. 1912. Zwei neue Pentatomiden-Gattungen (Hem.). Archiv Naturgeschichte 78
Abt. A 6:90-93.

Buckup, L. 1957. Pentatomideos neotropicais I — sobre o genero Agroecus Dallas, 1851, com
a descricao de duas especies novas (Hem.-Pentatomidae). Iheringia (Zool.) 6:1-20.
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Distant, W. L. 1880-1893. Insecta. Rhynchota, Hemiptera-Heteroptera. In: F. D. Godman
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Eger, J. E. 1978. Revision of the genus Loxa (Hemiptera:Pentatomidae). J. New York Ent.
Soc. 86(3):224-259.

Eger, J. E. 1980. Fecelia biorbis n. sp. (Heteroptera, Pentatomidae), a new species from Haiti.
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Heteroptera, Pentatomidae). Iheringia (Zool.) 35:45-49.

Grazia- Vieira, J. 1972. O genero Mayrinia Horvath, 1925 (Heteroptera, Pentatomidae, Pen-
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Grazia, J. 1976. Revisao do genero Fecelia Stcil, 1872 (Heteroptera, Pentatomidae, Penta-
tomini). Rev. Brasil. Biol. 36(l):229-237.

Grazia, J. 1978. Revisao do genero Dichelops Spinola, 1837 (Heteroptera, Pentatomidae,
Pentatomini). Iheringia (Zool.) 53:3-1 19.

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(Heteroptera, Pentatomidae). An. Soc. Ent. Brasil 9(1): 123-131.

Grazia, J. 1980b. Uma nova especie do genero Fecelia St&l (Heteroptera, Pentatomidae,
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Grazia, J. 1981. Novas consideracoes sobre Galedanta Amyot and Serville, 1843 com a
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9-19.

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N. America. J. New York Ent. Soc. 82(l):9-22.

McDonald, F. J. D. 1982. Description of the male genitalia of Holcostethus hirtus (Van Duzee)
with a revised key to North American species (Hemiptera: Pentatomidae). J. New York
Ent. Soc. 90(1): 5-7.

McPherson, J. E. 1982. The Pentatomoidea (Hemiptera) of Northeastern North America with
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Nelson, G. H. 1955. A revision of the genus Dendrocoris and its generic relationships. Proc.
Ent. Soc. Wash. 57(2):49-67.

Rolston, L. H. 1973a. The small Thyanta species of North America (Hemiptera:Pentatom-
idae). J. Georgia Ent. Soc. 7(4):278-285.

Rolston, L. H. 1973b. The genus Menecles St&l (Hemiptera:Pentatomidae). J. New York Ent.
Soc. 80(4):234-237 (1972).

Rolston, L. H. 1973c. A review of Hymenarcys (Hemiptera:Pentatomidae). J. New York Ent.
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Rolston, L. H. 1976. A new species and review of Sibaria (Hemiptera:Pentatomidae). J. New
York Ent. Soc. 83(4):2 18-225 (1975).

Rolston, L. H. 1978. A revision of the genus Mormidea (Hemiptera:Pentatomidae). J. New
York Ent. Soc. 86(3): 1 6 1-2 1 9.

Rolston, L. H. and F. J. D. McDonald. 1981. Conspectus of Pentatomini genera of the Western
Hemisphere— part 2 (Hemiptera:Pentatomidae). J. New York Ent. Soc. 88(4):257-272
(1980).

Rolston, L. H., F. J. D. McDonald and D. B. Thomas, Jr. 1980. A conspectus of Pentatomini
genera of the Western Hemisphere. Part 1. (Hemiptera:Pentatomidae). J. New York Ent.
Soc. 88(2): 120-1 32.

Ruckes, H. 1952. Two new species of Thyanta St&l (Pentatomidae: Heteroptera). Bull. Brook-
lyn Ent. Soc. 47:65-68.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Sailer, R. I. 1944. The genus Solubea (Heteroptera: Pentatomidae). Proc. Ent. Soc. Wash.
46(5): 105-127.

Sailer, R. I. 1957. Solubea Bergroth, 1891, a synonym of Oebalus StAl, 1862, and a note
concerning the distribution of O. ornatus (Sailer) (Hemiptera, Pentatomidae). Proc. Ent.
Soc. Wash. 59( 1 ):4 1—42.

Thomas, D. B., Jr. 1974. The genus Codophila Mulsant in North America (Hemiptera:
Pentatomidae). Pan-Pacific Ent. 50(4):44 1-442.

Thomas, D. B., Jr. 1983. The taxonomic status of the genera Chlorochroa St&l, Rhytidolomia
St&l, Liodermion Kirkaldy, and Pitedia Reuter, and their included species (Hemiptera:
Pentatomidae). Ann. Ent. Soc. Amer. 76(2):2 15-224.

Uhler, P. R. 1863. Entomological contributions. — No. 2. Proc. Ent. Soc. Philadelphia 2:361-
366.

Van Duzee, E. P. 1923. Expedition of the California Academy of Sciences to the Gulf of
California in 1921. The Hemiptera (true bugs etc.). Proc. Calif. Acad. Sci. (4)12(11):
123-200.

Westwood, J. O. 1837. In: F. W. Hope, A Catalogue of Hemiptera in the Collection of the
Rev. F. W. Hope, M. A. with Short Latin Descriptions of the New Species. Part 1.
London, 46 pp.

Received October 31, 1983; accepted January 23, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 87-92

TEMPORAL AND SPATIAL VARIATION IN SEX RATIO
AND MATING FREQUENCY IN SOLDIER BEETLES1

Kim Holder Bailey, David E. McCauley, and W. David Sissom
Department of Biology, Vanderbilt University, Nashville, Tennessee 37235

Abstract.— Previous studies of the mating system of the soldier beetle, Chauliognathus penn-
sylvanicus, have not considered spatial or temporal variation in sex ratio or the relative frequency
of mating and single adults. These two attributes of the mating system were sampled at mid-
day and again in the early evening thrice weekly throughout one adult flight season at two
localities in Tennessee. Sex ratios were equivalent within days but became increasingly male
biased as the season progressed. The proportion of all adults found copulating is about 5 times
greater in the early evening than at mid-day. Some between-site differences were found.

The soldier beetle, Chauliognathus pennsylvanicus (Coleoptera: Cantharidae), is
one of the most common old field insects in the eastern United States during late
summer and early autumn. A number of studies have examined mating behavior as
it relates to the operation of sexual selection in natural populations of this beetle
(Mason, 1972; McCauley and Wade, 1978; Mason, 1980; McCauley, 1981; McLain,
1981; Woodhead, 1981; McLain, 1982). Most of these studies have focused on
identifying those characteristics of individuals that determine the probability of being
found in copula when a random sample of single and mating adults is taken from a
field population. Body size, antennal morphology, population density, presence or
absence of interspecific competitors, and female reproductive status have all been
shown to influence the probability that an individual will be found mating.

The published studies of the Chauliognathus mating system are based on short
term observations. Typically, several collections are made at mid-day several days
apart. Any temporal or spatial variation in attributes of the mating system such as
the operational sex ratio or female receptivity that would be likely to affect the process
of mate selection would go undetected when data is taken in such a fashion. It is
unknown whether the mate selection process varies with time of day, between days,
or from locality to locality. A more complete assessment of the Chaulignathus mating
system would require knowledge of any such variation. The present study describes
one full season of mating activity of the beetles and analyzes daily and seasonal
variation in operational sex ratios and the frequency of mating in two natural pop-
ulations of C. pennsylvanicus located in middle Tennessee.

MATERIALS AND METHODS

The two study sites (A and B) in Davidson County, Tennessee were selected such
that their close proximity ensured nearly identical weather conditions on any given
census day. While field A was considerably larger than field B, they were similar in
the relative abundances and phenologies of the numerically dominant plants. Adult

1 Requests for reprints should be addressed to D. E. McCauley.

88

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

SAMPLING DAY

SAMPLING DAY

SAMPLING DAY

SAMPLING DAY

Fig. 1 (A-D). Percent males represented in samples of adult Chauliognathus pennsylvanicus
taken: A) at study site A at mid-day, B) at site A in the evening, C) at site B at mid-day, and
D) at site B in the evening. Least squares regression equations of percent males on (Y) sampling
day (X) are: A) Y = 32.5 + 4.4X; B) Y = 43.4 + 2.8X; C) Y = 55.9 - 2.8X; D) Y = 42.9 +
2.8X.

soldier beetles are pollen feeders and mate at the feeding sites. Goldenrod ( Solidago
spp.) provided the primary food resource at the beginning of the study in September
and was gradually replaced in October by an aster ( Helianthus sp.). Adult sex ratio
and percentage adults seen in copula were censused at each site. In Field A (the larger
field) the first 200 beetles encountered in a random transect of the field were sexed
and categorized as either “mating” or “solitary.” The same procedure was applied
to the first 1 00 beetles observed in Field B. Care was taken at both sites to look for
beetles throughout the vegetation (not just on the flowerheads where they tend to
aggregate) in order to prevent sampling bias in the sex ratios.

The populations of Chauliognathus were sampled during three mid-day surveys
(1 1:00 A.M.-1:00 P.M.) and three late afternoon surveys (5:00-7:00 P.M.) each week

1984

SEX RATIOS IN CANTHARIDAE

89

SAMPLING DAY

Fig. 2. Percent of sampled females found mating at study sites B (top) and A (bottom).
Dark bars represent mid-day samples; open bars evening samples.

beginning September 13, 1982. Field A was sampled until October 18; Field B until
October 4. Surveys were terminated at each site when adult beetles became very rare
or non-existent. When mid-day and late afternoon surveys could not be taken on
the same day, surveys taken on successive days were paired to facilitate temporal
and spatial comparisons.

RESULTS

The operational sex ratios estimated at the 2 study sites are presented in Figure 1
(A-D). At study site A the sex ratios became increasingly male biased as the season
progressed. Linear regression of percent males on time reveals this trend to be sta-
tistically significant for both the mid-day (P < 0.01) and evening samples (P < 0.05).
At study site B no such trend can be documented statistically (P > 0.10 in both
cases). The 4 regression lines were compared with respect to the equality of their
slopes and found to be highly heterogeneous (P < 0.001). There appears to be a real
difference between localities in temporal trends in the sex ratios. Testing of individual
samples for a goodness of fit to a 50:50 sex ratio reveals that not only are later samples
at site A significantly male biased but that the mid-day samples are actually slightly

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Table 1. Three factor analysis of variance of the percent of sampled females found mating.
With one replicate per treatment cell the A x B x C interaction term serves as the error mean
squares.

Source of variation

df

M.S.

F

Location, A

1

0.040

3.64

N.S.

Sample day, B

8

0.011

1.00

N.S.

Time of day, C

1

4.310

398.80

P < 0.001

A x B

8

0.011

1.00

N.S.

A x C

1

0.010

0.91

N.S.

B x C

8

0.050

4.55

N.S.

A x B x C

8

0.011

female biased at the onset of the adult flight season. As a group, the goodness of fit
G tests are highly heterogeneous as would be expected from the results of the regres-
sion analysis.

Given that sex ratios are male biased, at least some of the time, a study of temporal
variation in mating frequency requires that the proportion of males found mating
be analyzed separately from that of females. The proportions of females found to be
mating at the various censuses is presented in Figure 2. It is obvious that far more
females can be found mating in the early evening than at mid-day. The data were
analyzed by a three-factor analysis of variance in which the 3 factors were time of
day, day of season, and locality. Only time of day was found to be statistically
significant (Table 1). Pooling results across census days and fields, it was shown that,
on average, 13.2 percent of the adult females are mating at mid-day compared to
84.3 percent in the evening. A similar analysis of the proportion of males found
mating reveals that in addition to the highly significant effect of time of day, the day
of the season also has an effect (Table 2). This is not surprising given the observations
made on the sex ratio.

DISCUSSION

Despite the large number of studies of mate selection in C. pennsylvanicus, little
is known about those characteristics of its natural history and reproductive physiology
relevant to a more complete description of its mating system. Male biased sex ratios
have been shown previously (Wiener, 1974; Brown and Brown, 1984) but the extent
of the temporal trend in variation in the sex ratio has not been previously documented.
The mechanism by which populations become increasingly male biased is not clear
since these studies were not designed to partition sex-specific mortality effects from
emigration. Females could have a shorter life expectancy than males or they could
disperse from the old fields prior to oviposition. That sex ratios are not male biased
at all localities and are not male biased at the onset of the breeding season suggests
that the male bias does not result from unequal sex ratios at birth nor from differential
survivorship during the larval period. Whatever the mechanism, seasonal changes
in the operational sex ratio would be expected to exert an influence on the mating
system, probably intensifying competition among males for mates.

1984

SEX RATIOS IN CANTHARIDAE

91

Table 2. Three factor analysis of variance of the percent of sampled males found to be
mating. With one replicate per treatment cell the A x B x C interaction term serves as the
error mean squares.

Source of variation

df

M.S.

F

Location, A

1

0.00

0.00

N.S.

Sample day, B

8

0.04

3.64

P < 0.05

Time of day, C

1

2.70

245.45

P < 0.001

A x B

8

0.01

0.91

N.S.

A x C

1

0.01

0.91

N.S.

B x C

8

0.03

2.73

N.S.

A x B x C

8

0.01

It is also not clear what generates the daily cycle in the proportion of the population
found to be mating. Behavioral observations by McCauley and Wade (1978) suggest
that males will attempt to copulate with nearly any single female that they encounter.
Most females resist copulating with males, at least until the male has succeeded in
subduing the female with antennal stroking. Woodhead (1981) has shown that a
female’s receptivity seems to be predicted by the maturity of her eggs, at least at
mid-day. It would seem that female receptivity must further change with the time
of day, perhaps as a function of ambient temperature or incidental light. Depending
on the frequency and timing of oviposition and the manner in which sperm is
transferred and utilized, the success of a male of a given body size at mating at mid-
day could be a poor predictor of his overall reproductive success.

This paper is not intended to discredit those previous short term studies of C.
pennsylvanicus but rather to add to them by pointing out that temporal variation in
some rather obvious populational parameters could greatly influence our interpre-
tation of the mating system and what determines a fit male.

LITERATURE CITED

Brown, L. and V. Brown. 1984. Dispersal and dispersion in a population of soldier beetles,
Chauliognathus pennsylvanicus (Coleoptera: Cantheridae). Environ. Ent. (in press).

Mason, L. G. 1972. Natural insect populations and assortative mating. Amer. Midi. Natur.
88:151-157.

Mason, L. G. 1980. Sexual selection and the evolution of pair-bonding in soldier beetles.
Evolution 34:174-180.

McCauley, D. E. 1981. Application of the Kence-Bryant model of mating behavior to a natural
population of soldier beetles. Amer. Natur. 1 17:400-402.

McCauley, D. E. and M. J. Wade. 1978. Female choice and the mating structure of a natural
population of the soldier beetle, Chauliognathus pennsylvanicus. Evolution 32:771-775.

McLain, D. K. 1981. Interspecific interference competition and mate choice in the soldier
beetle, Chauliognathus pennsylvanicus. Behav. Ecol. and Sociobiol. 9:65-66.

McLain, D. K. 1982. Density dependent sexual selection and positive phenotypic assortative
mating in natural populations of the soldier beelte, Chauliognathus pennsylvanicus. Evo-
lution 36:1227-1235.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

Wiener, J. G. 1974. Unbalanced sex ratios in populations of Chauliognathus pennsylvanicus.
Environ. Ent. 3:572-573.

Woodhead, A. P. 1981. Female dry weight and female choice in Chauliognathus pennsyl-
vanicus. Evolution 35:192-194.

Received May 26, 1983; Accepted October 31, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(1), 1984, pp. 93-96

BOOK REVIEWS

Bark Beetles in North American Conifers.— Jeffrey B. Mitton and Kareen B. Sturgeon

(eds.). University of Texas Press, Austin. 527 pp. $17.50.

It is appropriate for a discipline to step back and examine itself occasionally, as
is done in this work. The volume consists of 1 0 chapters, each written by one or two
specialists in the areas treated, with chapters 1 and 1 0 by the editors. It appears that
camera-ready copy was prepared by the contributors in order to keep printing costs
minimal, but without stringent editorial control as judged by the irregularity in the
lower margin of each page and unused blank areas on several pages (e.g., p. 257). In
spite of these minor lapses, the articles are clear and informative, with surprisingly
little redundancy. Clerical errors vary from virtually none in one chapter to many
in another.

Each chapter reviews principal contributions to existing knowledge and attempts
to summarize the present status of the 1 0 fields treated. Several authors suggest areas
where future research would be appropriate. The chapters for the most part are
authoritative, but are unequal in quality, ranging from the very scholarly review of
Aggregation Pheromones by Borden to two that might cause one to wonder if the
authors had been exposed to literature of the past decade or two. In spite of this
inconsistency, the volume should be required reading for anyone engaged in bark
beetle research everywhere, not just in North America. However, the reader must
be informed and alert, because all that is written is not so.

If the volume is to be criticized, first on the list should be the lack of adequate
editorial and/or peer review prior to publication to remove archaic taxonomic no-
menclature, outright errors in species citations, and other erroneous information.
Second, with about three notable exceptions, the authors restricted their discussions
to American species on conifers when a worldwide perspective was needed to make
the point under consideration. No one called attention to the fact that Mexico, not
the United States, is and has been the center of Dendroctonus evolution, or that two
species, not just one member of this genus, occur in Asia. Third, a conspicuous void
in this volume is the absence of a chapter on mating systems and mating behavior,
recent advances of notable significance are not mentioned. Fourth, territorial be-
havior, including same-sex butting or dislodgement by attacking scolytids on the
bark surface, and courtship ritual by non-stridulating beetles are not mentioned.
Fifth, in the final chapter there are so many oversights, errors, and erroneous sup-
positions that its credibility is seriously undermined. For example, (a) scolytids are
unknown from the Triassic ( Paleoscolytus is an error in family placement and Pa-
leoipidus is unassignable to any family), (b) scolytids are primarily polyphagous (not
monophagous or oligophagous, as stated), (c) since few scolytid insect parasites prey
exclusively on scolytids, evolutionary radiations tied closely to scolytids are very
unlikely (parasitic mite radiations tied to scolytids are much more probable), (d)
speciation models (pp. 352-357) are based primarily on ecotypes (host races often
are examples) that are ephemeral populations that ordinarily do not form geographical
races (subspecies) or species. However, they occasionally can become important in
preadapting a population to a new environment which in turn could then lead to
speciation, (e) the lack of genetic (or biochemical) diversity in Colorado ponderosa

94

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

pine (as compared to great diversity in California) is attributed to interaction with
scolytids. However, pondersa pine did not occur in Colorado until about 12,000
years ago or within several hundred miles of Colorado as recently as 16,000 years
ago. It has occurred in California for a considerably longer period of time. Its recent
migration into Colorado alone could have resulted in the lack of genetic diversity
observed today. Considerably better evidence of bark beetle limitation of genetic
diversity on ponderosa pine must be documented before they can be identified as
significant factors in the suggested role.— Stephen L. Wood, Department of Zoology,
Brigham Young University, Provo, Utah 84602.

Biological Diversification in the Tropics: Proceedings of the Fifth International Sym-
posium for the Association for Tropical Biology.— G. T. Prance (ed.). 1982. Co-
lumbia University Press, New York. 714 pp. $60.00.

This large volume champions the thesis that the myriad of moist forest patches
(brejos) isolated during the Quaternary in several tropical areas of the world has
resulted in a good deal of allopatric divergence and, thus, diversity. The result of
such “refugia” are high degrees of endemism. This work is simply one of historical
biogeography, which is a province primarily of systematists. Since there are probably
more tropical insect species in the world than all other animals, it remains largely
for systematic entomologists to grapple with the problem as to how these faunas
originated. Although only 5 of the 37 papers in this book deal exclusively with insects
(and 4 of the 5 are on Lepidoptera), there is plenty of food for thought on the subject
provided by disparate disciplines. Unfortunately, the state-of-the-art on the historical
biogeography of the region emphasized in the book, the Neotropics, is not progressive,
let alone that of the other regions just casually covered.

Included are a section each on refugia theory and its applications, geology and
paleoclimatology, vegetation, insects, vertebrates, primates and anthropology, and
the Paleotropics. Redundancy throughout the book makes it much longer than is
necessary. The introductions of the majority of papers cite the few early works on
the subject and briefly review refugia theory despite the fact that a good introduction
is already provided in the first chapter by Haffer. Maps overlap considerably, too,
especially of the South American vegetation and climatic zones. Ab’Saber’s 1977
map of Pleistocene climates, for instance, is presented twice, each time on a full page.
A detailed map section in an appendix would serve for good general reference,
although many of the figures in each paper are very informative. Limiting each
contribution to four or five pages of original findings would not only have shortened
the volume, but would have prevented reiteration of what some five authors (at least
to my knowledge) have essentially published elsewhere.

The main accomplishment of the book is detailing endemism among various
Neotropical taxa. The botanists, in particular, are very lengthy in their descriptions
of floral regions. Some papers, such as those of Steyermark, Andrade-Lima, and
Huber on Neotropical plants, are mostly just big lists. Lamas, with the standard zeal
that lepidopterists have for geographic variation in wing patterns, lists subspecies of
mostly pierids, ithomiine nymphalids, and satyrids endemic to 48 Peruvian sites.
Although, as Strong comments, categorizations lead refugia theory to suffer because

1984

BOOK REVIEWS

95

they are without precise, falsifiable criteria, such thoroughness is nonetheless the right
beginning in understanding the biogeography of a traditionally neglected area.

Some authors do more than just detail endemism and they actually consider floral
and faunal relationships. However, as do Grubb (studying African forest mammals),
Walker (on the origin of the southeast Asian rain forests), and Magliazza (on linguistic
diversity in Amazonian languages), statements of evolutionary relationships are made
that rely upon inferences such as those endemic centers of greatest diversity being
the center of origin. One could very plausibly argue the opposite, that ancestral regions
should be relatively depauperate due to longer exposure to extinction. The obvious
shortcoming in most papers is a complete lack of phylogenetic thinking. In this
respect, the term refuge— supposedly the theme of the book— is prematurely used,
probably because it is such an attractive term to biogeographers. How does one
actually recognize a refuge? It is a relatively old area that has provided taxa that have
diversified subsequent to its isolation. Pragmatically, criteria and limits should be
defined to test, say, the requisite proportion of ancestral species found in a proposed
refuge. Such an approach is analogous to what many vicariance biogeographers are
doing based upon cladistic reconstructions of a taxon’s phylogeny (Patterson, 1981).

The vertebrate contributors are the most advanced in their approaches, which
perhaps reflects the status of vertebrate systematics in general. Heyer and Maxson
conclude that Leptodactylus frogs have diverged prior to the Quaternary in South
America, whereas Duellman shows that some refugia have influenced divergence in
two Hyla species groups. If improved, Heyer and Maxsons’ approach can be very
useful. However, their estimate of Leptodactylus divergence time is simply one of
the albumins on which the immunological distance matrices were based. With di-
rection given to their phenetic tree, and more loci incorporated, divergence time
could then be more accurately compared with the geological date for area divergence.
Probably the best conceived and written paper in the book is by Weitzmann and
Weitzmann, who (with several other authors) recognize the danger in scenario build-
ing based on predetermined notions of occasional dispersal events. They construct
a cladogram for each of two small fish genera ( Carnegiella and Nannostomus ), but
find little congruence between these and refugia proposed on independent grounds.
Dispersal to the present-day distributions is then suggested. Several authors, such as
Pearson (on birds) and Gentry (on woody angiosperms in Colombia) find that ende-
mism is sometimes a result of specific ecological conditions (due either to extinction
in inhospitable areas and/or dispersal to good areas). The preliminary indication is,
then, that dispersal has been important in the distribution of many taxa (hardly
surprising given the geographic scale of many groups); this should not, however, deter
us from adopting a vicariant test as the initial approach (Platnick and Nelson, 1978).

It is interesting that many of the proposed refugia do not overlap among taxa.
Exceptional is the very impressive superimposition, shown in Figure 16.36, of the
refugia based on ithomiine and heliconiine Nymphalidae subspecies distributions
(worked on by K. S. Brown) and the refugia compiled by Brown based on many
other data. Oren, at least, proposes that the average size of refugia varies among taxa
because of differences in the minimal critical population sizes (i.e., that number
needed to sustain a breeding population). This can partly explain a difference in
refugia distributions, but the matter as a whole is not attacked.

Dispute between some ecologists and the remainder of the authors provides some

96

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(1)

controversy to the topic. Endler antagonizes refugia theory with well constructed
arguments. However, his assertions are not well founded that many of the assump-
tions of refugia theory are unreasonable. For example: Many ‘refugiests’ do not
maintain allopatry is required for divergence, nor that it necessarily results in dif-
ferentiation. Generally agreed upon is that allopatric speciation is a ubiquitous, but
not universal, mode, and that sympatric speciation has certainly enjoyed no resound-
ing support in theory or fact despite the effort to prove otherwise (Futuyma and
Mayer, 1980; Jaenike, 1981). Benson, working on Heliconius communities, supports
Endlers’ parapatric model. The argument is that divergence is easily explained by a
cline of selection pressures, so much so that vicariance has little influence in distri-
butions. But, Bensons’ work deals with a small number of overt racial patterns with
simple genetic bases, the appearance of which are demographically mediated. Can
one make inferences of faunal origins based on studies of a simple and adaptive trait?
This line of argument would also claim that distinct populations of Biston betularia
have not been affected by any dispersal or vicariant event in England because the
appearance of color morphs accord so well with the presence or absence of air
pollution.

Erwin and Adis seem to be dealing more with habitat selection in carabids than
with the beetles’ biogeography. They attempt to reconcile this by stating “habitat
vicariance is simply geographic vicariance at a much finer resolution.” This seems
to suggest that sympatric speciation is the rule rather than exception for creatures of
low vagility, such as their arboreal Agra. The implication, of course, is that com-
munities can be inherently very unstable assemblages— a very unorthodox view, given
the persuasion of this symposiums’ participants.

Will we ever be able to reconstruct the effects of Quaternary forest fragmentation
on tropical diversity? Not unless better distributional and geological data is collected
and put into a phylogenetic perspective. At least with our present state of knowledge
on endemism, as stressed in the concluding chapters by Myers and by Lovejoy, we
can determine the best areas in which to establish natural preserves. For those who
feel that the biological diversity of the rapidly depleting tropics will be salvaged and
the origins of which are worth studying, I recommend gleaning this book for a few
salient references.— David Grimaldi, Department of Entomology, Cornell University,
Ithaca, New York 14853.

LITERATURE CITED

Futuyma, D. J. and G. C. Mayer. 1980. Non-allopatric speciation in animals. Syst. Zool. 29:
254-271.

Jaenike, J. 1981. Criteria for ascertaining the existence of host races. Amer. Nat. 117:830-
834.

Patterson, C. L. 1981. Methods of paleobiogeography. Pages 446-489 in: G. Nelson and D.
E. Rosen (eds.), Vicariance Biogeography: A Critique. Columbia University Press, New
York.

Platnick, N. I. and G. Nelson. 1978. A method of analysis for historical biogeography. Syst.
Zool. 27:1-16.

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Journal of the

New York Entomological Society

VOLUME 92 JANUARY 1984

NO. 1

CONTENTS

Directory of systematic entomologists and acarologists in the People’s Republic of
China Lloyd Knutson and W. L. Murphy 2-18

Vegetational complexity and parasitism of green peach aphids ( Myzus persicae (Sul-
zer) (Homoptera: Aphidae)) on collards David J. Horn 1 9-26

The Membracidae and other Homoptera described by Asa Fitch, 1851, and Ebenezer
Emmons, 1855: Historical perspective and analysis Jeffrey K. Barnes 27-34

Association of Paratrechina arenivaga (Hymenoptera: Formicidae), with nymphs of

Oecleus borealis (Homoptera: Cixiidae) C. R. Thompson 35-41

A new species of Sigara from western Oregon and Washington (Hemiptera: Corixidae)

Gary M. Stonedahl 42-47

Carvalhoma (Hemiptera: Miridae): Revised subfamily placement

Randall T. Schuh and Michael D. Schwartz 48-52

Life history and laboratory rearing of Euschistus ictericus (Hemiptera: Pentatomidae),
with descriptions of immature stages J. E. McPherson and S. M. Paskewitz 53-60

Laboratory rearing of Amaurochrous cinctipes (Hemiptera: Pentatomidae: Podopinae)
with descriptions of immature stages J. E. McPherson and S. M. Paskewitz 61-68

A conspectus of Pentatomini of the western hemisphere. Part 3. (Hemipera: Penta-
tomidae) L. H. Rolston and F. J. D. McDonald 69-86

Temporal and spatial variation in sex ratio and mating frequency in soldier beetles

Kim Holder Bailey, David E. McCauley, and W. David Sissom 87-92

Book Reviews

Bark beetles in North American conifers Stephen L. Wood 93-94

Biological diversification in the tropics: Proceedings of the Fifth International Sym-
posium for the Association for Tropical Biology David Grimaldi 94-96

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY

Editor: Randall T. Schuh, Department of Entomology, American Museum

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NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 97-120

REVISION OF AROCERA SPINOLA
(HEMIPTERA: PENTATOMIDAE)

F. J. D. McDonald

Department of Plant Pathology and Agricultural Entomology,

University of Sydney, Sydney, Australia 2006

Abstract.— The neotropical genus Arocera Spinola is revised and two keys are provided to
species, one based on color the other on the male genitalia. Two new species are described, A.
colombiana from Venezuela and Colombia and A. verdana from Panama and Costa Rica. The
following new synoymies are recognized: A. spectabilis (=A. apta Walker, =A.jalapensis Distant,
=A. patibula Distant, =A. melanopygai Stal, =A. crucigera Haglund); A. nigrorubra (=A. affinis
Distant, —A. altivola Distant, =A. chiriquensis Distant, =A. contralineata Piran, =A. placens
Walker, =A. protea Distant); A. elongata (=A. repleta Van Duzee).

Arocera Spinola occurs in Central and South America, with one species extending
into the Caribbean Islands. There is a great deal of synonymy in this genus due to
the fact that many color variants were described as species. The genus is fairly closely
related to Runibia Stal, the male genitalia of both genera being similar. However, in
Runibia the osteolar sulcus is plate-like extending less than half the distance between
the osteole and the margin of the pleuron, in Arocera the sulcus extends for two
thirds this distance and is narrow and curved cephalad. The first rostral segment
extends well past the bucculae in Runibia, whereas the first rostral segment is almost
the same length as the bucculae in Arocera. Nothing, unfortunately has been published
so far on the biology of this species. Rolston (1976) verified the generic assignment
of some species of Arocera. Distribution records indicate countries only, in the case
of larger countries, federal states are cited where possible to indicate smaller geo-
graphic regions. Froeschner (1981:68) gives a number of additional distribution rec-
ords for several species of Arocera in Ecuador.

Arocera Spinola, 1837

Arocera Spinola, 1837:316; Stal, 1867:529; Stal, 1872:37; Lethierry and Severin,

1893:158; Kirkaldy, 1909:109.

Ooedosoma Amyot and Serville, 1843:128. (Synonymized by Kirkaldy, 1909.)
Estphoria Gistel, 1848: VIII. (Synonymized by Kirkaldy, 1909.)

Type. Arocera aroleuca (Perty, 1833). Type not located.

Diagnosis. Brightly colored bugs patterned in a combination of yellow, orange or
red and black or brown, or mostly metallic green or brown. HEAD: Tylus and jugae
of equal length; jugae with margins raised at least at apex, in some species entire
margin raised, with diagonal striae. Antennae with segment 3 flattened, sometimes
grooved, flattening not very apparent in A. spectabilis ; some species also with either
2 flattened, or 4 flattened or both 2 and 4 flattened. Rostrum extending to hind coxae
or well beyond in A. elongata, first rostral segment projecting slightly beyond buc-

98

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

culae. THORAX: Prothorax with a small spine on each anterior angle. Osteolar
peritreme elongate, curved cephalad, and extending for % distance between osteole
and plural margin; evaporative area extensive, extending onto mesopleuron. AB-
DOMEN: Connexiva covered by hemelytra.

Male genitalia. Pygophore with small rounded tubercles or large horns covered
with spicules on margin. Claspers, simple L or T shaped with a small thumb-like
process in A. rufolimbata, A. verdana, A. colombiana, A. rufonotata and A. splendens.
Theca small, simple with well developed basal plates. Conjunctival appendages bi-
or tri-lobed apically, sclerotized to varying degrees; median penial lobes rod-like or
flattened disc-like structures fused basally; ejaculatory duct short, not protecting
beyond conjunctival lobes.

Female genitalia. External genitalia typically pentatomoid, dorsal surface of 8th
paratergites deeply concave in A. splendens, A. rufonotata, A. colombiana, n. sp., A.
verdana, n. sp., and A. rufolimbata, in the remaining species dorsal surface flat.
Spermatheca with well developed reservoir and pump; bulb of pump with an elongate
appendage in A. nigrorubra, A. spectabilis, and A. elongata, in the remaining species
bulb without appendages. Ring sclerites and accessory sclerites present in vulva
around spermathecal entrance.

key to the species of Arocera SPINOLA

1 . Dorsal surface* largely metallic green (sometimes blue), shades of metallic green or

dull brown, reddish brown, or brownish green 2

Dorsal surface colored in patterns of black or brown or yellow, orange or red 6

2(1). Dorsal surface metallic green (sometimes blue), with a narrow red band around
anterior margin of pronotum; or dorsal surface duller green or brown with abdominal

connexiva checked black and red 3

Dorsal surface dull red-brown or green-brown with narrow yellow or red band around

anterior margin of pronotum; abdominal connexiva unicolorous 5

3(2). Dorsal surface uniformly metallic green or greenish brown splendens (Blanchard)

Dorsal surface marked with red at least on scutellum 4

4(3). Scutellum with a brick red stripe centrally, outer margins of elytra with a brick-red

stripe from inner angle verdana McDonald

- Pronotum with a red spot centrally; red maculae on scutellum and hemelytra ....
rufonotata Stal

5(4). Dorsal surface cinnamomeus, margins of hemelytra and pronotum distinctly outlined

to varying degrees with black colombiana McDonald

- Dorsal surface brunneus or brown-green, unicolorous rufolimbata Stal

6(1). Nota and hemelytra with 5 longitudinal stripes of alternating yellow and black; black

stripes, uniting basally in membrane to form a horseshoe pattern, Fig. 9

acroleuca (Perty)

Color pattern otherwise 7

7(6). Dorsal surface dark to pinkish red with oblong black or brownish maculae on prono-
tum scutellum and hemelytra; antennal segments 2 + 3 usually distinctly flattened.

If background color is yellow or orange, then maculae on pronotum, scutellum and
hemelytra are a faded smoky brown rufifrons (Dallas)

* This term refers collectively to the pronotum, mesoscutellum and corium-clavus of the
hemelytra.

1984

REVISION OF AROCERA

99

- Species with yellow, brown or orange background, maculae on pronotum, scutellum

and hemelytra distinctly black; color if red not patterned as above 8

8(7). Dorsum patterned in distinct square maculae of black on yellow or orange (Fig. 1);
first antennal segment yellow; species usually larger than 11.0 mm long, 6.2 mm
wide (male) and 1 1.3 mm long and 6.5 mm wide (female) spectabilis (Drury)

- Color pattern variable; first antennal segment black 9

9(8). Dorsal surface bright orange red or black with red spots, well over 14 mm long . .

elongata Showalter

Dorsal surface with variable diffuse color patterns in brown or black with orange,
red or yellow; species usually no larger than 11.8 mm long and 6. 1 mm wide (male)
or 11.1 mm long and 6.4 mm wide (female) nigrorubra (Dallas)

KEY TO THE SPECIES OF ArOCera BASED ON THE MALE GENITALIA

1 . Dorsal margin without tubercles or plates. Claspers with a distinct finger-like process

on stem (Fig. 47) 2

Dorsal margin with tubercles or plates (Fig. 30). Claspers without finger-like process

on stem 6

2(1). Ventral margin produced into two stout arms one on each side of the proctiger and

an outer pair of plates, both structures covered in fine tubercles (Fig. 17) 3

- Ventral margin not so developed, lower plates absent 4

3(2). Claspers small triangular with a small process on stem (Fig. 20). Conjunctival ap-

pendages multi-lobed (Fig. 22) splendens (Blanchard)

Claspers L-shaped, process longer and curved (Fig. 53). Conjunctival appendages
bi-lobed (Fig. 54) rufolimbata Stal

4(2). Inferior ridge with small tubercles below the hom-like protuberances on either side

of the proctiger (Figs. 45, 67) 5

- Inferior ridge without tubercles colombiana, new species

5(4). Tubercles on inferior ridge arranged in a triangular patch below horns (Fig. 45).

Claspers distinctly L-shaped, apex blunt; process on stem elongate rufonotata Stal

Tubercles on inferior ridge arranged in a more linear fashion below horns (Fig. 67).
Claspers more triangulate with an acute apex, process small and stout (Fig. 47) . .
verdana, new species

6(1). Dorsal margin of pygophore with distinct plates fully visible; superior ridge not

developed acroleuca (Perty)

- Dorsal margin with tubercles partly concealed beneath the margin; superior ridge or

depression well developed (Fig. 2) 7

7(6). Dorsal margin of pygophore with a distinct depression above base of proctiger (Fig.

57). Claspers with a small tubercle on stem, apically blunt (Fig. 60)

elongata Showalter

- Dorsal margin of pygophore with a well developed superior ridge. Claspers without

tubercle, often acute apically (Fig. 40) 8

8(7). Claspers tri-lobed (Fig. 4). Ventral margin of pygophore almost straight (Fig. 3) . .

spectabilis (Drury)

Claspers otherwise. Ventral margin of pygophore either centrally concave or convex
9

9(8). Proctiger divided dorsally into two plates (Fig. 38). Apex of ejaculatory duct lying

well below outer margins of median penial lobes (Fig. 42) rufifrons (Dallas)

Proctiger not divided dorsally into two plates (Fig. 30). Apex of ejaculatory duct
reaching outer margins of median penial lobes (Fig. 34) nigrorubra (Dallas)

100

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Arocera spectabilis (Drury, 1782)

Figs. 1-8

Cimex spectabilis Drury, 1782:Index and p. 66, pi. 46.

Strachia spectabilis Dallas, 1851 :266.

Strachia generosa Stal, 1854:233.

Arocera spectabilis Stal, 1872:38; Kirkaldy, 1909:110; Grazia, 1977:163.

Arocera schumanni Distant, 1890:336, pi. 31; Kirkaldy, 1909:110; Rolston, 1976:3.
New Synonymy.

Strachia melanopyga Stal, 1858:437. New Synonymy.

Strachia apta Walker 1867:323. New Synonymy.

Arocera melanopyga Stal, 1862:107.

Arocera crucigera Haglund, 1868:157. New Synonymy.

Arocera apta Distant, 1880:73, pi. 7; Kirkaldy, 1909:109; Piran, 1963:107; Becker
and Grazia- Vieira, 1977:55.

Arocera jalapensis Distant, 1890:337, pi. 30; Kirkaldy, 1909:109. New Synonymy.
Arocera patibulata Distant, 1880:74, pi. 7. New Synonymy.

Types. The type of Cimex spectabilis was not located. However, from Drury’s
description and figure this is clearly A. spectabilis. The types of Strachia apta Walker
2; Arocera jalapensis Distant 2; Arocera schumanni Distant 6 and Arocera patibulata
Distant 2, all in the British Museum (Nat. Hist.) were examined. The type of Strachia
melanopyga Stal, A. crucigera Haglund were examined, both types are in the Na-
turhistoriska Riksmuseet, Stockholm.

Diagnosis. HEAD: Dorsal surface black basally, yellow apically; ventral surface
yellow. Eyes black. Antennae. First segment orange, remainder black; segments 2
and 3 slightly flattened. Rostrum surpassing mesocoxae, segments 1-3 orange, 4
black. THORAX: Pronotum yellow, with a black band between anterior angles and
basally two oblong black patches separated by a mesial strip of yellow. Scutellum
yellow with two oblong patches basally; confluent with those on the prothorax; two
further oblong black patches about three quarters length of scutellum separated from
basal patches by a band of yellow, apex yellow. Hemelytra yellow with two broad
oblong black patches in middle of coria confluent with patches on the scutellum.
Ventral surface orange, usually with large black spot in middle of each propleuron.
Evaporative area extending from base of mesopleuron to metapleuron. Legs. Coxae,
trochanters and proximal %, orange; femora orange; distal V6 fuscous; tibiae and tarsi
black. ABDOMEN: Connexiva orange; ventral surface orange and yellow.

Male genitalia. Pygophore with ventral margin smoothly and shallowly emarginate,
a crescent shaped depression found beneath centre of margin on central surface of
pygophore. Dorsal margin deeply arched with well defined superior ridge. A small
pair of triangular tubercles found on lateral inner margin, one on each side. Proctiger
oblong, box-like with unsclerotized mid-line. Numerous long fine setae found on
margins of pygophore and on proctiger. Clasper apically expanded into 3 lobes, dorsal
lobe finger-like, stem thick. A number of fine setae on outer surface of clasper.
Aedeagus. Theca cylindrical and very heavily sclerotized, basal plates large oblong.
Dorsal rim bearing a small membraneous bi-lobed process. One pair of conjunctival
appendages; small oblong membraneous structures slightly sclerotized apically. Me-

1984

REVISION OF AROCERA

101

1

5 -Omm

04 m m

Pt*9

Figs. 1-8. A. spectabilis. 1. Dorsal view. 2. Pygophore, dorsal view. 3. Ventral border of
pygophore. 4. Right clasper, ental view. 5. Aedeagus, lateral view. 6. Aedeagus, dorsal view. 7.
Female genitalia. 8. Spermathecal bulb. Conjunctival appendage (c.a.), first gonocoxa (lgx.),
median penial lobe (m.p.l.), process (pr.), paratergite 9 (pt.9), superior ridge (s.r.), tubercle (t.),
theca (th.).

dian penial lobes flattened disc-like. Ejaculatory duct lying entirely between median
penial lobes, short heavily sclerotized, slightly sinuous.

Female genitalia. Paratergites 8 triangular; paratergites 9 elongate, spatula-like. 1st
gonocoxae triangular meeting centrally. 2nd gonocoxae visible as a triangular section
above 1st gonocoxae. Paratergites and gonocoxae covered with long fine setae. En-
trance of spermathecal duct surrounded by a number of small sclerites and ring
sclerites. Spermatheca, typically pentatomoid with large spermathecal reservoir and
pump with distal and proximal flanges; one long process on bulb of pump.

Variations. The color of this species varies from pale cream to burnt sienna;
ventrally most specimens have large oblong black spots on the thoracic pleura on

102

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Figs. 9-16. A. acroleuca. 9. Dorsal view. 10. Pygophore, dorsal view. 1 1. Pygophore, ventral
border. 12. Left clasper, ectal view. 13. Aedeagus, dorsal view. 14. Female genitalia. 15. Entrance
of spermatheca. 16. Spermathecal bulb. Median penial lobe (m.p.l.), process (pr.), paratergite
9 (pt.9), ring sclerite (r.), tubercle (t.).

each side, but these vary in size and shape and some may be missing; the femora
have the dark brown color extending a V4-V2 way from distal end of segment.

Distribution. Colombia; Panama; Peru; Ecuador (Iquitos); Costa Rica; Brazil (Rio
de Janeiro, Mato Grosso, Obidos, Manaus, TefFe); Venezuela; Bolivia; Guatemala;
Guiana; French Guiana; Paraguay; Mexico.

Arocera acroleuca (Perty, 1833)

Figs. 9-16

Pentatoma acroleucum Perty, 1833:168, pi. 33.

Cimex acroleucus Burmeister, 1835:366.

Pentatoma aequinoxialis Westwood, 1837:33. (Synonymized by Distant, 1900b:823.)

1984

REVISION OF AROCERA

103

Ooedosoma acroleucum Amyot and Serville, 1843:128.

Arocera acroleuca Stal, 1872:37; Kirkaldy, 1909:109; Piran, 1966:86.

Type. Holotype Pentatoma acroleucum, Zoologische Museum, Munich. Type seen.

Diagnosis. HEAD: Dorsally and ventrally black except bucculae sometimes yellow
and extending from them a yellow band around base of head; bucculae distinctly
raised along outer margins, with lateral striae. Eyes black. Ocelli amber. Antennae
black, segments 3 and 4 flattened. Rostrum extending beyond metacoxae, yellow
except for apex smoky brown. THORAX: Yellow with broad horseshoe shaped black
mark, open anteriorly; black beginning on each side of pronotum basally, continuing
caudad across lateral portion of scutellum at base and coria excepting costal margin
into black membrane; anterolateral and anterior margins of pronotum, mesial stripe
on pronotum and scutellum, costal margin of coria, all yellow. Ventral surface of
thorax yellow. Legs. Coxae, trochanters yellow; basal half of femora yellow, apical
half dark brown; tibiae and tarsi dark brown. ABDOMEN: Connexiva yellow; sterna
yellow.

Male genitalia. Pygophore with ventral border biconcave on either side of a median
rounded projection, a shallow emargination found centrally beneath border. Dorsal
border broadly arched with a central collar forming a median emargination into
which the apex of the proctiger sits when at rest. A small flattened oval pygophoral
plate found one on each side laterally just adjacent to margin of border; apex of
claspers lying on top of this plate when at rest. Proctiger oblong. Claspers C-shaped,
apically broad, bi-lobed; basally with a small tubercle on inner dorsal margin; short
setae on tubercle and longer ones on outer margin of clasper. Aedeagus. Theca
cylindrical, moderately sclerotized; 1 pair membraneous conjunctival appendages,
small lobe-like. Median penial lobes small, spatulate, basally tapering; ejaculatory
duct small, straight, completely enclosed by median penial lobes.

Female genitalia. External genitalia very similar to A. elongata. A number of small
crescent shaped sclerites around ejaculatory duct opening (Fig. 15). Ring sclerites
present. Spermatheca. Bulb with a long and short process.

Distribution. Brazil (Goiaz, Encruzilhada; Vicosa; Sao Paulo; Teffe; Mato Grosso;
Barroso; Diamantino); Peru; Colombia; Surinam; Bolivia; Ecuador; Guatemala; Pan-
ama; Guyana.

Arocera splendens (Blanchard, 1841)

Figs. 17-25

Pentatoma splendens Blanchard, 1841:148.

Pentatoma splendida Dallas, 1851:256.

Pentatoma principalis Stal, 1855:182; 1856:58.

Arocera principalis Stal, 1862:107; Becker and Grazia- Vieira, 1971:12.

Arocera splendens Stal, 1872:38; Distant, 1880:pl. 7, figs. 13-17; Kirkaldy, 1909:
1 10; Becker and Grazia-Vieira, 1971:12.

Type. Type of Pentatoma splendens not located.

Diagnosis. HEAD: Dark metallic green or blue dorsally, reddish brown ventrally.
Eyes dark brown. Ocelli reddish brown. Antennae fuscous, segments 2 and 3 flattened.
Rostrum reaching hind coxae, dark brown. THORAX: Pronotum dark metallic green

104

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Figs. 17-25. A. splendens. 17. Pygophore, dorsal view. 18. Pygophore, caudal view. 19.
Pygophore, ventral border. 20. Right clasper, ental view. 21. Aedeagus, lateral view. 22. Ae-
deagus, ventral view. 23. Female genitalia. 24. Spermatheca. 25. Spermathecal bulb. Conjunc-
tival appendage (c.a.), dilation (d.), ejaculatory duct (e.d.), median penial lobe (m.p.l.), process
(pr.), paratergite 8 (pt.8), paratergite 9 (pt.9), ventral margin (v.b.).

with anterior and anterolateral margins outlined in red; scutellum dark metallic green.
Hemelytra, dark metallic green, coria with costal margin red basally. Pleura and
sterna red suffused with brown. Legs. Coxae reddish pink; trochanters basally red
darkening to brown apically; femora brown; tibiae and tarsi dark brown. ABDOMEN:

1984

REVISION OF AROCERA

105

Connexiva alternated red and dark brown; sterna red suffused with brown, lateral
margins of segments 3-7 with a bright red oblong spot.

Male genitalia. Pygophore with ventral margin divided into two sections, upper
margin consisting of two stout arms, one on each side projecting across the lateral
margins of the pygophoral opening; these arms covered on their apices and outer
margins with a series of small tubercles. Lower margin developed into two flat plates
meeting centrally in a deep U-shaped median emargination, between these plates lies
the inferior ridge which is broadly U-shaped with a small median notch forming the
ventral border to genital opening. Inner margins of lower plates covered with small
heavily sclerotized tubercles bearing fine setae. Dorsal margin broadly arched, form-
ing a small blunt hom-like process at its junction with ventral margin on each side.
Proctiger tube-like, distal margin produced into two upturned horns. Claspers small,
triangular, bearing a small thumb-like process half way along outer margin and
numerous fine setae on margins. Aedeagus. Theca small, weakly sclerotized. Con-
junctival appendages large, basally membraneous, apically divided into 4 finger-like
lobes, apical two broader and larger than basal two, all lobes moderately sclerotized,
upper 3 lobes covered in small fine teeth. Median penial lobes sclerotized and com-
pletely enclosed by conjunctival appendages, basally fused into a solid stem, apically
divided into two bluntly rounded curved flattened arms. Ejaculatory duct a small
sclerotized straight tube lying within median penial lobes.

Female genitalia. Externally similar to A. elongata, however 8th paratergites with
a vertical side on inner surface forming a concavity; 9th paratergites deeply impressed
centrally forming the anterior wall of concavity; 1 st gonocoxae ovoid, with a posterior
fringe of hairs; 2nd gonocoxae fused and lying vertically at right angles to 1st gono-
coxae, facing posteriorly. Ring sclerites and small accessory sclerites present round
entrance to spermathecal duct. Spermatheca. Reservoir apically spherical with a short
sclerotized sausage-like base. Pump with distal and proximal flanges; bulb simple,
dome-like.

Variations. Color varies from light metallic green to metallic blue or dark brown
with a slight metallic green sheen dorsally. Anterior margins of pronotum ventrally
outlined in yellow brown. Abdominal sterna may be dark brown or yellow brown,
sometimes with 3 patches of red or yellow brown centrally on each segment; lateral
spots may be yellow brown.

Distribution. El Salvador; Mexico; Guatemala; Ecuador; Colombia; Venezuela;
Panama; British Honduras; Tobago; Jamaica; Peru.

Arocera nigrorubra (Dallas, 1851)

Figs. 26-36

Strachia nigrorubra Dallas, 1851:267.

Strachia quadripunctata Signoret, 1851:337, pi. 10.

Arocera nigrorubra Stal, 1872:38; Kirkaldy, 1909:109; Rolston, 1976:3.

Arocera ajfinis Distant, 1880:74, pi. 7; Kirkaldy, 1909:109; Rolston, 1976:3. New

Synonymy.

Arocera altivola Distant, 1890:337, pi. 31; Kirkaldy, 1909:109; Rolston, 1976:3. New

Synonymy.

1 -4 m m

O -9 m m

1984

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107

Arocera chiriquensis Distant, 1890:337, pi. 30; Kirkaldy, 1909:109; Becker and Gra-
zia-Vieira, 1971:11; Rolston, 1976:3. New Synonymy.

Arocera contralineata Piran, 1955:82; Becker and Grazia-Vieira, 1971: 1 1. New Syn-
onymy.

Strachia placens Walker, 1867:316.

Arocera placens Distant, 1900:391; Kirkaldy, 1909:1 10; Rolston, 1976:3. New Syn-
onymy.

Arocera protea Distant, 1880:73, pi. 7; Kirkaldy, 1909:110; Rolston, 1976:3. New

Synonymy.

Types. Holotype Strachia nigrorubra Dallas 6, British Museum (Nat. Hist.). Type
seen. The following types in the British Museum (Nat. Hist.) were also examined:
Arocera affinis Distant $; Arocera altivola Distant <3; Arocera chiriquensis Distant <5;
Strachia placens Walker <3. The holotype of Arocera contralineata Piran could not
be obtained, however a paratype in the collection of the University Nacional de la
Plata was examined.

Comment. This proved to be an exceedingly difficult species to define. It consists
of a maze of color patterns which actually integrade with A. spectabilis. There are,
however, overall differences between these insects and their congeners in size and in
the male genitalia to warrant retaining species status for these forms. I suspect many
of the color patterns are local populations and a great deal of ecological work needs
to be done on this species to determine the limits of each color morph.

Diagnosis. HEAD: Dorsally and ventrally black; jugae swollen apically and raised.
Eyes brown. Ocelli red. Antennae black, segments 2, 3 and 4 flattened. Rostrum
black, apex exceeding hind coxae. THORAX: Pronotum with a broad black margin
around outer margins, central portion bright red with two black spots. Scutellum
black with two red spots one on each side near apex; elytra bright red with broad
black bands proximad, two small black spots in centre of distal red portion. Pleura,
sterna and legs black. ABDOMEN: Connexiva alternated red and black; sterna red
with a central broad black band and small black spots surrounding each spiracle;
further oblong black spots found in the disto-lateral comer of each segment.

Male genitalia. Pygophore with ventral margin almost straight with a shallow
median emargination, a triangular depressed area found below margin giving the
appearance of a vertical wall when pygophore is in situ. Fine setae on margins and
surface of pygophore. Dorsal margin with a superior ridge and a pair of small oval
well sclerotized tubercles, one on each side, lying beneath the border. Claspers L-shaped,
apex produced into a finger-like process, basally flattened and curved up, bearing a
tuft of long setae, fine setae and serrations on outer margin. Proctiger box-like, bearing
numerous fine setae. Aedeagus similar to A. spectabilis.

Female genitalia. Similar to A. spectabilis, paratergites 9 somewhat more slender;
1st gonocoxae with inner margins sinute (generally straight in A. spectabilis ). Sper-
matheca similar to A. spectabilis.

Figs. 26-36. A. nigrorubra. 26-29. Dorsal view showing color patterns. 30. Pygophore,
dorsal view. 31. Pygophore, ventral border. 32. Right clasper, ental view. 33. Aedeagus, lateral
view. 34. Aedeagus, ventral view. 35. Female genitalia. 36. Spermatheca. Conjunctival appen-
dage (c.a.), median penial lobe (m.p.l.), superior ridge (s.r.), tubercle (t.).

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Variations. There are several color patterns, the basic type of each will be described
although various integrades between each can readily be found.

1. Color pattern basically similar to that of A. spectabilis, color background may
vary from bright yellow to deep orange; head bicolored above and below, pattern
variable.

2. Prothorax with a broad orange T-shaped mark; scutellum with an orange cross;
distal portion of elytra with irregular orange macula. Abdominal sterna may lack
central maculae except on 7 (Fig. 28).

3. Prothorax and scutellum with a large cross outlined in brown; areas on each
side of cross orange or brown margining into dark brown across top of prothorax;
hemelyra with more orange or reddish orange areas. Abdominal sterna with small
central maculae and no maculae around spiracles. Some specimens have only a broad
brown band along the distal margin of each sternum (Fig. 27).

4. As above but intervening dark brown or black, leaving cross in yellow or red,
apical portion of cross swollen. Abdominal sterna without maculae around spiracles
(Fig. 29).

5. Dorsum uniformly yellow to red excepting black on part or all of head, anterior
margin of pronotum and small macula on disk of costal margin of coria.

Distribution. Peru; Mexico; Ecuador; Brazil (Rio de Janeiro, Bahia, Mato Grosso,
Obidos, Manaus, Teffe); Colombia; Cuba; Guatemala; Paraguay; Argentina; Panama;
Haiti; Jamaica; St. Lucia; Trinidad.

Arocera rufifrons (Dallas, 1851)

Figs. 37-43

Strachia rufifrons Dallas, 1851:267.

Arocera rufifrons Stal, 1872:38; Kirkaldy, 1909:110; Rolston, 1976:3.

Type. Two syntypes, both 9, British Museum (Nat. Hist.). Types seen. I select the
specimen with the label 85a as LECTOTYPE, the other specimen labelled 85b, is
the PARALECTOTYPE.

Diagnosis. HEAD: Dorsally, apical % scarlet, basal Vi dark brown, ventrally scarlet.
Jugae raised along outer margins with a series of diagonal striae (sometimes absent);
margins of jugae continued inwards to form a short ridge in front of each eye. Eyes
brown. Ocelli amber. Antennal segment 1 red, 2-5 dark brown; 2-4 flattened. Ros-
trum extending well beyond mid coxae, segment 1 red, 2-4 brunneus. THORAX:
Pronotum scarlet with black maculae in anterior angles; and two oblong black patches
in center of pronotum extending onto scutellum forming 2 large oblong maculae.
Scutellum scarlet with two further black maculae extending midway to near apex—
these merge with maculae on the hemelytra to form 2 bands separated centrally
across the mid portion of the species. Pleura mottled reddish brown. Legs. Coxae,
trochanters brunneus; remaining segments mid-brown. ABDOMEN: Connexiva and
sterna scarlet, with paired black spots on each side laterally on segments 3-6.

Male genitalia. Similar to A. spectabilis. Pygophore with ventral margin straight,
impressed below margin forming a vertical wall when pygophore in situ. Superior
ridge not as well defined as in A. spectabilis tubercles mainly concealed below dorsal
margin. Claspers, L-shaped with finger-like process apically; upper margin smoother

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109

Figs. 37-43. A. rufifrons. 37. Dorsal view. 38. Pygophore, dorsal view. 39. Pygophore,
ventral border. 40. Right clasper, ectal view. 41. Aedeagus, lateral view. 42. Aedeagus, dorsal
view. 43. Spermathecal bulb. Process (pr.), superior ridge (s.r.), tubercle (t.).

than in A. spectabilis. Aedeagus similar to A. spectabilis, median penial lobes more
rounded and club-like.

Female genitalia. External genitalia similar to A. spectabilis. Spermathecal bulb
with one process.

Variations. Background color may be orange or yellow brown. The proportion of
reddish and brownish color on head is variable. Some specimens have no maculae
on the abdominal sterna but spots at lateral margins; others have a large central spot
as well as lateral maculae. The color of the dorsal maculae can vary from black to
faded smoky-brown.

Distribution. Colombia; Peru; Belize; Mexico; Brazil.

Note. This species is very close to A. spectabilis and may with further biological
information prove to be only a color variant of that species.

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Figs. 44-50. A. rufonotata. 44. Pygophore, dorsal view. 45. Pygophore, caudal view. 46.
Pygophore, ventral border. 47. Right clasper, ectal view. 48. Aedeagus, lateral view. 49. Ae-
deagus, ventral view. 50. Spermatheca. Dilation (d.), dorsal lobe of conjunctival appendage
(d.c.a.), median penial lobe (m.p.l.), process (pr.).

Arocera rufonotata Stal, 1861
Figs. 44-50

Arocera rufonotata Stal, 1861:140; Distant, 1880:pl. 7; Kirkaldy, 1909:110.

Type. Holotype. Naturhistoriska Riksmuseet, Stockholm. Type seen.

Diagnosis. HEAD: Very dark metallic green dorsally, centrally dark brown with
patches of red on either side of bucculae. Jugae slightly raised apically with distinct
diagonal striae. Eyes brown. Ocelli amber. Antennae black, segments 2 and 3 flat-
tened. Rostrum reaching to or slightly past metacoxae, dark brown. THORAX:
Prothorax dark metallic green with red spots at anterior angles and lateral angles and

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11

one red spot centrally. Scutellum dark metallic green with an elongate red spot in
lateral angles. Hemelytra dark metallic green with an elongate red spot centrally at
distal margin of corium. Ventral surface of thorax black with red spots at anterior
and lateral angles of prothorax; base of proepistemum and proepimeron red; mes-
epistemum above metacoxae, red. Metepistemum with red patch above metacoxae
extending below evaporative area to posterior and lateral margins of segment. AB-
DOMEN: Connexiva alternated red and black, sterna black with red oblong spots
at lateral margins of segments 3-7 and a further two large red spots, one on either
side of the mid line on segments 3-6, widely separate on 3, less widely separated
on 6.

Male genitalia. Pygophore very similar to A. rufolimbata, shape of inferior ridge
somewhat different, deeper and without central notch. Clasper similar to A. rufo-
limbata, finger-like process longer, narrower; apex of clasper less acute than in A.
rufolimbata. Aedeagus. Conjunctival appendages tri-lobed; dorsal lobe bluntly round-
ed, sclerotized; middle lobe oblong, sclerotized and with minute spines over entire
surface; lower lobe acute, heavily sclerotized on upper surface; base of appendages
membraneous. Median penial lobes fused basally, apically developed into two broad
rounded lobes, heavily sclerotized. Ejaculatory duct short, straight, enclosed by me-
dian penial lobes.

Female genitalia. External genitalia similar to A. splendens. Spermatheca similar
to A. splendens, dilation with small thickened base somewhat differently shaped from
that of A. splendens.

Variations. Some specimens appear almost metallic dark brown and the dorsal
spots may be orange-brown; ventral surface may be orange-brown with no spotting
on the abdominal sterna. In other specimens the ventral surface is brown, the two
central spots of the abdomen become diffuse, giving the abdomen a mottled brown
and red appearance. Some specimens have 2 central red spots on abdominal ster-
num 7.

Distribution. Mexico.

Arocera rufolimbata Stal, 1872
Figs. 51-55

Arocera rufolimbata Stal, 1872:38; Kirkaldy, 1909:110; Monte, 1945:269; Grazia,
1977:163.

Type. Holotype. Naturhistoriska Riksmuseet, Stockholm. Type seen.

Diagnosis. HEAD: Dorsally greenish black overlaid with a metallic green sheen,
ventrally buff; jugal margins raised anteriorly, diagonal striae on jugae and tylus.
Eyes and ocelli brown. Antennae black, segments 2-4 flattened. Rostrum slightly
surpassing metacoxae; segment 1 buff, 3-4 dark brown. THORAX: Pronotum dark
greenish-black with slight metallic sheen; a yellow stripe running right round outer
margin of pronotum and extending down outer margins of elytra for lh their length.
Scutellum and elytra same color as pronotum. Pleura and sterna testaceous. Legs.
Coxae, trochanters, femora testaceous; tibiae and tarsi dark brown. ABDOMEN:
Connexiva testaceous; sterna yellow.

Male genitalia. Pygophore similar in some respects to A. colombiana. Ventral

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Figs. 51-55. A. rufolimbata. 51. Pygophore, dorsal view. 52. Pygophore, ventral border.
53. Left clasper, ectal view. 54. Aedeagus, lateral view. 55. Aedeagus, ventral view. Conjunctival
appendage (c.a.), median penial lobe (m.p.l.), process (pr.), superior ridge (s.r.).

margin developed into two large hom-like processes, one on each side, laterally
covered with small heavily sclerotized tubercles; centre of margin deeply emarginate
and crenulated. Inferior ridge shallow forming a flattened and somewhat depressed
area below ventral margin, bearing two distinct patches of tubercles one on each side
produced into two acute ridges centrally and separated by a short gap. Dorsal margin
with a deep oval depression one on either side of a narrow superior ridge. Numerous
long stout setae found on margins of pygophore. Proctiger box-like well sclerotized,
caudally expanded into two blunt projections. Claspers L-shaped, apically acute and
with a distinct thumb-like process near apex of stem. Aedeagus. Theca short sausage-
like. One pair of conjuntival appendages divided into two lobes, upper lobe oblong
sclerotized and covered with fine spines; lower lobe bluntly rounded, sclerotized
apically; basically both appendages membraneous. Median penial lobes Y-shaped,
basally fused, apically curved and cylindrical. Ejaculatory duct short, tubular and
sclerotized.

Female genitalia. Similar to A. splendens. Spermatheca also similar.

Variations. Some specimens may be dorsally distinctly reddish brown with ab-
dominal sterna black with 3 red spots centrally on segments 3-6; and 2 centrally on
7; lateral margins of segments 2-7 with broad red maculae. The abdominal sterna
of other specimens may be patterned in the same way with yellow or orange. Re-
ductions of this pattern are also found.

Distribution. Colombia; Venezuela; Brazil.

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113

Figs. 56-63. A. elongata. 56. Dorsal view. 57. Pygophore, dorsal view. 58. Pygophore, dorsal
view, right clasper removed. 59. Pygophore, ventral border. 60. Right clasper, ectal view. 61.
Right clasper, ventral view. 62. Aedeagus, lateral view. 63. Aedeagus, dorsal view. Conjunctival
appendage (c.a.), dorsal margin (d.m.), depression (dp.), ejaculatory duct (e.d.), median penial
lobe (m.p.l.), proctiger (pg.), process (pr.), tubercle (t.).

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Arocera elongata Showalter, 1929
Figs. 56-66

Arocera elongata Showalter, 1929:33, pi. V, and p. 40; Van Duzee, 1937:25-27;

Piran, 1962:5; Becker and Grazia- Vieira, 1971:11; Sabrosky, 1972:87-88.
Arocera repleta Van Duzee, 1931:94. New Synonymy.

Types. Lectotype No. 52107, U.S. National Museum (designated Sabrosky, 1972).
Type seen. The holotype of Arocera repleta Van Duzee, California Academy of
Sciences has also been examined.

Diagnosis. HEAD: Bright orange-red dorsally except for two triangular black patches
running across ocelli, ventrally orange-red; jugae slightly swollen apically and re-
curved; diagonal striae near base. Antennae. Segment 1 orange, 2-5 black, segments
2-4 flattened. Rostrum projecting well beyond hind coxae; segment 1 orange, 2-5
black. THORAX: Pronotum black; scutellum black with orange-red tip; hemelytra
with orange maculae at base and apical corners of corium; black in between; mem-
brane black; pleura black except for an orange band at lateral margin, sterna black;
legs black. ABDOMEN: Connexiva red; sterna orange with a black band at tip of
abdomen, covering segment 7 and small sections of 6.

Male genitalia. Pygophore with ental margin widely excavated forming two blunt
prominences one at each outer comer. Dorsal border deeply excavated bearing cen-
trally a crescent-shaped depression. Proctiger oblong, swollen basally, divided into
two plates down centre. Pygophore and proctiger bearing numerous fine setae. Inner
dorsal margin of cup bearing a small rounded club-like process one on each side.
Claspers L-shaped with broad base, bearing a number of setae on outer margin.
Aedeagus similar to A. spectabilis. Median penial lobes broadly hook-shaped, flat-
tened, sclerotized on outer dorsal margins, basally fused by a short cross bar.

Female genitalia. Similar to A. spectabilis.

Variations. Small black spots may be found on lateral margins of abdominal
segments 3-5. A distinct color variety exists which is dorsally bright orange-red or
orange-brown; some specimens have varying degrees of black forming maculae on
the prothorax/scutellum and black spots on hemelytra.

Distribution. Panama; Ecuador; Peru; Venezuela; Bolivia; Colombia. The bright
orange-red variety appears to be restricted to Panama.

Acrocera verdana, new species
Figs. 67-71

Description. HEAD: Dorsally metallic green, ventrally dark brown suffused with
red. Jugae only slightly raised anteriorly with numerous diagonal striae. Eyes black.
Ocelli amber. Antennae black, segments 2 and 3 flattened. Rostrum extending to
hind coxae, black. THORAX: Pronotum metallic blue-green with a broad brick-red
band running right around outer margin from lateral angles, this band extends down
along the outer margins of the hemelytra. Scutellum and remainder of hemelytra
metallic blue-green with an irregular brick-red broad stripe running centrally from
base of scutellum to the tip. Membrane metallic blue-green. Pleura iridescent black
with a broad brick-red band running along outer margins of propleura down the
lateral margins of the meso- and metapleura and along basal margin of metapleuron.

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115

Figs. 64-71. 64-66. A. elongata. 64. Female genitalia. 65. Spermatheca. 66. Spermathecal

bulb. 67-71. A. verdana. 67. Pygophore, caudal view. 68. Pygophore, ventral border. 69. Right
clasper, ectal view. 70. Aedeagus, lateral view. 71. Aedeagus, dorsal view. Ejaculatory duct
(e.d.), 1st gonocoxa (lgx.), process (pr.), paratergite 8 (pt.8), paratergite 9 (pt.9).

Legs dark reddish brown. ABDOMEN: Connexiva brick-red, sterna iridescent black
with a broad brick-red band along lateral margins of abdomen.

Male genitalia. Pygophore similar in many respects to A. rufonotata. Inferior ridge
not as deeply excavated and forming a shallower depression behind ventral margin.
A series of small tubercles running from large peg-like structures on lateral margin
inside this depression, position of these tubercles is different from those found in A.
splendens and A. rufonotata. Clasper very acutely pointed with a short thumb-like
process. Aedeagus. Conjunctival appendages tri-lobed as in A. rufonotata, ventral

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most appendages heavily sclerotized, more acute than in A. rufonotata. Median penial
lobes, slender arms fused to a broad base forming a Y; ejaculatory duct long, straight,
extending well beyond median penial lobes (unlike any other related species).

Female genitalia. External genitalia similar to A. splendens, spermatheca similar
to A. rufonotata.

Holotype. 6, labeled Panama: Chiriqui Pr., 3 km W. Cerro Punta, 80°5 1 'N, 82°36'W,
8-VI-77. Deposited in U.S. National Museum. Type No. 100413.

Paratypes. PANAMA: 9, Chiriqui Dist., Renacimiento, Sta. Clara, 17-V-77, R.
Hartman; 9, same data, except July 5, 1977; 9, same data, except 16-20-VI-77; 6,
same data, except 4-VII-77; 9, same data, except 17-V-77; 266, same data, except
5-VII-77; 9, Chirique, Boquette, 1,250 m, 8°48'N, 82°26'W, 14-X-1976, H. Wolda;
9, same data, except 26-V-1977; 9, Chirique Dist., Renacimiento, Oeste Clara, 5500',
16-20 Feb. ’77, lights, A. & E. Thurman; 299, Chiriqui Dist., Renacimiento, 5500',
18-22-V-77, at lights, Engleman and Matos; 9, Chiriqui Pr., 3 km W. Cerro Punta,
8°51'N, 82°36'W, 1,700 m, 8-VI-77 [In H. Dodge Engleman collection]. COSTA
RICA: 9, Guanacaste, V. 17. 1932, A. Alfaro, J. C. Butz Collection, 1961; 9, Juna
Vimas, Collection Wm. Schausz [In U.S.N.M.]. PANAMA: 6, Barriles, Chiriqui,
1-31-31, M. E. McLellan; 9, vicinity Boqueta, VIII-39, J. R. Slevin [In California
Academy of Sciences].

Arocera colombiana, new species
Figs. 72-79

Description. HEAD: Dorsally black with a slight metallic green sheen; ventrally
dark brown suffused with cream centrally; jugae with lateral angles slightly turned
up with horizontal striae. Eyes, ocelli brown. Antennae dark brown; segments 2, 3
and 4 flattened. Rostrum extending to hind coxae, first segment brown suffused with
yellow, segments 2-4 brown. THORAX: Pronotum anterior portion with a broad
yellow band extending along lateral margins, remainder black with a central brown
patch. Scutellum with a central brown patch surrounded by black. Hemelytra with
a yellow streak on costal margins of coria extending about xh way along, remainder
black surrounding a central triangular patch of brown. Propleura dark brown with
anterior margins yellow; meso- and metapleura dark brown. Legs chocolate brown.
ABDOMEN: Connexiva alternated yellow and brown; sterna chocolate brown with
4 large yellow spots one on each lateral margin and two on either side of mid line;
widely separate on segment 3, closer together on 7.

Male genitalia. Pygophore with ventral margin deeply excavated and bearing a well
developed inferior ridge forming an inner vertical wall facing caudad. Central margin
of inferior ridge deeply excised, laterally on each side developed into a stout well
sclerotized horn covered with tubercles and short setae. Dorsal margin broadly arched
bearing medianly a narrow ill-defined superior ridge. Patches of long setae on margins
of pygophore and on face of inferior ridge. Proctiger well sclerotized box-like, caudal
margin produced into two short acute tails. Claspers small triagular with a finger-
like process on the inner margins; claspers firmly embedded into wall of pygophore.
Aedeagus. Theca small squat. Conjunctival appendages tri-lobed, lobes apically bluntly
pointed, upper and middle lobes covered with small spines, lower lobes sclerotized
on inner surfaces. Sclerotized median penial lobes fused into a flattened collar-like

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117

Figs. 72-79. A. colombiana. 72. Pygophore, dorsal view. 73. Pygophore, ventral border. 74.
Right clasper, ectal view. 75. Aedeagus, lateral view. 76. Aedeagus, ventral view. 77. Female
genitalia. 78. Spermatheca. 79. Spermathecal bulb. Dilation (d.), dorsal lobe of conjunctival
appendage (d.c.a.), inferior ridge (i.r.), median penial lobe (m.p.l.), process (pr.), paratergite 9
(pt.9), ring sclerite (r.), ventral margin (v.b.).

structure surrounding base of ejaculatory duct, latter a short tube moderately scler-
otized.

Female genitalia. Paratergites 8 triangular. Paratergites 9 hook-shaped, deeply
impressed centrally giving the paratergite a twisted appearance. Swollen apically and
covered with long stout setae. 1st gonocoxae oblong heavily sclerotized, covered with
stout setae. 2nd gonocoxae fused into a trapezoid plate lying vertically above 1st
gonocoxae. Ring sclerites and accessory sclerites present internally. Spermatheca.
Dilation swollen into a bulb basally; spermathecal pump without appendages.

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Variations. Brown areas on pronotum scutellum and elytra may be extensive to
the extent of eliminating most of the black areas leaving only a thin black line around
scutellum and outer margins of elytra. Sterna may be completely orange-red with
evaporative areas black. Spots on abdominal sterna may be red with black maculae
centrally. Abdominal sterna may also be yellow with dark brown spots around spi-
racles and brown maculae centrally. Integrades between these types may also be
found.

Holotype. 6, labelled Venez. Deposited in the U.S. National Museum. Type No.
100414.

Paratypes. VENEZUELA: 9, (a) Merida, (b) W. Robinson bequest 1929; 9, H.
Pittier; 9, Rancho Grande Nr. Marcay, Ven. 8-V-1946. COLOMBIA: 6, (a) Pres, by
the Hono. Apolinar-Maria, (b) Fusagasuga, S. A. Rep. Col. 1915; 9, same data; 9, (a)
Muzo, S. A. Colombia, (b) Pres, by Hono Apolinar-Maria. [All in U.S.N.M.].

Nomina dubia

Arocera fasciativentris Breddin, 1901

The type of this species could not be located. From Breddin’s description it is
probably synonymous with A. spectabilis.

Arocera capitata Breddin, 1901

The type of this species could not be located. From Breddin’s descriptions this
species is probably synonymous with A. spectabilis.

ACKNOWLEDGMENTS

I should like to thank the following people for the loan of material and type specimens. W.
R. Dolling, British Museum (Natural History), London; C. A. Triplehom, Ohio State University,
Columbus; Paul H. Amaud, Jr., California Academy of Sciences, San Francisco; Richard C.
Froeschner, National Museum of Natural History, Washington, D.C.; Randall T. Schuh, Amer-
ican Museum of Natural History, New York; Luis de Santis, University Nacional de la Plata,
La Plata; A. Kaltenbach, Naturhistorisches Museum, Wien; Per Lindskog, Naturhistoriska
Riksmuseet, Stockholm; H. Dodge Engleman, Coco Solo, Panama Canal. I should also like to
thank Professor L. H. Rolston who suggested this study and carried out much of the initial
literature survey.

LITERATURE CITED

Amyot, C. J. B. and A. Serville. 1843. Histoire Naturelle des Insectes. Hemipteres. Roret,
Paris, LX VI + 675 + 6 pp„ 12 pis.

Becker, M. and J. Grazia- Vieira. 1971. Contribuicao as conhecimento da superfamilia Pen-
tatomoidea na Venezuela (Heteroptera). Iheringia (Zool.) 40:3-25.

Becker, M. and J. Grazia-Vieira. 1977. The Pentatomoidea (Heteroptera) collected in French
Guiana by the expedition of the Museum National d’Histoire Naturelle. Ann. Soc. Ent.
Fr. (N.S.) 13:53-67.

Blanchard, E. 1841. Histoire Naturelle des Insectes. Orthopteres, Neuropteres, Hemipteres,
Hymenopteres, Lepidopteres et Dipteres. Dumenil, Paris, 3 vols., 3 + 85-212 pp.

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Burmeister, H. C. C. 1835. Handbuch der Entomologie, Bd. 2. Berlin, ii + 400 + 4 pp., 2
pis.

Dallas, W. S. 1851. List of Specimens of Hemipterous Insects in the Collection of the British
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Distant, W. L. 1880-1893. Biologia Centrali-Americana. Insecta, Rhynchota, Vol. 1. London,
XX + 462 pp., 39 pis.

Distant, W. L. 1900a. Rhynchotal notes— iv. Heteroptera:Pentatominae (Part). Ann. Mag.
Nat. Hist. (7)5:386-397.

Distant, W. L. 1900b. Revision of the Rhynchota belonging to the family Pentatomidae in
the Hope collection of Oxford. Proc. Zoo. Soc. London (1900):800-824.

Drury, D. 1782. Illustrations of Natural History, Vol. II. B. White, London.

Froeschner, R. C. 1981. Heteroptera or true bugs of Ecuador: a partial catalog. Smithsonian
Contrib. Zool. 322:i-iv, 1-147.

Gistel, J. 1 848. Naturgeschichte des Thierreichs fur hohere Schulen. Zweite Auflage. Scheitlin
& Krais, Stuttgart, 216 pp., 32 pis.

Grazia, J. 1977. Revisao dos Pentatomineos citados no “Quarto catalogo dos insetos que
vivem nas Plantas do Brasil.” (Hemiptera— Pentatomidae— Pentatomini). Dusenia 10(3):
161-174.

Haglund, C. J. E. 1868. Hemiptera nova. Stett. Ent. 29:157.

Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera) Vol. 1. Cimicidae. Felix
Dames, Berlin, XL + 392 pp.

Lethierry, L. and B. Severin. 1893. Catalogue General des Hemipteres Vol. 1. Pentatomides.
Brussels and Berlin, IX + 286 pp.

Monte, O. 1945. Notas sobre alguns pentatomideos. Rev. Agr. Sao Paulo 20:269-273.

Perty, J. A. M. 1833. Delectus animalium articulatorium quae in itinere per Brasiliam. Fasc.
3. Monachii. Pp. 125-224, pis. 25-40.

Piran, A. A. 1955. Arocera contralineata, especie neuva de la fauna Argentina (Hemiptera,
Pentatomidae). Neotropica 1:81-84.

Piran, A. A. 1962. Hemiptera neotropica V. Notas sobre sistematica y Zoogeografia de Pen-
tatomidae. Acta Zool. Lilloana 18:5-10.

Piran, A. A. 1963. Hemiptera neotropica VIII. Especies neuvas o poco conocidas de las jauans
de Colombia, Peru, Brasil, Bolivia y Paraguay. Physis 24(67): 107-1 12.

Piran, A. A. 1966. Hemiptera Neotropica X. Rev. Soc. Ent. Arg. 28:85-89.

Rolston, L. H. 1976. An evaluation of the generic assignment of some American Pentatomini
(Hemiptera: Pentatomidae). J. New York Ent. Soc. 84(1): 2-8.

Sabrosky, C. W. 1972. The case of the elusive author. Proc. Ent. Soc. Wash. 74:87-88.

Showalter, W. J. 1929. Insects: exploring the wonders of the insect world. Nat. Geog. Mag.
(July): 1-90.

Signoret, V. 1851. Description de nouvelles especes d’Hemipteres. Ann. Soc. Ent. Fr. (2)9:
329-348.

Spinola, M. 1837. Essai sur genres d’insectes appartenants a l’ordre des Hemipteres L., on
Rhyngotes Fabr. et a la section des Heteropteres, Dujour. Chez Yves Gravier, Genes,
383 + 16 pp.

Stal, C. 1854. Nya Hemiptera. Ofversigt Svenska Vet.-Akad. Forh. 12:27-47.

Stal, C. 1855. Nya Hemiptera. Ofversigt Svenska Vet.-Akad. Forh. 12:181-192.

Stal, C. 1856. Hemipterologiska bidrag. Ofversigt Svenska Vet.-Akad. Forh. 13:51-68.

Stal, C. 1861. Miscellanea Hemipterologica. Stett. Ent. Zeit. 22:129-153.

Stal, C. 1 862. Hemiptera Mexicana enumeravit speciesque novas descripsit. Stett. Ent. Zeit.
23:81-118.

Stal, C. 1867. Bidrag till Hemipteranas systematik. Ofversigt Svenska Vet.-Akad. Forh. 24:
522-532.

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Stal, C. 1872. Enumeratio Hemipterorum. Svenska Vet.-Akad. Handl. 10(4):3-65.

Van Duzee, E. P. 1931. Four new neotropical Heteroptera. Pan-Pacific Ent. 8:93-95.

Van Duzee, E. P. 1937. A few new Hemiptera. Pan-Pacific Ent. 13:25-27.

Walker, F. 1867. Catalogue of the specimens of Hemiptera Heteroptera in the collection of
the British Museum. Brit. Mus. Publ. Pt. 1:1-240.

Westwood, J. O. 1837. A catalogue of Hemiptera in the collection of the Rev. W. F. Hope.
J. C. Bridgewater, London, 46 pp.

Received November 30, 1982; accepted August 29, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 121-124

PARATHROSCINUS, A NEW GENUS
OF BEETLES FROM SOUTHEAST ASIA
(LIMNICHIDAE: CEPHALOBYRRHINAE)

David P. Wooldridge

Department of Biology, The Pennsylvania State University, Ogontz Campus,
1600 Woodland Road, Abington, Pennsylvania 19001

Abstract. — The genus Parathroscinus is characterized. A new species, P. orientalis, is described
from the Philippine Islands and P. oculatus (Motschulsky) is transferred to the genus from
Byrrhinus.

The limnichid subfamily Cephalobyrrhinae (Champion, 1925) has heretofore con-
tained only the genus Cephalobyrrhus Pic (1923) from the Old World and Throscinus
LeConte (1874) from the New World. While attempting to determine a series of
limnichids from the Philippine Islands I discovered a few beetles that obviously
belonged to the subfamily but which agreed with neither genus. Recently, while
examining types of limnichids described by Motschulsky from India, I discovered
that the specimens he had described as Byrrhinus oculatus are actually not Byrrhinus
but belong to the Cephalobyrrhinae and are closely allied to the new Philippine
species. This paper describes the new taxa and transfers Motschulsky’s species to the
new genus.

Parathroscinus, new genus

Type-species. Parathroscinus orientalis Wooldridge, here designated.

General. Color black except legs red-brown, antennae and palpi dark red-brown.
Body elongate oval, shallowly convex. Overall vestiture short, dense, even, recum-
bent, with scattered erect longer hairs except on legs. Punctation variable from fine
to impressed, sometimes forming indistinct rows on elytra.

Head. Vertical, broad; eyes large, prominent, visible from above, separated on
vertex by more than their diameters; antennae inserted in shallow depressions on
front between eyes, segments 6 to 1 1 forming a slightly flattened, gradually enlarged
club.

Thorax. Pronotum narrowing from base, sinuate at base; disk evenly, strongly
convex, no sinuous depression; sides margined, margins continuous in outline with
elytral margins. Scutellum of moderate size. Prostemal process broad and flat. Meta-
coxae contiguous, transverse, flattened to about middle, grooved for reception of
femora.

Elytra. Narrowing gradually from humeri, attenuate toward rear; humeri promi-
nent; sides margined; epipleural folds continuous to apex; faint longitudinal grooves
from base onto disk of each elytron; elytral apices slightly projecting.

Abdomen. Closely punctate; first visible sternum with a low, curved, longitudinal
ridge on each side from the inner metacoxal margin to the posterior margin of the

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Figs. 1-4. Dorsal views showing relative shapes. 1. Cephalobyrrhus sp. 2. Throscinus sp.
3. Parathroscinus orient alis. 4. P. oculatus. Line equals 1 mm.

sternum; fifth visible sternum notched at tip, with a short longitudinal depression
extending forward from middle of notch.

This genus can be separated from Cephalobyrrhus by the form of the pronotum.
Cephalobyrrhus (Fig. 1) has a sinuous, depressed, more or less transverse groove near
the base of the pronotum; there is no sign of the groove in Parathroscinus. From
Throscinus, Parathroscinus differs in having the antennae set in shallow depressions
on the sides of the front (Fig. 5) rather than in deep pit-like excavations as in
Throscinus (Fig. 6). Throscinus also appears parallel-sided (Fig. 2) while Parathrosci-
nus narrows from about the humeral angles (Figs. 3, 4).

Parathroscinus orientalis, new species
Figs. 3, 7

Holotype. 8, Philippine Islands, Calian, Davao Prov., Mindanao, VII. 23. 30. C. F.
Clagg (FMNH).

Allotype. 2, same data as holotype.

Paratypes. 25 specimens, same data as holotype (22 FMNH, 3 in author’s collec-
tion); Philippine Is., 1910-1912, 3 spec. Acc. No. 8643, 2 spec. Acc. No. 8457, Lot
Bu. of Sci., P.I. (BMNH).

Diagnosis. This species has the bases of the elytra and pronotum less acutely sinuate
than oculatus (Fig. 3).

Description of holotype. Length 2.3 mm; greatest width 1.3 mm; elongate oval,
convex. Black. Overall vestiture of short variegated silver and yellow hairs and longer
gray hairs. Head with small, even punctation; eyes prominent, oval; antennae with
first segment small and short, 2nd and 3rd larger and longer, 2 > 3; 4 and 5 smaller
in diameter, 4 < 5; 6 to 1 1 gradually enlarged, slightly flattened, 11 > 10. Pronotum
with punctation somewhat impressed at sides, not impressed on disk; base shallowly
bisinuate. Elytra with regular, impressed punctation, punctures separated by 2-3 x

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NEW GENUS OF LIMNICHIDAE

123

Figs. 5-7. 5, 6. Frontal view of right side of head showing antennal insertions. 5. Para-

throscinus. 6. Throscinus. 7. Parathroscinus orientalis, male genitalia: a, dorsal view; b, para-
mere, lateral view. Overall length 0.45 mm.

their diameters; humeri prominent; each elytron with a small, broad, flat tooth at
apex. Ventral punctation minute, even. Prostemal process broad, hatchet-shaped,
margined at sides. Epipleura with a row of punctures on inner margin.

Aedeagus with parameres evenly converging to the rounded tips; penis broad and
flat, narrowing to an acute tip; basal piece about as long as parameres (Fig. 7).

Variation. Except for length (2.25-2.75 mm) there is little variation. In the female
the tooth at the tip of each elytron is broader than in the male. A single specimen
from Quan Nam Prov., Viet Nam, in the U.S. National Museum probably belongs
to this species but it differs somewhat in its punctation and it has not been included
as a paratype.

Parathroscinus oculatus (Motschulsky), New Combination
Fig. 4

Byrrhinus oculatus Motschulsky, 1858, p. 52. Ind. or. Lectotype here designated.

Type-material. There are three specimens of Motschulsky’s type-series mounted
on a single card in the Moscow Lomonosov State University Museum. These have
been numbered by me and the following labels are on the pin. 1. A small yellow
disk. 2. Handwritten on yellow: Byrrhinus/oculatus/Motsch./Ind. or. 3. On blue:
Spec. #1 Lectotype/Byrrhinus/oculatus/Motsch. 1858/by D. Wooldridge 1983. 4. On
blue: Lectoparatype/Byrrhinus/oculatus/Motsch. 1858/by D. Wooldridge/Spec. #2+3.
5. ?arathroscinus/oculatus/(Motsch.)/Det. D.P. Wooldridge.

Another specimen with only a yellow disk is among the types of Limnichus ori-
entalis Motsch. and has been labelled as P. occulatus by me.

Diagnosis. This species can be separated from orientalis by the acutely bisinuate
bases of the pronotum and elytra (Fig. 4). Because of this sinuation oculatus super-
ficially resembles a Byrrhinus. Motschulsky noted in his original description that it
should probably be in a new genus.

Description of lectotype. Length 2.4 mm, width 1.2 mm; elongate oval, convex.
Eyes large and prominent. Pronotum with fine, shallow punctation; base deeply and
acutely bisinuate on either side of scutellum. Elytra with shallowly impressed punc-
tation, punctures separated by 2-3 x their own diameters, punctures regular, forming

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vague longitudinal rows; bases bisinuate; strongly attenuate, tapering from humeri
to apex, humeri prominent. Prostemal process broad, margined at sides. Metaster-
num, metacoxal plates, and abdomen finely, closely, evenly punctate.

ACKNOWLEDGMENTS

I would like to thank Drs. Harry G. Nelson and J. Kethley, Field Museum of Natural History,
Chicago, 111. (FMNH) and Mr. R. D. Pope and Miss C. M. F. von Hayek, British Museum
(Natural History), London (BMNH) for the loan of specimens, and Mme. S. I. Kelevinikova,
Zoological Museum of the Moscow Lomonosov State University, USSR, for the loan of Mot-
schulsky’s types. The abbreviations are those used to designate the distribution of type material.

LITERATURE CITED

Champion, G. C. 1925. Some Indian (and Tibetan) Coleoptera (17). Ent. Mon. Mag. 61:169-
181.

LeConte, J. L. 1874. Descriptions of new Coleoptera chiefly from the Pacific Slope of North
America. Trans. American Ent. Soc. 5:43-72.

Motschusky, V. von. 1858. Insectes des Indes orientales. Etudes Entomologique 7:20-122.
Pic, M. 1923. Nouveautes diverses. Melanges Exotico-entomologiques 40:1-32.

Received November 9, 1983; accepted January 23, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 125-130

FLEAS (SIPHONAPTERA) FROM NESTS OF
WOODPECKERS IN ALASKA

Glenn E. Haas and Nixon Wilson

557 California Street, No. 7, Boulder City, Nevada 89005 and
Department of Biology, University of Northern Iowa, Cedar Falls, Iowa 50614

Abstract.— A total of 955 specimens came from 39 nests in 34 localities west of the Yukon
Territory, 1974-1978. Two Holarctic species were Ornithophaga anomala Mikulin with 371
specimens from 21 nests (mostly Downy Woodpecker and Three-toed Woodpecker) in 20
localities and Ceratophyllus zhovtyi Emel’yanova and Goncharov with 28 specimens from three
nests (two Flicker and an unknown species) in three localities. Two Nearctic species were C.
adustus Jordan with 168 specimens from 15 nests (mostly Three-toed Woodpecker) in 13
localities and C. rauschi Holland with 388 specimens from eight nests (seven Flicker) in six
localities.

Our survey of ectoparasites in nests of the Tree Swallow ( Trachycineta bicolor
(Vieillot)) and Violet-green Swallow (Tachycineta thalassina (Swainson)) yielded data
on fleas of swallows (Haas et al., 1981). The swallows sometimes built nests in tree
cavities made by woodpeckers. Thus, four species of little-known bird fleas were
discovered in old nests of woodpeckers, namely: Ornithophaga anomala Mikulin,
Ceratophyllus zhovtyi EmeTyanova and Goncharov, C. adustus Jordan and C.
rauschi Holland. The first two species were originally described from southcentral
USSR and the last two from northwestern Canada not far from the Alaska border.
Hopkins and Rothschild (1971) thought that O. anomala might be represented in
North America by the unique specimen of O. nearctica Holland and Loshbaugh from
Utah. Therefore, in Alaska, only the finding of C. zhovtyi could be considered un-
expected.

Some morphological anomalies in O. anomala and C. adustus were noted and
described (Haas, 1983). Our survey of fleas on birds found dead included one record
of C. adustus from Picoides tridactylus (Linnaeus) in southcentral Alaska (Haas et
al., 1980). The original description of C. adustus from a porcupine ( Erethizon dor-
satum (Linnaeus)) in British Columbia (Jordan, 1932) had obscured its relationship
with woodpeckers. The present report concerns records of 955 specimens of four
species of fleas from 39 nests of Picoides spp. and Colaptes auratus (Linnaeus) in 34
localities of Alaska west of the Yukon Territory, 1974-1978.

MATERIALS AND METHODS

In forested regions of Alaska west of the Yukon Territory, especially the interior
Copper River lowland and lowlands near upper Cook Inlet, dead stubs of spruce
(Picea glauca (Moench) Voss), poplar and cottonwood ( Populus spp.), birch ( Betula
papyrifera Marshall) and other trees were scanned from a distance for woodpecker
cavities. Stubs with cavities not being used by nesting birds were cut down or pushed
over to collect any nests present (Haas et al., 1981). Stubs that fell intact were cut

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open to remove nests with scraper, brush and forceps. The same tools were used for
retrieving nests that spilled from cavities when stubs broke open on impact with the
ground. Nests of woodpeckers, swallows (Haas et al., 1981), squirrels (Haas and
Wilson, 1982) and voles (Haas, 1982) were bagged separately. Nests of Colaptes
auratus were identified by the large cavity and presence of ant heads and flicker
feathers. Nests of Picoides spp. that contained small red-tipped feathers were classified
as P. villosus (Linnaeus) or P. pubescens (Linnaeus), not P. arcticus (Swainson) or P.
tridactylus. Nests of Picoides that were not of the last two species are indicated by
(NT) in the Records.

Five nests were held in plastic bags with moist towels for several weeks to rear
immature fleas. All nests of woodpeckers, except old ones that had become com-
pressed into a crust, were picked apart in a light-colored pan, and fleas were transferred
with brush or forceps to labelled vials. Specimens selected for permanent mounts in
Canada balsam were mostly those found in good condition, but all specimens of C.
zhovtyi were mounted despite severe structural damage. Specimens of C. rauschi
were deposited in the Canadian National Collection and collections of the authors;
specimens of the other species were retained by Haas.

RESULTS

Specimens totaling 955 (102 reared) of four species of fleas were obtained from
39 nests of woodpeckers ( Colaptes auratus and Picoides spp., mostly P. pubescens
and P. tridactylus ) in 34 localities in Alaska west of the Yukon Territory, 1974-1978.
Neither fleas nor fragments of them could be found in very old nests, and 14 cavities
without nests were negative for these species of fleas.

Ornithophaga anomala Mikulin, 1957

Specimens totaling 371 (15 reared) were obtained from 21 nests of woodpeckers
in 20 localities in two regions. All nests were of unidentified Picoides spp.; five
contained small red-tipped feathers. One nest appeared to be in an abandoned cavity
of C. auratus. The nests were in cavities an average of 3 m above ground. All cavities
were in dead stubs: 1 1 in birch, six in poplar and four in spruce.

Records. All from nests of Picoides spp. Chickaloon (mile 78 Glenn Highway):
two males, nine females (eight dead, one gravid), poplar, 8.IV.1976. Eklutna Lake,
3 km SE, 320 m: two males (dead), two females (also C. adustus ), poplar (NT),
18.X. 1974. Glennallen, 53 km W (mile 4.5 Lake Louise Road): three males (dead),
two females (dead) (also C. adustus ), spruce (probable cavity of C. auratus ), 1 4. VI. 1976.
Kasilof, 6.4 km S (Tustumena Lake Road): two females (one dead), under nest of
swallow, poplar, 31. VII. 1976. Kenai (city): one, sex unknown (left metepimeron
only), birch, 23. IV. 1976. Kenai Lake, west shore (mile 6 Snug Harbor Road): one
male, three females, birch, 29.VII.1976; (mile 7): two females (dead), under squirrel
nest, birch, same date. Palmer, 4.5 km NE (Clark- Wolverine Road): 23 males (eight
dead), 39 females (21 dead) (also C. adustus ), poplar, 13.IX.1975; 7 km E (Smith
Road): five males (one dead), 10 females (one dead, one gravid), poplar, 13.IX.1975;
7 km N (mile 52.8 Glenn Highway): one female (dead), under squirrel nest and vole
nest, poplar (NT), 30. IV. 1975; 8 km N (mile 53 Glenn Highway): four males, three
females (emerged from cocoons when disturbed), under nest of swallow, birch,

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127

3.VIII.1974; 14 km NW (mile 1.5 Edgerton Park Road): 68 males, 87 females (1
dead), birch (NT), 27. IX. 1975; 18.2 km SE (Knik River Road nr. Fox Lake): one
male (dead), nine females (dead) (also C. adustus ), spruce, 28.VII.1974; one female
(dead), same spruce, cavities interconnected. Sterling, 7.2 km NW (Sunken Island
Lake Road): 29 males, 39 females (one dead) (also C. adustus), birch, 30.VII.1976
(two males, six females reared 25. VIII. 1976); 22.4 km N (Swanson River Road): one
female (dead), under squirrel nest, spruce, 30.VII.1976. Talkeetna, 12.8 km NW
(mile 121.6 Parks Highway): one male, seven females (one dead), birch (NT), 1 3.V. 1 976
(one male, six females reared 8. VII. 1976); 24.1 km W (mile 12.2 Petersville Road):
two males, two females, birch, 28.IX.1975. Wasilla, 5 km E (Parks Highway): one
male, seven females (one dead), birch, 8. V. 1975; 5.5 km E (Parks Highway nr. Black
Lake): one female, birch (NT), 6.VIII.1974; 6 km NE (Lakeview Road nr. Kings
Lake): one female (dead) (also C. adustus), birch, 6. V. 1975.

Ceratophyllus zhovtyi Emel’yanova and Goncharov, 1966

Specimens totaling 28 were collected from three nests of woodpeckers in three
localities in two regions. Two nests were of C. auratus and one was of an unidentified
Picoides sp. that contained small red-tipped feathers. The nests were in cavities an
average of 3 m above ground. All cavities were in dead stubs: two in birch and one
in spruce.

Records. Chistochina, 22.5 km SW (mile 222.5 Glenn Highway): 15 males (dead),
1 1 females (eight dead) (also C. adustus and C. rauschi), nest of Picoides sp. (NT),
birch, 9. IV. 1976. Circle, 14 km SW (mile 151.3 Steese Highway): one female, nest
of C. auratus, birch, 1. VII. 1976. Nabesna Road, mile 31: one male (dead), nest of
C. auratus, spruce, 9. VIII. 1975.

Ceratophyllus adustus Jordan, 1932

Specimens totaling 168 (30 reared) were obtained from 15 nests of woodpeckers
in 1 3 localities in three regions. One nest was of C. auratus and 1 4 were of unidentified
Picoides spp. Two of the latter contained small red-tipped feathers. The nests were
in cavities an average of 2.5 m above ground. All cavities were in dead stubs: eight
in birch, four in spruce and three in poplar.

Records. All from nests of Picoides spp., except as indicated. Cantwell, 4.8 km NE
(Parks Highway south of Nenana River bridge): 29 males, 3 1 females, spruce, 3. IX. 1 974
(one male reared 10.X. 1974). Chistochina, 22.5 km SW (mile 222.5 Glenn Highway):
seven males (dead), 1 3 females (dead) (also C. zhovtyi and C. rauschi), birch (NT),
9. VI. 1976. Edgerton Highway, mile 10: one male (dead), poplar, 5. IX. 1975. Eklutna
Lake, 3 km SE, 320 m: six females (also O. anomala), poplar (NT), 18.X.1974.
Glennallen, 53 km W (mile 4.5 Lake Louise Road): 1 1 males (three dead), 17 females
(also O. anomala), spruce (probable cavity of C. auratus), 14.VI.1976. Gunsight
Mountain, 3 km SE (mile 119 Glenn Highway): two females, nest of C. auratus
apparently visited by Picoides sp., spruce, 8.VI.1976. Palmer, 4.5 km NE (Clark-
Wolverine Road): one male, three females (also O. anomala), poplar, 13.IX.1975
(one female reared 21.X.1975); 18.2 km SE (Knik River Road nr. Fox Lake): one
male (dead), one female (dead) (also O. anomala), spruce, 28.VII.1974. Skilak Lake,
north shore: three males (dead), four females (dead), under incomplete nest of swal-

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low, birch, 23. V. 1978. Sterling, 7.2 km NW (Sunken Island Lake Road): nine males
(one dead), 20 females (also O. anomala), birch, 30. VII. 1976 (eight males, 20 females
reared 25. VIII. 1976). Talkeetna, 2.9 km SE: one male, birch, 21. IV. 75. Wasilla, 4.6
km E (Matanuska Road): one male, three females (one dead), birch, 30. IV. 1975; one
female, cavity higher in same birch; 6 km NE (Lakeview Road nr. Kings Lake): one
male (dead), one female (also O. anomala), birch, 6. V. 1975; one female (gravid),
cavity higher in same birch.

Ceratophyllus rauschi Holland, 1960

Specimens totaling 388 (57 reared) were obtained from eight nests of woodpeckers
in six localities in two regions. Seven nests of C. auratus harbored 387 of the spec-
imens. These nests were in cavities an average of 4.6 m above ground. All cavities
were in dead stubs: four in spruce, two in birch and one in poplar. The last stub was
part of a live tree.

Records. All from nests of C. auratus except as indicated. Chistochina, 22.5 km
SW (mile 222.5 Glenn Highway): one male (dead) (also C. zhovtyi and C. adustus ),
nest of Picoides sp. (NT), birch, 9.IV.1976. Edgerton Highway, mile 7.5 (nr. Kenny
Lake): 58 males, 106 females, poplar, 11. VIII. 1974. Fairbanks, 35 km WSW (mile
336 Parks Highway): 1 5 males (one dead), 30 females (seven dead), birch, 29. VI. 1 976
(14 males and 23 females reared 9.VII-12.VIII.1976). Glennallen, 51 km W (mile
3 Lake Louise Road): four males, four females, spruce, 8.VI.1976; 47 males, 76
females, spruce, 27.IX.1976 (six males, 14 females reared 6.X-19.X. 1976). Nabesna
Road, mile 24.5: nine males, 34 females (one dead), spruce, 2. IX. 1975; one female,
cavity higher in same spruce. Nabesna Road-Glenn Highway Junction, 4.3 km W:
three females (dead), under squirrel nest, birch, 16.VIII. 1975.

DISCUSSION

The Northern Flicker and the Hairy, Downy, Black-backed and Three-toed Wood-
peckers are the five species of the family Picidae that breed in Alaska west of the
Yukon Territory (Gabrielson and Lincoln, 1959). The Hairy Woodpecker was con-
sidered as not abundant and the Downy and Black-backed Woodpeckers as uncom-
mon, but the Three-toed Woodpecker was referred to as one of the most common
yet as one that cannot be called abundant. Gabrielson and Lincoln (1959) could not
cite many records of nests of the five species in western Alaska. Recent data of the
Anchorage Audubon Society showed the status of the Black-backed Woodpecker as
rare and the other four species as uncommon in a well-studied area of the Cook Inlet
lowland (Klein et al., 1978). Only one nest of the Northern Flicker was collected in
the Cook Inlet region and it was uninfested. In contrast, nests of Picoides spp. were
easy to find in dead stubs in suitable habitat in the region. But collecting from only
dead stubs probably caused the nests of the Black-backed Three-toed and Hairy
Woodpeckers to be underrepresented in our data as both also nest in live trees and
the latter prefers them (Scott et al., 1977).

Ornithophaga anomala. The history of this Holarctic bird flea was reviewed by
Hopkins and Rothschild (1971). The original description was from one female from
P. tridactylus in Alma Ata Oblast, USSR, and Goncharov et al. (1966) provided

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129

supplementary descriptions that Hopkins and Rothschild (1971) thought were based
on two different species. The supplementary specimens consisted of seven males and
nine females from the owl Glaucidium passerinum Linnaeus from Irkutsk Oblast,
USSR, and one male and one female from Dendrocopos ( =Picoides ) leucotos Bechst
from Primor’ye Kray, USSR. Goncharov et al. (1966) thought that the latter two
specimens might represent a subspecies of the Alma Ata and Irkutsk species. Hopkins
and Rothschild (1971) thought that O. nearctica Holland and Loshbaugh from Den-
drocopos ( =Picoides ) pubescens leucurus (Hartlaub) in Utah (Holland and Loshbaugh,
1958) might represent a subspecies of O. anomala.

In Alaska, O. anomala was especially common in nests of Picoides spp. in the
upper Cook Inlet region, with 366 specimens infesting 20 nests. The only infested
nest (five dead fleas) not from this region was in the Copper River lowland about 53
km west of Glennallen. This flea might be more dependent than C. adustus on Hairy
or Downy Woodpeckers. The largest population (155 specimens) infested a nest that
was probably of the Downy Woodpecker.

Ceratophyllus zhovtyi. This bird flea was originally described from four specimens
from the owl G. passerinum and seven specimens from two woodpeckers in the
Irkutsk Oblast of the Soviet Union (Emel’yanova and Goncharov, 1966). One of the
woodpeckers was identified as a large motley-colored woodpecker. We presume it
was Picoides major (Linnaeus). Neither it nor G. passerinum occurs in North Amer-
ica. Two of our specimens were found alone in nests of Northern Flickers, but 26
specimens were present in a mixed population (C. adustus and C. rauschi) in a nest
of a Hairy or Downy Woodpecker. This probably means that breeding can occur in
nests of Picoides spp. More data are needed on host and nest relationships and
geographic distribution.

Ceratophyllus adustus. This northern Nearctic bird flea seemed to have a slight
preference for nests of P. tridactylus. Only two infested nests (20 and six flea speci-
mens) could be classified as not being of either species of three-toed woodpecker,
and the largest population (60 specimens) infested a nest that most likely was of P.
tridactylus. Six of the 1 5 nests infested by C. adustus were infested by O. anomala.
The highest proportion of these records, i.e., mixed populations in five of 10 nests,
was from the Cook Inlet region.

Ceratophyllus rauschi. This northern Nearctic flea of the Northern Flicker was
originally described from the nest of Colaptes sp. in the Yukon Territory (Holland,
1960). In Alaska it was found almost exclusively in nests of the Northern Flicker.
The only exception was the dead male in a nest of a Hairy or Downy Woodpecker.
Thus, all breeding populations were in nests of Northern Flickers. Five localities
were in the interior Copper River lowland and one was in the Yukon River watershed
(WSW of Fairbanks) about 358 km west of the US-Canada border. The type locality
(Stewart Crossing) is about 215 km east of the border on a tributary of the Yukon
River. In Alaska, C. rauschi probably infests nests of the Northern Flicker to the
northern and western limits of breeding by the bird.

ACKNOWLEDGMENTS

We thank F. G. A. M. Smit, former Custodian of the Rothschild Collection of Siphonaptera,
for confirming the identification of C. adustus and allowing an examination of specimens of

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

O. anomala, and Dr. G. P. Holland, Agriculture Canada, for confirming the identification of

C. rauschi and providing copies and translations of the descriptions of O. anomala and C.

zhovtyi and copy of the description of C. adustus.

LITERATURE CITED

Emel’yanova, N. D. and A. I. Goncharov. 1966. A new species of flea, Ceratophyllus zhovtyi
n. sp., from eastern Siberia (Suctoria, Ceratophyllidae). Izvest. Irkutsk. Protivochum.
Inst. 26:309-313 (tr. from Russian seen).

Gabrielson, I. N. and F. C. Lincoln. 1959. The birds of Alaska. Stackpole Co., Harrisburg,
Pennsylvania, and Wildlife Management Institute, Washington, D.C.

Goncharov, A. I., V. P. Gordeyeva and V. C. Sonin. 1966. Description of the male and a
supplementary description of the female of Ornithophaga anomala Mikulin, 1956 (Suc-
toria). Izvest. Irkutsk. Protivochum. Inst. 26:314-321 (tr. from Russian seen).

Haas, G. E. 1982. Fleas (Siphonaptera) from vole nests in subarctic Alaska. Can. J. Zool. 60:
2157-2161.

Haas, G. E. 1983. Anomalies of the head, thorax and abdomen in the order Siphonaptera.
Adv. Biosci. (in press).

Haas, G. E., T. Rumfelt, R. E. Barrett and N. Wilson. 1980. Reas from some Alaskan birds
(Siphonaptera). Pan-Pac. Ent. 56:105-106.

Haas, G. E., T. Rumfelt and N. Wilson. 1981. Reas (Siphonaptera) from nests of the tree
swallow ( Iridoprocne bicolor ) and the violet-green swallow ( Tachycineta thalassina ) in
Alaska. Wasmann J. Biol. 39:37-41.

Haas, G. E. and N. Wilson. 1982. Reas (Siphonaptera) from nests of the red squirrel ( Tami -
asciurus hudsonicus) and burrows of the arctic ground squirrel ( Spermophilus parryii) in
Alaska. Wasmann J. Biol. 40:59-65.

Holland, G. P. 1 960. Descriptions of two species of Ceratophyllus Curtis from Yukon Territory
(Siphonaptera: Ceratophyllidae). Can. Ent. 92:787-793.

Holland, G. P. and G. Loshbaugh, Jr. 1958. Two new species of fleas from Utah, with notes
on the genus Ornithophaga Mikulin (Siphonaptera). Can. Ent. 90:486-493.

Hopkins, G. H. E. and M. Rothschild. 1971. An illustrated catalogue of the Rothschild
Collection of fleas (Siphonaptera) in the British Museum (Natural History). Vol. V. British
Museum (Natural History), London.

Jordan, K. 1932. Siphonaptera collected by Mr. Harry S. Swarth at Atlin in British Columbia.
Novit. Zool. 38:253-255.

Klein, J., J. Pitcher, S. Scott and G. Tans. 1978. Birds of Anchorage, Alaska (Eagle River to
Portage Glacier). A Checklist. Anchorage Audubon Society, Inc., 8 pp.

Scott, V. E., K. E. Evans, D. R. Patton and C. P. Stone. 1977. Cavity-nesting birds of North
American forests. U.S. Dep. Agric., Agric. Handb. 511, 1 12 pp.

Received August 12, 1983; accepted November 2, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 131-149

SYNOPSIS OF OMMATIUS WIEDEMANN
(DIPTERA: ASILIDAE) FROM HISPANIOLA

A. G. Scarbrough

Department of Biological Sciences, Towson State University,

Baltimore, Maryland 21204

Abstract.— Eight species of the genus Ommatius Wiedemann are reported from Hispaniola
in the Caribbean Islands. Seven of the species are described as new ( O . haitiensis, O. nigellus,

O. russelli, O. gwenae, O. stramineus, O. cinnamomeus, O. hispaniolae ), O. vitreus Bigot is
redescribed, and illustrations and a key are included.

The genus Ommatius Wiedemann is apparently more widespread among islands
in the Caribbean than previously reported. Although the genus is poorly represented
in collections, nine species have been reported and most appear to be endemic to a
single island or in island group (Martin and Papavero, 1970; Scarbrough, 1983). For
example, 2 species are found in the Bahamas, 1 on Haiti, 4 on Jamaica, and 2 on
Puerto Rico and the Virgin Islands. Of these, only O. marginellus (Fabricius), which
is reported from Cuba southward to Brazil, appears to have a wide distributional
overlap with the other species. As more specimens accumulate in collections from
the islands and as taxonomic problems are resolved, so too should information on
species variation and distribution become more defined. This paper, one of several
that will appear on the Caribbean Ommatius, deals with the species found on His-
paniola (Haiti and the Dominican Republic).

Specimens for study were received on loan from the following institutions: Museum
of Comparative Zoology (MCZ), Cambridge; American Museum of Natural History
(AMNH), New York; United States National Museum (USNM), Smithsonian In-
stitution, Washington, D.C.; Canadian National Collection (CNC), Ottawa, Canada;
Institute of Jamaica (IJ), Kingston; and Hope Department of Entomology (OX),
Oxford University, Oxford, England.

The characters used in this study are well-known, traditional ones and are identified
by reference to Figures 1-3. I followed the nomenclature used by McAlpine (1981).
Unless otherwise stated, the paragraphs entitled “male” and “female” refer to the
description of a specimen as a holotype or an allotype. Those entitled “variation”
include only major characters of specimens in the type series that differ significantly
from those of the type specimen(s).

KEY TO THE SPECIES OF OmmatiUS FROM HISPANIOLA

1. Upper postocular bristles long, strongly proclinate and extending far forward above

eye (Fig. 1) 2

- Upper postocular bristles short, curved forward slightly only near their apices (Fig. 4)

5

2. Femora wholly black 3

- Femora with some reddish to yellowish coloration at bases 4

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3. Facial bristles wholly black; 2 or 3 strong marginal scutellar bristles, 1 or 2 anepimeral

bristles O. haitiensis, new species

- Facial bristles black and pale yellow; scutellar and anepimeral bristles absent

O. nigellus, new species

4. Most or all femoral bases with a small reddish spot or narrow band; epandrium slender,

somewhat narrowed apically (Fig. 11) O. russelli, new species

- All femora reddish to orangish on basal fourth or more; epandrium broad basally,

abruptly narrowed apically (Fig. 9) O. gwenae, new species

5. Meso- and metafemora wholly black or dark brown 6

- Meso- and metafemora yellowish to orangish on basal third or more 7

6. Facial bristles wholly pale (white or yellow), costal margin with slight bulge in males;
abdominal pollen yellow; epandrium with single, broad apical process (Fig. 13) ....

O. stramineus, new species

- Several dark facial bristles above mystax; abdominal pollen yellowish gray; costal bulge
absent; epandrium with two, slender, apical processes (Fig. 5); female with apical comers

of tergite 9 projecting posteriorly (Fig. 6) O. vitreus Bigot

7. Large species (20 mm); fore femur with apical third black; abdomen not noticeably

enlarged apically; stemite 8 with a moderately deep median apical notch and 2 lateral
protuberances (Fig. 14) O. cinnamomeus, new species

- Small species (11-14 mm); fore femur entirely black or nearly so; abdomen somewhat

enlarged apically; epandrium slender along its entire length (Fig. 7); stemite 8 without
a deep median apical notch, lateral protuberances widely spaced (Fig. 8)

O. hispaniolae, new species

Ommatius haitiensis, new species
Figs. 1-3

Description. MALE (Fig. 1): 13.5 mm. Head black; face and occiput gray tomentose
with traces of yellow along eye margin; tomentum of front generally brown, coppery
brown at some angles. Bristles and hairs of face, front, antenna and upper half of
occiput black, lower half (except 5-6 hairs below eyes) and base of proboscis with
long white pile; facial pile and bristles long and slender, 4 stout black bristles on
lower half, pile dense above and almost reaching base of antenna; several stiff ocellar
hairs. Occipital bristles thin, straight on sides, strongly proclinate near vertex and
long (ca. 3-4 times longer than those on sides), extending forward over eyes. Third
antennal segment three times longer than width and as long as first two segments
combined; style about one-third longer than the 3 segments combined.

Thorax black. Thoracic dorsum generally brownish pollinose; sides, grooves, pre-
scutellum and scutellum of scutum and pleura with yellowish gray to yellowish brown
pollen. Chaetotaxy of scutum black and abundant; hairs short anteriorly, much longer
and stronger posteriorly; bristles consist of 2 notopleurals, 2-3 supra-alars, 2 postalars
and a row of strong dorsocentrals which extend entire length of dorsum; scutellum
with abundant, slender, black vestiture covering entire dorsal surface and 2 strong
marginal bristles. Pleural pile abundant, long, strongest and primarily black on meta-
epistemum and laterotergite, sparse on anepimeron; meron bare. Row of pleural
bristles almost entirely black, 1-2 pale; 1-2 black anepimeron bristles.

Wing hyaline. Veins brown, lighter basally; costal margin without a bulge; crossvein
r-m at apical third of discal cell. Halter brownish, knob darkest.

Legs black. Coxae with pale yellowish gray pollen and mostly whitish bristly hairs,

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OMMATIUS FROM HISPANIOLA

133

Figs. 1-3. Ommatius haitiensis, head and terminalia. 1. Head, lateral view; Po = postocular
bristles. 2. Male terminalia, dorsal (a), lateral (b) and apical (c) views. 3. Female terminalia,
lateral (a) view and stemite 8 (b). Ce = cercus, Ep = epandrium, Gs = gonostylus, Gc = gono-
coxite, Hy = hypandrium, Vp = ventral plates, T = tergites, S = stemites.

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a few black hairs on fore and middle coxae. Femora moderately swollen and covered
with appressed yellowish or whitish pile, dense dorsally and posteriorly, shorter and
black apically; abundant, long, bristly hair below and apically (4-6), stronger hairs
and setigerous bristles black except for a few whitish ones basally on middle and
posteroventrally on fore femora. Tibiae with pale pile and long black bristly hairs
and bristles. Tarsi black with black bristles.

Abdomen dark brown to black, apical borders of segments brownish. Traces of
brownish pollen dorsally on most tergites, yellowish gray or gray pollen on sides and
on stemites; tergites 7-8 somewhat shiny. Long whitish pile on stemites and sides
of tergites, abundant and longest on first 5 segments. Abdominal bristles and ap-
pressed setae black, noticeably long and strong on tergites 1 and 7-8; tergite 2 with
a midlateral patch of long black, bristly hairs.

Terminalia (Figs. 2a-c) brownish with cercus and gonostylus orangish brown.
Cercus with pale weak hairs apically and stronger blackish ones laterally; black bristly
hairs elsewhere on terminalia; hairs longest near apex of epandrium and base of
gonocoxite. Gonostylus elongate with a flat preapical dorsal process and numerous
short, thick black setae and long hairs basally. Epandrium dark and somewhat inflated
basally, lighter and tapered apically. Hypandrium with rounded median point and
scattered black hairs. Ventral plates below cercus with pale hair apically and stronger
black bristly hairs basally.

FEMALE (Figs. 3a, b): 13.3 mm. The female differs from the male holotype as
follows: darker yellowish gray facial tomentum; scutellum with some pale pile in
addition to black pile. Femora slender, hind femur with pale yellow hairs and bristles
ventrally in addition to black. Abdomen slightly lighter brown than male, pollen
more yellow on sides of tergites; bristles and hairs of segments 1-6 pale yellow to
yellow; segment 7 with several long black hairs along apical margin. Tergite 9 hidden
by tergite 8, each apical corner wrapped around base of cercus, covering apical comer
of stemite below. Stemite 8 shiny black with several transverse wrinkles and a few
fine hairs basally; stemite paler apically with a sharp median pointed process. Cercus
shiny and with abundant fine yellowish pile.

Holotype. 6, Haiti, LaViste and Vie LaSalle Range, 5,000-7,000 ft., 16-23 Sept.
1 934 (M. Bates). Allotype 9 and paratype 6, same data as holotype. The type specimens
are deposited in the MCZ collection.

Variation. The paratype male (14.1 mm) differs from the holotype in that it has a
lighter (dark brown) ground color and fewer black bristles and hairs.

Etymology. The species is named for the country from which the species was
collected.

Diagnosis. This species is easily recognized by a black body and abundant black
vestiture; third antennal segment 3 times longer than wide, facial hairs wholly black
and abundant near base of antenna, several long black proclinate postocular bristles,
5-6 black hairs below eyes; 2 strong scutellar bristles, 1-2 anepimeral bristles, a patch
of long black pile on tergite 2; male with epandrium abruptly tapered apically and
an elongated gonostylus with a preapical dorsal process; female with a sharp median
pointed process on the apical margin of stemite 8.

Ommatius haitiensis is quite different from other Caribbean species. It resembles
O. nigellus, n. sp. which is also black but has black and light facial vestiture. Om-
matius haitiensis differs from the latter species by the presence of wholly black facial

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135

5 b

Figs. 4-6. Ommatius vitreus Bigot, head and terminalia. 4. Head, lateral view. 5. Male
terminalia, lateral (a), dorsal (b) and apical (c) views. 6. Female terminalia, lateral view (a) and
stemite 8 (b). Pg = process of gonocoxite, Ap = apical process of epandrium.

vestiture, black hairs below the eyes and on the upper half of the occiput, 2 marginal
scutellar and 1-2 anepimeral bristles.

Ommatius vitreus Bigot
Figs. 4-6

Ommatius vitreus Bigot, 1875:246; type locality Haiti, 2.

Ommatius marginellus: Hull, 1 962:435; listed O. vitreus as a synonym O. marginellus
(Fabricius).

Ommatius vitreus: Martin and Papavero, 1970:60; removed from synonymy.

Although Bigot (1875) indicated the existence of a female specimen in his descrip-
tion of O. vitreus, he neither designated a holotype nor indicated the number of
specimens in his possession. A female and a male of O. vitreus from Bigot’s collection
presently exist in the Hope Entomological Museum, Oxford, England. Both specimens

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bear a label with the specimen number (#779 in coll. Bigot) and a type number (either
2831/2 or 283 2/2). In order to clarify the confusion in this matter, I have selected
the best preserved specimen of these, a female with the type label #28 3 ¥2, as Lectotype.
The specimen is partially greased and lacks only the third antennal segment and
style. A description of the lectotype and the male follows.

Description. FEMALE (Figs. 4, 6a, b): 12.0 mm. Head brownish black; face yel-
lowish tomentose, front brown to yellowish brown. Mystax yellowish, 7 brownish
hairs above; 2 long brown ocellar bristles. Occiput somewhat whitish pollinose, pile
white with postocular bristles dark. Antenna brown with short dark bristles on seg-
ment 1 and 2; third segment with 1 or 2 short dorsal hairs, longer than wide, slightly
less than length of first and second combined; style less than twice length of all
segments combined.

Thorax with sparse pale pile, pile most abundant on prothorax, humeral callus,
disc of scutellum and on sides of scutum above wings. Chaetotaxy: 2 notopleurals,
1 supra-alar, 1 postalar, 4 dorsocentrals in prescutellar region; scutellar bristles absent
or at least not noticeably different from pile; row of pelural bristles yellowish. Thoracic
pollen mostly yellow to yellowish gray, brownish on postalar callus.

Legs. Coxae brown with yellowish gray pollen, pale bristles and pile. Femora
blackish brown, tibiae yellowish with brown apically; fore and middle tibiae with
narrow apical dark bands, hind tibia dark on apical half; tarsi brown with basal
tarsomere lightest, that of hind tarsus darkest. Pile of legs light yellow or whitish,
sparse posteriorly on fore and middle femora; some black pile dorsoapically on fore
and middle femora. Femoral bristles mostly pale yellow or whitish; fore femur with
black bristles and bristly hairs below and 2 short black hairs apically. Middle femur
with several strong bristles below on basal half, all light but 1; 4 additional black
bristles on anterior surface and 1 short black posteroapical bristle. Hind femur with
1 black bristle dorsoapically. Tibiae with appressed black setulae; tibial and tarsal
bristles black except for 1-2 on fore and middle tibiae and 2 on fore tarsus; middle
tibia with a ventral row of black hairs.

Wing hyaline, veins brown, lightest basally. Costal margin straight anteriorly, r-m
crossvein beyond middle of discal cell. Halter yellowish brown.

Abdomen dark brown to black, posterior borders somewhat lighter in color. Gray
to yellowish gray pollen and pale pile on sides of tergites and on sternites. Tergites
8 and 9 shiny black with long black hairs, apical corners of 9 extending some distance
posteriorly; cercus slightly lighter in color than tergites and with light pile; stemite
8 covered with meconium.

MALE (Figs. 5a-c): 1 1 .0 mm, differing from female as follows: dark bristles of
fore tibia only at apex, dark hairs of middle tibia absent. Wing with light brown in
anterior cells, r-m crossvein at or beyond middle of discal cell. Halter brownish.
Abdominal tergite 7 and 8 somewhat shiny with long, dark setae along apical margin,
slightly longer on sides.

Terminalia mostly dark brown. Hypandrium somewhat inflated, apically rounded.
Epandrium with basal two-thirds dark brown, greatly swollen and with dark setae;
base somewhat flat in apical view; apical third with two lighter colored, slender,
curved processes. Gonostylus somewhat oval in cross-section, yellowish brown. Cer-
cus brownish with fine pale pile; ventral plates with basolateral processes.

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OMMATIUS FROM HISPANIOLA

137

Lectotype. 2, #779, in coll. Bigot abt. 1845-93, Type 2831/2, Oxford Entomological
Museum.

Other specimens examined. HAITI: 8, #283 2/2, same data as lectotype; 8, Furcy,
July 25, 1950 (A. Curtiss); 288, 299, Kenskolf (nr. Port-au-Prince), 4,000-6,000 ft.,
August 8, 1934 (M. Bates); 299, Furcy, 4,000 ft., 10 Dec. 1956 (B. & B. Valentine).
DOMINICAN REPUBLIC: 2, Constanze, 3,000-4,000 ft., August 1938 (Darlington);
322, Sarabacoa, Nov. (?) 1950 (N. L. H. Drauss); 8, Constanze, 5 May 1959 (M. W.
Sanderson and T. H. Farr); 2, La Palma, 1.2 km E. El Rio, 2-13 June 1969 (Flint
and Gomez). Specimens are deposited in the Oxford Entomological Museum, USNM,
MCZ, AMNH, Ohio State University Museum (Columbus) and the collection of the
author.

The lectotype and the male in Bigot’s collection were examined.

Distribution. Hispaniola and Mona Island, Puerto Rico.

Variation. Only slight differences were noted in the series examined; 8 13.0-13.2
mm, 2 1 1.9-14.0 mm. The femora are invariable darker (black) and the facial to-
mentum is more white or gray than on the lectotype. The pleural and abdominal
pollinosity is sometimes slightly darker yellow than the lectotype. The dark apical
bands on the fore and middle tibiae are often absent or nearly so. In females, sternite
8 has abundant pale pile, a V-shaped row of stiff mostly black bristles and transverse
wrinkles which are often covered with sparse yellowish pollen; the apical third has
a low median ridge or line, a somewhat circular lighter colored (sometimes covered
with pollen or bare and shiny) depression to each side of the ridge, and 2 low, laterally
spaced, protuberances on the apical margin; middle of apical margin, with lateral
protuberances, sometimes noticably projecting.

Diagnosis. Ommatius vitreus is recognized by the wholly black femur, presence of
two ventral rows of whitish setigerous bristles on the hind femur, mystax light with
several dark hairs above, white to grayish facial tomentum, and an apically bifurcated
epandrium in males and the absence of a costal bulge and marginal scutellar bristles.

Ommatius vitreus will key to O. alexanderi Farr in Farr’s key (1965) but can be
recognized from that species by its wholly dark femora, apically bifurcated epandrium
and the absence of a costal bulge and a styliform gonostylus.

Ommatius hispaniolae, new species
Figs. 7, 8

Description. MALE (Figs. 7a-c): 11.1 mm. Head brown. Tomentum of face pale
yellowish white, yellow along eye margin. Front grayish or brownish gray tomentose
with 6 short, fine hairs and 2 long black ocellar bristles. Facial vestiture white with
4 stout bristles and longer bristly hairs. Occipital vestiture whitish, tomentum with
traces of yellow; postocular bristles straight or slightly curved near tips. Antenna
segments of equal length, style about twice length of all segments combined; segments
1 and 2 whitish tomentose, third segment somewhat oval, length slightly greater than
width and with 1 or 2 short hairs dorsally. Antennal bristles short, generally whitish
on segment 1 and brown on 2.

Thorax dark brown. Pollen of scutum and pleuron pale yellowish gray to gray. Pile
absent on scutum except on humeral callus and above wing base. Black scutal bristles

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Figs. 7, 8. Ommatius hispaniolae, terminalia. 7. Male, lateral (a) view, gonostylus (b) and
dorsal (c) view. 8. Female, lateral (a) view and stemite 8 (b).

consisting of 2 notopleurals, 1 supra-alars, 1 postalar and 6 slender dorsocentrals in
prescutellar region; several short hairs anteriorly. Scutellum with fine whitish pile,
none along margin conspicuously different from that on disc. Pleural pile whitish
and sparse or absent. Row of whitish pleural bristles above hind coxa.

Wing hyaline with slight costal bulge. Veins brown, light basally; crossvein r-m
before basal half of discal cell. Halter yellowish, knob brownish yellow.

Legs. Coxae brown with gray pollen and several whitish bristles and long pile.
Femora slender, dark brown or blackish, with basal third to fourth of middle and
hind femora yellow; fore femur slightly lighter brown with some yellow at extreme
base. Tibiae predominately yellow, apex of fore tibia narrowly brown banded, apical
fourth of midtibia and apical third to half of hind tibia brown. Tarsi primarily brown
with basal segments yellow to yellowish brown. Leg segments with appressed yel-
lowish pile, sparse on fore femur, some black pile at femoral apices; tibiae with short
dark setulae. Femoral bristles and hairs yellowish to yellowish white except as follows:
all femora with 2 short apical hairs; middle femur with 6 black bristles; hind femur
with 5-6 setigerous black bristles in ventroposterior row. Tibial and tarsal bristles
black except for 4 yellowish ones on fore tibia, 2 on middle tibia and 3 on fore tarsus.

Abdomen brown, apical margins of segments lighter. Segments 3 and 4 moderately
constricted with segment 5 expanded somewhat abruptly. Sides of tergites 1-6 and
all stemites with pale brownish white pollen, brown elsewhere; brown setae on darker
areas of tergites, yellowish white on lighter areas with 1 slightly longer hair at apical

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OMMATIUS FROM HISPANIOLA

139

comers of tergites 3-7; pile whitish, most abundant on basal five segments. Tergite
1 with 3 pale bristles.

Terminalia brown with weak pale pile. Epandrium slender, slightly wider basally;
apical margin somewhat oblique with ventral margin longer and abundant long pile.
Gonocoxite with 3 rather flat triangular processes, 2 below and 1 above. Gonostylus
somewhat flattened and slightly curved forward. Hypandrium with apical margin
sharply sloped toward middle. Cercus narrow with pale pile; ventral plates below
cercus fused, forming a broad M-shaped process.

FEMALE (Figs. 8a, b): 12.0 mm. The female differs from the holotype as follows:
tomentum and pollen of body with more yellow than male. Proclinate postocular
bristles black. Leg bristles primarily orangish yellow; black bristles in ventral row of
hind femur absent; 3 or 4 pale fore tarsal bristles. Crossvein r-m beyond middle of
discal cell; costal bulge absent. Halter brownish yellow. Segment 8 somewhat shiny
with abundant pale pile and a few hairs. Tergite 9 with only lateral comers exposed,
comers only slightly projecting posteriorly. Sternite 8 with transverse wrinkles, a
V-shaped row of pale bristles and a low mid-line ridge on apical third; a shiny
depression on each side of ridge; apical margin with 2 lateral, slightly projecting,
protuberances. Abdomen lacking constriction near middle.

Holotype. 6, Barahona, Dominican Republic, September 1938 (Darlington). Al-
lotype: 9, Port-au-Prince and vie., Haiti, March 18, 1934 (N. Bates). Paratypes: 9,
Dessorces, Haiti, alt. 100 ft., March 2, 1922, F. 4639 (Collector ?); 9, Diquini., Haiti,
date ? (W. M. Mann): 3<3<3, 499, Bois Caradeux, Port-au-Prince, Haiti, September 7-
10, 1936 (E. M. Ducasse), 9, Barahona Prov., Dominican Republic, July 13, 1967
(L. H. Rolston).

The holotype and allotype are housed at MCZ; and the paratypes at AMNH,
USNM, CNC, and the collection of the author.

Variation. Minor differences among the paratypes are as follows: lengths 6 12.1-
15.1 mm. 9 1 1.0-13.5 mm; one male with pale yellow or orange facial bristles, and
two black bristles in the antero ventral row of hind femur. Two females with white
tomentose faces. Apical margin of epandrium sometimes more rounded than on the
types.

Etymology. This species is named for the island from which the insect was collected.

Diagnosis. Ommatius hispaniolae is easily recognized by its dark brown fore femur
and an absence of marginal scutellar bristles; the slender epandrium, the slight bulge
in the costal margin of the wing and the spatulate abdomen of the male.

This species will run to O. jamaicensis Farr in Farr’s key (1965), but can be
separated from it by the virtual absence of orange-brown on the femora, pale to dark
brown tibiae and a sickle-shaped dististylus. Males of O. hispaniolae also have a
slender epandrium and a slightly spatulate abdomen.

Ommatius gwenae, new species
Figs. 9, 10

Description. MALE (Figs. 9a-c): 1 7.0 mm. Head dark brownish black. Face grayish
yellow tomentose, front yellow to brownish yellow. Mystax whitish with several long,
brown hairs above mystax; 4 brown ocellar bristles, 2 noticeably long. Occipital
tomentum whitish, pile white; postocular bristles black and proclinate, inner 3 or 4

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Figs. 9, 10. Ommatius gwenae, terminalia. 9. Male, lateral (a), dorsal (b) and apical (c)
views. 10. Female, lateral (a) and ventral (b) views.

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OMMATIUS FROM HISPANIOLA

141

long, curved above eyes. Antennal segments 1 and 2 with black hair, third with 2
short hairs dorsally and length twice its width; style slightly less than twice length of
all segments combined.

Thorax dark brown to black with some brown pollen. Scutum with mostly grayish
or whitish pollen on sides, prescutellar region, scutellum and pleuron; pale pile
abundant on prothorax, humeral callus and on anepistemum; pleural pile sparse or
absent elsewhere. Scutal bristles black as follows: 2 notopleural, 1 supra-alar, 1
postalar and 8 rather stout prescutellar dorsocentrals; numerous weaker bristly hairs
in prescutellar region, on side above wing and on postalar callus; several short dor-
socentrals anteriorly. Scutellum with long pale pile and bristly hairs, none on margin
conspicuously different in length or thickness. Vertical row of pleural bristles pale
and long. Halter reddish brown.

Legs. Coxae black with gray pollen and whitish bristles and pile. Femora, hind
tibia and tarsi primarily black; basal tarsomeres of fore and middle tarsi brown.
Femoral bases reddish yellow on basal third to half; fore and middle tibiae with
narrow apical dark bands and yellow above; hind tibia dark on apical half; reddish
yellow to reddish brown above. Legs with mostly yellowish to whitish pile, some
black pile on dark areas of femora and 1-2 short, black, bristly apical hairs. Fore
and middle femora with anterior bristles black and yellowish bristles below; 1 black
or pale preapical posterior bristle on middle femur. Hind femur with 4 whitish bristles
anteriorly; setigerous bristles in ventral rows primarily black with 1-3 pale yellow
apically. Fore and middle tibiae with only 1-2 black bristles beyond those at apex,
others yellowish. Hind tibial and fore tarsal bristles mostly black except for 1 or 2
yellowish bristly hairs and 4 pale yellow bristles, respectively.

Wing hyaline with veins dark brown apically, lighter basally; costal margin thick,
bulging greatly; anterior cells with ripples and slight brownish tint; r-m crossvein
before middle of discal cell.

Abdomen dark brown to black, with margins of most segments somewhat lighter.
Pollen primarily grayish to whitish with traces of brown on sternites 7-9. Pile pale,
longer and more abundant on basal segments. Tergite 1 with two pale bristly hairs;
segments 6-8 with black setae, longest along apical margins and comers.

Terminalia mostly black with abundant pale yellowish pile and hairs; some black
hairs on basal half of epandrium. Epandrium with length twice basal width, slightly
tapered on apical third, apical margin somewhat truncate. Hypandrium with nu-
merous transverse wrinkles, its apical margin tapered abruptly before middle, forming
an arched point; the latter preceded by a shallow transverse depression and a dense
row of fine black hairs on each side of middle. Gonocoxite with a shallow groove
leading to 2 tapered processes on each side of middle and a lateral flange. Gonostylus
reddish, gently arched forward, apex slightly curved laterally. Plates below cercus
fused, forming an M-shaped structure, the middle narrowly elongate and apically
truncate.

FEMALE (Figs. 9a, b): 1 7.0 mm. Slight differences include: facial tomentum slightly
darker yellow than male, pollen behind humeral callus golden in some light, yellowish
brown on sides of scutum. Femora less swollen than male; middle femur with several
fine black bristly hairs anteroventrally; hind femur with ventral rows of setigerous
bristles mostly pale, only 1-3 black. Fore and middle tibiae with several bristly black
hairs below. Wing without costal bulge; r-m crossvein beyond middle of discal cell.

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Tergite 8 somewhat shiny with black setae, longer along apical margins and apices.
Tergite 9 concealed by 8 above and wrapped around base of cercus. Sternite 8 with
yellow gray pollen basally, white setae and long hairs (the latter in a V-shaped pattern);
basal half with transverse wrinkles; apical half with a light colored, low mid-line
ridge narrowed to a point in apical margin; apical margin without usual lateral
protuberances, a shiny depression to each side of ridge.

Variation. Minor variations in the paratypes (3, 9) included; lengths 16.0 mm;
female with a few brown hairs above mystax; male front with fine brown hairs along
eye margin; 3 or 4 short pale or black bristles on posterodorsal surface of middle
femur; hind femur with posterior row of ventral bristles all black, 1 black bristle on
anterior surface. All tibiae with appressed black setulae.

Holotype. 6, Dominican Republic, Constanza, 1-10 June 1969 (Flint and Gomez).
Allotype: 9, same data as holotype. Paratypes: 6, Dominican Republic, La Toma, N.
of San Cristobal, 9-10 June 1969 (Flint and Gomez); 6, Santa Domingo, Dominican
Republic, date ?, Williston Collection.

The holotype and allotype are in the USNM and the paratypes in the AMNH and
the collection of the author.

Etymology. Ommatius gwenae is named in honor of my daughter, who developed
an interest in insects at an early age.

Diagnosis. Ommatius gwenae is recognized by a dark body; facial vestiture mostly
whitish with 6-7 brown hairs above mystax; inner postocular bristles black and
strongly proclinate; femora and hind tibia mostly black with basal third to half reddish
brown; scutellum with mostly pale pile, a conspicuous pair of scutellar bristles absent.
Males with a prominent costal bulge, apical margin of epandrium somewhat truncate,
a patch of fine black pile on the hypandrium, an M-shaped ventral plate and two
acutely tapered gonocoxal processes.

Ommatius gwenae is most similar to O. russelli, n. sp. but can be quickly recognized
by the characters described in the key and those discussed at the end of the section
on O. russelli, n. sp.

Ommatius russelli, new species
Fig. 1 1

Description. MALE (Figs. 1 la, b): 16.0 mm. Dark brown to black. Tomentum of
face, front and first antennal segment yellow to brownish yellow; occiput grayish
yellow tomentose. Mystax yellowish white; 6-8 brown hairs and several pale shorter
ones above mystax. Hairs of antenna, front and ocellar tubercle brown to black; 2
long ocellar bristles and several smaller ones. Upper postocular bristles dark, inner
2 or 3 on each side of vertex strongly proclinate; remaining postocular bristles slightly
curved or straight, shorter and pale yellowish white. First antennal segment longer
than either second or third segment, third longer than wide and with 3 inconspicuous
dorsal hairs; style almost twice length of all segments combined.

Thorax with mostly brownish pollen above, yellow to brownish yellow behind
humeral callus; yellowish to grayish yellow pollinosity in grooves, on sides, in pre-
scutellar region, on scutellum and on pleuron. Scutal bristles black (2 notopleural, 1
supra-alar, 1 postalar and 8-9 posterior dorsocentrals); black bristly hairs and pile
on sides and in prescutellar region, short anteriorly, longer posteriorly. Scutellum
with black bristly hair becoming longer toward posterior margin, 3 hairs on margin

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OMMATIUS FROM HISPANIOLA

143

Fig. 1 1 . Ommatius russelli, male terminalia; lateral (a) and apical (b) views.

Fig. 1 2. Ommatius nigellus, female terminalia; dorsal view (a), ventral plates (b) and stemite

8(c).

slightly longer than those on disc but otherwise not noticeably different. Pronotal
and pleural pile pale. Row of pleural bristles yellowish white. Halter reddish brown.

Legs. Coxae black with grayish yellow pollen and pale yellowish white bristles and
pile. Most or all femora primarily black with a narrow reddish basal ring or a small
reddish basal spot; tibiae yellow to reddish yellow except for dark apical third of
hind tibia and narrow dark apical ring on remaining tibiae. Tarsi dark brown with
basal tarsomeres slightly lighter. Legs with mostly yellowish short pile, some black
pile in dark areas; tibiae with black setulae. Black bristles on anterior surfaces of fore
(1) and middle femora (5); yellowish bristles and hairs below. Middle femora with
1 brownish yellow preapical bristle and 2-3 shorter dorsoposterior bristles. Hind
femur with yellowish or brownish bristles except for 3 or 4 black ones in posterior
ventral row; several slender pale hairs dorsally on basal half of hind femur. Bristles
of tibiae and tarsi black except for 3-4 on fore tarsus, 5 on fore tibia and 3 on middle
tibia. Fore and middle tibiae with a row of fine black hairs below.

Wing hyaline; costal margin thick, bulging greatly; anterior cells brownish especially
along margin of veins; r-m crossvein slightly beyond middle of discal cell.

Abdomen brownish black, margins of several segments slightly lighter. Abdominal

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pollen brown to yellowish gray; pale brown to brown short setae on dark areas of
tergites and long slender hairs in apices of tergites 7 and 8; pile and remaining bristles
pale yellow.

Terminalia mostly black with gonopods and apex of epandrium reddish; hair and
pile of basal half black, that of cercus and ventroapical margin of epandrium abundant
and pale. Epandrium length more than twice basal width, slightly swollen basally
and tapered apically to a point with a slight constriction before the apex. Hypandrium
with a dense, transverse tuft of black pile, apical margin forming a narrow, slightly
arched point at middle and a shallow transverse depression posteriorly. Gonocoxite
with a shallow basal groove leading to middle and connecting to a flat arched flange.
Gonostylus curved sharply outward on apical third, apex somewhat blunt. Plates
below cercus fused at middle forming a broad M-shaped structure, the middle short,
broad with apex truncate.

Female unknown.

Holotype. 6, Dominican Republic, La Palma, 12 km E. of El Rio, 1-13 June 1969
(Flint and Gomez). Paratype 6, same data. The types are stored in the USNM.

Variation. The paratype male differs as follows: 18.0 mm, 3 postalar bristles, 4-5
bristly dorsocentrals on each side, fore femur wholly black.

Etymology. Ommatius russelli is named after my son, who accompanied me on
numerous collecting trips.

Diagnosis. Ommatius russelli is recognized by a dark body, facial vestiture pri-
marily yellowish white with 6-8 brown hairs above mystax; inner postocular bristles
black and strongly proclinate; femora almost wholly black with only a narrow basal
reddish ring or spot; scutellum with abundant black bristly pile. Males with prominent
costal bulge, epandrium tapered to point, hypandrium with a complete row of fine
black pile and an M-shaped ventral plate.

Ommatius russelli is most similar to O. gwenae, n. sp. but differs from that species
by its almost wholly black femora, wholly black scutellar pile, a complete transverse
row of black pile on the hypandrium, a slender epandrium whose length is at least
twice or more than its widest width, and the absence of acute spine-like gonocoxal
process.

Ommatius nigellus, new species
Fig. 12

Description. FEMALE (Figs. 12a-c): 15 mm. Integument black. Tomentum of face
and occiput yellowish, that of front brownish. Hairs of lower half of mystax yellowish;
upper half with 3 strong, short black bristles, 5 or 6 longer black hairs and abundant
shorter black hairs. Hairs and bristles of antenna and front black; 9 ocellar hairs, 2
slightly longer than others. Postocular bristles black, 6 or 7 on each side of vertex
proclinate with inner 2-3 strongly so; a few weaker black hairs immediately behind
proclinate bristles with remaining occipital hairs white. Third antennal segment length
twice its width, slightly longer than basal segment, and with 2 short dorsal hairs;
style slightly more than twice length of the 3 segments combined.

Thorax dark brown to black. Pollen of scutum mostly brown; yellowish white to
yellowish brown in grooves and on sides; yellowish white behind, on scutellum and
pleuron. Scutal bristles and hairs black, weak pile primarily black, some pale; 2

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145

notopleural, 1 supra-alar, and 1 postalar bristles; abundant bristly hairs above wings
and between dorsocentrals; dorsocentrals extend full length of scutum, shorter an-
teriorly, stronger and more abundant posteriorly. Scutellum with abundant black
slender hairs, somewhat shorter and weaker basally, becoming gradually longer and
slightly thicker posteriorly on margin; marginal hairs not obviously different from
those on disc. Pleural pile mostly pale, scattered in thin patches or absent; some
weak bristly black pile on an- and katepistemum. Row of pleural bristles black on
upper half and pale yellow or whitish on lower half. Halter reddish brown.

Legs black. Coxae with yellowish white pollen, and yellowish bristles and hairs.
Femora and tibiae with primarily yellowish hairs and setae, bristles mostly black;
femora with 1-2 short apical bristly black hairs and some black apical setae; fore
femur with 1 short black bristle on basal fourth; middle femur with 4-5 black bristles
on basal half; 4 additional black ones on anterior face, and 1 posteroapical setigerous
black bristle. Hind femur with 1 black bristle on anterior face and most bristles on
antero ventral row black. Tibiae and tarsi with black bristly hairs and bristles except
for 1 yellowish one on fore tibia and tarsus and 2-3 on middle tibia.

Wing hyaline; veins dark brown, lighter basally; r-m crossvein beyond middle of
discal cell.

Abdomen black, apical borders of most segments brownish. Tergites with whitish
or yellowish white pollen on lateral margins and on sternites. Dark areas of tergites
with traces of brownish pollen and short setae; pale yellowish setae and hairs on
sides of tergites and stemite. Sides of tergites 1 and 2 with abundant pale hairs, tergite
1 with 2 black bristles. Tergite 8 shiny black with long black bristly hairs. Sternite
8 greatly arched upward on apical half and with V-shaped pattern of long black
bristles; apical margins with 2 apical protuberances and a median notch, an arched
ridge on apical third with shiny lateral depressions. Cercus with abundant pale pile
and a few scattered short bristly black hairs dorsally and apically. Ventral plates
below cercus with a short lateral lobe and several short spine-like bristles.

Male unknown.

Holotype. 2, Haiti, Furcy, July 25, 1950 (A. Curtiss). The holotype is stored in the
AMNH.

Etymology. Ommatius nigellus is named for its uniform black body.

Diagnosis. Ommatius nigellus is recognized by its black body, yellowish and black
vestiture of the face and legs, several strongly proclinate postocular bristles, a row
of black and white pleural bristles and abundant black scutellar hairs. This species
is similar to O. haitiensis, n. sp. but can be separated from it by the presence of
yellowish hairs in the lower half of the face, absence of anepimeral and strong marginal
scutellar bristles and the presence of a row of pleural bristles of which the upper half
is black and the lower half pale.

Ommatius stramineus, new species
Fig. 13

Description. MALE (Figs. 13a, b): 13.0 mm. Integument brown. Tomentum of
head bright yellow, yellowish brown at some light angles. Bristles and hairs of face
yellow, those of antennal segments, front, ocellar and postoculars brown, remaining

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y

>

Fig. 13. Ommatius stramineus, male terminalia; lateral (a) and ventral (b) views.

occipital vestiture yellowish; two long ocellar bristles and several (7) shorter ones.
First and third antennal segments about equal lengths and about one-third longer
than second; third segment length twice its width and with 1 or 2 dorsal inconspicuous
bristles; style almost twice length of the 3 segments combined.

Thorax dark brown, scutum with brownish pollen, yellow to light brownish yellow
pollen in grooves, on sides, behind, on scutellum and pleura. Scutal bristles dark
brown to black with 2 notopleurals, 1 supra-alar, 1 postalar and 4-5 weak yellowish
posterior dorsocentrals on each side. Thoracic pile weak and pale, most abundant
on prothorax, humeral callus and on anepistemum; few scattered hairs between
dorsocentrals and on pleura, meron bare. Scutellum with sparse, weak and yellowish
pile, bristles absent along margin. Row of yellowish bristly hairs above hind coxa.
Halter brownish yellow.

Legs. Coxae brown with light yellow tomentum, yellowish bristles and hairs. Fem-
ora, tarsi and apical third of hind tibia brown, a lighter narrow apical brown band
on anterior 4 tibiae. Legs with weak yellowish hairs and setae, some black on femoral
apices and black setulae on tibiae. One or 2 short black bristly hairs apically on
femora. Femoral bristles yellowish except black as follows: middle femur with 5
anteriorly and 1 posteriorly; hind femur with 3-4 anteriorly and all but 2 or 3 black
in both ventral rows. Tibial and tarsal bristles primarily black, only 2 or 3 yellow
bristles on anterior 4 tibiae and 4 on fore tarsus.

Wing hyaline. Veins brown, dark apically, lighter basally; costa slightly bulging,
anterior cells rippled and with a brownish tint, especially along veins in bulge; r-m
crossvein before middle of discal cell.

Abdomen yellowish brown, apical borders yellowish. Pollen yellow, abundant on
basal 4 segments and traces of brown above. Hairs and setae primarily yellow, some
brown on dorsum of last 3 tergites; hairs long and thin on basal segments with 2-3
bristly hairs on apical corners of tergites 5-8 and 2 yellow stiff bristles on tergite 1 .

Terminalia yellowish brown to dark brown; hairs and bristles yellow. Epandrium

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OMMAT1US FROM HISPANIOLA

147

Fig. 14. Ommatius cinnamomeus, female terminalia; lateral (a) and ventral (b) views.

dark brown, greatly inflated on basal three-fourths; apical fourth thin, yellowish
brown, strongly narrowed below, slightly truncate apically. Hypandrium dark brown,
apical margin straight or nearly so. Gonocoxal with short stiff bristly hair basally
and two slender projections, basally separated, apically converging. Gonostylus slight-
ly curved, slender, somewhat flattened laterally. Ventral plates below cercus fused,
forming a large single plate with two apical swollen ridges, the apex bearing yellow
pile.

Female unknown.

Holotype. 8, Haiti, Mt. Bouretta, 5,000 ft., 1 5 September 1934 (M. Bates). Paratype:
8, Dominican Republic, Constanze, 3,000-4,000 ft. (Darlington). The holotype is
stored in the USNM and the paratype in the MCZ.

Variation. The male paratype is identical to the holotype except as follows: length
14.0 mm; the body is darker, especially the femora, tarsi and abdomen; the proclinate
bristles are yellow, and additional black bristles are on the hind femur; the apical
margin of the epandrium is somewhat truncate.

Etymology. This species is named for the pale straw-colored tomentum of the face.

Diagnosis. Ommatius stramineus is easily recognized by its yellow facial tomentum
and vestiture, femora wholly brown, deep yellow pollinose abdomen, a slight bulge
in the costal margin, an epandrium with the apical margin somewhat truncate and
an absence of marginal scutellar bristles. This species is somewhat similar to O.
vitreus but differs from that species by a stronger body, a slight bulge in the costal
margin and the absence of an apically bifurcated epandrium.

Ommatius cinnamomeus, new species
Fig. 14

Description. FEMALE (Figs. 14a, b): 20.0 mm. Head brown to black. Tomentum
of face pale yellow, front yellowish brown, occiput pale yellowish to pale brownish
gray; 2 long ocellar and 6 proclinate postocular bristles black, remaining occipital

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hairs pale yellow. Facial bristles and hairs pale yellow. Antennal basal segment slightly
longer than second or third; third segment length slightly more than twice width;
antennal bristles dark to pale brown, third segment with two short inconspicuous
bristles dorsally; style length about one and one-half times that of all segments
combined.

Thorax dark brown. Dorsum with bright brown pollen, deep yellowish brown
behind humeral callus, lighter in grooves, on sides of and behind the scutum; pollen
of scutellum brownish, that of pleuron brownish to yellowish gray. Scutal bristles
black as follows: 2 notopleurals, 1 supra-alar, 1 postalar; 4 bristly dorsocentrals
posteriorly and several short weaker ones anteriorly. Several dark bristly hairs on
side above wing. Pale thoracic pile abundant on prothorax and humeral callus, less
abundant and sparse elsewhere. Scutellum with long pale pile, none on margin as
conspicuous pair. Row of pleural bristles whitish yellow. Halter brown with stalk
lightest.

Legs. Coxae brown with yellowish gray pollen and pale yellowish bristles and hairs.
Femora and tibiae mostly yellowish to brownish yellow; anterior legs lightest, dark
brown as follows: apical third to half of femora; a narrow apical dark band on fore
and middle tibiae and apical fourth to third on hind tibia. Tarsi dark brown to black
except basal tarsomeres slightly lighter. Pile and setae of legs mostly pale yellow with
1-2 black bristly apical hairs and black dorsoapical setae on femora, black setulae
on tibiae, fore tibia with a ventral row of long thin black hairs; femora with bristles
pale yellowish to orangish except as follows: middle femur with 5 black bristles
anteriorly, and 1 posterodorsal bristle; hind femur with 1 or 2 setigerous black, apical
bristles in each ventral row. Tibial and tarsal bristles black except for 1-3 yellowish
ones on fore and middle tibiae and fore tarsus.

Wing hyaline with a slight brownish tint, costal cell darker in part; costal margin
without bulge; r-m crossvein beyond middle of discal cell.

Abdomen dark brown with margins of segments lighter. Segments with yellowish
brown to brown pollen, sternites and sides of posterior 3 or 4 tergites somewhat
lighter. Dark brown setae on dark areas of tergites; thin pale hairs on lighter pollinose
areas, longest on basal 3 or 4 segments; bristles on basal segments yellowish. Apical
margins of tergites 7-8 with several black bristly hairs or bristles. Most of tergite 8
and all of 9 hidden from above by preceding tergites. Cercus with yellowish pubes-
cence, a few weak and bristly brown and yellowish hairs along margins. Sternite 8
with yellowish pubescence and a V-shaped pattern of black bristles; the apical margin
with a notch and lateral protuberances; a strong elevated ridge (in lateral view) and
a shallow, shiny depression on each side.

Male unknown.

Holotype. 2, Haiti, NE foothills. La Hotte, 200-400 ft., October 10-24, 1934
(Darlington). The holotype is in the USNM.

Etymology. This species is named for the brown pollen on the thoracic dorsum.

Diagnosis. Ommatius cinnamomeus is easily recognized by its large size, pale
yellow facial tomentum and hairs, brown scutal pollen and bristles, brownish tint of
the wing surface, legs mostly yellowish to yellowish brown and the absence of marginal
scutellar bristles. This species runs to O. tibialis in Curran’s key (1928) but differs
from that species in its larger robust body, more extensive yellow ground color of
the fore femur, the presence of only brown hairs on sternite 8 and the absence of

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OMMATIUS FROM HISPANIOLA

149

pale or yellowish scutal bristles. Females of O. tibialis typically have the anterior
surface of the fore femur black, occipital and pleural pollen white or gray and hair
of stemite 8 white or yellowish.

ACKNOWLEDGMENTS

I thank the following for the loan of specimens used in this study: Norman Woodley, Museum
of Comparative Zoology, Cambridge, Mass.; Pedro Wygodzinsky, American Museum of Natural
History, New York; Lloyd Knutson and Raymond Gagne, Systematic Entomology Laboratory,
USDA, Washington, D.C.; Charles A. Triplehom, Department of Entomology, Ohio State
University Museum, Columbus; Leif Lyneborg, Museum of Zoology, Copenhagen, Denmark;
Thomas Farr, Institute of Jamaica, Kingston: M. C. Birch, Hope Entomological Museum,
Oxford, England; D. M. Wood, Biosystematic Research Institute, Agriculture Canada, Ottawa,
Canada. Thanks are also due to the staff of the Diptera section in the Systematic Entomology
Laboratory, USDA, at the U.S. National Museum who so kindly assisted me with numerous
suggestions during this study; to L. Knutson, USDA, ARS, Systematic Entomology Laboratory,
Beltsville, Md. for reviewing and making helpful suggestions on an early draft of the manuscript;
to the Towson State University Faculty Research Committee for support of this study; and to
Keith Harris for preparing Figures 2, 3 and 9. The remaining figures were prepared by the
author.

LITERATURE CITED

Bigot, J. M. F. 1875. Dipteres nouveaux ou peu connus. 4e partie V. Asilides exotiques
nouveaux. Ann. Soc. Entomol. France (5)5:237-248.

Curran, C. H. 1928. New species of Ommatius from America, with key (Asilidae: Diptera).
Amer. Mus. Nov. 327:1-6.

Farr, T. H. 1965. The robber-flies of Jamaica (Diptera: Asilidae). Pt. 2. Bull. Inst. Jamaica
Sci. Ser. 13:5-36.

Hull, F. R. 1962. Robberflies of the world: the genera of the family Asilidae. Bull. U.S. Nat.
Mus. Pt. 2 224:443-446.

Martin, C. H. and N. Papavero. 1970. Family Asilidae. In: A catalogue of the Diptera of the
Americas South of the United States. Mus. Zool. Univ. Sao Paulo (35b): 1-1 39.
McAlpine, J. F. 1981. Morphology and terminology— adults. In Biosystematic Research In-
stitute (eds.), Manual of Nearctic Diptera. Agriculture Canada, Research Branch, Ottawa.
Monograph 27, Vol. 1:9-63.

Scarbrough, A. G. 1983. A new species of Ommatius Wiedemann from San Salvador, the
Bahamas. Proc. Entomol. Soc. Wash. 85:144-151.

Received June 29, 1983; accepted November 22, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 150-155

PROPLEURAL TRANSFORMATIONS WITH RESPECT TO
THE DISPOSITION OF PROPLEURAL SUTURE
IN ORDER HYMENOPTERA

Malkiat S. Saini

Department of Zoology, Punjabi University, Patiala- 147002, India

Abstract.— A distinct propleural suture, well developed proepistemum and considerably re-
duced proepimeron are clearly represented in Pamphiliidae, Xyelidae, Argidae, Tenthredinidae,
Diprionidae and Cimbicidae. In Cephidae, this suture has shifted slightly backward, thus, further
reducing the area of proepimeron. In Xiphydridae, the backward shift is more pronounced and
the posterior half of this suture has become marginal. In Siricidae, the entire propleural suture
has become almost marginal and the epimeral area if any is in the form of a thickened posterior
margin of propleural plate. In most of the apocritan families the same state is maintained. In
Formicidae even the thickened posterior margin of propleuron is absent. These observations
collectively help to establish a systematically changing pattern to which can be attached some
evolutionary significance within the order Hymenoptera.

The work that follows specifically deals with the positional variations and gradual
backward shifting of the propleural suture within the order Hymenoptera. The extent
and magnitude of the changes pertaining to the course of this suture as well as the
size of the epimeron are evolutionarily significant and bring to light some phylogenetic
relationships among different families of this insect order. The available literature is
completely devoid of such comprehensive studies excepting those of Snodgrass (1910)
and Matsuda (1970) which cover too limited a number of hymenopteran families.

However, a good deal of literature is available concerning the hymenopteran mor-
phology. Some important works include Crampton (1909, 1926), Snodgrass (1942),
Martin (1916), Weber (1927), Duncan (1939), Alam (1951), Arora (1953), Wong
(1963) and Dhillon (1966). These authors mainly worked on an ontological basis
rather than on the comparative basis, which is the main objective of the present
work. This work is based on the study of 22 different hymenopteran families.

MATERIALS AND METHODS

Most specimens of Apocrita were collected from the Punjab and Himachal Pradesh
during September and October 1975 and preserved in 80 percent alcohol. Except for
Megalodontidae, Orussidae and Pergidae, Symphyta were supplied by the Biosys-
tematic Research Institute, Canada, and the Zoological Survey of India. Since these
specimens were dry, they were softened in 2 percent KOH for 6 days. Drawings were
made with the help of a binocular microscope fitted with an ocular grid.

OBSERVATIONS AND DISCUSSION

The presence of a distinct propleural suture in some lower symphytans with its
gradual backward displacement among the higher symphytans till it takes up a mar-

1984

PROPLEURAL SUTURE

151

Figs. 1-12. Side view of the propleural plate of: 1 . Acantholyda maculiventris (Pamphiliidae).
2. Xyela bakeri (Xyelidae). 3. Arge clavicornis (Argidae). 4. Neodiprion abietis (Diprionidae).
5. Tenthredo verticalis (Tenthredinidae). 6. Xiphydria mellipes (Xiphydridae). 7. Cephus cinctus
(Cephidae). 8. Cimbex americana americana (Cimbicidae). 9. Sirex cyaneus (Siricidae). 10.
Netelia kashmirensis (I chneumonidae). 1 1. Trachysphyrus sp. (Ichneumonidae). 12. Sycoscapter
stabilis (Torymidae). Abbreviations (used also in Figs. 13-24): EPM— epimeron; EPST— epi-
stemum; PA— pleural articulation; PAP— pleural apophysis; PCX— procoxa; PS— pleural su-
ture; PST— prostemum.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

ginal course in the members of family Siricidae and all of the Apocrita, indicates a
trend which may have some evolutionary and phylogenetic significance. A coherent
account of all these modifications is as follows.

In Acantholyda maculiventris (Norton) (Fig. 1) (Pamphiliidae), Xyela bakeri Ko-
now (Fig. 2) (Xyelidae), Arge clavicornis (F.) (Fig. 3) (Argidae) Tenthredo verticalis
Say (Fig. 5) (Tenthredinidae), Cimbex americana americana Leach (Fig. 8) (Cim-
bicidae) and Neodiprion abietis (Harris) (Fig. 4) (Diprionidae) the propleural suture
is well represented. In all these cases this suture takes up a course which starts from
the pleural articulation of the first coxa, curves around the posterodorsal angle of the
propleural plate and ends up at the base of the concavity which receives the lateral
tip of the furcal arm of the prostemum. Internally, this suture is represented by a
distinct pleural ridge. The small area lying posterior to this suture is the proepimeron,
the large dominating area anterior to this suture being the proepistemum. Similar
conditions have also been observed in Onycholyda luteicornis (Norton) and Cephalcia
provancheri (Huard) (Pamphiliidae), Zarae inflata Norton (Cimbicidae), Pristiphora
cincta Newman, Pachyprotasis versicolor Cameron, Pachyprotasis brunetti Rohw.
and Eutomostethus assamensis (Rohw.) (Tenthredinidae). Such a condition has also
been described and similarly labelled by Snodgrass (1910), Wong (1963) and Dhillon
(1966) and under the name of cervicopleuron by Crampton (1909, 1926), Martin
(1916), Weber (1927) and Arora (1953).

Matsuda ( 1 970) in Arge sp. took the generalized propleural suture as the anapleural
suture and thus labelled the area anterior to it as the combination of pre-epistemum
and the kate-episternum while the small area lying posterior to this suture as the
anepistemum. According to him the actual propleural suture has become marginal
and consequently the area of proepimeron has been obliterated. In addition to the
anapleural suture, Matsuda showed the presence of a separate submarginal propleural
suture in Schizocerus sp. and Dolerus sp. — on which Weber (1927) and Crampton
(1926) worked, respectively— but did not label any separate submarginal suture.
However, on the basis of comparative study the present author is of the view that
there exists no anapleural suture in the propleuron of Symphyta. As the anapleural
suture of Matsuda (1970) bears some of the identification marks of the pleural suture,
i.e., its ventral end forms the pleural articular condyl which provides the pleural
articulation to the procoxa (a condition very much similar to meso- and metapleuron
and its dorsal end forms an articular facet which receives the lateral tip of the furcal
arm of the prostemum, a condition again similar to that usually met with meso- and
metafurcasternal arms which are generally associated with the pleural suture, par-
ticularly in Symphyta), so it is proposed to take this suture as the propleural suture.
To substantiate the above view it is further added that anapleural suture is not a
constant feature of all the symphytans (Saini and Dhillon, 1980), whereas, the pleural
suture is a constant feature within the entire range of Symphyta. Moreover, when
present, the anapleural suture takes up altogether a different and variable course than
that of the pleural suture (Matsuda, 1970; Saini and Dhillon, 1980). To confirm the
validity of Matsuda’s (1970) anapleural suture and a marginal pleural suture in Arge
sp. the present author studied some more species of Arge, viz. Arge simlaensis, A.
fumipennis, A. bipunctata, and A. xanthogastra, but failed to recognise any marginal
suture. However, if in the insect studied by Matsuda (1970) any marginal suture is

1984

PROPLEURAL SUTURE

53

Figs. 13-24. 13. Chrysis indogotea (Chrysididae). 14. Scolia quadripustulata ( Scoliidae). 15.

Scelephron intrudens (Sphecidae). 16. Stizus vespiformis (Sphecidae). 17. Eumenes dimidiate-
pennis (Eumenidae). 18. Calicurgus sp. (Pompilidae). 19. Vespa orientalis (Vespidae). 20. Xy-
locopa lemuiscapa (Xylocopidae). 21. Camponotus camelinus (Formicidae). 22. Tetraponera
rufonigra (Formicidae). 23. Dorylus labiatus (Formicidae). 24. Mutilla sp. (Mutillidae).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

present, that can be safely taken as the transepimeral suture as is the case in meso-
epimeron of Arge clavicornis (Saini and Dhillon, 1980).

The next stage in the evolutionary series can be observed in the members of family
Cephidae. In Cephus cinctus Norton (Fig. 7) due to the backward shifting of the
propleural suture the area of the epimeron is further narrowed. However, the iden-
tification marks of the suture are quite prominent and distinct.

In Xiphydria mellipes Harris (Fig. 6) (Xiphydridae) the backward shifting of this
suture is more pronounced and consequently the area of epimeron has been further
reduced. At this stage the basal half of this suture has become already submarginal.
This shows an inclination of this suture for becoming submarginal, a fact which is
evident in siricids. The identifying features of the pleural suture are quite clear.

In Sirex cyaneus F. (Fig. 9) (Siricidae) the propleural suture has become submar-
ginal and consequently the area of proepimeron has been almost obliterated. Sub-
marginal nature of the suture can be confirmed by the presence of a distinct sub-
marginal ridge, which on its ventral end gives rise to the pleural articular condyle,
that provides pleural articulation to the procoxa. However, the dorsal half of the
pleural ridge leading to the concavity which receives the furcal arm of the furcaster-
num has become almost inconspicuous. So, on the whole the propleural plate is
entirely represented by proepisternum alone. Similar conditions have also been de-
scribed by Snodgrass (1910) in Tremex columba (Siricidae).

In Hymenoptera Apocrita, the propleural suture has become completely marginal,
thus, losing its independent identity. Its ancestral presence is indicated only by the
thickened posterior margin of the propleuron. This thick margin is thought to contain
the remnants of the pleural ridge. The other remnants of the disappeared pleural
suture are the presence of pleural articular condyle and an association of the furcal
arm of the prostemum with this thickened posterodorsal margin. The above obser-
vations have been made on different apocritans which include Sycoscapter stabilis
(Walker) (Fig. 12) (Torymidae, Chalcidoidea), Netelia kashmirensis Cameron (Fig.
10) and Trachysphyrus sp. (Fig. 1 1) (Ichneumonidae), Chrysis indogotea Duf. et Pesr.
(Fig. 1 3) (Chrysididae), Mutilla sp. (Fig. 24) (Mutillidae), Scolia quadripustulata F.
(Fig. 14) (Scoliidae), Scelephron intrudens Smith (Fig. 15) Stizus vespiformis F. (Fig.
16) (Sphecidae), Vespa orientalis L. (Fig. 19) (Vespidae), Eumenes dimidiatepennis
Sauss (Fig. 17) (Eumenidae), Calicurgus sp. (Fig. 18) (Pompilidae) and Xylocopa
lemuiscapa Westwood (Fig. 20) (Xylocopidae). Similar conditions have been de-
scribed by Duncan (1939), Snodgrass (1942) and Alam (1951). However, in For-
micidae as seen in Dorylus labiatus Shuckard (Fig. 23), Camponotus camelinus Smith
(Fig. 21) and Tetraponera rufonigra (Jerdon) (Fig. 22) even the thickened posterior
margin of the propleuron is absent.

ACKNOWLEDGMENTS

I am grateful to the Director, Commonwealth Institute of Entomology, London and Dr. V.
K. Gupta of Delhi University, Delhi, for identifying some of the Hymenoptera. It is my special
pleasure to acknowledge the kind cooperation of Biosystematic Research Institute, Canada and
Zoological Survey of India, Calcutta, for supplying some direly needed Symphyta for this study.

1984

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155

LITERATURE CITED

Alam, S. M. 1951. The skeleto-muscular mechanism of Stenobracon deesae Cam. (Braconidae,
Hymenoptera)— an ectoparasite of sugarcane and juar borers of India. Part I, Head and
thorax, Alig. Musi. Univ. Publ. (Zool. Ser.) Ind. Ins. Typ. 3(1):74 pp.

Arora, G. L. 1953. The external morphology of Diprion pini L. (Symphyta, Hymenoptera).
Res. Bull. E. Punjab Univ. Hoshiarpur. Zool. 25:1-21.

Crampton, G. C. 1 909. A contribution to the comparative morphology of the thoracic sclerites
of insects. Proc. Acad. Nat. Sci. Philadelphia 61:1-53.

Crampton, G. C. 1926. A comparison of the neck and prothoracic sclerites throughout the
orders of insects. Trans. Amer. Ent. Soc. 52:199-248.

Dhillon, S. S. 1966. The morphology and biology of Athalia proximo Klug (Tenthredinidae,
Hymenoptera). Alig. Musi. Univ. Publ. (Zool. Ser.) Ind. Ins. Typ. 7:165 pp.

Duncan, C. D. 1939. A contribution to the biology of North American vespine wasps. Stanford
Univ. Publ. Biol. Sci. 8(1):272 pp.

Martin, J. F. 1916. The thoracic cervical sclerites in Insecta. Ann. Ent. Soc. Amer. 9:35-83.

Matsuda, R. 1970. Morphology and evolution of the insect thorax. Mem. Ent. Soc. Canada
76:431 pp.

Saini, M. S. and S. S. Dhillon. 1980. Changing course of the mesopleural suture in the order
Hymenoptera. J. Anim. Morphol. Physiol. 27(l&2):l-9.

Snodgrass, R. E. 1910. The thorax of the Hymenoptera. Proc. U.S. Nat. Mus. 39:37-91.

Snodgrass, R. E. 1 942. The skeleto-muscular mechanisms of the honeybee. Smith. Misc. Coll.
103(2): 120 pp.

Weber, H. 1927. Die Gliederung der Stemalregion des Tenthrediniden Thorax. Ein Beitrag
zur vergleichenden Morphologie des Insektenthorax. Z. Wiss. Insekt. Biol. Berlin 22:
161-198.

Wong, H. R. 1963. The external morphology of the adult and ultimate larval instar of the
larch sawfly, Pristiphora erichsonii (Htg.) (Hymenoptera: Tenthredinidae). Can. Ent. 95:
807-921.

Received November 25, 1981; accepted October 5, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 156-161

TOOL USE BY THE ANT,

NOVOMESSOR ALBISETOSUS (MAYR)

Philip McDonald

Department of Biopsychology, Hunter College,

City University of New York, New York, New York 10021

Abstract. — Soil-dropping behavior by the arid lands ants, Novomessor albisetosus (Mayr), is
considered as an example of tool use. Soil-dropping ceases to be a general response to any nest-
threatening liquid beyond a certain distance from the nest. As petri dishes of distilled water
and honey water are moved farther from the nest, ants stop dropping soil in water but continue
doing so in honey water. Since soil dropped in honey water is brought back to the nest, soil-
dropping serves the purpose of food retrieval. Questions concerning competitive adaptations
must be considered cautiously.

Alcock (1972) has provided an excellent review of tool use in feeding and proposed
the following definition: “Tool-using involves the manipulation of an inanimate
object, not internally manufactured, with the effect of improving the animal’s effi-
ciency in altering the position or form of some separate object” (p. 464). In what
may be the first reported case of tool use in a social insect, Lin ( 1 964-1965) described
the pavement ant’s ( Tetramorium caespitum) use of soil while attacking halictine
bees. Shultz (1982) enlarged on Lin’s findings, and Moglich and Alpert (1979) reported
similar behavior in a study of stone dropping by Conomyrma bicolor Wheeler to
possibly interfere with Myrmecocystus competition.

The first study of tool use by a social insect in retrieving food was made by Fellers
and Fellers (1976). Soil was placed by Aphaenogaster workers on jelly bait and later
retrieved to the nest. They concluded that tool using may increase the ability of
Aphaenogaster to compete directly with dominant species. A reconsideration of their
findings was presented by Fowler (1982) who found that most of the tools were not
retrieved, and concluded that tool use was more important in scramble competition
than in direct competitive interaction. Many years before these studies Wheeler (1910)
observed that many ants “throw pellets of earth or any other debris” on a “substance
that they cannot remove, such as a strong-smelling liquid.” He noted that liquids
more frequently evoke this behavior and that it may have evolved as a way of
protecting the nest against flooding.

Given this conjecture, it is not clear whether any kind of competition was involved
in the tool use response of the Feller and Feller (1976) or Fowler (1982) studies or
whether tool use to retrieve food was simply the coincidental outcome of a reflexive
response to a moist substance, signal of a potentially nest threatening liquid. I report
here on the reaction to liquids of Novomessor albisetosus (Mayr), an ant closely related
to Aphaenogaster, and the distribution of “tools” within the nest.

MATERIALS AND METHODS

Laboratory. Petri dishes of honey water (one part honey to 10 parts water) were
placed in the foraging arena (61 x 61 x 30 cm). Sand was collected from various

1984

TOOL USE

157

parts of the nest (3 plastic boxes joined by tubes in line from the foraging arena); it
was also collected from the floor of the foraging arena, midden pile, and petri dish.
Sand was analyzed for carbohydrate content by using the anthrone reaction method
slightly modified from that described by Scott and Melvin (1953). There were five
replications of each treatment. Two other liquids— maple sap (to test a sugar solution
of natural dilution) and distilled water— both with and without red food coloring,
were also placed in petri dishes in the foraging arena. Petri dishes of only one liquid
was available at a time, and sand was removed from the nests after each manipulation.
Lastly, water was poured directly into the nest near the exit to the foraging arena.

Field. Fifteen nests of N. albisetosus (Mayr) were selected within a 40,000 sq m
area about 1 km west of Portal, Arizona. Each was tested with both water and honey
water, once during the dry and once during the rainy season. Placement of petri
dishes was at various unmeasured distances for the 1st trial during the dry season.
Measurements were then taken and assigned randomly to the nests for the 2nd trial,
during the rainy season.

RESULTS AND DISCUSSION

Laboratory. Analysis of sugar content in sand from the nest boxes revealed the
presence of sugar in increasing quantities as the sand was sampled farther from the
nest entrance and closer to the brood (Fig. 1). Sand taken from the second of three
nest boxes, had the highest content of sugar (550 ± 5 yg per gram of sand). This was
where the queen and brood (especially feeding larvae) remained, and where chunks
of sand were first deposited by returning foragers. Although larvae were sometimes
found on the sand it was being fed on by workers of all ages. Sand from a stockpile
within the same nest chamber, but closer to the entrance, yielded a sugar content of
482 ± 2.2 yg. Samples taken from the next location closer to the entrance, in the
first chamber, had 189.7 ± 2.6 yg. The sand pile next to the nest entrance was littered
with dried mealworms, dead ants and other debris that were occassionally taken out
and placed on the midden heap. This had the least sugar content within the nest
(121 ± 4.3 yg ). Outside the nest, in the foraging arena, sand taken from the midden
heap yielded 94.3 ± 16 yg. Control samples of sand taken from three places in the
foraging arena revealed no detectible sugar content. A sample used as a standard for
comparison, taken from the petri dish had a sugar content of 568 yg, which was close
to the sugar content of sand first deposited in the nest by returning foragers. Following
a significant analysis of variance of samples taken from within the nest [F(4,20) =
3,775.17, P < 0.0001], comparisons of samples by the Newman-Keuls Multiple Range
Test were all significant at the 0.01 level of confidence. Apparently, as the food value
of the sand declined with its sugar content it was moved closer to the nest entrance,
and eventually removed to the midden pile.

Sand was dropped in both petri dishes of maple sap and distilled water. Both red
colored and uncolored sand from the maple sap was eventually taken into the nest,
but not sand placed in the water. Sand was also brought into the nest and deposited
on the water that had been spilled into the nest. As a further indication that it was
considered debris, sand taken from the pile with the least sugar content within the
nest was also placed on water flooding the nest.

Because of these results the field study was conducted. The possibility that the ants
were responding to any liquid, or treating the foraging arena as part of their nest,

158

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

DISTANCE FROM ENTRANCE WITHIN NEST (CM)

Fig. 1. The relationship between sugar content of sand and position of sand in the nest.
Sand brought to the brood at a point farthest from the entrance has the highest sugar content.
During stockpiling and nest maintenance, as sand is brought increasingly closer to the entrance,
the sugar content of sand decreases.

hence soaking up any nest-threatening liquid with sand, could not be overlooked.
Of course, this behavior itself could be viewed as tool use. However, I wanted to see
if returning to the nest with sugar-coated sand was not just coincidental to the ants’
dropping soil in any liquid and then discovering that the soil was sweet.

Field. The distances that water and honey water were placed from the nest en-
trances, both before and after the rainy season, are presented in Table 1. During the
dry season soil was regularly dropped into water up to 2.13 m from the nest, though
in diminishing quantities as water was placed farther from the nest. Recruitment to
water was observed up to 6. 1 m from the nest, through the response at this distance,
10 ants in 20 min, was about one-third the response to honey water at the same
distance. No ants returned to the nest with soil (pebbles and sand) that they had
dropped into the water. During the wet season (after mid-August in 1982, it usually
begins early in July) soil was dropped into water up to 1.22 m from the nest but not
beyond, although there was recruitment (15 ants in 30 min) up to 1.83 m from the
nest. Again, no ants returned to the nest with soil from the water. The Fellers (1976)
observed Aphaenogaster placing “tools” on jelly up to 152 cm from the nest. This
is well within the 2. 13 m range for dropping soil in water in the present study. What

1984

TOOL USE

159

Table 1 . The occurrence of soil-dropping in and recruitment to liquid, and returning to nest
with sand dropped in liquid.

Before rains

After rains

Water

Honey water

Water

Honey water

(m) from
nest

Soil

Re-

cruit

Return

Soil

Re-

cruit

Return

Soil

Re-

cruit

Return

Soil

Re-

cruit

Return

0.61

+

+

0

+

+

+

+

+

0

+

+

+

1.22

+

+

0

+

+

+

+

+

0

+

+

+

1.83

+

+

0

+

+

+

0

+

0

+

+

+

2.13

+

+

0

+

+

+

0

0

0

+

+

+

2.44

0

0

0

+

+

+

0

0

0

+

+

+

2.74

0

+

0

+

+

+

0

0

0

+

+

+

3.05

0

0

0

+

+

+

0

0

0

+

+

+

3.66

+a

0

0

+

+

+

0

0

0

+

+

+

4.27

0

0

0

+

+

+

0

0

0

0b

0

0

4.88

0

0

0

0C

0

0

0

0

0

0C

0

0

6.1

+d

+

0

+

+

+

0

0

0

0C

0

0

6.71

0

0

0

+

+

0e

0

0

0

+

+

+

9.15

0

0

0

+

+

+

0

0

0

+

+

+

12.2

0

0

0

+

+

+

0

0

0

+

+

+

15.24

0

0

0

-1-

+

+

0

0

0

+

+

+

a Placed 1 m from honey water, 6 pebbles dropped in.
b Undiscovered.

c Camponotus prevented access.
d 5 pebbles dropped in.
e Camponotus prevented return.

they observed may not have been tool use but simply a response to any moist
substance.

With but few exceptions soil-dropping and recruitment to honey water, and return
to the nest with the soil, was observed in both dry and wet seasons up to 15.24 m
from the nest. Unless hindered by a wall of feeding ants surrounding the petri dish
(many drank for long periods, up to 45 min), soil-dropping occurred shortly after
discovery. At times, ants would crawl over the backs of those feeding in order to
drop soil into the honey water. Even when recruitment was not heavy (sometimes
as many as 60 ants would recruit to honey water within 30 min) soil-dropping was
steadily pursued. Within 3 to 4 hr, petri dishes were filled with enough soil to soak
up all the liquid. By dawn most of the soil would be gone and a trickle of ants could
still be seen bearing pebbles and chunks of sticky sand back to their nests. (It was
noted that where Camponotus prevented N. albisetosus from access to honey water,
Camponotus did not drop soil into the liquid but fed from it and patrolled nearby).
This finding differs with Fowler’s (1982) study in which from 60-96 percent of the
tools were not recovered. The colonies he chose may have been smaller than mine,
with less of a demand for sugar.

Results from the field studies make it evident that, beyond a certain distance from
the nest, soil-dropping is not simply a general response to any liquid. Foragers were
selective. Within 2.13 m of the nest, however, the results were not so clear. Given

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

the laboratory ants’ behavior of dropping sand in water spilled into their nest, and
given that in the field ants drop soil into water from 0.6 1 to 2. 1 3 m from their nests,
but with diminishing intensity as the water approaches 2 m, liquid close to the nest
seems to pose a threat. A small puddle of rainwater, for instance, may become larger
with more rain and eventually flow into the nest entrance. Levelling surrounding
depressions indicated by small puddles may prove adaptive for nest protection.
Wheeler’s (1910) conjecture about the behavior evolving from a response to nest
threatening liquid seems to be reasonable.

Under crowded laboratory conditions, for example when placed in a single box
that serves as both nest and foraging space, many ant species, including N. albisetosus,
will drop debris from their midden piles into honey water. Though at first glance it
seems as if they are rejecting the honey water, this may be no more than a response
to a nest-threatening liquid by using the closest material available. In laboratory
nests, even those with foraging arenas, the debris is often not fed on for many days
if at all after it has been dropped in the honey water. This is in agreement with
Fowler’s (1982) finding that most tools are not recovered. Unfortunately, distances
from the nest are not given in his study. Further research will have to be done
examining the percentage of soil retrieved with increasing distance from the nest.

Since, in the field, the soil-dropping response continued only toward the honey
water as the two liquids were placed increasingly farther from the nests, it seems
apparent that nest protection ceases to be a possible reason for soil-dropping behavior
beyond a certain distance from the nest. The argument of chance has now been made
less tenable. That is, beyond 2.13 m it is not the case that the ants are responding
generally to a moist substance only to find that the soil which they drop into it
becomes laden with sugar, after which they take it back to their nests. Here a better
case for tool use can be made.

While the Fellers and Fellers (1976) study demonstrated the greater efficiency of
food retrieval by tool use when compared with internal transport their conclusions
about competition, as well as Fowler’s (1982), must be considered cautiously. Their
studies will have to be replicated at greater distances from the nest. Further, exper-
imental artifact, though it seems unlikely, cannot be overlooked. Though the Fellers
(1976) state that Aphaenogaster puts tools on squashed spiders, they do not say
whether the workers retrieved the tools to the nest. I have observed N. albisetosus
placing soil on a lizard squashed by a car, but not retrieving the soil to their nest.
During two summers of studying these ants in the field, I have never observed a
natural occurrence of tool use for food retrieval.

ACKNOWLEDGMENTS

I thank Dr. Michael Greenspan of Merck Research Laboratories, Rahway, New Jersey for
analysis of sugar content of soil samples. Special thanks are extended to Mr. G. Miller and Mr.
F. Richards of Portal, Arizona, who most kindly permitted me to conduct research on their
lands. This work was done while the author was supported in part by funds from the Theodore
Roosevelt Memorial Foundation and the Explorers Club Educational Fund.

1984

TOOL USE

161

LITERATURE CITED

Alcock, J. 1972. The evolution of the use of tools by feeding animals. Evolution 26:464-473.

Fellers, J. H. and G. M. Fellers. 1976. Tool use in a social insect and its implications for
competitive interactions. Science 192:70-72.

Fowler, H. G. 1982. Tool use by Aphaenogaster ants: a reconsideration of its role in com-
petitive interactions. Bull. New Jersey Acad. Sci. 27:81-82.

Lin, N. 1964-1965. The use of sand grains by the pavement ant, Tetramorium caespitum,
while attacking Halictine bees. Bull. Brooklyn Ent. Soc. 59-60:30-34.

Moglich, M. H. J. and G. D. Alpert. 1979. Stone dropping by Conomyrma bicolor (Hyme-
noptera: Formicidae): a new technique of interference competition. Behav. Ecol. and
Sociobiol. 6:105-113.

Schultz, G. W. 1982. Soil-dropping behavior of the pavement ant, Tetramorium caespitum
(L.) (Hymenoptera: Formicidae) against the alkali bee (Hymenoptera: Halictidae). J.
Kansas Ent. Soc. 55:277-282.

Scott, T. A. and E. H. Melvin. 1953. Determination of dextran with anthrone. Anal. Chem.
25:1656-1661.

Wheeler, W. M. 1910. Ants: their structure, development and behavior. Columbia University
Press, New York.

Received February 28, 1983; accepted October 17, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 162-168

FORAGING BEHAVIOR OF THE BEES
HALICTUS LIGATUS (HYMENOPTERA: HALICTIDAE) AND
CERATINA CALCARATA (HYMENOPTERA: ANTHOPHORIDAE):
FORAGING SPEED ON EARLY-SUMMER
COMPOSITE FLOWERS

Howard S. Ginsberg

Department of Ecology and Evolution, State University of New York,

Stony Brook, New York 11794

Abstract. — Pollen foraging Halictus ligatus and both pollen and nectar foraging Ceratina
calcar ata were timed on early-summer flowers on Long Island in New York State. Pollen-
collecting H. ligatus worked inflorescences of Erigeron annuus (Asteraceae) more rapidly (ave.
time 9.7 sec) than did C. calcarata (ave. time 18.3 sec). Handling time by nectar foragers did
not differ significantly from pollen collectors of C. calcarata. Handling time was longer on
young inflorescences of E. annuus that had numerous florets presenting pollen than on older
inflorescences with fewer florets. Halictus ligatus worked inflorescences of E. annuus faster than
the larger inflorescences of Chrysanthemum leucanthemum (Asteraceae). In addition to working
inflorescences faster, H. ligatus flew between inflorescences faster than did C. calcarata. This
suggests that handling time reflects the general pace of bee movement, which differs among bee
taxa.

The amount of time bees take to forage on flowers is currently of considerable
interest to students of pollination ecology. Indeed, handling time is an important
component of contemporary theoretical approaches to foraging (Schoener, 1971; Pyke
et al., 1977). However, the actual determinants of handling time on flowers are not
yet well understood.

Several factors that influence foraging speed have been identified. There is evidence,
for example, that bees that specialize on a flower species can work that flower more
rapidly than can generalists, as has been shown for Hoplitis anthocopoides (Megachili-
dae) foraging for pollen on its host plant, Echium vulgare (Boraginaceae) (Strickler,
1979). Even among generalized foragers, handling times vary considerably among
bee species on a given flower species. This undoubtedly results, in large part, from
morphological characteristics of the bees and flowers.

Some authors have suggested that nectar foraging speed is directly related to tongue-
length, bees with longer tongues being faster foragers (Brian, 1954; Holm, 1966;
Benedek, 1973; Inouye, 1980). Others have proposed that the fastest foragers are
those whose tongue-length best matches the corolla-length of the flower (Heinrich,
1976; Ranta and Lundberg, 1980). As might be expected, there is a great deal of
variability in the relationship between tongue-length and foraging speed and there
are several complicating factors. One is the influence of learning on foraging speed.
Individual bumble bees, for example, forage more “accurately” (Heinrich, 1979) and
thus more rapidly (Laverty, 1980) as they gain experience working a flower species.
Another is that bees take longer working flowers with large amounts of nectar than
flowers with little nectar (Thomson and Plowright, 1980; Hodges and Wolf, 1981).

1984

FORAGING BEHAVIOR

163

Presumably, ambient temperature can also influence the pace of foraging (Linsley,
1958).

These complicating factors may explain some of the variability in the relationship
between tongue-length and handling time (Ranta and Lundberg, 1980). However, it
may also be that factors unrelated to tongue-length are major determinants of foraging
speed. In the case of pollen-foraging bees especially, handling time may be quite
independent of tongue-length. Among pollen foragers, simple correlations of foraging
speed with morphological characteristics remain elusive. In the present study, I
compare foraging times of two native, nonspecialist bee species on early-summer
flowers in the northeastern United States, and examine some factors that may influ-
ence handling times on these flowers.

MATERIALS AND METHODS

I timed Halictus ligatus Say (Halictidae), a polyphagous, primitively eusocial, soil-
nesting bee (Little, 1977; Michener and Bennett, 1977), on freshly-cut daisy ( Chry-
santhemum leucanthemum, Asteraceae) flowers in soda bottles with water. I set up
5x5 and 7x7 square arrays of regularly distributed flower stalks (50 cm between
stalks) on a mowed field on the State University of New York campus at Stony Brook.
Halictus ligatus can be easily distinguished from most other local bees, but because
of their rapid movements in the field I may have mistakenly included a few moves
by H. rubicundus, a similar species. Also, the process of cutting flowers and placing
them in water undoubtedly influences nectar flow and may have modified handling
times. The H. ligatus foragers were all pollen collectors but they often probed for
nectar as well.

To avoid this problem for samples on fleabane ( Erigeron annuus, Asteraceae), I
found a dense patch of flowers and clipped unneeded flower stalks, leaving a 5 x 5
array (arranged as above) of intact flower stalks, presumably with undisturbed nectar
and pollen flow. This array was located in a clearing in an open woodland in Nisse-
quogue River State Park near Smithtown, Long Island, New York. I timed H. ligatus
and Ceratina calcarata foraging on this flower array, and on 18 June 1981 I used a
hand lens to count the number of florets presenting pollen in each inflorescence in
the array.

Ceratina calcarata Robertson (Anthophoridae) is a polyphagous, solitary species
that nests in plant twigs (e.g., sumac) that are hollowed-out by the females (Daly,
1973; Kislow, 1976). Female C. calcarata cannot be distinguished from C. dupla,
another local species. However, males of these species are easily distinguishable and
all males collected at the study site (N = 15) were C. calcarata. Thus, the females I
studied were probably also C. calcarata.

I sampled C. leucanthemum from 26 May to 8 June, and E. annuus from 13 to
26 June, 1981. I recorded times on and between flower heads to the nearest one-
tenth second with a Cronus digital sports timer. Air temperatures were recorded with
a Springfield outdoor thermometer, hung in the shade approximately 0.5 m above
the ground near the sample site.

I collected individuals of both bee species and placed them in Dietrich’s solution
for tongue-length measurements. I measured the labium of each bee under a stereo
microscope with an ocular scale. I also measured lengths of corolla tubes of florets

164

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

from heads of E. annuus at Nissequogue River State Park in 1 982. Voucher specimens
of the bee species were placed in the Cornell University Insect Collection, lot number

1114.

RESULTS

Disc florets of both flower species open in concentric circles around the center of
the inflorescence, the outermost florets opening first. Bees that are small relative to
the circle of florets follow the circle around the disc collecting nectar and/or pollen.
Relatively larger bees stand in the center of the disc and rotate around the central
axis, collecting resources from each floret. In general, bees make one revolution, but
visits of greater and of less than one revolution are common.

The Halictus ligatus foragers (all pollen collectors) worked Erigeron annuus inflo-
rescences more rapidly (mean = 9.7 sec, SD = 5.9, N = 57) than did pollen foraging
Ceratina calcarata (mean = 18.3 sec, SD = 12.1, N = 37) (Wilcoxon 2-sample test,
ts = 3.566, P < 0.05). In C. calcarata there was no significant difference (ts = 0.717)
in handling time between pollen and nectar foragers (mean for nectar foragers = 20.8
sec, SD = 14.8, N = 77). It is relevant that pollen foragers generally probed florets
for nectar as well. This probing may play a role in pollen collection. On intact stalks
of C. leucanthemum in 1982, 1 observed Ceratina foragers probing florets with their
proboscides, getting pollen dusted onto their heads and mouthparts, and then groom-
ing this pollen onto their scopal hairs. The speed of pollen collecting may thus be
related to the speed of nectar foraging in this genus.

The distance from the base of the prementum to the tip of the glossa is commonly
used as an estimate of effective tongue-length in bees (Heinrich, 1976; Harder, unpubl.
manu.). The prementum plus glossa of C. calcarata (mean length = 2.53 mm, SD =
0.177,N = 10) was slightly longer (t = 7. 12, N = 20, P < 0.01) than that of//, ligatus
(mean length = 2.04 mm, SD = 0. 125, N = 10). These bees also differ in labial mor-
phology; the glossa of C. calcarata is elongate, while that of H. ligatus (a “short-
tongued” bee) is short and relatively obtuse. The average corolla-length of E. annuus
was 1.90 mm (SD = 0.155, N = 50).

Handling time was not correlated with ambient temperature for either bee species
(H. ligatus, r = -0.015, N = 57, P > 0.9; C. calcarata, r = —0.152, N = 155, 0.1 >
P > 0.05; Rohlf and Sokal, 1981 Tables 12 and 25), although visits tended to be rapid
at high temperatures (Fig. 1). Temperature is probably correlated with time of day
and thus with resource levels in flowers, which may have influenced foraging speed.
Handling time was more directly related to the number of open florets on a flower
head (Table 1). Old inflorescences had fewer florets presenting pollen than did young
inflorescences, and bees ( H . ligatus at least) worked old flower heads faster than
young ones. Also, florets in old inflorescences are clustered at the center of the disc
while in young inflorescences they are scattered in a circle around the periphery of
the disc, so bees have to cover more ground to work a young inflorescence.

Handling time by H. ligatus differed on different flower species. The average time
on a daisy inflorescence was 12.7 sec (SD = 10.9, N = 25) while on fleabane it was
9.7 sec(SD = 5.9, N = 57; Wilcoxon 2-sample test, ts = 3.310, P < 0.001). This may
reflect differences in inflorescence size, resource levels, or the number of open florets
in each species, but note that the flower species were sampled at different times of

1984

FORAGING BEHAVIOR

165

90-

Ceratina ca lea rata

r = -0.152

80-

•

70-

•

-

60-

i

50_

: *

z

40-

•

, 1

[ t

o

l

t 1

1

•

30-

•

# •

U 1

w

1 • m

•

§

20-

!

1

1

•

h-

•

t

•

10-

•

1

•

2

t

•

•

1

•

20 21

22 23 24 25 26 27 28 29 30 31

32 33

40-1

30-

20-

10“

Halictus ligatus

r = -0.015

21

23

"I 1

32 33

TEMP. ( °C )

Fig. 1 . Handling times on inflorescences of Erigeron annuus at different temperatures.

the season. Therefore, differences between overwintering gynes and later-generation
workers may also have contributed to this difference in handling time.

Bees took longer to fly between inflorescences on different stalks than between
inflorescences on the same stalk (Table 2). Also, in addition to working inflorescences

166

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Table 1. Handling times of Halictus ligatus (Halictidae) and Ceratina calcarata (Antho-
phoridae) on Erigeron annuus (Asteraceae) as a function of age of inflorescence. (Time in
seconds; sample sizes in parentheses. NS = not significant [Kruskal- Wallis tests].)

Age of
inflorescence

H. ligatus
ave. time on

C. calcarata
ave. time on

Ave. # florets
presenting
pollen per
inflorescence

Young

13.8(13)

22.2 (37)

20.4 (61)

Intermediate

9.7(10)

19.9 (31)

19.5 (51)

Old

3.9 (7)

12.7 (12)

8.7 (34)

Significance

(alpha = 0.05)

*

NS

*

more slowly, C. calcarata flew between inflorescences more slowly than did H. ligatus.
Thus handling time is correlated with the general pace of movement by the bee.

DISCUSSION

The fact that Halictus ligatus spent less time both on and between inflorescences
than did Ceratina calcarata suggests that each species moves at a set pace that
determines the speed of foraging on flowers. In nectar foragers, this pace may be
partially set by the relationship between the tongue-length of the bee and the corolla-
length of the flower (Ranta and Lundberg, 1980). However, the pace may also be set
by any of a number of other factors, especially for pollen foragers. This general pace
of foraging may, in fact, be a genus- or higher-level characteristic of bees. Halictus
may move faster than Ceratina for physiological reasons unrelated to tongue-length.
Intergeneric comparisons should therefore be made only with great care.

Halictus ligatus is larger than C. calcarata (Mitchell, 1960, 1962) and may thus
be able to work an inflorescence more rapidly because it moves less between florets.
Thus body size may be a correlate of foraging speed. Alternately, H. ligatus may
simply work fewer florets per inflorescence than C. calcarata. My impression was
that both species worked about the same number of florets on an inflorescence, but
I have no quantitative data on this. Furthermore, this does not account for the
difference in flight speed between inflorescences.

Table 2. Time spent flying between inflorescences of Erigeron annuus by Halictus ligatus
and Ceratina calcarata. Time in seconds; standard deviation in parentheses. Significance of
differences between flight times in each category by Wilcoxon 2-sample tests.

Average time between inflorescences

On same
stalk

N

On different
stalks

N

Halictus ligatus

0.6 (0.3)

35

1.3 (0.6)

19

P < 0.001

Ceratina calcarata

1.1 (0.9)
P < 0.001

131

2.6 (1.3)
P < 0.001

51

P < 0.001

1984

FORAGING BEHAVIOR

167

The positive correlation of tongue-length with foraging speed found within the
genus Bombus (Holm, 1 966; Inouye, 1 980) does not seem to apply in this case because
the shorter-tongued bee in my study (H. ligatus) was the faster forager. Halictus
ligatus does have a closer match of tongue-length to corolla-length of E. annuus than
does C. calcarata, and this may explain its more rapid pace of foraging. However,
both species collected pollen as well as nectar so tongue-length may be only marginally
important. Nectar-collecting movements play a role in pollen collection by Ceratina
on daisies, but this may not be true for other bees or on other flowers. Furthermore,
my measure of tongue-length (length of prementum plus glossa) is not an accurate
estimate of the effective tongue-length of these bees because they differ as to head
width and labial structure. Halictus ligatus has elongate conjunctival thickenings
basal to the prementum (Michener, 1 944) that allow the mouthparts to swing forward,
so its effective tongue-length may be longer than that of C. calcarata (which has only
a short postmentum). Precise observations are needed on tongue-extension move-
ments of these bees.

Handling time apparently does not vary with temperature, at least within the range
I studied (Fig. 1), but it does differ on different flower species. It is not clear at this
point to what extent the foraging pace on a given flower species varies within a bee
species (in response to resource levels, etc.), and to what extent it is a fixed charac-
teristic of that bee species. Studies of intra- and inter-taxon variability in bee foraging
speeds on flowers with controlled amounts of nectar and pollen will help resolve this
issue.

ACKNOWLEDGMENTS

I thank G. C. Eickwort, L. D. Ginsberg, J. D. Thomson, and an anonymous reviewer for

constructive comments on early drafts of the manuscript. The staff members at Nissequogue

River State Park were always helpful. This is contribution no. 472 in Ecology and Evolution

at the State University of New York at Stony Brook.

LITERATURE CITED

Benedek, P. 1973. Relationship between the tripping rate and the tongue length of lucerne
pollinating wild bees. Z. Angew. Ent. 73:1 13-1 16.

Brian, A. D. 1954. The foraging of bumble bees. Bee World 35:61-67, 81-91.

Daly, H. V. 1973. Bees of the genus Ceratina in America north of Mexico (Hymenoptera,
Apoidea). Univ. Calif. Publ. Ent. 74:1-114.

Heinrich, B. 1976. Resource partitioning among some eusocial insects: bumblebees. Ecology
57:874-889.

Heinrich, B. 1979. “Majoring” and “minoring” by foraging bumblebees, Bombus vagans : an
experimental analysis. Ecology 60:245-255.

Hodges, C. M. and L. L. Wolf. 1981. Optimal foraging in bumblebees: why is nectar left
behind in flowers? Behav. Ecol. Sociobiol. 9:41-44.

Holm, S. N. 1966. The utilization and management of bumble bees for red clover and alfalfa
seed production. Ann. Rev. Ent. 11:155-182.

Inouye, D. W. 1980. The effect of proboscis and corolla tube lengths on patterns and rates of
flower visitation by bumble bees. Oecologia 45:197-201.

Kislow, C. J. 1976. The comparative biology of two species of small carpenter bees, Ceratina
strenua F. Smith and C. calcarata Robertson (Hymenoptera: Xylocopidae). Ph.D. thesis,
Univ. Georgia, Athens, Ga.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Laverty, T. M. 1980. The flower- visiting behaviour of bumble bees: floral complexity and
learning. Can. J. Zool. 58:1324-1335.

Linsley, E. G. 1958. The ecology of solitary bees. Hilgardia 27:543-599.

Litte, M. 1977. Aspects of the social biology of the bee Halictus ligatus in New York State.
Insectes Sociaux 24:9-36.

Michener, C. D. 1944. Comparative external morphology, phylogeny, and a classification of
the bees (Hymenoptera). Bull. Amer. Mus. Natur. Hist. 82:151-326.

Michener, C. D. and F. D. Bennet. 1977. Geographical variation in nesting biology and social
organization of Halictus ligatus. Univ. Kansas Sci. Bull. 51:233-260.

Mitchell, T. B. 1960. Bees of the Eastern United States. Vol. 1. N. Carolina Agr. Exp. Sta.
Tech. Bull. No. 141, 538 pp.

Mitchell, T. B. 1962. Bees of the Eastern United States. Vol. 2. N. Carolina Agr. Exp. Sta.
Tech. Bull. No. 152, 557 pp.

Pyke, G. H., H. R. Pulliam, and E. L. Chamov. 1977. Optimal foraging: a selective review
of theory and tests. Quart. Rev. Biol. 52:137-154.

Ranta, E. and H. Lundberg. 1980. Resource partitioning in bumblebees: the significance of
differences in proboscis length. Oikos 35:298-302.

Rohlf, F. J. and R. R. Sokal. 1981. Statistical Tables, 2nd edition. W. H. Freeman and Co.,
San Francisco, 219 pp.

Schoener, T. W. 1971. Theory of feeding strategies. Ann. Rev. Ecol. Syst. 2:369-404.

Strickler, K. 1979. Specialization and foraging efficiency of solitary bees. Ecology 60: 998-
1009.

Thomson, J. D. and R. C. Plowright. 1980. Pollen carryover, nectar rewards, and pollinator
behavior with special references to Diervilla lonicera. Oecologia 46:68-74.

Received April 19, 1983; accepted November 23, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 169-173

ON THE PSEUDOSCORPION-MIMICKING SPIDER
CHELIFER OWES (ARANEAE: SALTICIDAE)

Norman I. Platnick

Department of Entomology, American Museum of Natural History,

Central Park West at 79th Street, New York, New York 10024

Abstract.— The male of Cheliferoides segmentatus F. O. P.-Cambridge is redescribed and the
female is described for the first time; Cambridge’s hypothesis that these spiders mimic pseudo-
scorpions is supported by the discovery of specimens living with chemetids.

Jumping spiders are well known as mimics; many genera are extremely ant-like
in appearance and behavior (Reiskind, 1977), and others (such as the Neotropical
genus Cylistella ) are convincing impersonators of beetles. F. O. P.-Cambridge (1901),
in describing the male of a Guatemalan jumping spider which he aptly named Che-
liferoides segmentatus, indicated that a more unusual model might be involved: “One
would suspect by its general appearance that this spider mimics one of the Pseudo-
scorpions.” Cambridge’s hypothesis has remained uninvestigated (as has the species),
but it has recently gained support from the discovery of a male and juvenile specimens
living together with several chernetid pseudoscorpions of similar size and appearance
under the bark of mesquite in Maricopa County, Arizona; the pseudoscorpions belong
to Parachernes (Muchmore, in litt.). The flatness of the spiders, the modification of
the first legs into structures resembling pseudoscorpion palpal chelae, and the ab-
dominal pigmentation, which provides a strongly segmented appearance, all con-
tribute to the resemblance (Figs. 1, 2).

Since the original description of Cheliferoides segmentatus, two additional species
have been placed in the genus: C. longimanus Gertsch (1936) from Texas and C.
planus Chickering (1946) from Panama. These species do resemble C. segmentatus
in having an incrassate tibia I, but such modifications occur in a number of other
Neotropical genera, such as Bellota (see Galiano, 1 972a) and Chirothecia (see Galiano,
1972b), where they are generally accompanied by a ventral fringe of thick setae.
Comparison of the genitalia of C. longimanus and C. planus indicates that (as was
indicated for the first species by Richman and Cutler, 1978) neither of them is closely
related to C. segmentatus. Each belongs to a different (and possibly undescribed)
genus of the Bellota complex, but the problem of their proper assignment is beyond
the scope of the present paper. Both the genitalic structure and the presence of a
small caudal extension of the abdomen support instead Wanless’s (1978) hypothesis
of a close relationship between Cheliferoides and Marengo, a genus found in central
and southern Africa, Sri Lanka, Malaysia, Borneo, Java, and the Philippines in which
the front legs are similarly modified.

I am indebted to Dr. W. B. Muchmore for donating the specimens that initially
stimulated this paper. Further material, including the first known females of C.
segmentatus, was obtained from the collection of the American Museum of Natural
History (AMNH) and from Drs. F. Wanless of the British Museum (Natural History)

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Fig. 1. Cheliferoides segmentatus F. O. P.-Cambridge, male, dorsal view.

1984

PSEUDOSCORPION-MIMICKING SALTICID

171

Figs. 2-6. Cheliferoides segmentatus F. O. P.-Cambridge. 2. Cephalothorax and abdomen,
lateral view. 3. Palp, ventral view. 4. Palp, retrolateral view. 5. Epigynum, ventral view. 6.
Epigynum, dorsal view.

(BMNH), W. Pulawski of the California Academy of Sciences (CAS), G. B. Edwards
of the Florida State Collection of Arthropods (FSCA), and H. W. Levi of the Museum
of Comparative Zoology (MCZ). Drs. Wanless and Edwards, as well as Drs. B. Cutler,
M. E. Galiano, C. E. Griswold, and D. B. Richman, generously shared their knowledge
of salticids. The illustrations are by Dr. M. U. Shadab of the American Museum.
The format of the description follows that of Wanless (1978).

Cheliferoides segmentatus F. O. P.-Cambridge
Figs. 1-6

Cheliferoides segmentatus F. O. P.-Cambridge, 1901:254, pi. 22, figs. 12a-f (male
holotype from Guatemala, no specific locality, in BMNH, examined [the holotype
lacks both palpi; a single palp in the holotype vial does not correspond to Cam-
bridge’s figures or to the palpi of other specimens, and belongs to some other
spider]). Richman and Cutler, 1978:84.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Diagnosis. The modifications of leg I (Fig. 1 ), together with the basally widened
bulb and spiral embolus of males (Figs. 3, 4) and the posteriorly coiled epigynal ducts
of females (Figs. 5, 6), are diagnostic.

Male (Arizona). Carapace flattened, reticulate, with evenly distributed white setae,
dark brown, with area enclosed by front two eye rows and extending back along sides
of raised portion of pars cephalica to level of posterior eye row blackened, sides
orangeish except for dark lateral margins (Fig. 1); thoracic groove indicated only by
slight depression. Eyes in three rows, anteriors subcontiguous, with apices forming
distinctly recurved row, middle row slightly closer to posterior row than to anterior;
posterior row wider than anterior; quadrangle occupying almost half of carapace
length. Clypeus low, sloping backwards, with fringe of white squamous setae extending
back to level of coxae I. Chelicerae small, subvertical; promargin with two teeth,
retromargin with one; anterior surface with white squamous setae proximally. Endites
parallel, distally globose, with anterolateral serrula and anteromedian scopula. La-
bium slightly wider than long. Sternum oval, obtuse posteriorly, brown with scattered
darker pigment. Abdomen flattened, without dorsal scutum, light brownish yellow,
dorsum with dark brown markings providing segmental appearance, sides with scat-
tered dark brown spots, venter gray; posterior portion of dorsum with five telescoping
folds preceding elongated anal tubercle forming small caudal projection (Fig. 2);
spinnerets subequal in length, anteriors robust, medians and posteriors slender;
colulus apparently absent. Legs with first pair massive, femur, patella, and tibia
grossly enlarged, tibia with robust spines on well developed socket flanges but without
ventral fringe of stiff setae, metatarsus with distal prolateral spine robust, originating
from distinct cuticular lobe, opposing first tibial spine; remaining legs slender, formula
1432; leg I with femur and patella light brown, tibia dark brown, metatarsi and tarsi
orange, other legs yellow, femora and tibiae with prolateral dark stripes, patellae,
tibiae, and metatarsi with proximal and distal dark rings; spination: femora: II, III
dO-O-1, pO-O-1; IV dO-1-1; tibiae: I vO-3-3; II vO-2-1; metatarsi: I vO-2-2; II vO-1-
0; claw tufts present, scopulae absent. Palp with slender, apically bent tibial apophysis;
tegulum wide basally, with conspicuous spermophore duct and distally coiled em-
bolus (Figs. 3, 4). Dimensions: total length 3.38; carapace length 1.40, width 0.90;
abdomen length 1.84; eyes anterior row 0.78, middle row 0.72, posterior row 0.86,
quadrangle length 0.73. Ratios: AM:AL:PM:PL, 13:6:2:6; AL-PM-PL, 15-13.

Female (Nuevo Leon). As in male, except for the following. Chelicerae without
squamous setae. Abdomen with six posterior folds. Leg I slightly less massive than
in male, with stripes and rings as on legs II-IV; spination: tibia I v0-3-3; metatarsus
I vO-2-2. Palp yellow with prolateral dark stripes on femur and patella. Epigynum
pale (Figs. 5, 6). Dimensions: total length 3.02; carapace length 1.30, width 0.83;
abdomen length 1.69; eyes anterior row 0.71, middle row 0.69, posterior row 0.82,
quadrangle length 0.70. Ratios: AM:AL:PM:PL, 13:6:2:5, AL-PM-PL, 13-12.

Material examined. United States: ARIZONA: Maricopa Co.: Coons Bluff, along
Salt River, Tonto National Forest, Dec. 25, 1980, under bark of mesquite with
pseudoscorpions (L. Merkle, AMNH), 13. Pima Co.: Forestry Cabin, Baboquivari
Mountains, July 18-29, 1951, elevation 3,500 feet (W. S. Creighton, AMNH), 19.
Pinal Co.: Aravaipa Canyon, Mar. 8, 1970 (K. Stephan, FSCA), 13. TEXAS: Comal
Co.: Seancy Estates, New Braunfels, Apr. 12, 1936, tree trunk (S. E. Jones, MCZ),

1984

PSEUDOSCORPION-MIMICKING SALTICID

173

1<3. Frio Co.(?): Frio State Park, Mar. 5, 1952 (W. S. Creighton, AMNH), 16. Uvalde

Co.: no specific locality, May 20, 1938 (Robinson, AMNH), 16. Mexico: NUEVO

LEON: Villa de Santiago, Hacienda Vista Hermosa, June 19, 1940 (H. Hoogstraal,

MCZ), 1$. TAMAULIPAS: Mante, Apr. 17, 1963 (W. J. Gertsch, W. Ivie, AMNH),

16. Guatemala: ESCUINTLA: San Jose, Apr. 3, 1955 (E. I. Schlinger, E. S. Ross,

CAS), 16.

Distribution. Arizona and Texas south to Guatemala.

LITERATURE CITED

Cambridge, F. O. P.-. 1901. Arachnida. Araneida. In: F. D. Godman and O. Salvin, Biologia
Centralia-Americana.

Chickering, A. M. 1946. The Salticidae (spiders) of Panama. Bull. Mus. Comp. Zool. 97:1-
474.

Galiano, M. E. 1972a. Salticidae (Araneae) formiciformes. XIII. Revision del genero Bellota
Peckham, 1892. Physis 31:463-484.

Galiano, M. E. 1972b. Revision del genero Chirothecia Taczanowski, 1878 (Araneae, Saltic-
idae). Rev. Mus. Argentino Cienc. Nat., Ent. 4:1-42.

Gertsch, W. J. 1936. Further diagnoses of new American spiders. Amer. Mus. Novit. 852:1-
27.

Reiskind, J. 1977. Ant-mimicry in Panamanian clubionid and salticid spiders (Araneae:
Clubionidae, Salticidae). Biotropica 9:1-8.

Richman, D. B. and B. Cutler. 1978. A list of the jumping spiders (Araneae: Salticidae) of
the United States and Canada. Peckhamia 1:82-1 10.

Wanless, F. R. 1978. A revision of the spider genus Marengo (Araneae: Salticidae). Bull.
British Mus. Nat. Hist., Zool. 33:259-278.

Received November 22, 1983; accepted December 15, 1983.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 174-178

AUFEIUS IMPRESSICOLLIS (HEMIPTERA: RHOPALIDAE):
EASTERNMOST U.S. RECORD, HOST PLANT
RELATIONSHIPS, AND LABORATORY REARING

A. G. Wheeler, Jr.

Bureau of Plant Industry, Pennsylvania Department of Agriculture,
Harrisburg, Pennsylvania 17110

Abstract.— Aufeius impressicollis St&l (Hemiptera: Rhopalidae), a mainly southwestern U.S.
and Mexican species recorded only as far east as Columbus, Ohio, is reported from Baltimore,
Maryland, on smooth pigweed, Amaranthus hybridus L. (Amaranthaceae). A. impressicollis was
reared on smooth pigweed in the laboratory; its habits are described, and developmental times
of the immature stages are given. Ecological data from museum specimens and a review of the
literature support the hypothesis that amaranths (and possibly members of the related family
Chenopodiaceae) serve as host plants of this little studied rhopalid.

Aufeius impressicollis St&l, belonging to a monotypic genus, is a member of the
rhopalid subfamily Rhopalinae and New World tribe Harmostini, which includes
one other genus, Harmostes Burmeister (26 species) (Gollner-Scheiding, 1978). A.
impressicollis is easily distinguished from Harmostes spp. by the laterally dilated
abdomen and broadly exposed abdominal connexivum (Slater and Baranowski, 1978;
Hoebeke and Wheeler, 1982). It is widely distributed in the western United States
from Nebraska and South Dakota west to Idaho and Washington, and south to
California, Utah, Arizona, New Mexico, and Texas (Torre-Bueno, 1941; U.S. Na-
tional Museum collection); it ranges south through Mexico and into Guatemala
(Brailovsky and Soria, 1981). This rhopalid of probable Sonoran origin (Slater, 1974)
has not been collected frequently in the eastern part of its range. Arizona and Texas
records predominate in the USNM holdings, with considerable material from Col-
orado and California. Froeschner (1942) noted it was uncommon in Missouri; a few
specimens are known from Arkansas and Iowa (USNM).

The first record of A. impressicollis east of the Mississippi was that of Osborn and
Drake (1915), who reported “large numbers” at Columbus, Ohio. Blatchley (1926)
listed single specimens from Marion and Vigo counties, Indiana. No additional Ohio
records of A. impressicollis are available from the Ohio State University collection,
and Columbus remains the easternmost record.

On 14 July 1983 I collected 3 adults of A. impressicollis at Baltimore, Maryland,
a record that extends the distribution eastward more than 350 miles (560 km). The
specimens were swept from weeds in a vacant lot on Boston Street in an industrial
area along the Northwest Branch of the Patapsco River. Seven adults were observed
at the same site on 6 August 1983. Specimens have been deposited in the insect
collections of the Pennsylvania Department of Agriculture (PDA), Cornell University
(CUIC), and U.S. National Museum of Natural History (USNM).

It is possible that the Baltimore population is the result of a natural eastward
dispersal, even though no records are available for Ohio since 1915 and no specimens
have been recorded east of Columbus. Knowledge of heteropteran distributions in

1984

AUFEIUS- BIOLOGY AND DISTRIBUTION

175

North America is fragmentary, even for areas of the presumed well-collected eastern
states, and reports of range extensions of hundreds of miles are not uncommon.
However, it is equally likely that the population at Baltimore is adventive, the result
of an introduction with commerce. Oil terminals, warehouses, and numerous railroad
lines are present near the collection site of A. impressicollis. This area of the Port of
Baltimore (Canton) is notable for the large number of exotic plant species that have
been collected in ballast dumps along the port (Reed, 1964).

HOST PLANTS AND HABITS

Before returning to the original collection site, I reviewed the literature to determine
the host plants most likely to be used by Aufeius impressicollis in Baltimore. It soon
became apparent that little ecological information is available for this species. Schae-
fer and Chopra (1982) and Schaefer and Mitchell (1983) do not list A. impressicollis
in their review of coreoid host plants. This rhopalid has been collected by sweeping
bushes (Uhler, 1877), grasses (Osborn and Drake, 1915), borders of a timothy mead-
ow (Blatchley, 1926), and weedy fields (Froeschner, 1942). Some of the specific plants
mentioned in the literature may not represent true hosts: sugar beet, Beta vulgaris
L. (Chenopodiaceae) (Knowlton, 1933); another chenopod, Russian thistle, Salsola
iberica Sennen & Pau (Goeden and Ricker, 1968); and alfalfa, Medicago sativa L.
(Fabaceae) (Benedict and Cothran, 1975). Label data from western specimens in the
USNM collection provided three additional records from sugar beet, two from alfalfa,
and one from celery [Apium graveolens L.— Apiaceae], “beans” [probably Phaseolus
vulgaris L.— Fabaceae], and the chenopod Salsola kali L. (cited as S. pestifer). A
specimen from Texas in the Texas A&M collection was taken on cotton [Gossypium
hirsutum L.— Malvaceae]. Other labels on USNM specimens suggested a preference
for plants of the Amaranthaceae. Specimens from California (Chico and Lindsay)
had been collected on tumbleweed or tumble pigweed, Amaranthus albus L.; one
specimen from Victoria, Texas was labeled “amaranthus”; and two from Garden
City, Kansas had been taken on “pigweed” [probably Amaranthus sp.].

The vacant lot had been mowed when I returned to Baltimore on 6 August, and
Aufeius impressicollis was not collected by sweeping the cutover weeds. Various
unmowed plants growing along a brick wall and a building were then sampled in-
dividually by tapping vegetation over a small tray. Seven adults of the rhopalid were
collected by tapping flower spikes of smooth pigweed, Amaranthus hybridus L. Three
early-instar rhopalids that were observed but subsequently lost may have represented
this species. No other plants, including Chenopodium ambrosioides L. of the related
family Chenopodiaceae, yielded A. impressicollis.

Rhopalids, when collected by sweeping or when beaten onto a sheet or tray, initially
may be sluggish but usually within seconds become active. Similar behavior was
exhibited by the other Rhopalidae taken at the Baltimore site: Harmostes reflexulus
(Say), Liorhyssus hyalinus (F.), and Rhopalus ( Brachycarenus ) tigrinus1 (Schilling).

1 Maryland is a new state record for this Old World rhopalid recently recorded in North
America from New Jersey, New York (Long Island), and Pennsylvania (Hoebeke and Wheeler,
1982).

176

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Table 1 . Duration (in days) of the immature stages of Aufeius impressicollis reared in the
laboratory at 22-26°C.

Stage

No. of
observations

Range

Mean ± SE

Cumulative
mean age

Egg

22

8

8.0 ± 0

8.0

Nymphal stages
I

13

3-7

4.2 ± 0.27

12.2

II

12

2-4

2.8 ± 0.22

15.0

III

12

2-4

2.9 ± 0.15

17.9

IV

11

3-10

4.5 ± 0.58

22.4

V

7

5-8

6.3 ± 0.36

28.7

However, adults of A. impressicollis usually feigned death after being knocked from
pigweed onto a tray or when the contents of a sweep net were emptied.

In the laboratory under conditions of natural photoperiod (window light) and room
temperature (22-26°C), adults were placed in small plastic boxes with a water source
and excised flower spikes of smooth pigweed that included a few leaves. Mating and
oviposition readily occurred, and the biological notes that follow are based on a
limited number of observations on the resulting progeny. In a forthcoming paper on
immature stages of eastern Rhopalidae by E. R. Hoebeke and A.G.W., the egg and
fifth-instar nymph of A. impressicollis will be described and illustrated.

The premating period for each of the two pairs observed was 4 days. Mating pairs
assumed an end-to-end position that apparently is typical for the family (Paskewitz
and McPherson, 1983) and often remained in copula for 4 hours or longer. One of
the pairs mated 6 times; the other, 4 times. The preoviposition period was 2 days.
Eggs were laid mainly on the flower spikes and on the sides and bottom of the rearing
containers; a few were deposited on host foliage. One of the females laid 48 eggs
over an 8 -day period; the other, 34 eggs during 3 days. The incubation period was
8.0 days, and the total developmental period for the nymphal stages averaged 20.7
days (Table 1). Nymphs and adults fed only on floral structures, including seeds that
had dropped onto leaves or to the bottom of the rearing containers.

The laboratory rearing of Aufeius impressicollis on Amaranthus hybridus, coupled
with its collection on this plant at Baltimore and the records from amaranths in the
western U.S., seems to establish members of the Amaranthaceae as host plants. In
addition, I later discovered that Stegmaier (1950) collected 18 adults of A. impres-
sicollis (and observed additional specimens) on redroot or rough pigweed, Amaran-
thus retroflexus L., in Kansas during June-August 1949. Collections from alfalfa,
beans, celery, cotton, and other plants not closely related to the Amaranthaceae or
Chenopodiaceae may reflect “sitting” records, or possibly were made from weedy
amaranths growing in crop fields. Several Amaranthus spp., including A. hybridus
and A. retroflexus, are common weeds of arable land (Muenscher, 1980). Studies on
this rhopalid in western North America are needed to verify the apparent amaranth
feeding trend and to determine whether chenopods serve as hosts.

Among the Coreoidea, several coreine genera are associated with plants of the

1984

A UFEIUS— BIOLOGY AND DISTRIBUTION

177

order Caryophyllales, including Amaranthaceae, but no North American rhopalid is
known from this order (Schaefer and Chopra, 1982; Schaefer and Mitchell, 1983).
Species of Harmostes, the only other genus in the Harmostini, feed mainly on com-
posites (Schaefer and Chopra, 1982; Schaefer and Mitchell, 1983). Aufeius is closely
related to Harmostes but differs in several morphological characters (Chopra, 1967;
Schaefer and Chopra, 1982). It appears also that the two genera differ in their host
plant preferences and in one aspect of their behavior.

ACKNOWLEDGMENTS

For checking collections in their care for distribution records and host data for A. impressicollis
I thank H. Brailovsky (Instituto de Biologia, UNAM), T. J. Henry (Systematic Entomology
Laboratory, USDA, % U.S. National Museum, Washington, D.C.), J. C. Schaffner (Department
of Entomology, Texas A&M University, College Station), R. T. Schuh (Department of Ento-
mology, American Museum of Natural History, New York, N.Y.), and C. A. Triplehom (De-
partment of Entomology, Ohio State University, Columbus). R. J. Hill (PDA) kindly identified
plant specimens, and Henry and E. R. Hoebeke (Department of Entomology, Cornell University,
Ithaca, N.Y.) offered helpful comments on the manuscript.

LITERATURE CITED

Benedict, J. H. and W. R. Cothran. 1975. A faunistic survey of Hemiptera-Heteroptera found
in northern California hay alfalfa. Ann. Ent. Soc. Amer. 68:897-900.

Blatchley, W. S. 1926. Heteroptera or True Bugs of Eastern North America, with Especial
Reference to the Faunas of Indiana and Florida. Nature Publ. Co., Indianapolis, Indiana,

1116 pp.

Brailovsky, H. and F. Soria. 1981. Contribucion al estudio de los Hemiptera-Heteroptera de
Mexico: XVIII Revision de la tribu Harmostini Stcil (Rhopalidae) y descripcion de una
especie. An. Inst. Biol. Univ. Nal. Auton. Mexico 51, Ser. Zool. (1): 123-1 68.

Chopra, N. P. 1967. The higher classification of the family Rhopalidae (Hemiptera). Trans.
R. Ent. Soc. Lond. 119:363-399.

Froeschner, R. C. 1942. Contributions to a synopsis of the Hemiptera of Missouri, Pt. II.

Coreidae, Aradidae, Neididae. Am. Midi. Nat. 27:591-609.

Goeden, R. D. and D. W. Ricker. 1968. The phytophagous insect fauna of Russian thistle
(Salsola kali var. tenuifolia ) in southern California. Ann. Ent. Soc. Amer. 61:67-72.
Gollner-Scheiding, U. 1978. Revision der Gattung Harmostes Burm., 1835 (Heteroptera,
Rhopalidae) und einige Bermerkungen zu den Rhopalinae. Mitt. Zool. Mus. Berlin 54:
257-311.

Hoebeke, E. R. and A. G. Wheeler, Jr. 1982. Rhopalus ( Brachycarenus ) tigrinus, recently
established in North America, with a key to the genera and species of Rhopalidae in
eastern North America (Hemiptera: Heteroptera). Proc. Ent. Soc. Wash. 84:213-224.
Knowlton, G. F. 1933. Notes on Utah Heteroptera. Ent. News 44:261-264.

Muenscher, W. C. 1980. Weeds, 2nd Edition Reprinted. Cornell University Press, Ithaca,
New York, 586 pp.

Osborn, H. and C. J. Drake. 1915. Additions and notes on the Hemiptera-Heteroptera of
Ohio. Ohio Nat. 15:501-508.

Paskewitz, S. M. and J. E. McPherson. 1983. Life history and laboratory rearing of Arhyssus
lateralis (Hemiptera: Rhopalidae) with descriptions of immature stages. Ann. Ent. Soc.
Amer. 76:477-482.

Reed, C. F. 1964. A flora of the chrome and manganese ore piles at Canton, in the Port of

178

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Baltimore, Maryland and at Newport News, Virginia, with descriptions of genera and
species new to the flora of eastern United States. Phytologia 10:321-406.

Schaefer, C. W. and N. P. Chopra. 1982. Cladistic analysis of the Rhopalidae, with a list of
food plants. Ann. Ent. Soc. Amer. 75:224-233.

Schaefer, C. W. and P. L. Mitchell. 1983. Food plants of the Coreoidea (Hemiptera: Heter-
optera). Ann. Ent. Soc. Amer. 76:591-615.

Slater, J. A. 1974. A preliminary analysis of the derivation of the Heteroptera fauna of the
northeastern United States with special reference to the fauna of Connecticut. Mem.
Conn. Ent. Soc., 1974, pp. 145-213.

Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs (Hemiptera-Heter-
optera). Wm. C. Brown Co. Publ., Dubuque, Iowa, 256 pp.

Stegmaier, C. E., Jr. 1950. Insects associated with the rough pigweed, Amaranthus retroflexus
L. (Amaranthaceae). M.S. thesis, Kansas State Coll., Manhattan, 1 13 pp.

Torre-Bueno, J. R. de la. 1941. A synopsis of the Hemiptera-Heteroptera of America north
of Mexico. Part II. Families Coreidae, Alydidae, Corizidae, Neididae, Pyrrhocoridae and
Thaumastotheriidae. Ent. Amer. 21:41-122.

Uhler, P. R. 1877. Report upon the insects collected by P. R. Uhler during the explorations
of 1875, including monographs of the families Cydnidae and Saldae, and the Hemiptera
collected by A. S. Packard, Jr., M. D. Bull. Geol. Geogr. Surv. Terr. 3:355-475.

Received February 22, 1984; accepted April 10, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 179-183

COMPSIDOLON ( CONIOR T ODES) SALICELLUM
(HEMIPTERA: MIRIDAE): A PREDACEOUS PLANT BUG, NEW
TO THE UNITED STATES, FOUND ON FILBERT

John D. Lattin1 and Russell H. Messing

Systematic Entomology Laboratory, Department of Entomology,

Oregon State University, Corvallis, Oregon 97331

Abstract. — Compsidolon ( Coniortodes ) salicellum is recorded for the first time from in the
United States, feeding on aphids on cultivated filberts in western Oregon and Washington.

The natural range of Compsidolon ( Coniortodes ) salicellum (Herrich-Schaeflfer)
extends from Great Britain east to western Russia and Norway, south to the Med-
iterranean region (Kiritschenko, 1951; Wagner and Weber, 1964; Wagner, 1975).
Although the species is predaceous (Southwood and Leston, 1959; MacPhee and
Sanford, 1961; Wagner and Weber, 1964; and Kelton, 1982), it is associated with
relatively few plant species. The recorded plants include Corylus (Reuter, 1884;
Stichel, 1933; Southwood and Leston, 1959; Wagner and Weber, 1964; Wagner,
1975; and Kelton, 1982), Salix (Reuter, 1884; Stichel, 1933; and Kerzhner, 1964),
Lonicera (Stichel, 1933), Alnus (Southwood and Leston, 1959), Malus (Southwood
and Leston, 1959; MacPhee and Sanford, 1961; Sanford and Herbert, 1967; and
Herbert and Sanford, 1 969), sallow ( Salix spp.) and bramble ( Rubus spp.) (Southwood
and Leston, 1959) and raspberry (Rubus sp.) and thimbleberry ( Rubus parviflorus)
(Kelton, 1982). There are a number of other species of the subgenus Coniortodes in
Europe. These species occur on a variety of plants including species of Adenocarpus,
Cytisus, Salvia and Artemesia (Wagner, 1975).

Compsidolon ( Coniortodes ) salicellum overwinters in the egg stage as do most
mirids (Southwood and Leston, 1959). There is a single generation per year and the
adults are found from June to September (Wagner, 1975). Southwood and Leston
(1959) report the prey to be mites (e.g. Bryobia praetiosa Koch) and other small
animals. MacPhee and Sanford (1961) considered the mirid (reported as Psallus sp.)
to be predaceous, although they did not specify the exact prey species. Sanford and
Herbert (1967) in their work on the influence of spray programs for phytophagous
mites, including the European red mite, Pananychus ulmi (Koch), the blister mite,
Vasates schlectendali (Nal.) and the brown mite, Bryobia arborea (M. & A.), included
salicellum (reported as Coniortodes salicellus) among the list of predators. Herbert
and Sanford (1969) reported that salicellum (as Coniortodes salicellus) fed on the
apple rust mite, Vasates schlechtendali (Nal.), and Kelton (1982) reported salicellum
(as Psallus salicellus) feeding on aphids. The present paper reports salicellum as a
predator of the filbert aphid, Myzocallis coryli (Goetze) in western Oregon.

Specimens of C. (C.) salicellum were first collected from filbert orchards in the

1 Oregon Agricultural Experiment Station Paper No. 7104.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

vicinity of Corvallis, Oregon during the summer of 1981 when one of us (R.H.M.)
undertook a study of the predator complex on commercial filbert trees. The true
identity of the species was not worked out until early in 1982 (by J.D.L.). Because
of confusion with other mirid nymphs, the exact date of egg hatch is not known but
since the first adults appeared in early July, nymphs must have been present at least
from the middle of June. During the 1 982 season, Lattin, Gary M. Stonedahl (Oregon
State University) and Thomas J. Henry (USDA-ARS Systematic Entomology Lab-
oratory) collected adults and nymphs (third through fifth instars) on July 1 from
filbert trees in the vicinity of Corvallis. While the peak occurrence of the adults was
the end of July 1982, adults could be collected from the orchards until early October.

Adult bugs were observed feeding on the filbert aphid, and although no direct
feeding observations of the nymphs were made, single second or third instar nymphs
caged with aphids are believed to have consumed an average of 140 aphids each
during a fifteen-day period at 24°C. In one laboratory trial, five out of six third and
fourth instar nymphs reached the adult stage with only filbert leaf tissue for food.
Samples taken from two untreated orchards showed that the populations of adults
peaked in late July with about 5 individuals per tree sample, obtained by sharply
tapping three tree limbs over a 30 x 30 inch canvas beating sheet. An orchard treated
with Metasystox-R® in late April showed peak populations of less than one adult
per tree sample whereas one orchard treated with Guthion® in early July showed a
peak of 17 adults per tree sample at the end of July (Messing, 1983).

Compsidolon ( Coniortodes ) salicellum was first reported from North America from
Nova Scotia, Canada, by MacPhee and Sanford in 1961 (as Psallus sp.). It was
reported as Coniortodes salicellum from the same locality by Sanford and Herbert
(1967) and Herbert and Sanford (1969). Kelton (1982) reported it as Psallus salicellus,
citing the specific Nova Scotia locations referred to above and added Prince Edward
Island and British Columbia, Canada, localities, thus establishing the occurrence of
the species on both coasts. We have seen additional specimens from Oregon and
Washington.

Specimens examined. OREGON: Benton Co., Corvallis, 3 July 81, on Corylus
avellana, coll. R. Messing (OSU); Benton Co., 5 mi S. Philomath, 13 July 81, on
Corylus avellana, coll. R. Messing (OSU); Corvallis, 1 July 82, on Corylus avellana,
coll. T. Henry and J. D. Lattin (USNM: OSU); Corvallis, Oregon State University,
campus, 20 July 82, on Corylus, coll. G. M. Stonedahl (OSU). WASHINGTON:
Lewis Co., Chehalis, 110 Urquhart Rd., 22 July 79, on Corylus sp.) coll. G. M.
Stonedahl (GMS, OSU).

The comprehensive review of the Miridae of the Mediterranean region by E.
Wagner (1975) contains a thorough review of the genus Compsidolon Reuter including
the four subgenera Compsidolon, Apsinthophylus E. Wagner, Chamaeliops E. Wagner
and Coniortodes E. Wagner, complete with keys and many illustrations. His classi-
fication is followed here.

The taxonomic history of C. (C.) salicellum is complex. Carvalho (1958) presented
a synonymical bibliography complete to the date of his publication. Additional work
was done by E. Wagner (1952, 1975), E. Wagner and Weber (1964), and Kelton
(1982). A condensed taxonomic and biological bibliography is presented below to
summarize these prior contributions.

1984

COMPSIDOLON SAUCE LLUM

181

Fig. 1. Adult Compsidolon ( Coniortodes ) salicellum (Herrich-Schaeffer).

182

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

Compsidolon ( Coniortodes ) salicellum (Herrich-Schaeffer)

Capsus salicellus Herrich-Schaeffer, 1841:47, fig. 605.

Psallus salicellus: Fieber, 1861:305; Reuter, 1884: 146; Stichel, 1933:272; Kiritschen-
ko, 1951:206; Carvalho, 1958:130; Kelton, 1982:171-172; Kelton, 1983:140-142.
Eurymerocoris salicellus: Walker, 1873:151.

Psallus ( Coniortodes ) salicellum: E. Wagner, 1952:172; Southwood and Leston, 1959:
227, pi. 33, fig. 2.

Coniortodes salicellum: E. Wagner, 1954:9; Kerzhner, 1964:752, fig. 309, 328; San-
ford and Herbert, 1967:693-696; Herbert and Sanford, 1969:62, 64-65.

Psallus sp., MacPhee and Sanford, 1961:671-673.

Compsidolon salicellum: E. Wagner and Weber, 1964:489, fig. 258a, 259a-e.
Compsidolon ( Coniortodes ) salicellum: E. Wagner, 1975:151, fig. 729a, 731, 734g.

Compsidolon salicellum keys to Psallus (couplet 183) in Slater and Baranowski
(1978), but with some difficulty. Specimens are often badly rubbed, removing most
of the pubescence, at least on the dorsal surface. The head is rather strongly deflexed
downward and thus the tylus is not always visible from above. The tip of the rostrum
attains the middle of the abdomen and the sexes are not dimorphic. The width of
the vertex is greater than half the width of the head (4:7) and the length of antennal
segment II is slightly greater than the basal width of the pronotum (12.5:11). The
color pattern of small, regularly distributed brown spots on the clavi and coria of
the hemelytra are distinctive, together with the pattern on the membrane. As Wagner
and Weber (1964) and many others have pointed out, much work remains to be
done in clarifying the generic limits of Psallus and closely related genera.

Compsidolon is a large genus and the host range of its known species is wide. It
seems likely that additional species of this group of plant bugs will be detected in
the North American fauna, either as introductions or as naturally occurring species.
The western and southwestern portions of the United States, and particularly Cali-
fornia, would seem appropriate areas for their occurrence. The planned investigations
of the Phylinae of western North America by R. T. Schuh of the American Museum
of Natural History should clarify the matter.

ACKNOWLEDGMENTS

We thank Bonnie B. Hall for the illustration of the adult, the Oregon Filbert Commission
for financial assistance, M. T. AliNiazee for assistance and suggestions, Thomas J. Henry,
USDA-ARS, Systematic Entomology Laboratory for the loan of some literature and specimens,
and Gary M. Stonedahl for the loan of additional specimens from Washington state and for
technical assistance.

LITERATURE CITED

Carvalho, J. C. M. 1958. Catalogo dos Mirideos do Mundo. Parte II. Arquivos do Museu
Nacional, Rio de Janeiro 45:1-216.

Fieber, F. X. 1861. Die Europaischen Hemiptera. Halbfluger (Rhynchota Heteroptera). Vi-
enna, 444 pp.

Herbert, H. J. and K. H. Sanford. 1969. The influence of spray programs on the fauna of
apple orchards in Nova Scotia. XIX. Apple rust mite, Vasates schlechtendali, a food
source for predators. Can. Ent. 101:62-67.

1984

COMPSIDOLON SALICELLUM

183

Herrich-Schaeffer, G. A. W. 1841. Die Wanzenartigen Insecten. Numberg. Vol. 6:1-1 17.

Kelton, L. A. 1982. Description of a new species of Plagiognathus Fieber, and additional
records of European Psallus salicellus in the Nearctic Region (Heteroptera: Miridae).
Can. Ent. 114:169-172.

Kelton, L. A. 1983. Plant Bugs on Fruit Crops in Canada. Heteroptera: Miridae. Agriculture
Canada, Monograph No. 24, 201 pp.

Kerzhner, I. M. 1 964. In: Bei-Bienko, G. Y. (ed.), Keys to the Insects of the European USSR.
Volume I. Apterygota, Paleoptera, Hemimetabola. Academy of Sciences of the USSR,
Zoological Institute, Moscow-Leningrad, 874 pp. (In Russian).

Kiritshenko, A. N. 1951. True Hemiptera of European USSR. Academy of Sciences of the
USSR, Zoological Institute, Moscow-Leningrad, 423 pp. (In Russian).

MacPhee, A. W. and K. H. Sanford. 1961. The influence of spray programs on the fauna of
apple orchards in Nova Scotia. XII. Second supplement to VII. Effects on beneficial
arthropods. Can. Ent. 93:671-673.

Messing, Russell H. 1 983. The biology of the predator complex of the filbert aphid, Myzocallis
coryli Goetze, in Western Oregon. M.S. thesis, Oregon State University, 115 pp.

Reuter, O. M. 1884. Hemiptera Gymnocerata Europae. Part I. Acta Soc. Sci. Fennicae 13:
1-568.

Sanford, K. H. and H. J. Herbert. 1967. The influence of spray programs on the fauna of
apple orchards in Nova Scotia. XVIII. Predator and prey populations in relation to
miticides. Can. Ent. 99:689-696.

Slater, J. A. and R. M. Baranowski. 1978. How to Know the True Bugs. W. C. Brown Company,
Dubuque, Iowa, 256 pp.

Southwood, T. R. E. and Dennis Leston. 1959. Land and Water Bugs of the British Isles.
Frederick Wame and Company, Ltd., London, 436 pp.

Stichel, W. 1933. Illustrierte Bestimmungstabellen der Deutschen Wanzen (Hemiptera-Het-
eroptera). Berlin, Lieferung 9:243-274.

Wagner, E. 1952. Blindwanzen oder Miriden. Teil 4 1 . In: F. Dahl, Die Tierwelt Deutschlands.
Jena, 218 pp.

Wagner, E. 1954. Psallus Fieb. subgen. Coniortodes nov. subgen. (Heteroptera Miridae). Soc.
Sci. Fennicae, Comm. Biol. 14(3): 1-10.

Wagner, E. 1975. Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen
Inseln (Hemiptera, Heteroptera). Teil 3. Entomologische Abhandlungen Staatliches Mu-
seum Tierkunde Dresden. Supplement Vol. 40:1-483.

Wagner, E. and H. H. Weber. 1964. Faune de France. No. 67. Heteropteres. Miridae. Paris,
589 pp.

Walker, F. 1873. Catalogue of the specimens of Hemiptera Heteroptera in the collections of
the British Museum. London. Vol. 6:1-210.

Received March 5, 1984; accepted March 20, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 184-187

NOTES AND COMMENTS

NOTES ON THE DISTRIBUTION OF DERMAPTERA IN
SOME COSTA RICAN CACAO PLANTATIONS

Annual cycles of wet and dry seasons in the tropical rain forest zone of eastern
Costa Rica alter the distribution and abundance of leaf litter-inhabiting insects in
cacao plantations (Young, A.M. 1982. J. Appl. Ecol. 19:47-63; 1983a. J. Appl. Ecol.
20:in press; 1983b. J. New York Ent. Soc. 9 1 :in press; 1983c. Sociobiology 8:51-
76). Few data, however, are available on the effects of tropical seasonality on litter-
inhabiting Dermaptera. In this note I summarize some recent observations on earwigs
associated with rotting sections (slices) of banana tree trunks in some Costa Rican
cacao plantations.

Earwigs in rotting sections of banana tree trunks distributed on the ground (each
piece about 6-8 cm high x 15-18 cm dia.) were studied during 1981 and 1982 in
these cacao plantations on the Atlantic or eastern watershed of Costa Rica: “La
Tigra”, near La Virgen (10°23'N, 84°07'W; 220 m elev.), Heredia Province; “Tur-
rialba” or CATIE, in Turrialba (9°54'N, 83°41'W; 600 m elev.); “La Lola”, near
Siquirres (10°06'N, 83°30'W; 50 m elev.). Between 200 and 300 freshly-cut slices of
banana tree trunks were placed in piles of 20-30 pieces each, in the leaf litter (details
of these studies also given in Young, 1982, 1983a, op. cit.) three times each year.
Earwigs were collected from the rotten slices along with occasional observations on
these insects in leaf litter.

Two families, Carcinophoridae and Labiidae, were represented in the samples, the
former by four genera and seven species, and the latter by one genus and species
( Marava triquetra Hebard). Although a total of seven species were collected from La
Lola, three from Turrialba, and two from La Tigra, no one genus or species occurred
at all three localities (Table 1). Carcinophora was represented by three species, and
all other genera by one or two species each. A total of 146 individuals (all species
combined) were collected for all three plantations and dates, with three species having
similar proportions in abundance, ranging from 22-27 percent. These species are:
Euborellia annulipes Lucas (about 21.3 percent), Anisolabis maritima Bonelli (about
24.0 percent), and M. triquetra Hebard (about 27.3 percent). A fourth species, Car-
cinophora gagatina Klug, comprised another 12.3 percent of the sample, and all
remaining species were far less numerous (Table 1). Interestingly, and in spite of the
very small sample size and the equal sample sizes between seasons, more than five
times the number of individuals (114) were collected in the dry season (Fig. 1 ) samples
for all localities combined, and these were distributed among six species. By contrast,
only 25 insects were collected in the wet season samples, representing five species.
Of the very diverse dry season samples, about 35 percent were M. triquetra, 30
percent A. maritima, and 14 percent C. gagatina, for all three plantations combined.
One species, Isolabis howardi Burr, was found only in the Turrialba dry season
samples (Table 1). Collections of leaf litter during the wet season yielded 1-5 indi-
viduals of various species, for a total of five 1 x 1-m x 10 cm deep samples of cacao

Table 1. Distribution and abundance of earwigs (Dermaptera) in rotting cut slices of banana tree trunks at three Costa Rican cocoa plantation
localities during rainy and dry seasons.

1984

NOTES AND COMMENTS

185

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The mark “ — ” means that these species were not found in samples collected at the indicated locality.

186

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

1980 1981

SUCCESSIVE MONTHS

1984

NOTES AND COMMENTS

187

leaves collected each time (N = 4 collections), while similar collections in the dry
season produced only 0-1 earwigs.

These data strongly suggest that “diversity” of earwigs in rotting sections of banana
tree trunks, that is, numbers of individuals (abundance) weighted by the number of
species, greatly increases during the tropical dry season in these cacao plantations,
even though the dry season is short and irregular (Fig. 1). The immature stages of
Diptera such as Ceratopogonidae exhibit very high densities in rotting slices of banana
tree trunks during the dry season in these cacao plantations (e.g., Young 1983a, op.
cit.). Quite possibly, these microhabitats remain moist as the surrounding leaf litter
dries out, thereby establishing mini-ecological refugia for insects such as earwigs.

This research is a by-product of grants from the American Cocoa Research Institute
of the United States of America. I thank Drs. Gustavo A. Enriquez and J. Robert
Hunter for logistical field assistance. Dr. H. Steinmann (Zoological Department,
Hungarian Natural History Museum, Budapest) made the determinations of the
Dermaptera cited in this paper, and voucher specimens are deposited in his collec-
tions. Tammy McCarthy prepared Figure 1 .—Allen M. Young, Invertebrate Zoology
Section, Milwaukee Public Museum, Milwaukee, Wisconsin 53233.

Fig. 1. Monthly patterns of rainfall at three localities in eastern Costa Rica. There is a
tendency for drier conditions to prevail during the period January through April each year at
these localities, although the “dry season” is very variable from year to year in this region of
Costa Rica.

NEW YORK ENTOMOLOGICAL SOCIETY
92(2), 1984, pp. 188-192

BOOK REVIEW

Naturalists’ Handbooks.— 1983. Cambridge University Press, New York. 65 pp. and

$16.95 each.

1 . Insects on Nettles by B. N. K. Davis.

2. Grasshoppers by Valerie K. Brown.

3. Solitary Wasps by Peter F. Yeo and Sarah A. Corbet.

4. Insects and Thistles by Margaret Redfem.

In keeping with the great tradition of British natural history, a series of handbooks
has been initiated to promote original field studies of insects. Written by professional
entomologists, the books are aimed at “sixth-formers” (students in their twelfth and
thirteenth year who are studying to gain entrance into a university— roughly equiv-
alent to American college freshmen and sophomores), and others without university
training. They are intended to “draw attention to regions on the frontiers of current
knowledge where amateur studies have much to offer” and to show that “natural
history need not be a spectator sport.” The Naturalists’ Handbooks are of two types:
guides to the fauna associated with common plants (nettles, thistles) or reviews of
specific taxonomic groups (grasshoppers, solitary wasps). With the emphasis on nat-
ural history in British schools, there is a ready market for the series.

But North American entomologists should not overlook these handbooks. Because
our fauna contains a substantial Old World element— naturally Holarctic species and
those introduced with man’s commerce— the North American reader is likely to
encounter familiar insects, although some of the British common names may seem
strange (e.g. “Cuckoo-spit insect” for the meadow spittlebug, Philaenus spumarius).
Federal and state regulatory personnel and others concerned with detecting exotic
insects should find useful information on habits of common British species and
illustrations helpful in recognizing potential additions to our fauna. High school
teachers and 4-H leaders might use the books to introduce students to fascinating
field studies involving insects, and college students taking introductory courses in
biology or entomology will find helpful suggestions for research projects. Even profes-
sional entomologists interested in insect-plant interactions might find information
useful in conducting surveys of particular plant species.

Insects on Nettles, the first book of the series, contains introductory material on
stinging nettle, Urtica dioica L., and small nettle, U. urens L., and relates them to
other plants of the order Urticales. The reader is given insight into the plants’ mor-
phology and habitat preferences and is told that stinging nettle, a dioecious species,
hosts seed-feeding insects that occur only on female plants and pollen feeders that
are found on male plants. Small nettle has separate male and female flowers on the
same plant. This first chapter also introduces the binomial system of nomenclature
and briefly discusses insect orders and their identification. In tabular form the orders
and numbers of species associated with nettles are summarized; of 107 nettle asso-
ciates, 3 1 are “restricted” species more or less confined to stinging nettle and related
plants. The introduction concludes with the advice that students “concentrate initially
on the most common species as these are likely to be part of the true nettle fauna
and not just chance visitors.”

1984

BOOK REVIEW

189

Chapter 2, “Keys to the Insects on Nettles,” forms a major portion of the book.
First, hints are given for using artificial, dichotomous keys. The author is well aware
that “many other insects which are not included in the keys will be collected from
nettles; these are probably just sheltering amongst the dense vegetation or they may
be insects which are really associated with other low plants or trees growing nearby.”
He cautions that the reader “ must therefore be prepared to reject a specimen as being
merely a casual visitor on nettles if it does not fit any of the possibilities given, though
the keys may allow you to place it in a family or genus.” Notes on insect structure,
size, color, seasonal occurrence, and sex differences make the keys easier to use.

The major groups of nettle insects then are keyed and separate keys are presented
for Lepidoptera adults and larvae; Coleoptera; adult Heteroptera and families of
heteropteran nymphs (which the British usually term “larvae”); Homoptera; free-
living larvae of Diptera, Coleoptera, and Neuroptera; leafmining and gall-making
Diptera; and parasites. The keys, replete with simple but effective line drawings
presented with most couplets, should enable the amateur to identify most nettle-
associated insects— parasites and predators as well as herbivores. For most nettle
associates a summary of life history is given in the keys by indicating the months
when lepidopteran and dipteran larvae can be found on the plants, and the occurrence
on nettles of adults belonging to other groups. Also useful are five plates of adult or
immature insects, including four color plates. The author notes that certain groups
are taxonomically difficult, and includes comments on several mirid species that may
be confused with those occurring on nettles.

In general the keys seem workable, although it will be apparent to specialists of
certain groups that they could “break out” taxa more easily and eliminate the need
to repeat certain characters. I was disappointed that anthocorids were considered as
cimicids when most specialists place these heteropterans in the separate family An-
thocoridae. In a few instances species are keyed and referred to by common name
but no family name is indicated. For example, the name Nitidulidae is not mentioned
in connection with the flower beetles Brachypterus glaber and B. urticae, and Chrys-
omelidae is not associated with the flea beetles listed in the key.

Chapter 3, “Biology,” categorizes the nettle fauna by feeding habits (phytophagous
species, parasites, predators, scavengers) and summarizes the plant parts used by
plant feeders. Nettle specialists and species feeding on other plants of the Urticales
in addition to nettles are listed; other tables given the principal predators and parasites
found on stinging nettles. Various patterns of voltinism and interesting life cycle
strategies are discussed. Several areas in which the amateur can make worthwhile
contributions to the knowledge of nettle insects are suggested.

Presented in Chapter 4, “Techniques and Approaches to Original Work,” are
sections on collecting, keeping live insects, storing and displaying insects, drawing
and dissecting specimens, sampling and experimenting, preparing the published pa-
per, and locating books and journals. Again, suggestions for further research are
made: influence of shade, soil type, and altitude on host plants and their associated
fauna, and effect of temperature on numbers of insects captured in sweep-net samples.
The sampling methods needed to assess the impact of such variables are briefly
presented. Obviously, complex topics such as sampling theory can be given only
cursory treatment, but the reader can obtain more information from “References

190

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

and Further Reading.” The 50 references listed include recent papers on biology and
taxonomy of various British insect groups and titles of general text and reference
books. A list of some British suppliers of entomological equipment and an appendix
giving some snails and spiders that may occur on stinging nettles are also included.
An index to common and scientific names and a glossary of 75 technical terms
complete the handbook.

Insects and Thistles treats the fauna of creeping thistle, Cirsium arvense L., and
spear thistle, C. vulgare L. The author notes that the different plant parts support a
characteristic fauna, with flower heads (Chapter 2) representing a rich food source
that contains “the most varied, specific and well-known insect fauna of any part of
the plant.” The biology of Urophora stylata (F.), a gall-inducing tephritid, is covered
in some detail, including comments on adult behavior, oviposition, phenology, nat-
ural enemies, and an introduction to life table analysis. Other tephritids and the
Cecidomyiidae associated with thistle heads are discussed, and a simplified food web
is presented; various insects using the heads for shelter are mentioned. Chapter 3
summarizes the life cycles of stem borers and leafminers, mainly Agromyzidae,
Tephritidae, and Curculionidae, and discusses mortality factors. Several nonspecific
stem-boring and leaf-tying Lepidoptera are briefly noted. Sapfeeding herbivores,
defoliators, and flower feeders are covered in Chapter 4. Because thistles have been
accidentally introduced to North America and have become serious pests, the thistle-
associated herbivores of Britain and continental Europe have been investigated to
assess their potential for use in biological control. In Chapter 5 the most promising
candidates for introduction to North America are discussed: tephritids, chrysomelids,
and curculionids.

To identify an insect from thistle, its location on the plant must be known. Thus,
in Chapter 6 keys are presented for species occurring in the heads; larvae and pupae
found within stems and roots; immature stages in mines, folded leaves, or silken
webs; and adults and immatures found on the outside of the plant. For species in
the last category there are keys for identifying adult and nymphal Heteroptera, adult
Lepidoptera and Diptera, and coleopteran adults and larvae. The author stresses that
determinations of parasitic Hymenoptera and agromyzids should be checked by a
specialist if the names are to be published; she also lists taxonomic works that may
facilitate identifications in these groups. Chapter 7 gives a brief discussion of tech-
niques. As further reading 75 journal articles and books are suggested.

Although the information on insect communities presented in Thistles is similar
to that in Nettles, there are several differences. In addition to the emphasis on
microhabitats found on the plant, author names are supplied in the keys rather than
referring the reader to the checklist of British insects by Kloet and Hincks and, instead
of having a terminal glossary, technical terms, as they appear in the text, are defined
in marginal notes. These are welcome changes. I also approve of using “nymph”
rather than “larva” to refer to young of exopterygote insects. Thistles also is notable
for its excellent line drawings and superb color plates, including one of the host plants.
A few minor points: the heteropteran family name Piesmatidae should have been
used for Piesmidae (p. 45), Degeer (pp. 37, 57) should be written DeGeer, Fallen
(pp. 38, 53) should be Fallen, and both thread-like (p. 56) and threadlike (p. 57) were
used.

1984

BOOK REVIEW

191

Grasshoppers treats the 21 British members of the Acrididae, “true” or “short
horn” grasshoppers, and Tettigoniidae, “bush crickets” or “long horn grasshoppers,”
enabling the reader to identify all common species and to learn about their habits.
After an introductory chapter that gives a table of Orthoptera occurring in Britain
(including author and common names and also listing species of Tetrigidae, Gryllidae,
and Gryllotalpidae), there are chapters on “Life History and Reproductive Strate-
gies,” “Adaptations and Habits,” “Identification,” “Distribution and Habitat Pref-
erence,” “Relevant Techniques,” and “References and Further Reading” (101 titles).
In Chapter 4 the keys to families of British Orthoptera and to species of acridids and
tettigoniids are accompanied by line drawings and common names, including the
curious designation “wart-biter” for the tettigoniid Decticus verrucivorus L., a very
local species of dry heathland and dry grassland that supposedly bites off warts. The
chapters on life history and habits contain numerous figures and tables, and through-
out the reader is encouraged to contribute to the many poorly known aspects of
acridid and, especially, tettigoniid biology. There are four excellent color plates fea-
turing 8 species of Acrididae and 9 of Tettigoniidae.

Solitary Wasps, covering the British Sphecoidea, Scolioidea, and solitary species
of Vespoidea, is intended to encourage original work on these fascinating insects and
to allow the species under observation to be identified. An introductory chapter
provides a glimpse into the habits of solitary wasps and compares their morphology
and behavior to those of other hymenopteran groups. Chapter 2, “Natural History,”
discusses the behavior of species in three sphecoid genera —Passaloecus, Cerceris,
and Ammophila— and demonstrates that “behavioural patterns are worth studying
at two levels: the broad strategy that matters to the wasp, and the fine detail that
matters to the evolutionist.” A checklist reminds field workers to ask key questions
about a wasp’s nest site and architecture, prey selection and capture, egg-laying
behavior, nest associates, and other aspects of adult behavior (mating, feeding, and
possible cleptoparasitism and nest sharing). In Chapter 3 the reader is furnished
several identification aids: a novel “Guessing Guide” that allows approximate generic
placement of wasps observed in the field, providing their general coloration, nest
site, and prey type are known; a “Quick-Check Key” that allows dead or anaesthetized
individuals to be placed to genus; and keys that should enable the amateur to de-
termine most British solitary wasps to species. Line drawings placed in the keys’
margins, and 8 plates (4 in color) illustrating adult habitus facilitate the identifications;
included in the keys and plates are several bee species that could be confused with
solitary wasps. Chapter 4 is a short discussion of techniques and approaches to original
work. Also included are further reading (29 references), an alphabetical checklist of
species (without author names) and their British distribution, and an index.

In summary, all four books are well written, contain illustrated keys and color
plates, and emphasize the role of amateurs in making significant contributions to
our knowledge of insects. The paper quality is excellent, and the attractive covers
feature color illustrations of insects. The volumes appear sufficiently durable to with-
stand field use; they are remarkably free of typographical errors, and in my copies
only a few widely scattered broken letters mar the printing quality. These short books,
although devoted to the British insect fauna, deserve a wide audience— the numerous
suggestions for careful biological work by amateurs are equally relevant to studies of

192

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(2)

North American insects. For beginning students especially, the Naturalists’ Hand-
books seem certain to arouse a curiosity about insects and their habits.— A. G. Wheel-
er, Jr., Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harris-
burg, Pennsylvania 17110.

INSTRUCTIONS TO AUTHORS

The Journal of the New York Entomological Society is devoted to the advancement and
dissemination of knowledge of insects and related taxa. The costs of publishing the Journal are
paid by subscriptions, membership dues, page charges, and the proceeds from an endowment
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consider for publication manuscripts of any length dealing with original research in entomology.
Longer papers will be printed as articles, shorter ones as “scientific notes.” Book reviews will
be solicited by the Book Review Editor.

Manuscripts should be submitted in duplicate to: Dr. Randall T. Schuh, Editor, Journal of
the New York Entomological Society, c/o Department of Entomology, American Museum of
Natural History, Central Park West at 79th Street, New York, New York 10024. All material
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Longer manuscripts intended for submission as articles should be accompanied by a brief
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Authors will receive page proofs, with the original manuscripts, directly from the printer.
Corrected proofs should be returned promptly to the Editor to avoid publication delays. Re-
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Society members will be charged a fee of $10.00 per printed page for the 1984 publication
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who do not have institutional funds may petition to the Society for waiver of page charges on
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Authors will receive a reprint order blank with the proofs. Reprints are ordered directly from
the printer with no benefit accruing to the Society.

Journal of the

New York Entomological Society

VOLUME 92 APRIL 1984

NO. 2

CONTENTS

Revision of Arocera Spinola (Hemiptera: Pentatomidae) F. J. D. McDonald

Parathroscinus, a new genus of beetles from Southeast Asia (Limnichidae: Cephal-
obyrrhinae) David P. Wooldridge

Fleas (Siphonaptera) from nests of woodpeckers in Alaska

Glenn E. Haas and Nixon Wilson

Synopsis of Ommatius Wiedemann (Diptera: Asilidae) from Hispaniola

A. G. Scarbrough

Propleural transformations with respect to the disposition of propleural suture in
order Hymenoptera Malkiat S. Saini

Tool use by the ant, Novomessor albisetosus (Mayr) Philip McDonald

Foraging behavior of the bees Halictus ligatus (Hymenoptera: Halictidae) and Cer-
atina calcarata (Hymenoptera: Anthophoridae): Foraging speed on early-summer
composite flowers Howard S. Ginsberg

On the Pseudoscorpion-mimicking spider Cheliferoides (Araneae: Salticidae)

Norman I. Platnick

Aufeius impressicollis (Hemiptera: Rhopalidae): Easternmost U.S. record, host plant
relationships, and laboratory rearing A. G. Wheeler , Jr.

Compsidolon ( Coniortodes ) salicellum (Hemiptera: Miridae): A predaceous plant bug,
new to the United States, found on Filbert

John D. Lattin and Russell H. Messing

Notes and Comments

Notes on the distribution of Dermaptera in some Costa Rican cacao plantations

Allen M. Young

Book Review

Naturalists’ Handbooks A. G. Wheeler, Jr.

97-120

121-124

125-130

131-149

150-155

156-161

162-168

169-173

174-178

179-183

184-187

188-192

'9S, /Ob 73

Vol. 92 JULY 1984 No.

Journal

of the

New York

Entomological Society

(ISSN 0028-7199)

Devoted to Entomology in General

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY

Editor: Randall T. Schuh, Department of Entomology, American Museum
of Natural History, Central Park West at 79th Street, New York, New
York 10024

Book Review Editor: Quentin D. Wheeler, Department of Entomology,

Cornell University, Ithaca, New York 14853

Publications Committee: Louis Trombetta, St. Johns University, Queens,

New York, Chairman; Alfred G. Wheeler, Jr., Pennsylvania State De-
partment of Agriculture, Harrisburg; Joseph M. Cerreta, St. Johns Uni-
versity, Queens, New York.

The New York Entomological Society
Incorporating The Brooklyn Entomological Society

President: Gerard Iwantsch, Department of Biological Sciences, Fordham

University, Bronx, New York 10458

Vice President: Henry M. Knizeski, Jr., Department of Biology, Mercy
College, Dobbs Ferry, New York 10522
Secretary: Irene E. Matejko, Science Department, The New Lincoln School,
New York, New York 10021

Assistant Secretary: Dennis J. Joslyn, Department of Biology, Rutgers

University, Camden, New Jersey 08102
Treasurer: Louis Sorkin, Department of Entomology, American Museum

of Natural History, New York, New York 10024
Trustees: Class of 1984— Joseph Cerreta, St. Johns University, Queens,

New York; Durland Fish, Fordham University, Bronx, New York;
Class of 1985— Peter Chabora, Queens College, New York; Charles
Porter, Fordham University, Bronx, New York.

Annual dues are $18.00 for established professionals with journal, $10.00 without journal,
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Subscriptions are $27.00 per year domestic and $30.00 foreign. All payments should be made
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Meetings of the Society are held on the third Tuesday of each month (except June through
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Street, New York, New York.

Mailed December 21, 1984

The Journal of the New York Entomological Society (ISSN 0028-7199) is published quarterly (January, April, July,
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NEW YORK ENTOMOLOGICAL SOCIETY
92(3), 1984, pp. 193-306

GENUS IRBISIA REUTER
(HETEROPTERA: MIRIDAE)

MICHAEL D. SCHWARTZ

Curatorial Assistant, Department of Entomology,

American Museum of Natural History,

Central Park West at 79th Street,

New York, New York 10024

Abstract.— The black grass bug genus Irbisia Reuter is revised. Over 20,000 specimens were
examined and information concerning the biology, host plant and distributional relationships
of the species is compiled. In addition to external characters, the male parameres and vesica,
and the female dorsal labiate plate of the bursa copulatrix and sclerotized rings are used to
distinguish the species. Thirty-four characters selected from male genitalia, and external mor-
phology are cladistically analyzed. Twenty-three species in five species groups are recognized
in the genus. Two keys to the species are presented. Two new species, Irbisia bliveni and I.
cascadia, are described. The following synonymies are proposed: I. brachycera (Uhler) = I.
gorgoniensis Bliven, I. paeta Van Duzee, I. tejonica Bliven and I. vestifera Bliven; I. californica
Van Duzee = I. eurekae Bliven, I. paenulata Bliven and I. umbratica Bliven; I. elongata Knight
= I. retrusa Bliven; I. mollipes Van Duzee — I. upupa Bliven; I. setosa Van Duzee = I. ustricula
Bliven; I. silvosa Bliven = /. paulula Bliven; I. sita Van Duzee = I. neptis Bliven; I. solani
(Heidemann) = I. inurbana Bliven and I. lacertosa Bliven.

Irbisia species diversity is greatest in the Coastal, Peninsular, Sierra and Transverse Ranges
of California and decreases northward, northeastward and eastward. Major centers of endemism
are located in California and the Rocky Mountains northwest of the Wyoming Basin; a minor
center is located in the Siskiyou Mountains of southwestern Oregon. The distribution of the
Irbisia species appears to be delineated by climatic conditions; late winter and early spring
precipitation is required for growth of their cool season grass hosts. Over twenty native and
introduced species of grasses are utilized by nymphal and adult stages of Irbisia in the Pacific
Northwest. Larval feeding is observed on four species of nongrass monocots, and species of
Lupinus and Lathyrus (Fabaceae). There appears to be no grass host specificity among the Irbisia
species. However, all the grass species utilized are in the subfamily Pooideae; genera from the
tribes Avenae, Poeae and Triticeae predominate. Irbisia species are the earliest mirids to
consume the grass resource, completing their life cycle before grass seed is mature. Many Irbisia
species, upon maturity, leave grasses and congregate on conspicuous nearby shrubs and trees.
Their dispersal is correlated with the blossoming of these plants; adult feeding on pollen, nectar
or honeydew is suspected.

TABLE OF CONTENTS

Introduction 194

Literature Review 195

Materials and Methods 197

Biology 198

Discussion of Taxonomic Characters 202

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Taxonomy

Description of the Genus Irbisia Reuter 206

Keys to the Species 218

Descriptions of the Species

Irbisia bliveni, new species 225

Irbisia brachycera (Uhler) 227

Irbisia calif or nica Van Duzee 232

Irbisia cascadia, new species 235

Irbisia castanipes Van Duzee 238

Irbisia cuneomaculata Blatchley 239

Irbisia elongata Knight 241

Irbisia fuscipubescens Knight 243

Irbisia incomperta Bliven 245

Irbisia knighti Schwartz and Lattin 247

Irbisia limata Bliven 248

Irbisia mollipes Van Duzee 250

Irbisia nigripes Knight 252

Irbisia oreas Bliven 254

Irbisia pacifica (Uhler) 256

Irbisia panda Bliven 259

Irbisia sericans (Stal) 261

Irbisia serrata Bliven 264

Irbisia setosa Van Duzee 269

Irbisia shulli Knight 271

Irbisia silvosa Bliven 273

Irbisia sita Van Duzee 278

Irbisia solani (Heidemann) 279

Methods of Classification 283

Discussion of Species Groups 285

Biogeography 287

Acknowledgments 296

Literature Cited 298

Appendices 304

INTRODUCTION

The species of the genus Irbisia Reuter are predominately Nearctic in distribution.
One species, I. sericans (Stal), occurs on the southern coastline of the Kamchatka
Peninsula, and Commander Islands of the U.S.S.R. Another species, I. brachycera
(Uhler), is distributed in the northwestemmost region of the Great Plains. The rest
of the species are found in or west of the Rocky Mountains. Within this large geo-
graphical area the species are primarily associated with grassland, steppe or open
woodland situations. As a result of this study, there are 23 recognized species of
Irbisia.

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195

The majority of the species in the family Miridae, the largest family in the order
Heteroptera, are plant associates and phytophagous. A distinct assemblage of these
species comprising several lineages feeds, to a variable degree of specificity, on the
plant family Poaceae. The tribes Stenodemini and Mirini, of the subfamily Mirinae
contain the majority of grass-feeding mirid species in the Nearctic. The Stenodemini
are exclusively phytophagous on grasses, and non-grass relatives, the rushes and
sedges. The Mirini contain few exclusive grass-feeding species. The species of Irbisia
and, its tentative sister group, Capsus Fabricius are the only native Nearctic grass-
feeding members of the tribe Mirini. Two grass-feeding Palearctic species of the
Mirini, Stenotus binotatus (Fabricius) and Calocoris norvegicus (Gmelin), were ap-
parently introduced into the United States (Knight, 1941).

Irbisia species have long been known to attack and damage natural and cultivated
native or introduced rangeland grasses (Knowlton, 1951; Haws, 1978). Under certain
conditions, many agricultural crops, particularly in California, have also received
injury from these bugs (Essig, 1 927). Despite the fact that the most destructive species
were recognized over a century ago, published life history studies of the species of
Irbisia are absent. The mating and oviposition habits of these bugs are practically
unknown.

Previous to the present study, there were three published keys to the identification
of the species of Irbisia. Van Duzee (1921a) dealt with the complex of species which
occurred primarily in California, Knight (1968) provided a key to those species with
a predominately intermountain distribution and Kelton (1980) presented a key to
the species of the prairie provinces of Canada. Bliven (1961, 1963) doubled the
number of known species in the genus, but did not include a key or attempt to place
his newly described species within any previous construct of the genus. Even though
the majority of Bliven’s new species are distributed in California, one species, I.
serrata Bliven, occurs throughout the Intermountain West. This species will key to
I. brachycera in the previous keys but has a completely different habitus.

LITERATURE REVIEW

Reuter (1875b) erected the genus Irbisia with Leptomerocoris sericans Stal 1858
the type species by monotypy. This description by Reuter was only one line in length.
Reuter (1896a) published the first full diagnosis of the genus. Leptomerocoris Kirsch-
baum 1855, was placed in synonymy with the genus Orthonotus Stephens 1829, by
Carvalho (1959). This genus is in the subfamily Phylinae and tribe Phylini.

Uhler (1 872) described Rhopalotomus brachycerus and R . pacificus from specimens
collected from Colorado and Idaho respectively. Reuter (1875a:21) placed the genus
Rhopalotomus Thomas 1843 in synonymy with Capsus Fabricius 1803, in a paper
dealing with the Miridae of Europe. He did not treat the Uhler species. He defined
Capsus as possessing a second antennal segment which was “clavatoincrassato”; R.
brachycerus and R. pacificus have cylindrical second antennal segments. Uhler (1886),
in a checklist of Heteroptera of North America, placed both brachycerus and pacificus
in the genus Capsus. Uhler (1894) erected the genus Thyrillus (type species, T. pa-
cificus) for these two species after examining specimens of them from southern Cal-
ifornia. Reuter (1896a) placed Thyrillus in synonymy with Irbisia. The close rela-

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tionship of these two genera was supported by the examination of the female and
male genitalia by Slater (1950) and Kelton (1959) respectively.

Capsus solani Heidemann 1910 was placed in the genus Irbisia by Van Duzee
(1914) and considered a variety of I. brachycera. Subsequently, Van Duzee (1917b)
placed solani in the genus Lygus, but he (1921a) finally elevated it to species status
as Irbisia solani.

The rest of the species placed in the genus Irbisia have not had extensive nomen-
clatorial changes. The remaining species were described by Knight (1925a, 1941),
Blatchley (1934), Bliven (1961, 1963) and Schwartz and Lattin (1984). Two new
species, I. bliveni and I. cascadia, are described herein.

Species of Irbisia were included in state or regional surveys of Heteroptera or
Miridae as follows: Stal (1858), Uhler (1871, 1875, 1886, 1894), Sahlberg (1885),
Cockerell (1893, 1910), Gillette and Baker (1895), Reuter (1896b), Ashmead (1898),
Heidemann (1900), Breddin (1902), Oshanin (1910), Van Duzee (1914, 1917b),
Knight (1921, 1968), Parshley (1922), McAtee (1923), Kiritshenko (1926), Downes
(1927), Harling et al. (1977) and Kelton (1980).

There are two references to the structures of the male and female genitalia which
include species of Irbisia. Three species were included by Slater (1950) in his inves-
tigation of the female genitalia; emphasis was placed on the posterior and dorsal wall
of the bursa copulatrix and the sclerotized rings. Two species were considered by
Kelton (1959) in his study of the male genitalia. There are no investigations of the
nymphal stages of the Miridae which include species of Irbisia.

Biological studies of this genus are limited to 19 reports of an economic nature,
which may include short descriptive features of the species and their feeding damage,
life histories, distributions, food plants and control. These studies are: Vosler (1 9 1 3),
Childs (1914), Essig (1915, 1927), Herms (1926), Lockwood (1933, [1937 not veri-
fied]), Tavemetti (1933), Knight (1941), Essig and Hoskins (1944), Lange (1941),
Ryan (1945 not verified), Lockwood and Glammon (1949), Knowlton (1951, 1961a,
b, 1967, 1968) and Knowlton and Roberts (1971).

Irbisia brachycera, I. fuscipubescens Knight, I. pacifica and I. shulli Knight have
been cited numerous times in the U.S. Department of Agriculture, Cooperative
Economic Insect Report (in the years 1953, 1957, 1958, 1960-1 975) and its successor
the Cooperative Plant Pest Report (1976-1981). The reports estimate Irbisia density
and economic losses to forage crops in Alberta, California, Idaho, Montana, Nevada,
Oregon, Utah and Washington.

Some information on I. brachycera and I. pacifica is contained in Haws (1978), a
thorough presentation of the accumulated knowledge of Labops hesperius Uhler on
range grasses (another species of grass-inhabiting mirid). All three species frequently
occur together in large numbers causing economic damage to rangeland grasses.
Hewitt and Burleson (1975, 1976) studied the arthropod fauna associated with both
planted and natural rangelands and found very low populations of Irbisia brachycera.
McKendrick and Bleicher (1980), studying I. sericans near Homer, Alaska, provide
quantitative analyses of the composition of damaged and adjacent undamaged blue-
joint reedgrass plants. Hall (1959) investigated the pathology of an entomophthor-
aceous fungus attacking I. solani in Riverside County, California.

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197

MATERIALS AND METHODS

Just over 20,000 specimens from throughout the known range of Irbisia were
examined during the course of this study. Specimens of the Capsus cinctus (Kolenati)
were also examined. A listing of the collections, curators, and abbreviations used in
the text is contained in the Acknowledgments.

The recording of the label data for each specimen was not attempted because of
the large number of specimens examined. However, each specimen received an
institutional label indicating where they are retained. The specimens of 7. pacifica
were separated from the rest of the material because of their ease of identification.
The two groups of material were initially sorted by country, state or province, county
and locality. The next phase of the sorting process involved separating the geograph-
ically arranged specimens into species level taxa, on the basis of the types, the original
descriptions and my own understanding of the aggregations of characters available.
This sorting process was initiated with material representing the northern and eastern
limits and proceeded toward the California specimens. Once sorted in the above
manner, the geographically arranged groups of taxa were consolidated on the basis
of their aggregate characters. I wrote the descriptions and redescriptions as species
concepts emerged from the taxon groupings.

At this point, it was not difficult to arrange the specimens alphabetically and record
the locality data. Hosts, collection dates and elevations of adult specimens were
recorded and summarized. Localities that could not be located were listed after the
county label data. Names of the collectors were omitted for the redescriptions. All
label data were recorded in the new species descriptions. The label data accompanying
each examined holotype was recorded as given. All specimens including type spec-
imens and many paratypes, as well as critical specimens examined by Bliven, Knight
and Van Duzee, that I was able to identify satisfactorily have been individually
labeled with appropriate determination labels. The allotypes were treated as para-
types. All Knight, many Van Duzee, and some Bliven paratypes were conspecific
with the associated holotypes. All data pertaining to these specimens were incor-
porated in the synonymy portion of the species treatments. For a complete accounting
of the label data and depository of the holotypes placed in synonymy and their
associated secondary types refer to my master’s thesis at Oregon State University,
Corvallis, Oregon.

In addition to intensively collecting specimens from Benton and adjoining counties
of Oregon, I conducted several extended collecting excursions to: central and south-
west Oregon; northwest, central and southern California; and west and central Wash-
ington. These trips were taken during the springs of 1979 and 1980 for the purpose
of obtaining specimens from the localities listed in the original descriptions of Bliven
(1961, 1963), and to gather information on the distribution, host plants and biology
of Irbisia. Live nymphs and adults were collected and placed in cages containing
potted sod of natural grasses. The live material was observed in the laboratory for
a period of several weeks.

Biological information was gained from both field and laboratory observations.
Many grass and non-graminoid hosts were collected, pressed with locality data, and

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identified to genus by myself and to species by the staff of the Oregon State University
Herbarium. The host data contained on the labels of the specimens were corrected
according to Munz and Keck (1973).

In order to plot the distributions for the species, each unique locality, for each
species, was located on road maps, for each state, and a signal dot was placed on the
map. Because of the great number of specimens from California, Oregon and Utah,
separate maps were used for several species. At this point, because of the high
resolution of the road maps, the potential natural vegetation (Kiichler, 1966, 1977)
and ecoregions (Bailey, 1976) were recorded and summarized for each species. Rec-
ords made on the state road maps were combined by species and are summarized
in the distribution maps which follow the species descriptions.

Male and female genitalia of specimens from different portions of the range of
each species were examined. The number of specimens dissected varied according
to the variability of the other characters, extent of the distributional range, and the
availability of material. Only dried specimens were used for the study of the genitalia.
Teneral specimens were avoided whenever possible. The method employed for the
dissections has been established and recorded by Slater (1950) and Kelton (1959)
and perfected by Razafimahatratra (1980). After dissection and study, the dissected
parts were placed in a plastic microvial containing glycerin and attached to the pin
bearing the appropriate specimen.

Illustrations were drawn using a Leitz Wetzler stereomicroscope with an ocular
grid. The figures were drawn on grid paper and later inked on velum. Unless otherwise
indicated, the illustrations were drawn to the same scale. All measurements are given
in millimeters and obtained from an ocular micrometer. Various magnifications were
used depending on the structure being measured, 160x or 64 x for genitalia, 40 x
for head and thorax, 20 x for body width and 1 0 x for body length.

The description or redescription of each species takes into account the size, struc-
tural, and color variations encountered. A range is given for each measurement.
These sections are therefore composite. The descriptions or redescriptions summarize
specific differences or additions not found in the detailed generic description. The
form of each redescription is uniform; the taxonomic characters are presented in the
same order. The diagnoses will separate the species of concern from other taxa to
which it is superficially similar or sympatric. Polymorphic or polytypic variation of
color or structure, as well as any taxonomic consideration, is included in the dis-
cussion of each species.

A list of the abbreviations used in the specimens examined section of the species
treatments is found in Appendix B.

BIOLOGY

During the spring, species of Irbisia are common, conspicuous inhabitants of
chaparral, steppe, open woodland and certain grassland situations in North America
west of approximately the 1 00th meridian. Although primarily grass feeders, which
disperse to many non-graminoid plants late in the season, several species of Irbisia,
in the nymphal stage, feed extensively on non-grass species of plants.

The eggs of Irbisia are elongate, slightly curved, with a flattened cap on one end.
Oviposition occurs from late March in southern California to late May and early

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June at higher elevations of the Pacific Northwest and the Intermountain states.
These bugs diapause in the egg stage, protected through the summer and winter until
the occurrence of the succulent regrowth of their host plants in late winter and early
spring. The duration of diapause is typically nine to ten months regardless of the
geographic location of the species; the initiation and termination dates, of course,
are variable. The oviposition sites apparently vary according to habitat. Taylor (un-
publ. data) observed that the dead and prostrate stems of blue elderberry ( Sambucus
caerulea Rafi), in the Davis area, were "... filled with thousands of insect eggs.” The
stems had a pithy internal structure and woody outside surface. Upon “... sticking
the stems in water for a few minutes to get them wet then putting them in an incubator
the eggs started development and emergence was then sure. I reared many hundreds
of them to the adult stage so am sure that the eggs were of this genus [Irbisia].” The
distributional range of blue elderberry is extensive, occupying open places up to
10,000 feet elevation in montane coniferous forests and foothills. The co-occurrence
of many species of Irbisia within the range of this and other species of Sambucus
may only be coincidental, but is interesting. Many other shrubby plants (listed in the
following species redescriptions) may provide the necessary architecture and phe-
nology to be utilized as oviposition substrates.

Mr. Jeff B. Knight, at the University of Nevada/Reno is conducting research on a
degree-day model for 7. brachycera in the Reno area. He successfully induced 7.
brachycera to oviposit in the laboratory in the culms of Elymus sp. (pers. comm.,
1980). This is similar to the oviposition habits of Labops hesperius which lays eggs
in the dry culms of Agropyron desertorum (Fisch.) Schult. and Poa secunda Presl.,
from the previous year (Todd, 1973). Knight (pers. comm., 1981) has also found
that 7. brachycera will oviposit in the ligule of the lower leaves of Sitanion hystrix
(Nutt.) J. G. Sm. and the sheaths of the lower leaves of Elymus cinereus Scribn. &
Merr. The eggs are aligned parallel to leaf veins. The eggs after a period of diapause,
in situ, when placed at approximately room temperature will achieve peak hatch
within ten days. McNeill (1971) relates that the females of Leptopterna dolobrata
(L.) oviposit directly into the base of the present-year’s grass culm.

As quoted above, Taylor found that moisture was the key to the hatching of the
eggs. After winter rainfall, the bugs would emerge during the coldest period of the
year, February. Refrigeration without moisture did not induce emergence in the
laboratory. His observations (which are collaborated by my field collections) indicate
that the species of Irbisia in Davis, California emerge from the egg stage in this order:
7. solani, I. californica Van Duzee and 7. pacifica. In Corvallis, Oregon, 7. sericans
emerges before 7. cascadia, n. sp.

It is interesting to note that oviposition sites or physiology of 7. brachycera and 7.
pacifica must be different. Haws (pers. comm., 1979) indicated that the harsh snowless
winter of 1976 greatly reduced many populations of the latter species in traditional
outbreak localities in Utah. The former species and Labops hesperius were largely
unaffected.

Observations in the field and laboratory indicate that molting occurs directly on
the grass blades. Successful molting is easily fouled by excessive moisture in rearing
cages. I have no detailed data on the duration of the five nymphal instars but, after
the first observation of early nymphal stages, in Benton County, Oregon, under field
conditions, approximately four weeks elapse before the adult stage is reached. Males

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usually mature before the females; female specimens are often the only sex encoun-
tered late in the season. There is a temporal stratification of the height at which both
nymphal and adult stages of I. cascadia and I. sericans feed. It is possible to collect
many more specimens with the same effort by sweeping grass at night than during
the day. This may indicate an adaptation to reduce desiccation by avoiding low
relative humidity encountered in the higher horizons of a grass meadow during
daylight hours. Further, wind speeds may be lower during the evening hours.

Although the majority of species of Irbisia I have observed fed on a variety of
grasses as nymphs, several (/. californica, I. cascadia, I. serrata and I. solani) do feed
as nymphs on the following non-graminoids: Allium spp., Brodiaea sp., Iris tenax
Dougl., Lathyrus spp. and Lupinus spp. In Arroyo Seco, California I observed nymph-
al stages feeding on Lupinus sp. in an area that had grasses without any apparent
feeding damage. This observation raises the possibility that grass may not be an
obligatory host for all species in all situations. I have raised late instar I. cascadia
nymphs to adulthood on caged Lupinus plants in the laboratory. The females even-
tually became gravid (as did female specimens from the source populations, which
did not feed on Lupinus).

Both immature and adult stages of all species of Irbisia produce diagnostic feeding
damage on all the species of grasses I have encountered as hosts in the field. Feeding
is always located on the grass blade. The method of feeding is typical of mirids; the
cells of the plant are lacerated by the stylets of the rostrum, and the cell contents are
flushed out by saliva, then the resulting “soup” is sucked up. The dead cells produce
the pale white or chlorotic spots on the leaf blade. In a severe infestation, the blade
may appear completely white. This type of feeding may cause stunted plants and
reduced forage yields (Haws, 1978). The pale spots are, in practically all cases,
accompanied by dark droplets of anal exudate. Stenodemine mirids do not produce
feeding damage similar to that just described. Species of tribe Stenodemini, when
sympatric with species of Irbisia, are usually phenologically later, and tend to con-
centrate their feeding on the culm or seed head (Kamm, 1979). I. cascadia was
observed to produce similar chlorotic spots and dark droplets on the flowers and
young, apical leaves of Lupinus sp. in Skamania County, Washington.

Much of the early literature concerning Irbisia deals with the later phase of its
feeding biology; dispersing from dried grass fields to other non-grass plants. The
species I. solani is most often cited; it has caused damage to artichokes (Tavemetti,
1933); lettuce, radishes, onions and rhubarb (Vosler, 1913); peaches (Lockwood,
1933); and cherries and plums (Lockwood and Glammon, 1949). An interesting note
by Herms (1926) states that the possibility exists for honey bee poisoning, if the bee
feeds on droplets of the sap of California buckeye (Aesculus californica [Spack] Nutt.)
exuding from punctures on the young twigs, leaves and blossom buds which were
caused by I. solani. Besides these plants, I have collected practically all the species
of Irbisia from the blossoms of many additional shrubs or trees including: Ceanothus
spp., Cercocarpus spp., Pinus spp., Quercus spp., Simmondsia sp. (male) and Yucca
spp. Many of the specimens are heavily covered with pollen grains. This demonstrates
that these individuals may have been moving around the blossoms, and more sig-
nificantly, that they were seeking nutrients from the plants. It is possible that the
pollen grains are mixed up in saliva and sucked up (Kullenberg, 1 944). It would be
a simple matter to dissect the gut of freshly “pollen fed” individuals and verify this

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201

hypothesis. Many of the previous plants are also heavily covered with honeydew,
another desirable insect food source. Another interesting correlation is evident; the
phenology of the flowering of many of previous plants coincides with the drying of
the early spring grasses upon which Irbisia initially develop. The species of the
secondary plants that are frequented, of course, varies according to the floristic
association of the surrounding area, i.e. Pinus at high elevations, Ceanothus in the
chaparral, Quercus over a wide range, and Yucca in southern California.

In two excellent papers, McNeil (1971, 1973) explored the energetics and dynamics
of a population of Leptopterna dolobrata in Berkshire, England. He concluded that
this grass-feeding mirid, which does not switch to non-grass hosts, changes its feeding
habits from leaf feeding on Holcus mollis L. (low nitrogen) to seed feeding on H.
lanatus L. (high nitrogen) when the bug is experiencing maximum growth and the
initiation of gonad development. This might be the case with Irbisia. The necessary
nitrogen required for egg production (Mattson, 1980) may have to be supplemented
by non-grass sources. In the Mediterranean climate of California, any species of grass
may be nutritionally depleted too quickly to be an assured resource.

There appears to be resource partitioning between the species of Irbisia and the
stenodemine species which are abundant in the range of Irbisia. Irbisia is phenolog-
ically earlier, feeds on the blades of the grass plant, and can leave the grass for
nutritional supplements. The stenodemines feed on the culms of the grass plant and
seek out grass seed to complete their development. McNeill (1973) pointed out that
individuals of Leptoterna which feed only on leaves risk death by taxing already low
reserves, by increased metabolic rates in warm summer weather, and dehydration
at the growing grass tips during moult. Irbisia may avoid these dangers by completing
its life cycle rapidly. I have observed that late in the season (April and May) in
southern California, Irbisia is readily collected on grasses in the shade of surrounding
trees.

Another phenomenon which I have observed in several locations is the aggregation
of female Irbisia on prominent vegetation in grassy situations. Female I. cascadia
were aggregated on a single Artemisia cana Pursh plant at Siskiyou Pass, Jackson
County, Oregon; female I. fuscipubescens aggregate on Lupinus sp. in Oregon and
Washington; female I. sericans aggregate on Epilobium sp. near Homer, Alaska
(Bleicher, pers. comm., 1980), and female I. shulli and I. fuscipubescens aggregate
on Balsamorhiza spp. in Oregon and Washington. The only apparent Irbisia feeding
damage appeared on the Lupinus. In the Siskiyou Pass instance, female I. serrata
occupied adjacent Quercus foliage, and were not on the sagebrush. Perhaps these
occurrences are just simple dispersal to a prominent landmark because the males
were absent. I have not observed mating in the field or laboratory. However, Oman
(pers. comm., 1981) has observed Irbisia spp. in copula on balsamroot in central
Washington.

In the Corvallis, Oregon area, three species of Irbisia are separated by an apparent
habitat preference. I. sericans is early, and found in moist tall grass ( Festuca arun-
dinacea Schreb.) in low lying situations. I. cascadia is somewhat later, and is usually
found on exposed south-facing hillsides. I. solani has the same phenology as I.
cascadia, and may be found on hillsides, but is more often found at the lowest
elevations in the Willamette Valley, and is associated with Bromus sp.

There are no quantitative studies on the distributions and abundances of Irbisia

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species coexisting in the same grassland. Gibson (1979, 1980) throughly investigated
a five species complex of grass-feeding Stenodemini at Wytham Hill, England. Per-
haps the relationships of the Irbisia species may be similar in some respects to the
dynamics of this group of closely related stenodemine mirids. Gibson concluded that
each of the stenodemine species fed on several foodplants at once and that this host
spectrum changed seasonally and with the age of the bugs. It was also found that
there was considerable overlap between the hosts of different bug species and that
ecological separation was achieved by emergence timing and/or feeding on different
parts of the same grass species.

Dispersal from rangeland grasses to cultivated grains has been noted by Knowlton
(1951) for I. pacifica. Records of the Nevada Cooperative Economic Insect Report
(Anonymous, 1972) implicate I. brachycera in dispersing to bluegrass (Poa sp.) lawns
in the Reno area. Fuxa (1 975) determined that the number of dispersing I. brachycera
caught on sticky traps within a crested wheatgrass field decreased with the height of
the trap; the most bugs were captured below 0.61 meters. The peak flight period
lasted for six days and was maximal at 3 p.m. Fuxa concluded that I. brachycera
dispersed more readily than Labops hesperius.

I observed few predators of the species of Irbisia in the field; a nabid and crab
spider were the only examples. Few parasites of Irbisia have been confirmed; after
I treated a female specimen of I. serrata from Modoc County, California, for genitalic
dissection a dipteran larva was found within the bug’s abdomen. The larva may be
of the family Tachinidae. Entomophthora erupta (Dustan) is a known fungal parasite
of/, solani (Hall, 1959).

DISCUSSION OF TAXONOMIC CHARACTERS

Characters discussed below have proved to be most useful in distinguishing the
species of Irbisia and are utilized in the following keys, descriptions and redescrip-
tions. The characters are from adult specimens of both sexes. Ratio and measurement
values are presented as ranges which adequately cover the variation encountered in
the specimens examined. An asterisk indicates when certain characters show value
as synapomorphies of species groups; thirty-four of these characters compose Ap-
pendix A.

Total length of body. The total length is measured from the tip of the tylus to either
the posterior tip of the membrane in macropterous species, or to the posterior tip of
the terminal abdominal segment in brachypterous species. It is measured in dorsal
view. The length varies greatly between, and within some, species. The range, in
conjunction with other characters, is useful for separating some species.

Total width of body. The total width is measured across the greatest width of the
hemelytra, in dorsal view. The body width alone will separate few species.

Head. The structure, ratios and measurements of the head are particularly useful
in distinguishing species groups and species. MEASUREMENTS. Width across eyes—
maximum width between the lateral margins of the eyes; measurement in dorsal
view. * Vertex— minimum width between the interior margins of the eyes; measure-
ment taken at posterior margin of head in dorsal view. Dorsal width of eye— max-
imum width in dorsal view; varies little between species. Distance from tylus to
ventral margin of eye— measured in lateral view with the tip of the tylus and the

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ventral margin of the eye in simultaneous focus; a minimum width measurement.
Height of eye— maximum measurement in lateral view (measurement of I. pacifica
is taken in frontal view, at the junction of the eye and the frons, because of the
dorsally projected lateral eye margins of this species); this measurement varies little
between the species. Maximum interocular width— measured in cephalic view; place-
ment of this measurement is dorsal of the antennal insertions in the concave portion
of the eyes across the frons. RATIOS. ’•‘Length of antennal segment I : vertex. Distance
from tylus to ventral margin of eye : height of eye. Maximum width of pronotum :
width across eyes.

STRUCTURES. Shape of head— triangular or elongated anterior of eyes (ratio of
width of head across eyes and head length is 1:0.75). Surface of head— smooth,
asperate or shagreened; typically examined on the frons. Temporal areas— distinct
or indistinct, may or may not be demarcated from the frons by faint sulci; the surface
structure may differ from the frons; the areas may be bordered basally (at the vertex)
by sulci of variable strength; the area is usually glabrous. Median depression— absent
(or indistinct), broad (deep or shallow), or reduced to a longitudinal furrow. Basal
carina— either abruptly produced or declivous with respect to the level of the vertex;
the dorsal surface is either acute or rounded in cross section; the width may be narrow
or broad; the posterior margin is either straight or arcuate posteriorly; the carina
may or may not form the posterior margin of the head. Shape and insertion of eyes—
either large and broadly joining the frons, or small and narrowly joining the frons;
the latter type has an obvious pedunculate eye form; the posterolateral portion of
the eyes are either straight or arcuate in an anterior or posterior direction. Clypeus—
basal portion may be convex; standing above the frons, or not projecting above the
frons; the anterodorsal surface is convex, giving a flattened appearance to the surface
between the antenna insertions, or the surface is gently curved. Posterior margin of
head— may or may not be formed by the basal carina, and/or the posterior margin
of the eyes; regardless of the makeup, the posterior margin of the head is either
straight, or arcuate in an anterior or posterior direction. COLORATION. Several
species have light colored markings on the ventral portion of the head. The sutures
separating the genae, juga and lora, and variable portions of these regions may be
testaceous or fulvous.

Rostrum. The length is taken from the junction of the epipharynx and tylus to the
apices of the stylets. The coloration is quite variable within the species ranging from
testaceous to black.

Antennae. The length of segment I is measured from the center of the rounded
basal insertion to the apex. The remaining segments, II-IV, are maximum measure-
ments. The smallest measurements of segments III and IV were taken from specimens
with shrivelled segments. The lengths of segments I and II are quite useful for dis-
tinguishing several species. *The ratio of the length of segment I to the width of the
head across the eyes is a useful phylogenetic character. The coloration of segment I
is variable between and within species, ranging from entirely testaceous to black.
Most intraspecific variation involves light coloration on the apex and base of this
segment.

Pronotum (and remainder of thorax). MEASUREMENTS. All are maximal.
Length— measurement taken on midline, from anterior margin of collar to posterior
margin of disk. Anterior width— measurement taken between the anterior angles; on

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species with prominent anterior angles the apices of the angles are the end points of
the measurement; on species with rounded, broadly curved or obsolete anterior angles
the anteromedial comers of the pronotum are the end points of the measurement.
Maximum width— measurement taken at the posterior margin of the pronotal disk;
the extreme posterolateral comers are the end points. STRUCTURE. The shape of
the pronotum is either conical, subconical, or trapezoidal. The pronotal disk is usually
broadly convex, but on some species, the disk is strongly flattened. The surface texture
of the pronotal disk for the genus may be smooth, with faint, shallow, deep, or
confluent punctures, mgulopunctate, rugulose, transversely rugulose or rugose. The
posterior margin is either straight or arcuate posteriorly; the posterolateral margins
may be slightly or broadly curved. The lateral margins are either straight, slightly or
strongly concave. The junction of the propleura and pronotal disk may be slightly
carinate, rounded or broadly rounded. The anterior angles of the pronotum, in dorsal
and lateral view, are good diagnostic characters for species. The anterior angles range
from strongly produced (rounded or broadly rounded) to indistinct (broadly rounded).
The anterior angles, in lateral view, vary in the development of the sulci on the
anterior portion of the propleural-pronotal lateral junction dorsal or the coxal clefts.
The sulci are either strongly (the pronotum is extended “roof-like” over the ridge
running dorsad of the coxal cleft) or slightly developed or absent. The calli vary in
the degree of convexity, size, the degree of anterior confluence and surface texture.
The width of the collar varies. The scutellum and underside of the thorax exhibit
great similarity of structure among the species. Most of the species are uniformly
black on the ventral surface, except for thin light “trim” surrounding the pro and
mesocoxae, on the anterior ventral one quarter of the propleura, and the ostiolar
peritreme. More extensive light coloration on the underside of the thorax is a good
distinguishing character for several species. On these species ivory, testaceous, fulvous
or fuscous coloration is prevalent on these additional regions: ventral one third to
one half of the propleura, front margin of the prostemum, basalar plate, and variable
portion of epistemum and epimeron.

Legs. The color patterns of the femora and the tibiae display great variation between
and within species. Once the range of the within species variation was determined
it was possible to use leg coloration as a diagnostic character for several species. The
great majority of the species have short, suberect black setae on the tibiae, and short
suberect fulvous setae on the femora. Several species, additionally, have much longer
erect fulvous setae on the tibiae (these setae are longer than the tibial bristles); the
femoral setae are longer as well. The coxae and trochanters also show a great diversity
in coloration. The tarsal structure, including the claws, is very uniform among the
species.

Hemelytra. The embolar margin varies from nearly straight to arcuate and, in
conjunction with the general body form, is a useful diagnostic character. The surface
texture and the structure of the regions of the hemelytra are of minimal value for
the separation of the species. The insertion may be lightly colored in several species.
The hemelytra are either macropterous or brachypterous; the hind wings of the
brachypterous species and forms are brachypterous.

Venter. The venter is uniform among the species: black, with short appressed or
suberect testaceous or fulvous setae. Some species have patches of light coloration

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205

on the dorsolateral regions of the second or third stemite. The setae along the midline
of the stemites may be longer in some species.

Parameres of male. *The structure of the parameres is very useful in differentiating
the species groups and species of Irbisia. The parameres show very little intraspecific
variation. I have adopted the terminology of Kelton (1959) with some additions.
The regions of the left paramere used herein are: sensory lobe, outer surface of arm
(region between sensory lobe and medial angle of the paramere), the angle (the outer
surface of the curved, angle bearing region of the paramere), shaft (the remaining
portion of the paramere apical of the angle), and apex. All five of these regions show
interspecific variation useful for delimiting the species. The variation is as follows:
sensory lobe— prominent or moderately developed (standing greatly or moderately
above the curvature of the arm) or indistinct, either with or without stout tubercles
or spines; outer surface of arm— either with or without tubercles or spines; angle—
’C’, ’L’, *U* or ’V’ shaped; shaft— long or short tubercles; apex-sharp or distally
expanded (either truncated perpendicular or parallel to the axis of the shaft). The
arm, angle, and shaft may bear either single widely spaced tubercles or a series of
tubercles. The tubercles (blunt) may give way to spines (sharp) on the shaft.

Vesica of male. ’“This structure has great utility in differentiating species groups or
species. The terminology follows Kelton (1959). The membranous lobes may be
simple or have several associate lobes, and are more or less rounded or greatly
elongated and pointed. The sclerotized process is highly diverse: straight or curved,
acute or blunt, unexpanded, expanded (and/or concave on the interior surface); with
or without a surface of spicules on the outer or inner surface; with or without larger
spicules on the margin of the inner surface; and with or without large curved spines
on the inner concave surface. All references herein to areas on the sclerotized process
are based on dissected preparations of the vesica; the vesica is oriented so the gono-
pore faces the observer. The species show all combinations of these character states.
Only mature and fully inflated specimens can be used for comparison of the mem-
branous lobes. It is possible, however, to determine the structure of the sclerotized
process on mature totally uninflated specimens, by carefully pulling the vesica out
of the phallotheca with fine forceps.

Posterior wall of bursa copulatrix of female. This structure is of limited value in
determining species. I have followed the terminology of Davis (1955) and Slater
(1950). The posterior wall consists of the wing-like inter-ramal sclerites (’A’ structure
of Slater) which show slight variation in height and width. The sigmoid process (’B’
structure of Slater) and *C* structures are attached to the dorsomedial portion of the
’A’ structure, and show slight interspecific variation (but also great intraspecific
variation).

Sclerotized rings of females. The rings and their supporting structures are very
useful for some species group and species separation. The sclerotized rings vary in
size and shape between species. There is some intraspecific variation, but it is usually
less than the interspecific variation. Of equal importance in distinguishing species
groups and species is the shape and extent of the dorsal labiate plate (’F structure
of Slater). The rings may be small, large, oval or compressed and vary in the distance
between them. The dorsal wall may be narrow or broad, expanded anterior to the
rings, and variably concave medially.

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Coloration. As mentioned previously, the basic color for the species of the genus
Irbisia is black. Only one species has light markings on the dorsum (the base of the
cuneus of I. cuneomaculata Blatchley is testaceous). The light color on the ventral
side of a few other species is always in the form of a light colored vitta. The specific
areas of the head, thorax and abdomen of concern are described in the previous
sections dealing with those regions of the body.

Vestiture. The type, length, and density of the setae on the dorsum are good
taxonomic characters for species discrimination. Most of the types of setae are fulvous
or fuscous; the stiff long bristles which are present in varying amounts on several
species are black. The simplest vestiture, found in some species, consists of a single
type of short, appressed setae. Several species have a single type of densely distributed,
medium length, suberect setae. Many species possess setae of two types; densely
appressed or woolly sericeous setae in combination with the previous two types. A
few species have a vestiture of the last type which is intermixed with long upright
bristle-like or thin setae. Because of the vexing ease with which these bugs lose their
vestiture it is necessary to examine a series of specimens to accurately determine the
character of the setae.

Irbisia Reuter

Irbisia Reuter, 1875b, p. 548; 1879, p. 57; 1880, p. 509; 1890, p. 253; 1896a, pp.
11-12, 347-348; 1896b, p. 157; 1909, p. 13; 1910, p. 1 58. Atkinson, 1890, p. 125.
Hueber, 1898, p. 80. Kirkaldy, 1906, p. 142. Oshanin, 1910, p. 760. Van Duzee,
1916a, p. 207; 1916b, p. 38; 1917a, p. 324; 1921a, pp. 151-152. Slater, 1950, pp.
35-36. Carvalho, 1952, p. 89; 1955, p. 88; 1959, p. 104. Kelton, 1959, pp. 17-
18; 1980, p. 83. Bliven, 1963, pp. 68-69. Knight, 1968, p. 185.

Thyrillus Uhler, 1894, p. 266 (synonymy by Reuter, 1896a, p. 1 1). Van Duzee, 1916a,
p. 207; 1916b, p. 38; 1917a, p. 325.

Type species. Leptomerocoris sericans Stal, 1858 (monobasic).

Diagnosis. The possession of the following characters of the subfamily Mirinae
will distinguish the species of the genus Irbisia from the superficially similar genus
Orthocephalus (Miridae: Orthotylinae): parempodia flattened and divergent apically;
claws rather strongly bent, with sub-basal, ventral pulvilli; rounded pronotal collar
strongly developed and conspicuous; vesica of male with rim of gonopore circular
in outline, simulating a coiled spring; sclerotized rings of female ovoid with strongly
sclerotized margins that are not infolded on the lateral margins; posterior wall of
female with highly differentiated sigmoid process. Specifically, the species of Irbisia
never possess flattened scale-like setae, and numerous stout black bristles as found
on O. coriaceus (Fabricius) or O. saltator (Hahn).

Several species of Irbisia (fuscipubescens, cascadia, knighti, nigripes, sericans, ser-
rata, shulli and solani) are sympatric with Capsus cinctus (Kolenati) in these provinces
and states: Alberta, British Columbia, Colorado, Idaho, Montana, Oregon, Utah,
Washington and Wyoming. All the species listed above are predominately black, but
the Irbisia species never have any light coloration on the vertex and frons of the
head, and have a light ringlet bordering the posterior margin of the eye. In addition,
the second antennal segment of the Irbisia species always is linear and never swollen

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207

apically or pre-apically. The male and female genitalic structures of the two genera
are diagnostic. The vesica of the male Irbisia does not possess a very long and slender
spiculum, and the ductus seminis is not constricted medially. The female sclerotized
rings of the species of Irbisia are never subrectangular and one third to one half the
diameter of the sclerotized rings of C. cinctus. The sigmoid process of the Irbisia
species is always attached to the sclerotized portion of the inter-ramal sclerites and
never to the membranous portion. The inter-ramal sclerites of the species of Irbisia
are never bisected by membranous sutures.

Description. SIZE: length 3.80-8.20, width 1.60-3.20. COLORATION: dorsum
usually black, either polished and shining, asperate, shagreened, or with a bronze
luster; propleura and pleura black, in most species ivory or testaceous narrowly
bordering the pro and mesocoxae, the ventral one quarter of the propleura and the
ostiolar peritreme; venter shining black. The light coloration of the ventral aspect is
more extensive on several species; a light vitta extending from the head, at the tylus,
posteriorly across the pleura, terminating on the second or third stemites; posterior
margins of eyes bordered by narrow testaceous or ivory band. VESTITURE: usually
of similar structure and density on head thorax and hemelytra; simple— consisting
of either appressed short or suberect medium length, fulvous or fuscous setae; com-
plex-consisting of a mixture of the simple type together with either upright, stiff,
long, black bristles and/or appressed or woolly sericeous setae; eyes with sparse minute
pubescence between facets; venter with short or long appressed or upright setae.
BODY: oval, suboval or elongate, dorsum smooth, either with faint, shallow, deep
or confluent punctures, or rugulopunctate, rugulose, transversely rugulose or rugose;
female usually more robust than male. HEAD: triangular, or pedunculate in dorsal
view, region anterior of the eyes sometimes elongated (ratio of width to length of the
head ranging from 1:0.50 to 1:0.75); front moderately or strongly convex, smooth,
asperate or shagreened; basal carina abruptly produced or declivous with respect to
the level of the vertex, dorsal surface acute or rounded in cross section, width narrow
or broad, posterior margin straight or arcuate posteriorly; posterior margin of head
sometimes formed by basal carina and/or posterior margin of the eyes; collum faintly
or strongly separated from carina (depending on structure of the carina); eyes either
large and broadly joining the frons, or small and narrowly joining the frons; clypeus,
juga, and lora slightly or greatly tumid, the sutures separating these regions always
deep; bucculae, gula and genae well defined; antennal sockets either ventrad or dorsad
of the eyes. ROSTRUM: barely reaching bases of mesocoxae to just surpassing apices
of metacoxae. ANTENNAE: cylindrical, inserted below, above or at the level of the
anterior margin of the eye; with densely distributed, fine appressed or suberect,
fuscous or black, short or medium length setae; segment I with several suberect,
stout, long, bristle-like black setae; segment I just reaching tylus or surpassing tylus
by one eighth of the length of the segment; segment II gradually, but only slightly,
thickening toward apex (thickened apex is never greatly evident). PRONOTUM:
conical, subconical, or trapezoidal, broader at base than long (ratio of width to length
of the pronotum ranging from 1:0.54 to 1:0.66); disk broadly convex or, in some
species, strongly flattened; collar convex, well differentiated, forming a complete ring,
widest dorsomedially; posterior margin either straight, slightly or strongly concave;
junction of propleura and disk slightly marginate, rounded or broadly rounded;

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Figs. 1-6. 1, 2. Lateral view of head. 1. 7. pacifica. 2. 7. californica. 3-6. Anterodorsal view

of left side of head (centered at angle of frons and eye). 3. 7. pacifica. 4. 7. brachycera. 5. 7.
California. 6. 7. solani.

Figs. 9-14. a. Dorsal view of left paramere. b. Dorsal view of right paramere. 9. 7. bra-
chycera. 10. 7. elongata. 11.7. pacifica. 12. 7. sericans. 13. 7. californica. 14. 7. s//a.

anterior angle, in dorsal view, either indistinct, moderately or strongly produced,
sulcus on anterior angle, in lateral view, either absent, slightly or strongly developed;
calli slightly or strongly convex, smooth or asperate, narrowly or broadly confluent
anteriorly, sometimes with one or two fovate depressions on each callus; the calli
sometimes reaching the anterolateral margin of pronotum; mesoscutellum narrowly
or widely exposed, smooth, gradually or steeply sloping to scutellum; scutellum
weakly or strongly convex, and transversely rugulose; propleura with ventral and
posterior margins sinuous. HEMELYTRA: lateral margins arcuate or subparallel;
embolium explanate anteriorly; rugulopunctate or rugulose; cuneus slightly or mod-
erately deflected; fracture deep; membrane— macropterous species and forms, sur-
passing the genital segments in both sexes, with two well developed areoles— bra-
chypterous species and forms, membrane is reduced, and sometimes slightly surpassing
apex of cuneus, the genital segments are exposed, areoles are either developed or

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REVISION OF IRBISIA

Fig. 7a. Dorsal view of body, 1. pacifica.

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Fig. 7b, c. b. Dorsal view of body, I. knighti. c. Dorsal view of pronotum, I. sericans.

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REVISION OF IRBISIA

Fig. 8. Dorsal view of body, I. cascadia.

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Figs. 15-31. a. Dorsal view of left paramere. b. Dorsal view of right paramere. 15. 7.
cuneomaculata. 16. 7. oreas. 17. 7. incomperta. 18. 7. solani. 19. 7. knighti. 20. 7. limata. 21.
7. nigripes. 22. 7. shulli. 23. 7. serrata. 24. 7. castanipes. 25. 7. bliveni. 26. 7. silvosa. 27. 7.
cascadia. 28. 7. fuscipubescens. 29. 7. mollipes. 30. 7. panda. 31.7. setosa.

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213

Figs. 32-43. Ventral veiw of vesica. 32. I. brachycera. 33. I. elongata. 34. I. pacifica. 35.
I. sericans. 36. I. californica. 37. I. sita. 38. I. cuneomaculata. 39. I. oreas. 40. I. incomperta.
41. /. solani. 42. I. knighti. 43. I. limata.

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Figs. 44-54. Ventral veiw of vesica. 44. I. nigripes. 45. I. shulli. 46. I. serrata. 47. I.
castanipes. 48. I. bliveni. 49. I. silvosa. 50. I. cascadia. 51. I. fuscipubescens. 52. I. mollipes.
53. I. panda. 54. I. setosa.

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215

Figs. 55-66. Posterior view of sclerotized rings and dorsal labiate plate. 55. I. brachycera.
56. 1. elongata. 57. 7. pacifica. 58. 7. sericans. 59. 1, californica. 60. 1. sita. 61.7. cuneomaculata.
62. I. oreas. 63. 7. incomperta. 64. 7. solani. 65. 7. knighti. 66. 7. limata.

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Figs. 67-77. Posterior view of sclerotized rings and dorsal labiate plate. 67. /. nigripes. 68.
I. shulli. 69. I. serrata. 70. I. castanipes. 71. I. bliveni. 72. I. silvosa. 73. I. cascadia. 74. I.
fuscipubescens. 15. I. mollipes. 16. I. panda. 11. I. setosa.

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217

Figs. 78-100. Posterior view of inter-ramal sclerites. 78. I. brachycera. 79. /. elongata.
80. I. pacifica. SI. I. sericans. 82. 1, californica. 83. I. sita. 84. I. cuneomaculata. 85. /. oreas.
86. I. incomperta. 87. I. solani. 88. I. knighti. 89. I. limata. 90. I. nigripes. 91. I. shulli. 92. I.
serrata. 93. /. castanipes. 94. 1. bliveni. 95. /. silvosa. 96. /. cascadia. 97. 1. fuscipubescens. 98.
/. mollipes. 99. /. panda. 100. /. setosa.

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not; hind wings are reduced in brachypterous species or forms. LEGS: moderately
long, hind femora not reaching apex of abdomen; testaceous, fulvous, fuscous, ru-
fuscent or black; femora with short suberect fulvous setae; tibiae with short suberect
black setae; some species have longer erect fulvous setae on the femora and tibiae;
tibiae with four rows of suberect black bristles, and numerous apical black bristles;
femora with a pair of apical black bristles; tibial comb present on apices of protibiae,
formed by a row of short fine setae on ventral side and bordered laterally by a pair
of bristles; three tarsal segments subequal in length; claws are not cleft; puvilli present;
parempodia lamellate and diverging. GENITALIA: Male: left paramere: acutely or
broadly angled, with or without prominent sensory lobe; either with or without
tubercles or spines on angle, arm and shaft; apex either sharp, securiform or truncated;
right paramere: cylindrical; the preapical portion sometimes expanded, if not, the
paramere is of equal diameter throughout; apically with a single simple spine or with
a double faceted apex; vesica: with two membranous lobes either rounded or conical
and with or without accessory lobes of various forms; with a furcate sclerotized
process, the larger process either simple, expanded (with or without a concave furrow)
or linear, this process is acute or blunt apically, and is ornate with various types of
spines, the smaller process possessing a pointed membranous lobe. Female: posterior
wall of the bursa copulatrix with two types of inter-ramal sclerites (that differ in
shape), a prominent sigmoid process with wide and convoluted C structures; scler-
otized rings are oval or compressed, and either open or closed; the dorsal labiate
plate is either narrow throughout or expanded anteriorly.

KEYS TO THE SPECIES OF IRBISIA

I have constructed two keys to the adults of the species of Irbisia; the first attempts
to distinguish the species on external characters alone; the second relies predominately
on the structure of the male and female genitalia. The species of Irbisia display great
similarity in practically all characters. Positive identification of some species is fa-
cilitated by the examination of the internal genitalia of both sexes.

KEY I

1.

1'.

2(1).

2'.

3(2).

3'.

4(2)'.

4'.

5(4).

5'.

6(1').

Brachypterous 2

Macropterous 6

Males 3

Females 4

Pronotum shagreened and transversely rugulose (Fig. 7c) sericans (St&l)

Pronotum shining and shallowly punctate (Fig. 7b) .... knighti Schwartz and Lattin
Distributed west of Cascade Range in Oregon and Washington and along coast
of Alaska and British Columbia (also in Columbia R. Gorge west of The Dalles,

Oregon) 5

Widely distributed east of Cascade Range in British Columbia, Oregon and Wash-
ington; and Rocky Mountain regions of Colorado, Idaho, Montana, Utah and

Wyoming shulli Knight

Pronotum shagreened and transversely rugulose (Fig. 76) sericans (St&l)

Pronotum shining and shallowly punctate (Fig. 7a) knighti Schwartz and Lattin

Antennal socket below ventral margin of eye; eye rounded in lateral view (Figs.

1, 3,4)

7

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REVISION OF IRBISIA

219

6'.

7(6).

7'.

8(7').

8'.

9(6').

9'.

10(9').

10'.

1 1(10).
11.

12(10').

12'.

13(12').

13'.

14(13').

14'.

15(14').

15'.

16(15').

16'.

17(16).

17'.

18(9').

18'.

Antennal socket above ventral margin of eye, or with at most only ventral portion

of socket below eye; eye oval or reniform (Figs. 2, 5, 6) 9

Length of antennal segment II much greater than maximum width of pronotum

(almost 2 x as great) pacifica (Uhler)

Length of antennal segment II less than 1.25 x the maximum width of pro-
notum 8

Tibial setae long, fulvous or white; length of antennal segment I equal to width

of vertex (ratio of vertex to segment I 1:0.90 to 1:1.00) elongata Knight

Tibial setae short, black; length of antennal segment I less than width of vertex

(ratio of vertex to segment I 1:0.60 to 1:0.80 brachycera (Uhler)

Distribution outside of California 10

Distribution in California and Baja California Norte (or in woodlands of Mari-
copa, Pinal and Yavapai Counties, Arizona and Washington County, Utah) .... 18
Dorsum with single type of setae; either sparsely distributed fine short, suberect
or appressed fuscous to golden setae; or moderately or densely distributed woolly

sericeous, recurved, longer golden setae 11

Dorsum with two types of setae; mixture of moderately to densely distributed
fine short setae and recurved (perhaps woolly) sericeous setae (sericeous setae

most noticeable on clavus) 12

Dorsum dull black with bronze luster, and moderately to densely distributed,

woolly sericeous recurved longer golden setae male, sericans (Stfil)

Dorsum shining black with sparsely distributed fine, short, suberect or appressed

fuscous to golden setae fuscipubescens Knight

Ostiolar peritreme (except for fuscous extreme posterior margin) and propleuron

(except for very narrow fuscous trim on ventral apex) black nigripes Knight

Ostiolar peritreme and ventral one quarter of propleuron ivory 13

Ventral surface of head and pleura variably marked with testaceous or ochraceous

patches; female venter light; distributed in southwestern Oregon silvosa Bliven

Ventral surface of head and pleura black with, at most, light colored trim near

coxae 14

Lateral margins of pronotum strongly concave; femora with fuscous spots; dis-
tributed in southwestern Oregon calif or nica Van Duzee

Lateral margins of pronotum straight; femora without fuscous spots 15

Anterior angle of pronotum rounded and indistinct in dorsal view; female femora
rufescent or extensively testaceous; distribution restricted to Willamette Valley
in Oregon, not collected above 300 m. (1000 feet) elevation . . . solani (Heidemann)
Anterior angle of pronotum prominent and distinct in dorsal view; female femora
black or fuscous; distribution widespread (including Willamette Valley, Oregon),

but usually collected above 300 m. (1000 feet) elevation 16

Pronotum shining black, with distinctly separated punctures; calli prominent,
polished; distribution practically restricted to Oregon (collected in Modoc County,

California and Skamania County, Washington) cascadia, new species

Pronotum shagreened, with confluent punctures; calli weakly convex, asperate;

distribution extensive outside of Oregon 17

Width of vertex of males 0.45 to 0.59; females macropterous serrata Bliven

Width of vertex of males 0.68 to 0.75; females brachypterous shulli Knight

Dorsum covered with one type of setae; either sparsely distributed minute ap-
pressed gray setae; or sparsely distributed short, fine, suberect, testaceous, fulvous

or fuscous setae; or densely distributed long, white to golden setae 19

Dorsum covered with two types of setae; mixture of moderately or densely dis-
tributed short to long, suberect to erect, light to black setae and recurved (perhaps
woolly) sericeous setae (sericeous setae most noticeable on clavus) 23

220

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

19(18).

19'.

20(19').

20'.

21(20).

21'.

22(20').

22'.

23(18').

23'.

24(23').

24'.

25(24).

25'.

26(24').

26'.

27(26').

27'.

28(27).

28'.

29(27').

29'.

30(20').

30'.

31(30').

31'.

Dorsum dull black, with obvious bronze luster and sparsely covered with minute

appressed gray setae castanipes Van Duzee

Dorsum shining or polished black, without bronze luster and minute appressed

gray setae 20

Pronotum with very faint, shallow punctures; dorsum sparsely covered with short,

fine suberect setae 21

Pronotum with obvious deep punctures, or confluently punctate, rugulose or

rugose; dorsum with densely distributed, long setae 22

Antennal segment I and legs flavescent; basalar plate shining black . . . limata Bliven

Antennal segment I and legs fuscous to black; basalar plate ivory panda Bliven

Width of vertex ranging from 0.56 to 0.64 in males, and from 0.60 to 0.70 in
females; ventral femoral setae densely distributed and erect; protibial setae long

and fulvous bliveni, new species

Width of vertex ranging from 0.40 to 0.50 in males, and from 0.48 to 0.55 in
females; ventral femoral setae usually sparsely distributed and suberect; protibial

setae usually short and black mollipes Van Duzee

Cuneus with basal margin testaceous cuneomaculata Blatchley

Cuneus black 24

Vestiture of apical portion of hemelytra moderately or densely distributed long
(projecting far above sericeous setae), erect, bristle-like fulvous to black setae and

densely distributed woolly sericeous setae 25

Vestiture of apical portion of hemelytra never with long erect setae that project

far above sericeous woolly setae 26

Tibiae densely covered with long (longer than tibial bristles) suberect fine white

setae oreas Bliven

Tibiae moderately covered with shorter (much less than length of tibial bristles)

suberect fuscous setae setosa Van Duzee

Ventral surface of head and pleura variably marked with testaceous or ochraceous;

female venter light silvosa Bliven

Ventral surface of head and pleura black with, at most, light colored trim near

coxae 27

Lateral margins of pronotum strongly concave 28

Lateral margins of pronotum straight, or, at most, very slightly concave 29

Femora with fuscous spots californica Van Duzee

Femora without fuscous spots sit a Van Duzee

Basalar plate with variable portion of posterior margin ochraceous; median
depression of head obvious, either distinct longitudinal furrow or broad sunken

area; female femora with fuscous spots incomperta Bliven

Basalar plate black; median depression of head shallow or indistinct; femora

without fuscous spots 30

Dorsal vestiture a mixture of densely distributed, long setae that are erect on head
and pronotum and suberect on hemelytra; and woolly sericeous setae; occurs in

Kern County, California bliveni, new species

Dorsal vestiture a mixture of moderately distributed, medium length, suberect
setae; and woolly sericeous setae; occurs widely outside of Kern County, Cali-
fornia 31

Anterior angle of pronotum rounded and indistinct in dorsal view; female femora

may be rufescent; distribution widespread in California solani (Heidemann)

Anterior angle of pronotum prominent and distinct in dorsal view; female femora
black or fuscous; distribution in northern California 32

1984

REVISION OF IRBISIA

221

32(31). Pronotum shining black, with distinctly separated punctures; calli prominent,

polished cascadia, new species

32'. Pronotum shagreened, with confluent punctures; calli weakly convex, asperate .

serrata Bliven

KEY II

1 . Antennal socket, in lateral and cephalic view, below ventral margin of eye; eye

round in lateral view (Fig. 1),; ratio of distance between tylus and ventral margin
of eye to height of eye less than or equal to 1:0.50; eyes narrowly joined to
frons— angle at junction of frons and ventral margin of eye, in frontal view, less

than or equal to 135° (Figs. 3, 4) 2

1 '. Antennal socket, in lateral and cephalic view, at most with only ventral portion
below ventral margin of eye; eye oval or reniform in lateral view (Fig. 2); ratio
of distance between tylus and ventral margin of eye to height of eye greater than
1:0.50 (usually ratio greater than 1:0.60, only female specimens of I. shulli with
a ratio approaching 1:0.50, but are distinguished by brachyptery); eyes broadly
joined to frons— angle at junction of frons and ventral margin of eye, in frontal

view, greater than 135° (Figs. 5, 6) 4

2(1). Length of antennal segment II almost 2 x maximum width of pronotum; length
of antennal segment I greater than width of vertex— ratio 1 : 1 .00 to 1:1 .40; head
anterior of eyes elongate— ratio of width of head across eyes to head length 1:

0.75 (Figs. 1, 3), eyes in lateral view projecting above basal carina; body length

5.75 to 8.20 pacifica (Uhler)

2'. Length of antennal segment II less than or equal to 1.25 x the maximum width
of pronotum; length of antennal segment I, at most, equal to width of vertex—
ratio 1:0.60 to 1:1.00; head anterior of eyes somewhat elongate— ratio of width
of head across eyes to head length 1:0.50 to 1:0.60 (Fig. 4), eyes in lateral view

not projecting above basal carina; body length 4.90 to 6.25 3

3(2'). Tibial spines long, fuscous; body elongate, parallel-sided, width 1.70 to 2.00
males, 1.08 to 2.40 females; length of antennal segment I subequal to width of
vertex— ratio ranging from 1:0.90 to 1:1.00; ratio of distance between tylus and
ventral margin of eye to height of eye 1:0.50; pronotum conical . . .elongata Knight
3'. Tibial spines short, black; body stocky, ovate, width 1.85 to 2.30 males, 2.21
to 2.78 females; length of antennal segment I less than width of vertex— ratio
ranging from 1 :0.60 to 1 :0.80; ratio of distance between tylus and ventral margin

of eye to height of eye greater than 1:0.50 (Fig. 4); pronotum trapezoidal

brachycera (Uhler)

4(1'). Male parameres as in Figures 12a, b to 14a, b, with the following combination
of characters: left paramere— sensory lobe straight, length greater than or equal
to length of arm, and distally expanded with apex truncated at right angles to
axis of shaft; angle sharp, ’V’ shaped (less than 45°); right paramere— cylindrical
throughout, slightly curved mesad; apex with double faceted spine directed at
right angle to long axis of paramere. Females either: 1) brachyperous, with
transversely rugose pronotal disk; distributed in Oregon and Washington west
of Cascade Range, as far south as Eugene, Lane County, Oregon, and in British
Columbia and Alaska in coastal grasslands as far west as Kamchatka Peninsula
of U.S.S.R.; or 2) with lateral margins of pronotum transversely rugose; vestiture
a mixture of long, golden to white setae and woolly sericeous white setae; femora
with fuscous spots; distributed throughout California within and east of the

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

4'.

5(4).

5'.

6(5').

6'.

7'.

8(7').

8'.

9(8).

9'.

10(8').

10'.

11(10').

Coastal Ranges; or 3) dorsum polished shining black; pronotal disk distinctly
and closly punctate; femora fulvous to castaneous (without fuscous spots); tibiae
without long setae; venter black; vestiture without long bristle-like setae; basalar

plate entirely black 5

Male parameres as in Figures 1 5a, b to 3 1 a, b, without above combination of
characters: left paramere— sensory lobe small or indistinct, i.e. barely produced
above outline of arm, shaft curved toward sensory lobe, length less than or equal
to length of arm, and distally expanded with apex truncated parallel to axis of
shaft; angle broad, ’U’ shaped (60-90°); right paramere— cylindrical, but ex-
panded distally; apex with simple spine directed in line with long axis of par-

amere. Females without the previous arrangement of characters 7

Females and some males brachypterous; pronotum trapezoidal, lateral margin

straight sericans (St&l)

Both sexes macropterous; lateral margin of pronotum concave 6

Femora with fuscous spots; head long ventrad of eyes— ratio of distance between
tylus and ventral margin of eye to height of eye 1:0.68 to 1:0.86 for males and
1:0.60 to 1:0.80 for females; dorsum shining black; pronotal disk rugulose or
confluently punctate; femora testaceous, fulvous or fuscous; body length 4.75-

7.30 calif or nica Van Duzee

Femora without fuscous spots; head shorter ventrad of eyes— ratio of distance
between tylus and ventral margin of eye to height of eye 1 :0.96 to 1 : 1 .04 for
males and from 1:0.85 to 1:1.04 for females; dorsum polished shining black;
pronotal disk distinctly and closely punctate; femora fulvous to castaneous; body

length 4.25-5.40 sit a Van Duzee

Dorsum shining black without obvious bronze luster; vestiture single type of
long upright or short appressed setae or mixture of fine setae; woolly sericeous

setae or long bristle-like setae 8

Dorsum with very faint, shallow punctures; dorsum sparsely covered with short,
fine suberect testaceous, fulvous or fuscous setae; antennal segment I flavescent,

or fuscous and black with testaceous apex 9

Pronotal surface sculptured with obvious, deep punctures, or confluently punc-
tate, rugulose or rugose; dorsum black with longer single type of setae or mixture
of fine setae, woolly sericeous setae, and long bristle-like setae; antennal segment

I never entirely flavescent 10

Antennal segment I and legs flavescent; basalar plate shining black; left paramere
without tubercles or spines (Fig. 20a); axis of sclerotized process of vesica of
male roughly parallel with main axis of vesica (Fig. 43); sclerotized rings of

bursa copulatrix of female oval and open (Fig. 66) limata Bliven

Antennal segment I fuscous to black; legs fuscous or piceous, femora rufescent
medially; basalar plate ivory; left paramere with tubercles or spines (Fig. 30a);
axis of sclerotized process of vesica deflected 30° to left of main axis of vesica
(Fig. 53); sclerotized rings of bursa copulatrix compressed, truncated on the
medial margin and closed (Fig. 77) panda Bliven

Cuneus with basal margin testaceous cuneomaculata Van Duzee

Cuneus black 11

Vental surface of body variably marked with testaceous or ochraceous patches;
females with venter light (dorsolateral region of stemites may be infuscate);
males predominately black with light coloration on jugal-loral sutures; sclero-
tized process truncated distally, distal one third of frontal margin with rows of
large spines, decreasing in length apically, frontal surface without deep concavity
(Fig. 49) silvosa Bliven

1984

REVISION OF IRBISIA

223

11'.

12(11').

12'.

13(12).

13'.

14(12').

14'.

15(14').

15'.

16(15').

16'.

17(16').

Ventral surface of body of both sexes black (jugal-loral sutures black); sclerotized
process rounded distally, arrangement of spines different than above (Figs. 39-

42, 44-48, 52, 54); if with large spines then frontal surface with deep concavity

(Figs. 19, 20) 12

Dorsal vestiture a mixture of moderately or densely distributed long, erect
fulvous to black bristle-like setae projecting above a layer of densely distributed,
woolly sericeous setae; tibiae may be densely covered with long (subequal to

length of tibial bristles) suberect testaceous or white setae 13

Dorsal vestiture never with long erect setae projecting above sericeous woolly

setae 14

Tibiae densely covered with long (subequal to tibial bristles) suberect, fine,
testaceous or white setae; dorsum densely covered with long fulvous setae, erect
on head, pronotum, and scutellum, suberect on hemelytra; body length— males
3.83 to 4.75, females 4.50 to 5.10; right paramere with apical spine minute,

barely projecting beyond apical surface of paramere (Fig. 16b) oreas Bliven

Tibiae moderately covered with shorter (length much less than length of tibial
bristles) suberect fuscous setae; dorsum moderately covered with uniformly
erect, long black or fuscous setae; body length greater— males 4.70 to 5.40,
females 4.70 to 5.45; right paramere with apical spine prominent, projecting

well beyond apical surface of paramere (Fig. 31) setosa Van Duzee

Legs entirely black; propleura and pleura entirely black on most specimens,
narrow fuscous trim present or not on ventral apex of propleura, pleura (bor-
dering coxae) and posterior margin of ostiolar peritreme; anterior angle of prono-
tum very prominent in dorsal and lateral view, angle slightly explanate and

directed anterad; pronotal disk somewhat flattened nigripes Knight

Legs not entirely black; trim on propleura and pleura and ostiolar peritreme
extensive, ivory or testaceous; anterior angle of pronotum either prominent, but
rounded, or reduced in dorsal and lateral view; pronotal disk broadly arcuate 1 5
Both sexes brachypterous; width of vertex of female 0.66 to 0.70; dorsum mod-
erately covered with single type of white setae; distributed on coastal flatland

of Pacific Northwest knighti Schwartz and Lattin

Both sexes usually macropterous (if female brachypterous then: width of vertex
of female 0.73 to 0.85; dorsum densely covered with combination of suberect
white, fulvous or fuscous setae and woolly sericeous setae; distributed east of

Cascade Range in Oregon and Washington 16

Densely distributed long black tibial setae; width of vertex— males 0.68 to 0.75,
females 0.73 to 0.85; females brachypterous; pronotum rugulopunctate; scler-
otized process as in Figure 45, with posterior half greatly expanded to left side
of the vertical axis of the process— expansion is 30 percent of height of process
in posterior view (height of process measured from apex to insertion of process
on vesica); spines of sclerotized process of one type, and bordering lateral margin

of the lateral expansion shulli Knight

Sparsely or moderately distributed short black or fuscous tibial setae; width of
vertex— males, 0.40 to 0.63; females 0.48 to 0.70; females macropterous; scler-
otized process either not expanded or with two types of spines (if process is
slightly expanded then pronotum distinctly punctured) (Figs. 40, 41, 46-48, 50-

52,54) 17

Dorsum densely covered with single type of suberect, long, shining pale white
to pale golden setae (if long setae mixed with woolly, sericeous setae then,
pronotal surface strongly rugose with distinct punctures; calli reaching antero-
lateral margin of pronotum, broadly confluent anteriorly, strongly asperate;
anterior angle blunt, not produced, and rounded in dorsal view) 18

224

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

17'.

18(17).

18'.

19(17').

19'.

20(19').

20'.

21(20').

21'.

22(21').

22'.

Dorsum moderately covered with short, fine fuscous setae, or dorsal surface

moderately or densely covered with two types of setae 19

Dorsum densely covered with single type of setae; dorsum polished shining
black; pronotum conical in dorsal view; width of vertex— males 0.40 to 0.50,
females 0.48 to 0.55; ratio of width of vertex to length of antennal segment I—
males 1:1.30 to 1:1.70, females 1:1.35 to 1:1.53; lateral membranous lobes of

the vesica tapering distally (Fig. 52); sclerotized rings closed (Fig. 75)

mollipes Van Duzee

Dorsum with either one or two types of setae; pronotum trapezoidal; vertex of
head wider— males 0.56 to 0.64, females 0.60 to 0.70; ratio of width of vertex
to length of antennal segment I— males 1:1.10 to 1:1.30, females 1:0.90 to 1:

1 .05; lateral membranous lobes of the vesica rounded (Fig. 48); sclerotized rings

open (Fig. 71) bliveni, new species

Basalar plate of pleuron with dorsal margin, to variable degree, light colored,
either ivory, testaceous, fulvous or fuscous; female femora with fuscous spots
ventrally; temporal areas of head distinct, slightly flattened with obvious trans-
verse sulcus on posterior margin; median depression of head obvious— either
distinct longitudinal furrow or broad sunken area; body length, male 4.30 to
4.90, female 4.87 to 5.50; width, male 1.70 to 1.95, female 1.90 to 2.25; form
of shaft of left paramere slender, slightly expanded and curved (toward sensory

lobe) distally, ventral surface of apex without obvious flanges (Fig. 17a)

incomperta Bliven

Basalar plate black; female femora never with fuscous spots; temporal areas of
head rounded, without obvious transverse sulcus on posterior margin; median
depression of head indistinct; body length, male 3.80 to 6.10, female 4.60 to
6.00; width, male 1 .70 to 2.80, female 2. 10 to 3.00; shaft of left paramere stocky,
strongly expanded and curved (toward sensory lobe) distally, ventral surface of

apex with large flange (Figs. 18, 23, 27, 28) 20

Anterior angle of pronotum rounded and indistinct in dorsal view; anterior
angle, in lateral view, broadly rounded at junction of pronotum and dorsal-most
portion of ridge posterior to coxal cleft of propleuron; female femora sometimes
extensively rufescent; left paramere with weak spines (Fig. 18); dorsal labiate

plate of the bursa copulatrix not strongly expanded anteriorly (Fig. 64)

solani (Heidemann)

Anterior angle of pronotum prominent and distinct in dorsal view; anterior
angle, in lateral view, distinctly concave at junction of pronotum and dorsal-
most portion of ridge posterior to coxal cleft of propleura (anterior angle ov-
erhanging the ridge); femora black or fuscous, never rufescent; left paramere
with strong spines (Figs. 23, 27, 28); dorsal labiate plate of the bursa copulatrix

strongly expanded anteriorly 21

Dorsum with sparsely distributed, single type of fine, short suberect or appressed

fuscous to golden setae fuscipubescens Knight

Dorsum with moderately to densely distributed mixture of short or medium,
suberect or erect, white, fulvous or golden setae and recurved sericeous setae . . 22
Pronotum shining black, with punctures separate; calli prominent, polished;
ratio of width of vertex to length of antennal segment I ranging from 1:0.77 to
1:0.93 for males, and 1:0.72 to 1:0.87 for females; sclerotized process with very

large and prominent spines (Fig. 50) cascadia, new species

Pronotum shagreened, punctures confluent; calli weakly convex, asperate; ratio
of width of vertex to length of antennal segment I ranging from 1 : 1 . 10 to 1 : 1 .24
for males, and 1 : 0.8 9 to 1 : 1 .07 for females; sclerotized process with small spines
(Fig. 46) serrata Bliven

1984

REVISION OF 1RBISIA

225

Irbisia bliveni, new species
Figs. 25a, b, 48, 71, 94; Map 17

Diagnosis. Irbisia bliveni can be separated from most of the other species of Irbisia
by the structure of the vestiture. Both I. bliveni and I. mollipes Van Duzee are densely
covered with a single type of suberect, long shining pale white to pale goldish setae.
I. bliveni differs from I. mollipes in the former species’ wider vertex— vertex to
antennal segment I ratio for I. bliveni ranging from 1:1.07 to 1:1.30 for the male,
and from 1:0.88 to 1:1.05 for the female. The ratio for I. mollipes ranging from 1:

I. 29 to 1:1.70 for male and from 1:1.36 to 1:1.53 for the female. The sclerotized
process of the vesica of the male I. bliveni is unique among the species of the genus.
It is long and narrow throughout, with minute serrations on the thin frontal margin.

Description. Macropterous male. Length 5.70-6. 10, width 2.20-2.60, shining black,
densely clothed with long, erect light setae. HEAD: width across eyes 1.26-1.35,
vertex 0.59-0.65, dorsal width of eye 0.33-0.38; triangular, smooth; temporal area
glabrous; basal carina prominent, abruptly rising from vertex, sublinear; median
depression broad, sometimes deep; eyes large broadly joined to frons, posterior
margin straight, distance from tylus to ventral margin of eye 0.63-0.65, height of
eye 0.53-0.58, maximum interocular width 0.90-0.91. ROSTRUM: length 1.95-
2.03, black, reaching apex of mesocoxae. ANTENNAE: black; I, length 0.64-0.73;

II, 1.79-1.93; III, 0.95-1.05; IV, 1.20-1.28. PRONOTUM: length 1.16-1.20; anterior
width 0.93-0.98; maximum width 2.98-2.03; trapezoidal, disk broadly convex, ru-
gose with distinct punctures, posterior margin straight, lateral margins straight or
slightly concave, rounded at junction with propleura, anterior angles blunt or round-
ed, not prominent in dorsal view, but sulcate dorsad of coxal cleft; calli slightly
elevated, confluent anteriorly, asperate, with fovate impression on inner anterior
angle, calli reaching anterolateral margin of pronotum. LEGS: black; tibiae fuscous;
apices black. VESTITURE: dorsum with dense, long, erect white to fulvous setae,
intermixed (in variable degree) with fine, recurved or prostrate sericeous white setae;
venter, ventral surface of femora and tibiae with long testaceous setae. GENITALIA:
Left paramere (Fig. 25a): sensory lobe slightly projecting above arm; angle broad;
length of shaft less than length of arm; arm and shaft with prominent spines; arm
diameter with distal two thirds constricted; apex moderately securiform; with nu-
merous bristles. Right paramere (Fig. 25b): diameter of shaft expanded distally; apical
process with double faceted spine; with a few short lateral setae. Vesica (Fig. 48):
two smooth distally broad membranous lobes; sclerotized process with proximal two
thirds of narrow diameter, distal one third gradually expanded parallel to the axis of
the sclerotized process, ventral surface with numerous small serrations.

Macropterous female. Slightly more robust than male, but similar in color, vestiture
and structure. Length 5.90-6.55, width 2.25-2.95. HEAD: width across eyes 1.29-
1.40, vertex 0.60-0.70, dorsal width of eye 0.35-0.38, distance from tylus to ventral
margin of eye 0.65-0.73, height of eye 0.53-0.60, maximum interocular width 0. 95-
LOO. ROSTRUM: length 1.95-2.23, reaching or surpassing mesocoxae. ANTEN-
NAE: I, length 0.60-0.68; II, 1.48-1.84; III, 0.90-1.13; IV, 1.02-1.08. PRONOTUM:
length 1.13-1.20, anterior width 0.90-1.10, maximum width 1.85-2.28. GENITA-
LIA: sclerotized rings (Fig. 71): area within rings oval; dorsal labiate plate uniformly
narrow anterior of rings; width of one ring 0.25-0.30, maximum width across rings
0.81-1.15. Posterior wall (Fig. 94): A-structure curving, ventral margin broadly

226

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

rounded; dorsal margin straight. B-structure protruding posteriorly, without inden-
tations on ventral posterior surface; median process gradually expanded dorsad,
maximum length 0.25-0.33, maximum width 0.56-0.75.

Holotype. Male. Hat Creek, California, [Shasta Co.] Lassen National Forest, 24
June 1961, S. L. Wood, J. B. Karren and D. E. Bright, BYU; CAS type no. 15182.
Retained at the California Academy of Sciences, San Francisco. Two 86 and four 99
paratypes with the same label data, and two 88 and two 9$ with Hat Crk., UCB are
also at CAS.

Etymology. Named for Brunson P. Bliven, an avid and reclusive student of the
Hemiptera. He named nineteen new Irbisia taxa, six of which I believe deserve the
status of valid species1.

Taxonomic consideration. The specimens from north of Kern County, California
are very similar to the original description and homogeneous. Specimens from the
southern part of the range of this species are slightly different from the more northern
populations. In the southern populations the sclerotized rings of the female are closer
together (width across the rings is 0.85 in the south and 1.10 in the north), the
sclerotized process of the vesica of the male is less constricted (giving the appearance
of a teneral specimen). Narrower vertices and more abundant sericeous setae on the
dorsum also characterize these southern specimens. Additional material from the
Tehachapi and San Bernardino ranges would aid in determining if two species are
actually involved.

Remarks. I. bliveni has a distribution which is restricted to the Sierra Forest

1 Brunson P. Bliven, the “city entomologist of Eureka, California since 1937” in his first
work (1954) published descriptions of eight new species of Miridae and Cicadellidae in the
Bulletin of the Brooklyn Entomological Society. This article was the only one of his thirteen
papers, a total of 192 pages establishing 124 “new” species, which was not published privately.
“The Occidental Entomologist” (1957-1973) and its two predecessors “Insects of the Redwood
Empire I” (1955) and “New Hemiptera from the Western States” (1956) were published and
edited by Bliven himself. The circumstances which precipitated Bliven to publish privately
surrounded the review process of his manuscript, the placing of holotypes from his first paper
at the United States National Museum and the subsequent questioning of the validity of some
number of these species by other hemipterists.

Apparently Bliven desired to conduct his research in private to avoid the critique of his peers.
His specimens were not available for loan to other researchers during his lifetime. Kelton (196 1)
in his synopsis of the Nearctic species of Stenodema Laporte, had what may have been the
only working relationship with Bliven. Kelton had his manuscript read and the included key
checked with three Bliven species in the Eureka entomologist’s office.

Bliven passed away in the fall of 1980 at his home. He was found, amid piles of dusty,
unopened correspondence by the postal carrier delivering his mail-ordered food stuffs. Shortly
after he died his research collection and library were advertised for sale by his heirs in the
Bulletin of the Entomological Society of America. After considerable bargaining the California
Academy of Sciences and California State Department of Agriculture joinly acquired Bliven’s
tangible legacy. The type material is now retained at the California Academy. My revision of
Irbisia represents the first publication in which Bliven material was freely available for study.
I want to personally thank Dr. Paul H. Amaud, Jr. for allowing me to examined the type
specimens, soon after their arrival to the Academy, in time to complete this revision when it
was a thesis manuscript.

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REVISION OF IRBISIA

227

province, and ponderosa shrub forest section of the Intermountain sagebrush prov-
ince (Bailey, 1976). Within the mixed conifer forest of the Sierras, this species occurs
in floristic associations which have pines as a component. Adult specimens have
been collected on Pinus contorta Dougl. ex Loud., P. ponderosa Dougl. ex P. & C.
Lawson, P. sabiniana Dougl. and Pinus sp. Circumstantial evidence (specimens from
Siskiyou County, California, densely covered with pollen) indicates that adults may
feed on pine pollen. The primary grass hosts of this species are unknown but are
probably an understory component of pine forests. Adult specimens were collected
from 28 April to 10 July, and from 381 to 1676 m. (1250 to 5500 feet) elevation.

Specimens examined. 54 additional specimens were examined from the following localities
[specimens from localities which typify southern populations are marked with an asterisk] (Map
17). UNITED STATES: CALIFORNIA: Colusa Co., Wilbur Hot Spgs., CL 1622, 21 April
1980, J. T. Polhemus, JTP (a). Fresno Co., Boy Scout Cmp.— Coalinga, 19 February 1931, E.
P. Van Duzee, CAS (3); ““Converse Basin— near Hume, Lake Dist., Sequoia Nat. For., 1 1 June
1961, E. Ball, CAS (9); Dalton, Rgr. Stn., 5000 ft., 2 May 1920, H. Dietrich, USNM (3). *Kern
Co., Antelope Cyn.— Tehachapi Mts., 15 May 1976, J. D. Pinto, UCR (6 33, 9); Tehachapi Mts.,
4800-5000 ft., 15 May 1976, G. Clark, UCR (366, 499) and 14 May 1976 (6); 3 mi. W Woffard
Hts., 4200 ft., 28 April 1964, W. Turner, UCB (499) and R. L. Langston (5); 4 mi. W Woffard
Hts., 4200 ft., 14 April 1965, G. L. Jensen & W. J. Turner, ex Pinus sabiniana, UCB (6).
Madera Co., Oakhurst, 19 May 1942, C. Kennett, sweeping, UCB (9). Mariposa Co., Miami
Rgr. Stn., 16 May 1972, beating, ex Pinus ponderosa, UCB (9); *Yosemite, 16 May 1931, 3880-
4000 ft., D. W. Clancy, UCR (9) and *27 May 1931, UCB (9); *Yosemite Nat. Pk., 1 1 June
1932, USNM (9). Napa Co., N side Howell Mt.-2 mi. NNE Angwin, 1300 ft., 8 April 1980,
H. B. Leech, ex foliage of Pseudotsuga menziesii, CAS (3). Plumas Co., Meadow Valley, 3500-
4000 ft., 6 June 1924, E. C. Van Dyke, CAS (9). *San Bernardino Co., S of Cmp. Angeles, 4100
ft., 1 1 May 1978, J. D. Pinto, UCR (299); above Mt. Home— San Bernardino Mts., 440 m., 1 1
May 1978, J. D. Pinto & R. T. Schuh, AMNH (9); Wrightwood, 5000 ft., 25 May 1956, H.
Ruckes, Jr. & B. J. Adelson, ex Pinus monophylla, UCB (9). Shasta Co., Redding, 4 April 1966,
WSU (6, 9) and 6 April 1966, B. A. Freeman, WSU (9). Siskiyou Co., 1 mi. W Bartle, 1220 m.,
7 July 1979, R. T. & J. Schuh, AMNH (3). Tehama Co., Saddle Cmp., 31 May 1951, H. H.
Keiffer, ex Pinus sp., F&A (6, 499). Tuolumne Co., Bald Mt.— nr. Long Bam, 5600 ft., 20 April
1934, E. P. Van Duzee, CAS (9); Pine Crest, E. P. Van Duzee, CAS (9). *Sand Flat, 5500 ft., 7
June 1930, E. O. Essig, UCB (9), 1 1 June 1930, UCB (9) and D. W. Clancy, UCR (9). (UCB).
NEVADA: “Nev.”, P. R. Uhler collection, USNM (9). OREGON: Jackson Co., Buckhom
Mineral Spgs. — 1 1 mi. ESE Ashland, Emigrant Crk., 2800 ft., 19 May 1960, OSU (3). Josephine
Co., 10 mi. W Selma, 1250 ft., 13 May 1972, J. Sawbridge, OSU (9). WASHINGTON: Yakima
Co., Signal Pk.— Mt. Adams, [37 mi. W Toppenish], 10 July 1927, M. W. Stone, OSU (9).

Irbisia brachycera (Uhler)

Figs. 4, 9a, b, 32, 55, 78; Map 1

Rhopalotomus brachycerus Uhler, 1872, p. 416 (new species); 1875, p. 319.

Capsus brachycerus : Uhler, 1886, p. 19. Atkinson, 1890, p. 105. Cockerell, 1893, p.
363 (misspelled br achy corns).

Thyrillus brachycerus: Uhler, 1894, p. 267. Gillette and Baker, 1895, p. 39.

Irbisia brachycerus: Van Duzee, 1916b, p. 38 (in part); 1917a, p. 325 (in part).

Parshley, 1922, p. 15. Carvalho, 1959, pp. 104-105 (in part).

Irbisia arcuata Van Duzee, 1921a, pp. 148-149, 151 (first synonymized by Knight
1941, p. 79).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

Irbisia paeta Van Duzee, 1921a, pp. 150-151, 152. New Synonymy.

Irbisia brachycera: Knight, 1941, p. 79; 1968, p. 185. Haws 1978, pp. 100, 101, fig.

34. Kelton, 1980, p. 85, map 24.

Irbisia vestifera Bliven, 1961, pp. 46-^17; 1963, p. 85, pi. 7, figs. 4, 4a. New Synonymy.
Irbisia gorgoniensis Bliven, 1961, pp. 47-48; 1963, p. 85. pi. 7, figs. 5, 5a. New

Synonymy.

Irbisia tejonica Bliven, 1961, p. 48; 1963, p. 85, pi. 7, figs. 6, 6a. New Synonymy.

Diagnosis. Regardless of the type of vestiture, surface features of the pronotum
and size, all individuals of this species are readily separated from the majority of the
Irbisia species by their “pedunculate” eye structure. Both I. pacifica and I. elongata
exhibit somewhat similar eye structure to I. brachycera. I. brachycera may be seg-
regated from these two species on the basis of these antennal characters: segment II
does not exceed the width of the base of the pronotum (as in I. pacifica) and segment
I is much shorter than the width of the vertex (in I. elongata, segment I is subsequal).

Description. Macropterous male. Length 4.90-5.90, width 1.85-2.30, shining black,
covered with a combination of short to long, white, sericeous, appressed setae and
short to long, white, suberect to erect setae. HEAD: width across eyes 1.23-1.45,
vertex 0.64-0.84, dorsal width of eye 0.29-0.34; smooth or asperate; temporal areas
distinct or undifferentiated; basal carina weakly or strongly rounded, wide, posteriorly
arcuate mesad, always declivous to level of vertex; median depression shallow; eyes
small, roundish, slightly pedunculate; posterior margin of head (including dorsal
margin of eyes) straight in dorsal view, distance from tylus to ventral margin of eye

0. 70-0.78, height of eye 0.41-0.43, maximum interocular width 0.88-1.05; lora in
some specimens with pale spot at base; antennal sockets below ventral margin of
eye. ROSTRUM: length 1.85-2.20, infuscated, fulvous or pale, reaching metacoxae.
ANTENNAE: black; I, length 0.48-0.75; II, 1.30-1.95; III, 0.63-1.05; IV, 0.86-0.98.
PRONOTUM: length 1.00-1.23; anterior width 0.88-1.03; maximum width 1.58-
1.88; trapezoidal, disk convex, rugulose, rugulopunctate or mostly punctate, posterior
margin nearly straight, lateral margins straight, or slightly to strongly concave, broadly
rounded at junction with propleura, anterior angles blunt, evident but not prominent;
calli barely produced, confluent anteriorly, smooth or asperate. LEGS: black, darkly
infuscated, castaneous or fulvous; tibiae always paler; coxae pale apically on some
dark legged specimens, or dark basally on some light legged specimens; tarsi always
black. VESTITURE: dorsum clothed with sericeous, appressed, white setae and/or
longer, suberect or erect white setae; venter with long appressed or upright setae.

Macropterous female. More robust than male, but similar in color, structure and
vestiture. Length 4.90-6.20, width 2.21-2.78. HEAD: length 0.63-0.83, width across
eyes 1.30-1.53, vertex 0.68-0.86, dorsal width of eye 0.30-0.35, distance from tylus
to ventral margin of eye 0.71-0.90, height of eye 0.43-0.45, maximum interocular
width 0.95-1.18. ROSTRUM: length 2.10-2.35, reaching metacoxae. ANTENNAE:

1, length 0.44-0.69; II, 1 .20-1 .83; III, 0.80-1 . 1 3; IV, 0.88-1 .28. PRONOTUM: length
1.08-1.40, anterior width 0.93-1.15, maximum width 1.65-2.08.

Types. Described from a series from Weld County, Colorado. In 1979, R. C.
Froeschner located a female specimen, without legs or terminal antennal segments
and designated it as one of the syntypes of I. brachycera. This female was the only
specimen located from the original series; it bears these labels: PR Uhler Collection;

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REVISION OF IRBISIA

229

Thyrillus brachycerus, Weld Co., Uhler Col.; Thyrillus brachycerus Uhler, Weld Co.,
Det. Uhler; USNM type no. 1129. I have examined this specimen and designated it
the lectotype of Rhopalotomus brachycerus. The lectotype is retained at the United
States National Museum, Washington, D.C.

Taxonomic consideration. I. brachycera is a morphologically variable species with
a wide geographic distribution. The following external morphological characters ty-
pify specimens from most of its range: vertex to antennal segment I ratio, males 1 :

0. 79 to 1:0.85, females 1:0.71 to 1:0.80; pronotal surface rugulose, rugulopunctate
or scabrous; dorsum covered with a preponderance of slightly thickened, long, se-
riceous, white setae, which are upright and antrorse on the head, and dense, woolly
and appressed on the pronotum and hemelytra; legs fulvous to castaneous or darkly
infuscate. This widespread form is the nominate species, I. brachycera. A localized
form from Trinity County, California, with concave lateral pronotal margins was
described by Bliven (1961) as I. vestifica Bliven. Specimens within and south of the
Transverse Ranges in Kern, Los Angeles, Orange and Riverside Counties as well as
within the Peninsular Ranges of San Diego County, California are of two forms. One
form is similar to the previous type in all characters except vestiture. It possesses a
predominance of thin, long, upright, white setae and is also thinly clothed with shorter,
appressed, sericeous white setae; the legs tend to be quite dark on most specimens.
This form was described as I. tejonica by Bliven (1961). The other southern Cali-
fornian form is recognized by the following set of characters: vertex to antennal
segment I ratio, males 1:0.71 to 1:0.75, females 1:0.64 to 1:0.67; pronotal surface
punctate; dorsum covered with a predominance of thin, long, upright, white setae
which overlie sparse appressed, sericeous, white setae; leg color whitish testaceous
to flavous. This form has received species status twice as I. paeta Van Duzee (1921a)
and I. gorgoniensis Bliven (1961).

I have synonymized all of the forms above with I. brachycera. This conclusion is
based on all of the forms possessing identical male and female genitalia, and the
existence of several populations with intermediate states of the previous characters.
But, where the typical form and the smaller paeta form are sympatric they are
apparently separated by different plant associations. In San Diego County, California
specimens of the typical form were collected from isolated patches of California
prairie and juniper-piny on woodland. In Riverside County, California specimens of
the form paeta were collected from chaparral and southern yellow pine forests.

Remarks. Short economic reports of a descriptive nature (Vosler, 1913; Child,
1914; Essig, 1915) mistakenly identified I. solani as I. brachycera. These errors were
rectified by Essig (1926). The specimens upon which these reports were based have
been verified as I. solani and correctly labeled by me. Haws (1978) notes that feeding
damage by I. brachycera and I. pacifica on Intermountain range grasses is very similar
to that of Labops hesperius. Cockerell (1893) is the only citation for which I am
unable to determine if the specimens of concern are conspecific with I. brachycera.
I have not seen specimens determined by the workers mentioned by Cockerell. Both

1. brachycera and I. serrata occur in Weld County, Colorado.

This species has a wide distribution. The easternmost and northernmost records
are from Sioux City, Iowa and Medicine Hat, Alberta respectively (Knight, 1941). I
have not verified the identification of the former. No specimens from the states of
Washington or New Mexico were located in the extensive material on hand. Although

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the distribution of I. brachycera is great, encompassing eight ecoregional provinces
(Bailey, 1976), individuals of this species are restricted to open woodland and chap-
arral situations. Within these habitats, the occurrence of the active life stages of the
bug is correlated with the availability of winter and spring moisture regenerating the
cool season grass hosts.

Collection dates for the majority of the specimens are from mid May to late June.
The earliest date is 10 February from Berkeley, Alameda County, California; the
latest is 1 October from Monarch Pass, Gunnison County, Colorado. Specimens have
been taken from sea level to 3448 m. (11, 312 feet) elevation.

Adult specimens were collected from the following grass hosts: Agropyron cristatum
(L.), A. spicatum (Pursh) Scribn. & Sm., Agropyron sp., Avena fatua L., Bromus mollis
L., B. rigidus Roth, B. rubens L., Bromus sp., Elymus triticoides Buckl., Festuca rubra
L., Festuca sp., Hordeum leporinum Link., Hordeum sp., Secale cereale L., Sitanion
hystrix (Nutt.) J. G. Sm. and Triticum sp. Adult specimens were also collected from
the following non-grass plants: Artemisia tridentata Nutt., Balsamorhiza sagittata
(Pursh) Nutt., Balsamorhiza sp., Ceanothus integerrimus H. & A., Chrysothamnus
sp., C. nauseosus (Pall.) Britton., Erigeron sp., Eriodictyon sp., Juglans californica
Wats., Lasthenia sp., Lotus sp., Lupinus sp., Medicago sp., Phacelia sp., Pinus jeffreyi
Grev. & Balf. in A. Murr., Platystemon californicus Berth., Quercus chrysolepis Liebm.,
Salix sp., Salvia sp. and Sambucus caerulea Raf. The majority of the plants in the
latter list are probably casual records. However, I observed other Irbisia species
feeding on the flowers and young leaves of Lupinus species. Sambucus caerulea is a
verified oviposition site for several Irbisia species (E. J. Taylor, pers. comm.)

Specimens examined. 1,530 specimens were examined (Map 1). The following localities are
for the typical form: CANADA: ALBERTA: Conrad; Coutts; Elkwater Pk.; Irvine; Manyberries;
Medicine Hat; One Four; Pakouki L. - Manyberries; Walsh (all CNC). BRITISH COLUMBIA:
Oliver & Seymour Lk. (CNC, USNM). SASKATCHEWAN: Assiniboia; Great Sand Hills;
Lisieux; Rockglen; St. Victor; Val Marie; Wood Mt. (all CNC). UNITED STATES: ARIZONA:
Apache Co., Lukachukai Mts. (USNM). Coconino Co., Bright Angel Pt.— Grand Canyon Nat.
Pk. (USNM); Hualapai Indian Rsv. (USU); Oakridge (USU). CALIFORNIA: Alameda Co.,
Berkeley (UCB). Contra Costa Co., Byron (UCB); 4 mi. S Pittsburg (OSU). El Dorado Co.,
Byron (UCB); 4 mi. S Pittsburg (OSU). El Dorado Co., El Dorado (F&A); Pine Hill (UCD); 3
mi. S Shingle Spgs. (F&A). Fresno Co., 12 mi. W Coalinga (UCB). Humboldt Co., Dinsmores
(CAS). Kern Co., Antelope Cyn.— Tehachapi Mts. (UCR); 27 mi. SE & 4 mi. NE Bakersfield
(OSU & UCR); Cuyama (F&A); Edison (UCR); Lebec (UCB), Tejon Pass (UCB). Lassen Co.,
10 mi. S Doyle (OSU); Hallelujah Jet. (UCD). Los Angeles Co., Browns Flat (UCB); Claremont
(LACM, UCB); Covina (F&A); Desert Spgs. (UCB); Elizabeth Lk. Cyn. (LACM); Elys Park
(LACM); Glendale (LACM); Gorman (CAS); Little Harbor-Sta. Catalina Is. (UCB); Lwr. Shake
Cmpgd. — L.A. Nat. For. (OSU); Mint Cyn. (CAS); Palmdale (CAS, UCR); Pine Cyn. Rd.—
L.A. Nat. For. (OSU); Sta. Monica Mts. (LACM); Solemint (CNC); Westwood Hills (KU);
Whittier (LACM). Mendocino Co., Bell Spgs. (UCB); U.C. Hopland Exp. Sta. (CAS). Modoc
Co., 7 mi. NE Alturas (OSU); Buck Crk. Rgr. Stn. (UCB); 10 mi. S Canby (OSU); Rush Crk.
(UCB); 20 mi. NE Tulelake (AMNH). Monterey Co., 6 mi. W Greenfield (UCB); 1 mi. S
Jamesburg— Sta. Lucia Mts. (UCB). Orange Co., Arch Beach (CAS); 5 mi. W Irvine (UCD);
Newport (LACM); Salinas (USU); Orange (CAS); San Clemente (UCR); 10 mi. E San Juan
Capistrano (AMNH). Riverside Co., Alberhill (UCR); Bautista Cyn. (UCR); Corona (UCB); 2
mi. N Poppet Flat (UCR); Riverside (LACM, UCB, UCR, USNM). Sacramento Co., Folsom
(F&A). San Benito Co., Mercy Hot Spgs. (UCD); 2 mi. NE New Idria (UCB); 10 mi. S Pinnacles

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REVISION OF IRBISIA

231

Jet. (UCB); Pinnacles Nat. Mon. (OSU). San Bernardino Co., Cajon Pass (OSU); Colton (UCR);
Highland (UCB, UCR); 10 mi. N Fontana (UCB); 4 mi. E Mentone (AMNH); Ontario (USNM);
10 mi. SW Redlands (UCD); Sta. Ana River (UID); Scissors Crossing— Valle de San Felipe
(UCR); SE Sunshine Smt. mp. 7.7 rt. 79 (OSU); Palm Cyn. — Borrego Valley (LACM). San
Joaquin Co., Lone Tree Cyn. (F&A). S.L.O. Co., Cholane (CAS); Dune Lks. — 3 mi. S Oceano
(UCB); 2 mi. N Harmony (OSU); 10 mi. E Morro Bay (OSU); San Luis Obispo (OSU). Sta.
Barbara Co., Figueroa Mt. (LACM); Oso Cmpgd. (UCD). Sta. Clara Co., Los Gatos (F&A).
Shasta Co., 8 mi. W Harrison Gulch Rgr. Stn. (OSU); Whiskeytown (UCD). Siskiyou Co., 2
mi. SW Lava Beds Nat. Mon. HQ (OSU). Solano Co., 1 1 mi. S Dixon (UCD). Sutter Co.,
Marysville Buttes (CAS). Tehama Co., 21 mi. NW Red Bluff (USU). Tulare Co., Fairview
(UCB). Trinity Co., 9 mi. E Forest Glen (OSU); 6 mi. NE Hayfork (UCB); Indian Dick Rgr.
Stn. (UCB); Van Duzen Rd. (CAS). Ventura Co., Chuchupate Rgr. Stn.— Frazier Mt. (UCB);
Grade Valley— 9 mi. SW Stauffer (UCB); Pirn Crk.— Alamo Mt. (UCB); Wheller Spgs. (AMNH).
Yolo Co., Putah Cyn. (UCD). COLORADO: Albert Co., 0.5 mi. W Bijou Crk. (AMNH & JTP).
Archeluta Co., 17 mi. SE Pagosa Spgs. (TA&M). Boulder Co., Boulder & 5 mi. S (WSU, CNC,
USNM); Flagstaff Cyn. — Boulder (CNC); Sunset (CAS); Valmont Butte— Boulder (CNC). Doug-
las Co., Chaffield St. Pk. (JTP); 1 mi. S Parker (JTP). Eagle Co., State Bridge (CNC). El Paso
Co., Colorado Spgs. (UWY). Grand Co., Berthoud Pass (USNM). Gunnison Co., Monarch Pass
(USNM). Jefferson Co., Golden (USNM). La Plata Co., Durango (USU); 9 mi. E Mancos
(TA&M). Larimer Co., Ft. Collins & 9 mi. NW (KU, USNM & TA&M); S Estes Pk. (USNM).
Montezuma Co., Mesa Verde Nat. Pk. (AMNH). Pueblo Co., 12 mi. W Pueblo (JTP). Mis-
hawauka (KU). “Colo.” (KU, UN & USNM). IDAHO: Bannock Co., Pocatello (USNM). Camas
Co., 16.5 mi. N Fairfield (UID). Elmore Co., 31 mi. W Hill City (MSU); Mayfield (OSU).
Gooding Co., Bliss (USNM); Hagerman (USNM). Idaho Co., Grangeville (UID). Jerome Co.,
Jerome (UID); 4 & 10 mi. NE Hazelton (OSU & KU). Nez Perce Co., Lenore (UID). Oneida
Co., Black Pine Cyn., Holbrook; Meadow Brook Crk., Rock Crk., Salyer Cow Camp., Twin
Spgs. (all USU). IOWA: Woodbury Co., Sioux City (CNC). MONTANA: Cascade Co., Cascade
(MSU); Great Falls (USU); Vaughn (MSU). Richland Co., Sidney (USNM). Stillwater Co.,
Reedpoint (MSU). Teton Co., Shelby (TA&M). Yellowstone Co., Custer (MSU). NEBRASKA:
Dawes Co., 10 mi. E Chadron (OSU). Hooker Co. (USNM). Sheridan Co., Hay Spgs. (UN).
Sioux Co., War Bonnet Cyn. (UN). NEVADA: Carson City (CAS). Douglas Co., Holbrook
(UCD). Elko Co., Carlin (KU); 10 mi. N Wells (UCB). Eureka Co., Roberts Mts. (UCR). Lander
Co., Big Crk. Cyn. (USNM). Lyon Co., 7 mi. NW Wellington (F&A). Nye Co., nr. Shoshone
Pk., Nevada Atomic Test Site (AMNH). Washoe Co., Reno (F&A, UCD, USNM). NORTH
DAKOTA: Billings Co., 0.5 mi. S Medora (UCB), Theodore Roosevelt Rch. (NDS). Bowman
Co., (NDS). Dunn Co., 5 mi. W Killdeer (NDS). Golden Valley Co., Beach (USNM). Grant
Co., Caron (NDS). Mercer Co., (NDS). Morton Co., Mandan (NDS, SHF & USNM). Mountrail
Co., (NDS). Sioux Co., (NDS). Stark Co., 3 mi. NW Dickinson (UCB). OREGON: Deschutes
Co., Bend (KU); Deschutes R.— nr. Redmond (OSU); 2 & 3 mi. NE Sisters (OSU). Harney
Co., 12 mi. E Riley (CAS). Jefferson Co., 10 mi. N Redmond (OSU); 2.5 mi. N Warm Spgs.
(OSU). Klamath Co., 9 & 13 mi. W Beatty (OSU), 4 mi. E Bly (OSU), 30 mi. E Klamath Falls
(OSU). Malheur Co., Jordan Valley (OSU), 3 mi. S Vale (OSU). Marrow Co., 7.5 mi. N Hardman
(OSU). Wasco Co., 10 mi. N Warm Spgs. (OSDA). SOUTH DAKOTA: Jones Co., Capa (CAS
& SDSU). Pennington Co., Rapid City (USNM). Shannon Co., Pine Ridge (USNM). UTAH:
Box Elder Co., Blue Crk. (UCD); Cedar Crk., Cedar Hill, Curlew Jet.; Kelton; Kelton Pass;
Park Valley; 4 mi. W & 17 mi. SW Snowville (all USU). Cache Co., Beaver Dam; Blacksmith
Fork Cyn.; Green Cyn.; Logan Cyn.; Paradise (all USU); Wellsville Cyn. (OSU). Davis Co.,
Ogden (USU). Duchesne Co., Indian Cyn. (USNM). Garfield Co., Ruby’s Inn (USU). Grand
Co., Moab (USNM). Iron Co., Cedar City (USU); Iron Mite Site (USU); Kanarraville (USNM,
USU); Summit (UCD, USU). Juab Co., Levan (UCD); Mt. Nebo Loop (USU); Nephi (USNM).
Kane Co., Alton (SHF, USNM, USU); S Alton (SHF, TA&M); Glendale (USU); Glendale Mesa

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(USU); Johnson Cyn. (USU); 20 mi. N Kanab (BYU); Long Valley Jet. (SHF, USU); Mt. Carmel
(SHF, USNM). Millard Co., Cove Fort (USU); Fillmore (USNM, USU); Fool Crk. Pass (USU);
Meadow (UCD); Oak Crk. Cyn. (USU); Scipio (USNM). Morgan Co., Porterville (USU). Salt
Lake Co., Dry Cyn.— S.L.C. (USU); Granite (USU); Lake Pt. (USU); Salt Lk. City (USNM).
San Juan Co., Pack Crk. -La Sal Mts. (USU); Monticello (USNM, USU); 1 1 mi. SE Monticello
(TA&M). Sanpete Co., Flat Cyn. (USU); Fountain Green (USU). Sevier Co., Cove Mt. (USU);
Monroe Mt. (USU). Tooele Co., Johnson Pass (BYU, CAS). Utah Co., Blackett’s Pasture (USU);
Lehi (ASU); Payson (ASU); Pleasant Grove (USU); Provo (USU); Rock Cyn. (BYU); Spring
Lk. (USU). Washington Co., Grass Valley, New Harmony; Pine Valley (all USU). Copper
Mts.— Tecoma Rng. (BYU). WYOMING: Converse Co., Glenrock (UWY). Goshen Co., (UWY).
Laramie Co., High Plains Rsrh. Stn. — E Cheyenne (UWY). Niobrara Co., Lusk (UWY); Man-
ville (UWY). Platte Co., Wheatland (USNM). Weston Co., 1 mi. N Newcastle (OSU).

The following localities are for the small “ paeta^ form of I. brachycera. MEXICO: BAJA
CALIFORNIA NORTE: 3.0 mi. S El Condor (AMNH). UNITED STATES: CALIFORNIA:
Riverside Co., Banning Hts. (UCR); de Cuir Ranch— Beaumont (UCR, CAS); 8 mi. N Deep
Crk. & Horsethief Crk. (UCR); Dutch Flat— San Jacinto Mts. (CAS); Hemet Res.— San Jacinto
Mts. (CAS, UCB); Herkey Crk. -San Jacinto Mts. (UCB); Idyllwild (AMNH, CAS, UCB); Keen
Cmp.— San Jacinto Mts. (LACM, UCB, UCD); Pinon Flat— San Jacinto Mts. (CB). San Ber-
nardino Co., Running Spgs. — San Bernardino Mts. (UCB); Yucaipa (UCR, USNM). San Diego
Co., Anza-Borrego Desert St. Pk. (TA&M); Banner Grade (ASU); Couser Cyn. (UCR); Del Mar
(ASU); 3 mi. SE, & El Cajon (UCB & CAS); Green Valley Mdw.-Cuyamaca St. Pk. (UCB);
Julian (SDNM, UCR); Kitchen Crk.— Laguna Mts. (SDNM); Laguna Mts. (KU); Lakeside
(USNM); La Mesa (USNM); Miramar (UCB); Mission Gorge (ASU, UCB, UID); Monument
Pk.- Laguna Mts. (SDNM); Mt. Palomar (UCB); Pala (UCB, USNM); Pine Valley (CAS,
SDNM); Ramona (CAS); San Diego (CAS); 1.3 mi. S San Felipe (OSU); 9.3 mi. NW Sissors
Crossing (OSU); Warner Spgs. (SDNM); Viejas Crk. (UCB).

Irbisia calif ornica Van Duzee
Figs. 2, 5, 13a, b, 36, 59, 82; Map 1 1

Irbisia sericans: Childs, 1914, p. 220. Van Duzee, 1917b, p. 264.

Irbisia brachycerus: Essig, 1915, p. 213, fig. 188.

Irbisia californica Van Duzee, 1921a, pp. 146-147, 152 (new species). Essig, 1926,
p. 361. Blatchley, 1934, p. 13. Essig and Hoskins, 1944, pp. 71-72. Carvalho,
1959, p. 105.

Irbisia eurekae Bliven, 1961, pp. 45-46; 1963, p. 85, pi. 7, figs. 1, la. New Synonymy.
Irbisia paenulata Bliven, 1961, p. 46; 1963, p. 85, pi. 7, figs. 2, 2a. New Synonymy.
Irbisia umbratica Bliven, 1963, pp. 69-70, 85, pi. 7, figs. 7, 7a. New Synonymy.

Diagnosis. The material on hand adheres to the habitus of the holotype despite
the great range in the size of the specimens. The characters which make I. californica
distinctive are: dorsum densely covered with a mixture of long golden to white setae,
and sericeous white setae; relatively narrow anterior pronotal width; highly concave
lateral pronotal margins; straight posterior head margin; testaceous or fulvous col-
oration on legs with fuscous spots on the femora; robust, abruptly angled left paramere
of male with a distal triangular expansion and truncated apex; series of obvious,
sharply pointed spicules on the sclerotized process of the male vesica; open and oval
sclerotized rings, and relatively small posterior wall of female. The previous features

1984

REVISION OF IRBISIA

233

will adequately separate I. calif ornica from I. sericans and I. sita Van Duzee; all
three have superficially similar male genitalic structures.

Description. Macropterous male. Length 4.75-7.30, width 1.80-2.60, shining black
densely clothed with a mixture of long suberect golden to white setae, and sericeous
woolly white setae. HEAD: width across eyes 1.15-1.46, vertex 0.51-0.75, dorsal
width of eye 0.28-0.38; triangular, asperate; temporal areas indistinct but smooth;
median depression broad but shallow, or indistinct; basal carina straight, moderately
prominent, acute and abrupt or rounded and declivous to level of vertex; eyes broadly
joined to frons, but protruding laterally, posterior margin straight in dorsal view,
distance from tylus to ventral margin of eye 0.58-0.80, height of eye 0.45-0.56,
maximum interocular width 0.79-1 .06, base of clypeus convex in lateral view. ROS-
TRUM: length 1.73-2.48, fulvo-testaceous, apex and base sometimes black, reaching
meso- or metacoxae. ANTENNAE: black; I, length 0.58-0.99; II, sometimes fuscous
medially, 1.53-2.60; III, 1.05-1.60; IV, 1.03-1.68. PRONOTUM: length 0.88-1.25,
anterior width 0.80-1.09, maximum width 1.50-2.08, conical, broadly convex, ru-
gulose or confluently punctate, posterolateral margin arcuate, lateral margins dis-
tinctly concave, broadly rounded at junction with propleura, anterior angles slightly
produced but small and rounded in dorsal view, slightly or not sulcate in lateral view,
disk depressed immediately posterior of calli; calli moderately convex, asperate,
confluent anteriorly, reaching lateral margins of pronotum. LEGS: testaceous, fulvous
or fuscous; coxae black basally; femora with fuscous spots or variably sized patches;
tarsi black. VESTITURE: dorsum densely clothed with long golden to white setae,
erect on head and pronotum, suberect on hemelytra; intermixed with woolly, recurved
sericeous white setae; venter sparsely clothed with thin erect fulvous to white setae.

Macropterous female. More robust than male, but similar in structure, color and
vestiture. Length 5.15-7.25, width 2.13-3.00. HEAD: width across eyes 1.17-1.56,
vertex 0.55-0.8 1 , dorsal width of eye 0.28-0.38, distance from tylus to ventral margin
of eye 0.60-0.85, height of eye 0.48-0.56, maximum interocular width 0.82-1.10.
ROSTRUM: length 1 .83-2.38, reaching apices of mesocoxae. ANTENNAE: I, length
0.58-0.87; II, 1.55-2.41; III, 1.00-1.56; IV, 0.88-1.67. PRONOTUM: length 0.98-
1.35, anterior width 0.88-1.20, maximum width 1.73-2.33.

Types. Male holotype. Hills back of Oakland, Cal, IV- 12-08; Coll, by E. C. Van
Dyke; HOLOTYPE califomica, [red], CAS type no. 800. Retained at the California
Academy of Sciences, San Francisco. All the paratypes I examined (39 specimens)
are conspecific with the holotype and are retained at CAS and USNM.

Taxonomic consideration. A clear clinal series exists in this species. Specimens
from populations north of approximately 36° 30' north latitude are larger, and have
the dorsum densely covered with a mixture of long golden setae and woolly sericeous
setae. Specimens south of this line tend to be smaller, and possess shorter white
vestiture. The difference in stature is most noticeable in the head measurements, i.e.,
for male specimens from Coos County, Oregon to Monterey and San Benito Counties,
California the range of the width of the vertex is from 0.78 to 0.60 mm.; then to
Baja California Norte the range is from 0.65 to 0.51 mm. The smallest specimens
encountered in the material on hand occur south of the Transverse Ranges in Los
Angeles, Riverside, San Diego and San Bernardino Counties, California.

Prior to the erection of the species I. California, specimens of the species were

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placed in 7. sericans (Childs, 1914), and due to a nomenclatural error, specimens
were identified as I. brachycerus [a] (Essig, 1915). The preceding action was the result
of examining the specimens which precipitated the misidentifications. The specimens
of concern are retained in the California Academy of Sciences, San Francisco.

I have examined the types of 7. eurekae Bliven, I. paenulata Bliven and I. umbratica
Bliven and placed all in synonymy with 7. calif ornica. Both of the original descriptions
fit within the previous redescription of 7. californica in all respects: structure, col-
oration and vestiture.

Remarks. The distribution of this species is bounded on the south, in Baja Cali-
fornia Norte, by the Sonoran Desert; the southernmost record is San Vicente. Within
California, south of the San Francisco Bay Area, the distribution of 7. californica is
restricted to the California chaparral province; no localities penetrate the Coast or
Transverse Range. The northernmost portion of the distributional range is in Coos
County, Oregon and is continuous south to the Bay Area within the Pacific Forest
province. The vegetational associations encountered by the range of 7. californica
are: redwood, mixed evergreen, mixed hardwood, blue oak-digger pine and southern
oak forests, as well as, chaparral and coastal prairie shrub mosaic.

Adult and immature stages have been collected from these grasses: Anthoxanthum
aristatum Boiss., Avena fatua L., Avena sp., Bromus rigidus L., B. rubens L., Bromus
sp., Dactylis glomerata L., Elymus sp., Festuca arundinacea Schreb., Hordeum le-
porinum Link., Hordeum sp., and Poa sp. Adult specimens have been collected from
the following non-graminoid plants: Amsinckia intermedia F. & M., Artemisia cal-
ifornica Less., A. douglasiana Bess, in Hook., Astragalus nuttalli (T. & G.) J.T. Howell
var. virgatus (Gray) Bameby, Baccharis sp., Brassica sp., Cardaria sp., Ceanothus
arboreusG reene, C. crassifolius T orr. , C. integerrimus H. & A., Cer cocarpus ledifolius
Nutt., Citrus sp., Cryptantha intermedia (Gray) Greene, Encelia farinosa Gray, Lath-
yrus laetiflorus Greene ssp. alefeldii (White) Brads., Lotus scoparius (Nutt, in T. &
G.) Ottley, Lupinus albifrons Benth., L. polyphyllus Lindl., L. nanus Dougl. in Benth.,
L. succulentus Dougl. ex Koch., Malva parviflora L., Marah macrocarpus (Greene)
Greene, Medicago sativa L., Phacelia distans Benth., Prunus lyonii (Eastw.) Sarg.,
Quercus agrifolia Nee, Q. chrysolepis Liebm., Q. engelmannii Greene, Q. wislizenii
A. DC., Ranunculus californicus Benth., Ranunculus sp., Ribes indecorum Eastw.
Salix laevigata Bebb., Salix sp., Salvia apiana Jeps., S. mellifera Greene, Sambucus
caerulea Rafi, Scrophularia sp., Sisymbrium sp., Solanum douglasii Dunal in DC.,
S. xanti Gray and Yucca schidigera Roezl ex Ortgies.

The earliest record of occurrence is 2 1 February at Riverside and the latest is 27
July. The elevational range is from 180 to 2000 m. (600 to 3600 feet).

Specimens Examined. 2,295 specimens were examined from the following localities (Map
1 1). MEXICO: BAJA CALIFORNIA NORTE: 6.3 mi. S El Condor (AMNH); 7 mi. S Ma-
neadero (UCB)); San Vicente, & 10 mi. N (UCR & UCB). UNITED STATES: CALIFORNIA:
Alameda Co. (17 localities). Contra Costa Co. (24). Del Norte Co., mp. 14.39 on rt. 101 (OSU).
Humboldt Co. (7). Los Angeles Co. (42). Marin Co. (16). Mendocino Co., 1 mi. SE Piercy (UCB).
Monterey Co. (24). Napa Co. (6). Orange Co. (8). Riverside Co. (30). Sacramento Co., 1.5 mi.
S Fulsom (CAS). San Benito Co. (6). San Bernardino Co. (13). San Diego Co. (26). San Francisco
Co. (3). S.L.O. Co. (13). Sta. Mateo Co. (13). Sta. Barbara Co. (23). Sta. Clara Co. (17). Sta.
Cruz Co. (2). Solano Co. (3). Sonoma Co., Hacienda (CAS); Petaluma (UCB); Petaluma Marsh
(JTP); Sonoma Co. (AMNH, CAS). Ventura Co. (11). Yolo Co. (3). Channel Islands: East,

1984

REVISION OF IRBISIA

235

Middle & West Anacapa, San Miguel, San Nicolas, Santa Catalina & Santa Cruz. All specimens
are retained at AMNH, CAS, F&A, LACM, OSU, SBNH, SDNH, UCB, UCD, UCR & USNM.
OREGON: Coos Co., 0.4 mi. NE of mp. 9 on rt. 42S— nr. Riverton (OSU). Curry Co., Humbug
Mt. St. Pk. (CAS, OSU); T34S R14W Sec. 6 on Hwy. 101 (OSU).

Irbisia cascadia, new species
Figs. 8, 27a, b, 50, 73, 96; Map 23

Diagnosis. This species is very similar to I. fuscipubescens but is distinguished
from it by the mixed vestiture, shorter antennal segment I and genital structure of
both sexes. I. cascadia and I. solani are sympatric in the Willamette Valley of north-
western Oregon. Within this region specimens of the former species are wider between
the tylus and the ventral margin of the eye. The ratio of this measurement to the
height of the eye for both species is: for I. cascadia from 1:0.74 to 1:0.86 for males,
and from 1:0.70 to 1:0.78 for females; for I. solani the ratio ranging from 1:0.88 to
1:0.96 for males, and from 1:0.84 to 1:0.88 for females. The anterior angles of the
pronotum of I. cascadia are broadly prominent and evident in lateral view, whereas
this region of I. solani is rounded and indistinct. The femora of the female of the
latter species are extensively rufescent. The genital structures of both sexes are quite
different. The left paramere of the male of I. cascadia is strongly tuberculate; and
the sclerotized process of the vesica is expanded preapically and strongly spined. In
I. solani, the paramere is slightly tuberculate, and the process is not expanded and
has a linear series of small spicules on the inner margin. The dorsal labiate plate of
the female of I. cascadia is wide and broadly arcuate anteriorly near each sclerotized
ring.

I. cascadia is sympatric throughout its entire range with I. serrata. The former
species is distinguished by a smaller antennal segment I, forming a vertex to antennal
segment I ratio that ranging from 1:0.77 to 1:0.93 for males, and from 1:0.72 to 1:
0.87 for females. The range of this ratio for I. serrata is from 1:1.01 to 1:1.24 for
males and from 1:0.89 to 1:1.07 for females. Additionally, I. cascadia is separated
from I. serrata by the former species’: more shining dorsum, distinct punctation of
the pronotal disk, polished calli, and less dense vestiture. The vesica of I. cascadia
is obviously different from I. serrata ; the latter species does not have large, curved
spines on the sclerotized process, and has a membranous lobe attached medially to
the outer surface of the process. The female structures are very similar; only the
sclerotized rings are subtly different between the species.

Description. Macropterous male. Length 4.70-5.60, width 2.30-2.70, shining black,
clothed with recurved sericeous white to golden setae. HEAD: width across eyes
1.16-1.28, vertex 0.55-0.60, dorsal width of eye 0.28-0.31; triangular, mostly smooth;
temporal area shining; median depression faint or absent; basal carina broad, mod-
erately prominent, sometimes declivous or abrupt to level of vertex posteriorly ar-
cuate, with lateral portion anterior of posterior margin of head; eyes broadly joined
to frons, posterior margin straight in dorsal view, distance from tylus to ventral
margin of eye 0.58-0.66., height of eye 0.45-0.50, maximum interocular width 0.83-
0.93. ROSTRUM: length 1.60-1.85, fuscous to black, just reaching mesocoxae or,
in some specimens, metacoxae. ANTENNAE: black; I, length 0.48-0.53; II, 1.35-
1.60; III, 0.73-0.95; IV, 1.01-2.00. PRONOTUM: length 1.00-1.18, anterior width

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

0.94-1.04, maximum width 1.84-1.94; trapezoidal, disk broadly convex, distinctly
punctate, posterior margin broadly arcuate, lateral margins straight, rounded or broadly
margined at junction with propleura, anterior angles broadly prominent, sometimes
directed anteriorly, rounded or flattened, quite evident in lateral view; calli clearly
defined, convex, confluent anteriorly, polished, shining, with fovate impression on
anteromedial angles, sometimes reaching anterolateral margins of pronotum. LEGS:
black; tibiae in varying degree testaceous to fulvous medially. VESTITURE: head,
pronotum and scutellum with suberect to erect shining golden setae, pronotum usually
denuded, hemelytra with white or golden recurved or appressed sericeous and short
straight setae; venter with sparse, thin, fulvous setae. GENITALIA: Left paramere
(Fig. 27a): sensory lobe slightly projecting above arm; angle broad; length of shaft
less than length of arm; arm and shaft with prominent spines; arm diameter with
distal two thirds constricted and considerably more constricted subapically; apex
strongly securiform; with numerous bristles. Right paramere (Fig. 27b): diameter of
shaft expanded distally; apical process with double faceted spine; with a few short
lateral setae. Vesica (Fig. 50): two smooth membranous lobes each with a small
preapical side lobe; sclerotized process cone-shaped with a deep furrow and many
very large spines on the left surface.

Macropterous female. Slightly more robust than male, but similar in structure,
color and vestiture. Length 5.15-5.60, width 2.55-2.80. HEAD: width across eyes
1.23-1,33, vertex 0.58-0.68, dorsal width of eye 0.29-0.35, distance from tylus to
ventral margin of eye 0.57-0.71, height of eye 0 . 4 8-0 .53, maximum interocular width
0.90-0.98. ROSTRUM: length 1.73-1.85, attaining apex of mesocoxae. ANTEN-
NAE: I, length 0.46-0.55; II, 1.38-1.63; III, 0.78-1.00; IV, 0.95-1.25. PRONOTUM:
length 1 .08-1 .20, anterior width 0.98-1 .05, maximum width 1 .95-2.08. LEGS: apices
of femora fuscous. GENITALIA: sclerotized rings (Fig. 73): area within rings broadly
oval; dorsal labiate plate very broad directly anterior and anterolateral of rings;
strongly constricted medially; width of one ring 0.25-0.30, maximum width across
rings 0.88-0.98. Posterior wall (Fig. 96): A-structure curving, ventral and dorsal
margins flattened. B-structure protruding posteriorly, without indentations on ventral
posterior surface; median process gradually expanded dorsad, maximum length 0.20-
0.29, maximum width 0.70-0.75.

Holotype. Male. Oregon, Benton Co., Findley National Wildife Refuge— Bald Hill,
9 May 1979, M. D. Schwartz, ex Bromus rigidus & Festuca arundinacea\ CAS type
no. 15183. Retained at the California Academy of Sciences, San Francisco. Paratypes
same data, AMNH, CAS, CNC, OSU, USNM (17<$<$, 1 199).

Etymology. Named for the physiographic region in which this species is abundant.

Remarks. The main distribution of this species is in Jackson and Josephine Coun-
ties of southwestern Oregon. The vegetational associations which predominate in
this area are the Oregon oakwoods ( Quercus garryana ) and mixed conifer forest
{Abies- Pinus-Pseudotsuga). The range of I. cascadia extends north through the hill-
sides of the Willamette Valley, and to southern Skamania County, Washington. The
southernmost limits of its distribution are the Warner Mountains of Modoc County,
California. The vegetation associations of this southern area are marked by the
intergradation of Sierran montane forest (Abies-Pinus) with sagebrush steppe (Agro-
pyron- Artemisia). Adult and immature specimens have been collected from the fol-
lowing grass hosts: Avena barbata Brot., Bromus mollis L., B. rigidus Roth, Elymus

1984

REVISION OF IRBISIA

237

glaucus Buckl., and Festuca arundinacea Schreb. (alta fescue). Other plants on which
adult specimens were collected are Abies procera Rehd., Artemisia cana Pursh., A.
tridentata Nutt., Balsamorhiza sagittata (Pursh) Nutt., Chrysothamnus nauseosus
(Pall.) Britton and Symphoricarpus sp. This is a moderate to high altitude species
with collected specimens ranging from 760 to 1860 m. (500 to 6100 feet) elevation.
High altitude record is from Fandango Pass, Modoc County, California. Dates of
occurrence are 29 April to 10 August.

Specimens examined. 827 additional specimens were examined from the following localities
(all OSU unless otherwise noted, Map 23). UNITED STATES: CALIFORNIA: Modoc Co.,
Ceder Pass, 7 mi. W Cederville, 29 June 1955, D. L. Dahlsten, UCD (3); E Fandango Pass,
6000 ft., 22 June 1971, J. Sawbridge (5, 525); 20 mi. NE Tulelake, 5 June 1972, ex Balsamorhiza
sp., J. Schuh, AMNH (3). OREGON: Benton Co., all Corvallis, 13 June 1899 (3); 27 May 1900
(3); 6 mi. SW, 30 May 1973, L. Russell (2); Animal Science Pasture, 0.6 mi. N Harrison on
53th rd., 12 May 1979 (43 33, 2422) and 29 May 1979 (533, 722), M. D. Schwartz, ex Bromus
mollis & B. rigidus-, 0.4 mi. N on Glen Ridge Drive, 29 April 1979, M. D. Schwartz, ex Festuca
arundinacea (833, 322); intersection of Harrison & 53th rd., 1 May 1979, M. D. Schwartz, ex
Festuca arundinacea, reared from nymph (2); “IV Hill”, 8 May 1980, J. D. Lattin (322); McDonald
Forest, 20 May 1979, M. D. Schwartz (3) and Misty Point (Dimple Hill), 19 May 1979, M. D.
Schwartz, ex Avena barbata, Bromus mollis, B. rigidus & Elymus glaucus (1936, 522); Vineyard
Mt., 0.6 mi. N Lewisburg Ave., 30 April 1979, M. D. Schwartz, ex Bromus rigidus (3 33, 222)
and 1 1 May 1979 (35 33, 1722). All Mary’s Peak (14 mi. W Corvallis), R7W T12S Sec. 19 SE
Va , 3250 ft., 3 July 1979, G. M. Cooper, ex Abies procera (2); Mary’s Peak Road, 7.8 mi. N rt.
34, 17 May 1979, M. D. Schwartz, ex Festuca rubra, reared from nymphs (1 1 specimens) and
1 June 1979, K. West & G. M. Stonedahl (3333, 2122); meadow, 1 mi. below campground, on
shoulder, 3350 ft., (No. 71-3), 1 July 1971, J. D. Lattin (733, 722); north spur at top, 3500 ft.,
10 August 1971, J. D. Lattin (322); summit, 3700 ft., 1 July 1971, (No. 71-4) (3, 722) and 10
August 1971, (No. 71-2) (3, 2), J. D. Lattin, ex sweeping meadow, picnic area, 4000 ft., 13 July
1956, J. D. Lattin (1133, 522); 4000 ft., 9 June 1957, B. Malkin (433, 522), 27 June 1962, R.
W. Matthews (1033, 822) and 30 July 1966, W. Gagne & J. Haddock, UCB (34 33, 1422); 3800
ft., 24 June 1970 (2), 15 July 1969, P. Oman (233, 222) and S. Rose & J. McClay (1333, 322);
3000 ft., 18 July 1968, P. Oman (2); 6 June 1951, V. Roth (233, 2); 24 June 1962, D. & L. May
(733, 822); 30 June 1896, No. 190 (3 33); 4 July 1934, J. Schuh (2); 6 July 1956, F. F. Hasbrouck
(233); 12 July 1956, J. Capizzi, OSDA (233); 12 July 1958, J. D. Lattin (1933); 13 July 1941
(2); 15 July 1969, P. Oman (2); 25 July 1959, S. Radinovsky (1733, 1822); 9 August 1955, R.

F. Koontz (233); 10 August 1956, J. D. Lattin (3, 322). Douglas Co., Myrtle Creek, 30 May
1961, L. G. Gentner, ex sweeping (633, 622); 3 mi. N Roseburg, roadside, 18 May 1960, F. A.
Michel (3, 2), J. D. Lattin (333, 422) and D. R. Smith (2); 2 mi. N Tiller, 16 May 1962 (633,
422); Umpqua River— 2 mi. W Tiller, 1000 ft., 22 May 1960, J. D. Lattin (2). Jackson Co., 23
mi. NE Ashland, 1 June 1972, P. Oman (533, 822); Condie Creek, 26 June 1974, SOS (2); Dead
Indian Soda Springs, 26 June 1974, R. G. Lynch, SOS (2); near Dead Indian Soda Spring, 23
June 1978, J. Schuh, ex Symphoricarpus sp., AMNH (433, 722); 29 mi. ESE Eagle Point, 4450
ft., 23 June 1978, N. L. Herman, AMNH (3 33, 822); Medford, (S.S. 25357), 2 May 1945, ex
clover (45-9842), USNM (2); N Medford, 1 May 1970, P. Oman (233, 422); all Mt. Ashland,
4600 ft., 24 June 1971, G. Steyskal, USNM (233, 2); 20 June 1975, L. Russell (2); 4600 ft., 17
June 1970, P. Oman (3 33, 922); 4100 ft., 27 May 1970, P. Oman, ex Chrysothamnus nauseosus
(3, 222); 5 mi. W Pinehurst, 24 May 1958, B. Ainscough (233 322) and 4526 ft., J. D. Lattin
(433, 622); Rouge River, 16 May 1952, A. B. Black, OSDA (2) and (no. 124), 13 May 1962, F.

G. Bowes, SOS (2); all Sam’s Valley, 18 May 1968, P. Oman, ex Arctostaphylos sp. (2); 1 mi.
SE, 10 May 1969, M. Stock (222); 3 mi. E, 18 May 1962 (3, 222). Shady Grove, 15 May 1972,
J. Schuh, AMNH (2); Siskiyou, 14 June 1959, Kelton & Madge, CNC (2833, 2822); Siskiyou

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Summit, 3 June 1975, B. Summey, SOS (9) and 24 June 1974, K. W. Beatty, C. Oakley, M.
Swisher & N. Turner, SOS (599); 0.5 mi. S Siskiyou Summit on old rt. 99, 1350 m., 26 June
1979, R. T. & J. Schuh, AMNH (599); 4300 ft., 27 June 1979, M. D. Schwartz, ex Artemisia
cana (4699). Josephine Co., Grants Pass, 8 May 1963, D. B. Goroon, SOS (6); Grave Creek, 30
May 1952, B. Malkin & VER., CAS (266, 299); all Merlin, 1 May 1970, P. Oman (1556, 1099)
and F. A. Rushmore, ex Alta fescue (266); 9 May 1969, P. Oman (1356, 1 199); 13 May 1972,
C. Musgrave, ex grasses (299) and 12 May 1972, J. Sawbridge (299); 27 May 1970, P. Oman
(5); 2 mi. E Merlin, 1480 ft., 13 May 1972, J. D. Lattin (399). Lake Co., Cottonwood Reservoir,
T38S R13E Sec. 18, 20 June 1980, P. Oman (365, 899). Linn Co., Monument Park, 8 mi. ESE
Gates Summit, 4725 ft., 16 June 1960, J. D. Lattin (6, 299); Peterson Butte, 2.7 mi. rt. 34 on
Steckley Rd., 1 May 1979, M. D. Schwartz, ex Bromus mollis & B. rigidus (766, 1799) and south
slope, 16 June 1977, G. Eulensen (6). Linn/Lane Co., H. J. Andrews Experimental Forest,
Carpenter Mtn., access rd. 1501, 6.6 mi. N jet. FS rds. 1502-1501, 4800 ft., 6 July 1977,
Eulensen & Searles, ex sweeping (6, 299). Yamhill Co., McMinnville, Peavine Ridge, 7 July
1952, J. F. Bock, OSDA (9). WASHINGTON: Skamania Co., Dog Mtn. on Pacific Crest Trail,
250-500 ft., 18 May 1979, M. D. Schwartz, ex Balsamorhiza sagittata & grasses (466, 699).

Irbisia castanipes Van Duzee
Figs. 24a, b, 47, 70, 93; Map 18

Irbisia castanipes Van Duzee, 1921a, pp. 145-146, 151 (new species). Carvalho 1959,

p. 105.

Diagnosis. I. castanipes may be separated from all other species of Irbisia by the
dull black color with bronze luster, and sparsely distributed short, appressed, light
setae on the dorsal surface.

Description. Macropterous male. Length 5.40-6.10, width 2.10-2.60, dull black
with bronze luster, sparsely clothed with minute appressed, light colored setae. HEAD:
width across eyes 1.23-1.30, vertex 0.53-0.60, dorsal width of eye 0.33-0.38; tri-
angular, shagreened, temporal area asperate, basal carina sharp in cross section,
declivous to level of vertex, posteriorly arcuate mesad; vertex flattened, median
depression very shallow; eyes large broadly joined to frons, posterolateral margin
arcuate caudad in dorsal view; distance from tylus to ventral margin of eye 0.65-
0.68, height of eye 0.53-0.58, maximum interocular width 0.85-0.95. ROSTRUM:
length 2.03-2.35, black, reaching or surpassing apices of metacoxae. ANTENNA:
black; I, length 0.65-0.68, base and apex sometimes fuscous; II, 1.80-2.04; III, 1.05-
1 . 1 0; IV, 1 . 1 0-1 . 1 5. PRONOTUM: length 1 .03-1 .20; anterior width 0.84-0.96; max-
imum width 1.74-2.03; trapezoidal, with sparse long bristles anterolaterally; disk
broadly convex, rugose or transversely rugose and/or punctate, posterior and lateral
margins straight, lateral margin rounded at junction with propleura, anterior angles
rounded and indistinct in dorsal and lateral view; calli inconspicuous, shagreened,
confluent anteriorly, reaching anterolateral margin of pronotum. LEGS: castaneous
to fuscous; apices of femora and tibiae lighter; apices of coxae and base of trochanters
light. VENTER: polished black. VESTITURE: dorsum clothed with minute, sparse,
appressed light setae; setae longest on head; venter with sparse, long, upright setae.

Macropterous female. More robust than male, but structure, color and vestiture
are similar. Length 5.00-6.30, width 2.25-2.50. HEAD: width across eyes 1.33-1.40,
vertex 0.58-0.63, dorsal width of eye 0.36-0.40, distance from tylus to ventral margin
of eye 0.70-0.75, height of eye 0.56-0.60, maximum interocular width 0.95-1.00.

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239

ROSTRUM: length 2.33-2.55, surpassing apices of metacoxae. ANTENNA: I, length
0.68-0.78; II, 1.80-2.01; III, 1.08-1.18; IV, 0.85-1.11. PRONOTUM: 1.03-1.23,
anterior width 0.95-0.98, maximum width 2.00-2.10.

Types. Male holotype. Keen Camp Cal., Riverside Co., June 6-12, 1917; Pine; EP
Van Duzee Collector; HOLOTYPE castanipes [red]; CAS type no. 798. I have ex-
amined the holotype and all paratypes; all are conspecific and retained at the Cali-
fornia Academy of Sciences, San Francisco or the Canadian National Collection,
Ottawa.

Remarks. This species appears to be the southern counterpart to I. bliveni. Both
species occur in pine associations of the Sierran forest province. I. castanipes is also
found at higher elevations in the Laguna, San Jacinto and Tehachpi Mts. within the
California chaparral province. Adult specimens have been collected on Pinus jejfeyi
Grev. & Balf. in A. Murr. and P. ponderosa Dougl. ex P. & C. The localities range
from 1 372 to 2225 m. (4500 to 7300 feet) elevation. Collection dates for adult species
are 18 May to 10 July.

Specimens examined. 46 specimens were examined from the following localities (Map 18).
UNITED STATES: CALIFORNIA: Fresno Co., Florence Lk. (UCB); Huntington Lk. (CAS).
Kern Co., Antelope Cyn.— Tehachapi Mts. (UCR); Mt. Pinos (UCD). Riverside Co., Idyllwild
(UCB, UCD); Keen Cmp. — San Jacinto Mts. (CAS); Poppet Flat (UCB); Tahquitz Valley— San
Jacinto Mts. (UCB). San Diego Co., Green Valley Mdw.— Cuyamaca St. Pk. (UCB); Mt. Laguna
(UCB). Tulare Co., Kennedy Mdw. (UCD). Tuolomne Co., Kennedy Mdw. (UCD).

Irbisia cuneomaculata Blatchley
Figs. 15a, b, 38, 61, 84; Map 13

Irbisia cuneo-maculata Blatchley, 1934, p. 13 (new species). Carvalho, 1959, p. 105
(misspelled cuneomaculata )„

Diagnosis. This species is readily identified by the presence of pale markings on
the base of the cuneus of both sexes.

Description. Macropterous male. Length 3.80-450, width 1.60-2.00, shining black
densely covered with a mixture of recurved, appressed, sericeous white setae and
long erect fuscous setae. HEAD: width across eyes 0.92-1 .09, vertex 0.43-0.50, dorsal
width of eye 0.25-0.28; triangular, asperate; tylus rounded in dorsal view; temporal
areas distinct, smooth; median depression absent; basal carina prominent, rounded
or acute in cross section declivous to level of vertex posteriorly arcuate, eyes broadly
joined to frons, posteromedial margin posteriorly arcuate; distance from tylus to
ventral margin of eye 0.45-0.50, height of eye 0.38-0.45; maximum interocular width
0.65-0.78; clypeus and juga entirely, to not at all, fulvous or fuscous ventrad of
antennal insertions. ROSTRUM: length 1.28-1.45, fuscous to black, base and apex
usually darkly infuscated, reaching bases and sometimes slightly surpassing apices
of mesocoxae. ANTENNAE: fuscous; I, length 0.31-0.41, black, apex and, in some
specimens, apical half fuscous or fulvous; II, 0.90-1.13, apex and base sometimes
black, extreme apex sometimes fulvous; III, 0.53-0.70; IV, 0.53-0.71; III and IV
usually darkly fuscous or black. PRONOTUM: length 0.75-0.88, anterior width 0.75-
0.88, maximum width 1.43-1.68; conical, broadly convex, or slightly flattened con-
fluently punctate or rugose, posterolateral margins arcuate, lateral margins concave

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to strongly concave, slightly margined at junction with propleura, anterior angles
produced or slightly rounded but still evident, in dorsal view, sulcate in lateral view;
calli barely convex, but distinct, asperate, narrowly confluent anteriorly, not reaching
anterolateral margins of pronotum; ventral third of propleura, prostemum narrowly
bordering coxae and possibly attaining front margins, basalar plate, median portion
of epistemum, epimeron bordering coxae, and dorsolateral portion of second and
third stemites of abdomen testaceous or ivory. LEGS: fulvous, fuscous or black;
apices and bases of coxae testaceous or ivory; femora with black or dark fuscous;
apices and bases sometimes darkly infuscated; apices of tibia and tarsi black. HE-
MELYTRA: inner basal region of cuneus, in variable degree, testaceous; some spec-
imens with black cuneus. VESTITURE: dorsum densely clothed with a mixture of
recurved sericeous white setae and long fuscous setae; sericeous setae are woolly on
head and pronotum, appressed on hemelytra; fuscous setae erect on head, pronotum
and basal portion of hemelytra, suberect on remainder of hemelytra; venter and legs
moderately covered with long testaceous setae.

Macropterous female. More robust than male, with more extensive pale coloration
and somewhat oval, otherwise similar in structure, and vestiture. Length 3.80-5.23,
width 1.60-2.00. HEAD: width across eyes 0.98-1.19, vertex 0.48-0.59, dorsal width
of eye 0.23-0.29, distance from tylus to ventral margin of eye 0.48-0.58, height of
eye 0.38-0.48, maximum interocular width 0.70-0.86; some specimens with entire
ventral portion of head, below antennal insertions, excluding buccula and variable
portion of gena, testaceous or fulvous. ROSTRUM: length 1.31-1.56, reaching bases
and sometimes slightly surpassing apices of mesoxocae. ANTENNAE: I, length 0.33-
0.46, sometimes entirely testaceous; II, 0.80-1.16 sometimes extensively testaceous
or fulvous; III, 0.59-0.70; IV, 0.60-0.80. PRONOTUM: length 0.70-1.04, anterior
width 0.76-1.00, maximum width 1.45-2.00. VENTER: testaceous, or fulvous, both
sides with medial black line of variable thickness, vaginal exterior sometimes black.

Type. Female holotype. Los Angeles, Cal., W.S.B., 2-26-28; Purdue Blatchley
Collection; TYPE [red]. No other specimens were collected by Blatchley. The holotype
is deposited at the Purdue Entomological Research Collection, Purdue University,
West Lafayette, Indiana. Blatchley (1934) states in the original description that the
holotype was collected on a white flowered shrub near Sunland [Los Angeles]. The
material on hand is within the species limits of Blatchley’s description. He does state,
however, that the holotype is brachypterous. All the specimens of both sexes ex-
amined are clearly macropterous. Mr. Arwin V. Provonsha, curator at the Purdue
collection, has examined the holotype and verified that it is macropterous and female.

Remarks. The distribution of this species is entirely within the California chaparral
and Sierran forest provinces. The floristic associations occupied are: mixed hard-
woods, blue oak-digger pine, and southern oak forests as well as chaparral. Adult
specimens have been collected on these grasses: Agropyron sp., Bromus sp., Elymus
sp. and Hordeum vulgare L. Other non-graminoids plants on which adult specimens
have been collected are: Arctostaphylos sp., Artemisia sp., Ceanothus cuneatus (Hook.)
Nutt., C. intergerrimus H. & A., C. megacarpus Nutt., Cercocarpus ledifloius Nutt.,
Keckiella sp., Quercus sp., Rhamnus crocea Nutt, in T. & G., Salvia apiana Jeps.
and S. mellifera Greene. The range of occurrence is 1 5 February (San Diego County)
to 7 May, and altitudinal range is from 560 to 980 m. (1850 to 3200 feet).

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241

Specimens examined. 92 specimens were examined from the following localities (Map 13).
UNITED STATES: CALIFORNIA: Alameda Co., Cedar Mtn. (UCB). Butte Co., Oroville
(CAS). Fresno Co., Jacalitos Cyn.— Coalinga (CAS). Kern Co., Lk. Isabella— main dam cmpgd.
(OSU). Los Angeles Co., Claremont (USNM); Los Angeles (USNM); Nicholas Beach (LACM);
Santiago Canal & Sta. Ana (USNM). Monterey Co., 5 mi. N Escondido Cmpgd. — Sta. Lucia
Mts. (UCB); Paraiso Spgs. (CAS); Stone Cyn.— W Jolon (CAS). Orange Co., Trabuco Cyn.
(F&A). Riverside Co., Bautista Cyn. (T6S R2E Sec. 18) (UCR); Keen Cmpgd. — San Jacinto
Mts. (UCB); Menifee Valley (UCR); 1 mi. SE Radec (UCB); Railroad Cyn.— 4 mi. E Elsinore
(UCB); Riverside (UCR); Sage (UCB); 5.5 mi. & 5.6 mi. S Sage (T7S R1E Sec. 32) (UCR);
Temecula (F&A); mp. 21.29 on rt. 243— S Banning (OSU). San Benito Co., Panoche Pass
(UCB); Pinnacles (UCB). San Diego Co., Cuyanaca Lk. (UCB); Descanso Rgr. Stn., & 4 mi.
NW (UCB); Jacumba (SDNM); Me Cain Valley (SDNM); 2 mi. S Pine Valley (UCB); Pala
(USNM); mp. 7.7 on rt. 79 — SE Sunshine Summit (OSU). S.L.O. Co., Temblor Range— 12 mi.
E Simmler (UCB). Sta. Barbara Co., Los Prietos (UCB). Sta. Clara Co., Colorado Crk. & Mines
Rd. (UCB); S end Mines Rd. (CAS). Stanislaus Co., W Adobe Crk. (UCB).

Irbisia elongata Knight
Figs. 10a, b, 106, 33, 56, 79; Map 2

Irbisia elongata Knight, 1941, pp. 77-78 (new species); 1968, p. 185. Carvalho, 1959,

p. 105. Kelton, 1980, p. 85, map 24, 86.

Irbisia retrusa Bliven, 1963, pp. 71-72, 85 pi. 7, figs. 9, 9a. New Synonymy.

Diagnosis. Individuals of this species may be distinguished from most of the other
species of Irbisia by the strongly pedunculate eyes, antennal sockets below the ventral
margin of eye and long and narrow body form. I. elongata differs from I. pacifica
and I. brachycera in antennal structure. The length of segment II is subequal to the
width of the pronotum (much greater in I. pacifica ), and the vertex to segment I ratio
is from 1:0.96 to 1:1.04 (from 1:0.64 to 1:0.85 in I. brachycera ). The ratios of the
distance between the tylus and the ventral margin of the eye to the height of the eye
also differ from I. brachycera. The ratio for I. brachycera is greater than 1 :0.50; ratio
for I. elongata is equal to or less than 1:0.50.

Description. Macropterous male. Length 5.20-5.90, width 1.70-2.00, black, shin-
ing, predominately clothed with thin, white, woolly sericeous setae. HEAD: width
across eyes 1.18-1.31, vertex 0.68-0.75; dorsal width of eye 0.27-0.29; asperate;
temporal areas smooth and glabrous; basal carina indistinct, posteriorly arcuate mes-
ad; median depression broad and shallow; eyes small, roundish, pedunculate, pos-
terolateral margin slightly arcuate anteriorly in dorsal view; distance from tylus to
ventral margin of eye 0.76-0.80, height of eye 0.39-0.40, maximum interocular width
0.88-0.90, lora with pale spot at base; antennal sockets below ventral margins of
eyes. ROSTRUM: length 1.98-2.13, testaceous, with dark infuscations (especially
last two segments), surpassing mesocoxae, sometimes reaching apices of metacoxae.
ANTENNAE: fuscous; I, length 0.73-0.80; short light setae, black, apical half some-
times light; II, 1.71-1.94; III, 1.05-1.20; IV, 0.90-1.08. PRONOTUM: length 1.08-
1.25; anterior width 0.85-0.96; maximum width 1.58-1.75; somewhat conical, disk
convex, rugulopunctate, posterior margin straight, lateral margins slightly concave,
very broadly rounded at junction with propleura, anterior angles blunt, inconspic-
uous, sulcate dorsad of coxal cleft, in lateral view; calli barely produced, confluent

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anteriorly, with short transverse depression anteriorly, asperate. LEGS: whitish tes-
taceous to flavous with black markings; procoxae black basally, meso- and metacoxae
black in middle; femora black basally, terminating in a series of fuscous or black
spots; tibia black on base and apex; tarsi black. VESTITURE: dorsum clothed with
erect, long, white, shining setae, intermixed with sericeous, recurved setae; setae on
head antrorse, woolly on anterior portion of pronotum, somewhat appressed on
remainder of dorsum; venter with dense, shining white setae.

Description. Macropterous female. Slightly more robust than male, but similar in
struture and vestiture; coxae and xyphus with more extensive white coloration. Length
5.50-6.25, width 2.08-2.40. HEAD: width across eyes 1.25-1.38, vertex 0.70-0.82,
dorsal width of eye 0.28-0.30, distance from tylus to ventral margin of eye 0.80-
0.88, height of eye 0.38-0.41, maximum interocular width 0.91-1.00. ROSTRUM:
length 2.06-2.25, barely surpassing mesocoxae. ANTENNA: I, length 0.75-0.80; II,
1.98-2.18; III, 1.13-1.30; IV, 1.03-1.92 (a population from Grant Co., Oregon con-
tains several specimens with smaller lengths; I, 0.68; II, 1.66; III, 1.08; IV, 0.95).
PRONOTUM: length 1.06-1.25, anterior width 0.95-1.08, maximum width 1.60-
1.92.

Types. Male holotype. Sundance, Crook Co. Wyoming, 30 July 1927, H. H. Knight.
Retained at the United States National Museum, Washington, D.C. I have examined
the holotype and paratypes (22 specimens); all are conspecific and are deposited at
BYU, CNC, JCS, and USNM.

Taxonomic consideration. The junior synonym, I. retrusa Bliven, was described
from a single specimen. I have examined this male and as mentioned in the original
description the apical two thirds af the first antennal segment is orange. This antennal
condition may be observed on several specimens from Alberta and Montana. I.
retrusa is synonymized with I. elongata on the basis of their similar external and
internal morphology.

Remarks. This species has a northern and montane distribution extending from
approximately 41° to 52° 30' north latitude between 1219 and 2438 m. (4000 and
8000 feet) elevation. In British Columbia, specimens were collected from the interior
plateau and Fraser basin within the northern prolongation of the Palouse Prairie.
Grassland and sagebrush steppe vegetation, which border woodland situations, char-
acterize the floristic associations of the rest of the range of I. elongata. I have collected
adult specimens from two grass hosts: Agropyron desertorum (Fisch.) Schult. and
Stipa thurberiana Piper. Some specimens from Alberta were collected on A. inter-
medium (Host) Beauv. Collection dates are from 12 May to 3 August.

Specimens examined. 234 specimens were examined from the following localities (Map 2).
CANADA: ALBERTA: Conrad; Cowley; Frank; Lethbridge (USNM); Spring Pt.; Waterton Nat.
Pk. (all CNC). BRITISH COLUMBIA: Chase (UBC); Chilcotin (UCB, USNM); Kamloops
(UBC); Lac du Bois- Kamloops (UBC); nr. Lytton (UBC); Pavilion Lk. (CNC); Soda Crk.
(UBC); Vaseaux Lk. & White Lk.- Oliver (CNC, USNM). UNITED STATES: CALIFORNIA:
Humboldt Co., Dinsmores (CAS). Modoc Co., Rush Crk. — 9 mi. NE Adin (UCB). COLORADO:
Eagle Co., Muddy Pass Rd. (JTP). IDAHO: Bonner Co., 4 mi. S Elmira (UID); Big Smokey
Grd. Stn. (UID). Kootenai Co., Farragut St. Pk. (UID). Lemhi Co., 14.4 mi. N Cobalt (UID).
Oneida Co., Black Pine Cyn. (USU). MONTANA: Broadwater Co., Winston (MSU). Fergus
Co., mi. SW & 6 mi. N Lewiston (MSU); Moulton (MSU). Gallatin Co., Bozeman (MSU);
Springhill (MSU); Willow Crk. (USNM). Glacier Co., 7 mi. NW Kiowa (CAS). Judith Basin

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Co., SW Utica (MSU). Latah Co., Genesee (USNM). Ravalli Co., Florence (USNM). Teton
Co., 25 mi. W Chateau (MSU). Wheatland Co., 8 mi. N Harlowton (MSU). NEVADA: Elko
Co., E slope Spruce Mt. (UCB). OREGON: Baker Co., 30 mi. NE Baker (OSU); Haight Rch.—
Halfway (OSU); 6.5 mi. E Richland (OSU). Crook Co., 5.6 mi. NE rt. 25— Ochoco Crk. (OSU);
20 mi. E Prineville (OSU); Ochoco Nat. For. (T14S R18E Sec. 1 1) (OSU). Grant Co., 13 mi.
S Dale (OSU); 1 1 mi. N Seneca (OSU). Wallowa Co., Lone Spg.-32 mi. N Joseph (OSU).
Wheeler Co., 4.5 mi. S Mitchell (AMNH, OSU). UTAH: Box Elder Co., Both well (USU). Cache
Co., Blacksmith Fork (OSU); East Cyn. (USU); Green Cyn. (USU); Logan Cyn. (USU); Ricks’
Spg. — Logan Cyn. (USU); Smithfield Cyn. (AMNH). Rich Co., Bear Lk. lookout— Smt. Logan
Cyn. (USU). WASHINGTON: Adams Co., 13 mi. S Ritzville (OSU). Grant Co., 7 mi. NE
Coulee City (OSU). Whitman Co., Colton (WSU). WYOMING: Park Co., Lk. Crk. Cmpgd.-
13 mi. S Cooke City, Montana (AMNH). Sheridan Co., Sheridan (SHF, UWY).

Irbisia fuscipubescens Knight
Figs. 28a, b, 51, 74, 97; Map 22

Irbisia fuscipubescens Knight, 1941, pp. 16-11 (new species). Carvalho, 1959, p. 105.

Kelton, 1980, p. 84, map 23.

Diagnosis. I. fuscipubescens is distinguished from other species of Irbisia by the
sparsely distributed fine, short, erect or appressed fuscous to golden setae on the
polished, shining dorsum. Irbisia cascadia also has prostrate, sericeous white setae,
but positive separation of it and I. fuscipubescens requires the careful examination
of genitalic structures. The sclerotized rings of female I. fuscipubescens are strongly
compressed dorsoventrally, and not completely open interiorly; I. cascadia has oval,
open sclerotized rings. The outer edge of the concave ventral surface of the sclerotized
process of the male of the former species does not have a row of strong spicules; the
latter species has a clearly defined row of strong spicules on the outer edge that are
separated from the major strongly curved spines of the inner edge of the frontal
concavity. The left paramere of the male of these two species is diagnostic; I. fus-
cipubescens has tubercles which are confined to the shaft.

Description. Macropterous male. Length 5.20-6.10, width 2.20-2.80, polished,
shining black, sparsely covered with short, thin fuscous setae. HEAD: width across
eyes 1.13-1.30, vertex 0.53-0.60, dorsal width of eye 0.28-0.35; triangular, polished;
temporal areas indistinct; median depression usually absent or a single faint impressed
line; basal carina abruptly produced, bluntly prominent, straight or slightly posteriorly
arcuate mesad; eyes broadly joined to frons, posterior margin straight in dorsal view,
distance from tylus to ventral margin of eye 0.58-0.66, height of eye 0.48-0.58,
maximum interocular distance 0.83-1.08. ROSTRUM: length 1.65-2.00, black or
fuscous, apical half sometimes fulvous, reaching apex of mesocoxae, sometimes
approaching and attain apex of metacoxae. ANTENNAE: black; I, length 0.53-0.70,
apical third to quarter sometimes fulvous; II, 1.48-2.00; III, 0.90-1.10; IV, 1.03-
1.25. PRONOTUM: length 1.03-1.15, anterior width 0.85-1.08, maximum width
1.73-2.01; trapezoidal, broadly convex, rugulopunctate or distinctly punctate, pos-
terior margin broadly arcuate, lateral margins straight, rounded at junction with
propleura, anterior angles bluntly prominent, usually rounded, sometimes protruding
slightly anteriorly, evident in lateral view; calli clearly defined, convex, broadly con-
fluent anteriorly, shining smooth, with fovate impression on anteromedial angles,

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reaching anterolateral margin of pronotum; ostiolar peritreme on some specimens
black. LEGS: black; tibiae except apices and bases, and apices of femora, testaceous;
metatibia frequently with basal third black. VESTITURE: dorsum covered with
sparse thin fuscous to golden setae; upright on head, suberect and conspicuous on
pronotum, practically appressed, inconspicuous, thin on hemelytra; basal portion of
hemelytra in some specimens, with sericeous, recurved setae; venter with sparse thin
fulvous setae.

Macropterous female. More robust than male, but similar in structure, color and
vestiture. Length 4.90-6.00, width 2.50-2.90. HEAD: width across eyes 1.19-1.34,
vertex 0.56-0.68, dorsal width of eye 0.28-0.35, distance from tylus to ventral margin
of eye 0.63-0.74, height of eye 0.48-0.55, maximum interocular width 0.87-0.98.
ROSTRUM: length 1.80-2.00, reaching and sometimes surpassing mesocoxae to
apex of metacoxae. ANTENNAE: I, length 0.53-0.70, sometimes extensively tes-
taceous or fulvous; II, 1.45-1.91; III, 0.73-1.13; IV, 0.98-1.25. PRONOTUM: length

l. 10-1.18, anterior width 0.87-1.08, maximum width 1.85-2.10. LEGS: light areas
of femora and tibiae testaceous, rufescent or fulvous.

Types. Male holotype. Tampico, [Yakima Co.], Wash., May 21, 1932, A. R. Rolfs.
Retained at the United States National Museum, Washington, D.C. I have examined
38 paratypes and the holotype, all are conspecific. The paratypes are deposited at
BYU, JFS and USNM.

Taxonomic consideration. Specimens from a population of I. fuscipubescens col-
lected 15 miles north of Weiser, Washington County, Idaho, on Balsamorhiza sp.,
have vestiture which is similar to the condition which typifies I. cascadia. The
specimens are typical of I. fuscipubescens in the vertex to antennal segment I ratio
and genital structure. These two species are sister groups and are mostly allopatric.
Specimens of I. fuscipubescens from Chelan County near Dryden, have dark ostiolar
peritremes and long fuscous setae on the hemelytral border. Such specimens can be
confused with I. nigripes Knight.

Remarks. I. fuscipubescens has a northern and montane distribution which is
bordered on the west by the east slopes of the Cascade Range where the vegetational
associations are an intergradation of western ponderosa pine forst and sagebrush
steppe. The northern limit of its range is 50° 30' north latitude. The vegetation
associations occupied by I. fuscipubescens, south and east of this limit, consist of
fescue-wheatgrass grassland (Palouse Prairie) and Douglas fir and western ponderosa
pine forests. The far eastern limit of its range remains within the Douglas fir forest
of Rocky Mountains of western Wyoming. Its elevational range is from 210 to 2570

m. (700 to 8435 feet) and dates of adult occurrence are 20 April to 25 August. Adult
specimens have been collected from the following plants: Agropyron sp., A. repens
(L.) Beauv., Balsamorhiza sagittata (Pursh) Nutt., Betula sp., Chrysothamnus sp.,
Poaceae, Helianthus sp., Lathyrus sp., Lupinus sp., Medicago sp., Pinus contorta
Dougl. ex Loud., Potentilla sp., Salix sp., Smilacina racemosa (L.) Desf. and Trifolium
pratense L.

Specimens examined. 860 specimens were examined from the following localities (Map 22).
CANADA: ALBERTA: Banff (CNC, KU, USNM); Blairmore (CNC); Cameron Lk. Rd.-
Waterton Lks. Nat. Pk. (CNC); Coleman (CNC); Frank (CNC): Gorge Crk. (UALB); Jasper
Nat. Pk. (CNC); Kananaskis-Hwy. 25, Rds. 20 & 27 (CNC); Lundbreck (CNC); Waterton

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(CNC, UALB). BRITISH COLUMBIA: Aspen Grove (CNC); Brand Forks (UBC); 7 mi. W
Bridesville (UCB); 4 mi. E Christina Lk. (UBC); road to Copper Mt.— nr. Nelson (UBC); Eva
Lk. Trl. (CNC); Femie & Lizard Crk. (UBC); Glacier Nat. Pk. (CNC); Hall Crk. (UBC); Hedley-
Nickel Plate City & mine (CNC); Hosmer (UBC); Kamloops (UBC) & Paul L. (CNC); Kootenay
Prov. Pk.— Vermilion R. (CNC); Loma (USNM); Manning Prov. Pk.— Allison Pass, Blackwall,
Gibson Pass (CNC); Grand Forks (CNC); Mara (UBC); Merritt -Midday Valley (USNM), &
10 mi. S (CNC); Midway (CNC); Monaskie Pass (CNC); Mt. Revelstoke (CNC); Nicola (UBC);
Oliver— 7 mi. E & Meyers Rat (CNC); Osoyoos— Anarchrist Mt. (CNC); Penticton (USNM);
Robson (UBC); 8 mi. W Rossland (CNC); Salmo (UBC); Sicamous (CNC); Trinity Valley
(CNC); Vernon (CAS, UBC); Yoho Valley- Yoho Nat. Pk. (AMNH). UNITED STATES:
IDAHO: Blaine Co., Alturas Lk. (AMNH). Bonneville Co., Palisades Dam (UCD). Boundary
Co., Brush Lk. (UID). Camas Co., Fairfield (UID). Caribou Co., 8 mi. E Wayan (UID). Kootenai
Co., Athol (UID); 12 mi. N Coeur d’Alene (KU). Latah Co., Moscow Mt. (USNM). Lemhi
Co., Salmon River (KU). Valley Co., No Business Lookout (UID). Washington Co., 15 mi. N
Weiser (OSU). MONTANA: Glacier Co., 10 & 15 mi. W Babb -Glacier Nat. Pk. (OSU); Glacier

N. P. (UALB); Swift Current Pass-Glacier N. P. (AMNH). Lake Co., Swan Lk. (USNM).
Lincoln Co., 1 1 mi. E Libby (MSU). OREGON: Baker Co., 5 mi. NW Ballards Landing (T5S
R78S Sec. 31 SE Va SW »/4) (OSU). Deschutes Co., 6 mi. NW & E Sisters (AMNH, OSU); Three
Crks. Mdw. (OSU). Hood River Co., Hood R. (USNM). Wallowa Co., Minam St. Pk. (OSU).
Wasco Co., Cherry Hts. Rd.-2.5, 3.6 & 5.5 mi. S The Dalles (OSU); 5 mi. N & 1 1 mi. S Dufur
(JS, OSU); Mayer St. Pk. (OSU); Mosier (JS); Rowena Crest (OSU); Rowena Loop Rd. — 5, 5.5
& 7 mi. E Mosier (OSU); 2 mi. S & 7 mi. N Simnasho (OSU); Skyline Dr.— The Dalles (OSU);
The Dalles (JS). WASHINGTON: Chelan Co., 8.4 mi. SW & Dryden (OSU & USNM), 0.4
mi. W Leavenworth (OSU). Ferry Co., W Fk. Sanpoil River (OSU). Grant Co., Vantage (AMNH).
Klickitat Co., 1.4, 2 & 3 mi. E Bingen (OSU); 14 mi. W Goldendale (OSU); 3 & 4.5 mi. S
Husum (OSU); 2 mi. NW & 21.7 mi. NE Lyle (OSU); Maryhill (OSU); 3.7 E Wahkiacus (OSU);

O. 4 mi. E of mp. 70 on rt. 830 (OSU). Okanogan Co., Nespelem (OSU). Skamania Co., Dog
Mt. — Pacific Crst. Trl. (OSU). Stevens Co., Northport (UCB, USNM). Yakima Co., Clear Lk.
(SHF); 13, 13.5 mi. W & 15 mi. WNW Naches (OSU); Rimrock (WSU); Tampico (BYU);
Tieton (OSU); Tieton Cyn. (SHF); 27.6 & 30 mi. S & Toppenish (OSU, USNM). WYOMING:
Teton Co., 1 1.2 mi. W Jackson (OSU); Teton Pass (USNM).

Irbisia incomperta Bliven
Figs. 17a, b, 40, 63, 86; Map 7

Irbisia incomperta Bliven, 1963, pp. 73-74, 86, pi. 8, figs. 1, la (new species).

Diagnosis. This species can be distinguished from allied species on the basis of
these characters: slightly shagreened frons; obvious, transverse sulcus on posterior
margin of temporal area; deep and broad median depression on head; narrow vertex
(vertex to antennal segment I ratio: for males from 1 : 1 . 1 3 to 1 : 1 .35; for females from
1:1.00 to 1:1.04); concave lateral margins of pronotum; ochraceous basalar plate;
sclerotized rings and dorsal labiate plate of the female are small, compressed, barely
open and narrow, respectively; the left paramere of the male is broadly curved, the
outer surface with a series of moderately sized pointed tubercules, the shaft is not
constricted preapically, only slightly curved toward the sensory lobe, the apex barely
securiform. In addition to the above characters, I. incomperta differs from I. solani,
in the female of the former with fuscous spotted femora. I. incomperta does not
possess densely distributed long suberect fulvous setae on the tibia which additionally
distinguish this species from I. oreas Bliven.

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Description. Macropterous male. Length 4.20-4.90, width 1 .70-1 .95, shining black,
moderately clothed with a mixture of recurved sericeous white setae and thin suberect
fuscous setae. HEAD: width across eyes 1.04-1.16, vertex 0.43-0.48, dorsal width
of eye 0.28-0.33; triangular, smooth or slightly shagreened; tylus rounded in dorsal
view; temporal areas smooth, distinct, slightly flattened with obvious transverse
sulcus on posterior margin; median depression evident, sometimes broad or a lon-
gitudinal furrow; basal carina moderately prominent, moderately acute or rounded
in cross section, declivous to level of vertex, straight; eyes broadly joined to frons,
protruding laterally, posterolateral margin slightly arcuate postad in dorsal view;
distance from tylus to ventral margin of eye 0.48-0.54, height of eye 0.48-0.53,
maximum interocular width 0.70-0.83; jugal-loral suture sometimes fulvous. ROS-
TRUM: length 1.60-1.68, black or piceous, medial region fulvous, reaching bases of
mesocoxae, sometimes surpassing apices of mesocoxae in some specimens. ANTEN-
NAE: black; I, length 0.51-0.65; II, 1.38-1.63; III, 0.73-0.93; IV, 0.86-1.00.
PRONOTUM: length 0.83-0.95, anterior width 0.80-0.88, maximum width 1.45-
1.63; conical broadly convex, distinctly punctate on disk, lateral and posterior mar-
gins confluently punctate, posterolateral margin arcuate, lateral margins slightly con-
cave, broadly rounded at junction with propleura (sometimes appearing slightly
margined) anterior angles broadly rounded in dorsal view, not sulcate in lateral view;
calli slightly to moderately convex, mostly smooth, broadly confluent anteriorly,
reaching anterolateral margins of pronotum; basalar plate and insertion of hemelytra
ochraceous. LEGS: black; apices of coxae and margins of trochanters testaceous;
apices of femora and tibiae fulvous or fuscous; apices and bases of tibiae and tarsi
black. VESTITURE: moderately clothed with a mixture of recurved, appressed se-
riceous white setae, and thin white to fuscous setae, thin setae are erect on head,
pronotum and scutellum, suberect on hemelytra; venter sparsely clothed with thin
suberect white to fulvous setae; eyes very sparsely pubescent between facets.

Macropterous female. Slightly more robust than male, but similar in structure,
color and vestiture. Length 4.85-5.50, width 1.90-2.25. HEAD: width across eyes
1.09-1.19, vertex 0.50-0.53, dorsal width of eyes 0.28-0.31, distance from tylus to
ventral margin of eye 0.53-0.58, height of eye 0.46-0.53, maximum interocular width
0.78-0.85; cly peal, jugal and loral sutures sometimes testaceous. ROSTRUM: length
1.58-1.80, reaching bases of metaxocae. ANTENNAE: black or fuscous, I, length
0.50-0.56, sometimes entirely fulvous except for bases; II, 1.18-1.45; III, 0.70-0.95;
IV, 0.90-1.03. PRONOTUM: length 0.90-1.03, anterior width 0.85-0.95, maximum
width 1.63-1.81. LEGS: fulvous; medial portion of coxae, preapical one fourth of
femora, bases and apices of tibiae, and tarsi, fuscous; femora with fuscous spots
ventrally.

Types. Male holotype. Woody, Kern Co., California, Mar. 5, 1961; HOLOTYPE,
Irbisia incomperta, 1963, B. P. Bliven [red]; CAS type no. 13853. I have examined
the holotype and all 26 paratypes. All are conspecific and retained at the California
Academy of Sciences, San Francisco.

Remarks. The distribution of this species is fairly widespread throughout the blue
oak-digger pine forest floristic association of the Californian chaparral and Sierra
forest provinces. Specimens may be found in the northern yellow pine forest floristic
associations in Shasta County, and chaparral associations in Tulare and San Diego
Counties. Adult specimens have been collected from the following plants: Ceanothus

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cuneatus (Hook.) Nutt., Prunus subcordata Benth., Quercus douglasii H. & A., Q.
mslizenii A. D.C., Rhamnus crocea Nutt, ex T. & G., and Yucca brevifolia Engelm.
in Wats. This species has also been collected from several unidentified Poaceae. The
dates of adult occurrence are 10 March to 10 May. The elevational range of the
collection sites is from 655 to 1070 m. (2150 to 3500 feet).

Specimens examined. 176 specimens were examined from the following localities (Map 7).
UNITED STATES: CALIFORNIA: Butte Co., Oroville (CAS). El Dorado Co., Clarksville
(F&A); Cool (UCB). Fresno Co., 12 mi. SW Coalinga (UCB); Jacalitos Cyn.— Coalinga (CAS).
Kern Co., Kemville (CAS, UCB); Miracle Spgs. (UCB); 1 mi. E Woody (UCB). Mendocino Co.,
Ukiah (USNM). Monterey Co., Bryson (CAS). Napa Co., Monticello (F&A). San Benito Co.,
Big Panoche Crk. (UCB); Panoche Pass (UCB); Pinnacles Nat. Mon.— W entr. (OSU); 5 mi.
W San Juan Bautista (UCD). San Diego Co., 2 mi. S Pine Valley (UCB). S.L.O. Co., 4 & 10
mi. S & 10 mi. SE Creston (UCB); Pozo & 3 mi. E (UCB); Temblor Rng.— 12 mi. E Simmler
(UCB). Shasta Co., Whiskeytown St. Pk.— on rt. 299 (OSU). Stanislaus Co., Del Puerto Cyn.
(UCB, UCD). Sutter Co., Sutter Buttes (UCD). Tehama Co., 21 mi. NW Red Bluff on rt. 36
(OSU). Tulare Co., 9 & 1 1 mi. NW Calif. Hot Spr. (UCB); Fairview (UCB); 2 mi. E Johnsondale
(UCB).

Irbisia knighti Schwartz and Lattin
Figs. 7b, 19a, b, 42, 65, 88; Map 8

Irbisia knighti Schwartz and Lattin, 1984, pp. 413-417, figs. 1-8 (new species).

Diagnosis. Similar to I. solani in size and surface features but is consistently
recognizable by brachyptery of both sexes and white vestiture. I. knighti is separated
from the brachypterous form of I. sericans by the slightly punctate pronotum and
white vestiture of the former species (Figs. 1, 2).

Description. Brachypterous male. Length 3.90-4.63, width 2.18-2.30, shining black,
moderately covered with, shining white setae. HEAD: width across eyes 1.25-1.30,
vertex 0.59-0.63, dorsal width of eye 0.35-0.36; triangular, smooth; temporal areas
distinct, bordered basally by transverse short sulcus, glabrous; median depression
indistinct; basal carina moderately prominent, slightly rounded, declivous to level
of vertex, straight eyes broadly joined to frons, posterolateral margins arcuate pos-
teriorly in dorsal view; distance from tylus to ventral margin of eye 0.6 1-0.63, height
of eye 0.50-0.53, maximum interocular width (anterior view of frons) 0.90-0.95.
ROSTRUM: length 1.70-1.83, black, surpassing mesocoxae to just attaining apices
of metacoxae. ANTENNAE: black; I, length 0.50-0.55, apices fuscous; II, 1.45-1.55;
III, 0.78-0.93; IV, 1.02-1.20. PRONOTUM: length 0.86-0.88, anterior width 0.93-
0.95, posterior width 1.45-1.55; subconical, slightly flattened, distinctly or con-
fluently punctate, lateral margins straight, broadly rounded, at junction with pro-
pleura, anterior angles rounded indistinct in dorsal view, gently sulcate in lateral
view; calli slightly convex, smooth, narrowly confluent anteriorly, with transverse
depression on inner, and foveate depression of outer anterior angles reaching antero-
lateral margins of pronotum. LEGS: black; coxae and femora with apices testaceous;
tibiae testaceous with bases of bases black; tarsi black. HEMELYTRA: membrane
reduced, extending slightly beyond apex of cuneus, cells not developed; apex of
abdomen exposed in dorsal view. VESTITURE: dorsum clothed with suberect white
setae; head and pronotum sparsely covered with thin setae; hemelytra moderately

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covered with sericeous setae basally, these setae grading into and being replaced by
thin setae apically; venter moderately clothed with white setae; eyes with sparse
minute pubescence between facets.

Brachypterous female. More robust than male, but very similar in color, structure
and vestiture. Length 4.40-4.95, width 2.40-2.50. HEAD: width across eyes 1.35-
1.36, vertex 0.66-0.68, dorsal width of eye 0.35-0.38, distance from tylus to ventral
margin of eye 0.66-0.68, height of eye 0.50-0.54, maximum interocular width 0.96-

0. 98. ROSTRUM: length 1.88-1.93, barely reaching apices of metacoxae. ANTEN-
NAE: I, length 0.55-0.58; II, 1.53-1.55; III, 0.85-1.00; IV, 1.03-1.15. PRONOTUM:
length 0.93-0.96, anterior width 1.02-1.04, maximum width 1.65-1.70.

Types. Holotype male. Washington, Pacific Co., Nahcotta Oyster Research Station,
14 June 1979, M. D. Schwartz, ex Agropyron repens\ CAS type no. 15024. Retained
at the California Academy of Sciences, San Francisco. Paratypes are retained at
AMNH, CAS, CNC, OSU and USNM.

Discussion. This species is narrowly distributed along the coastal flatlands of the
Pacific Northwest. It occupies a very restricted portion of the Vancouveran Zone as
defined by Van Dyke (1939). Adult specimens were collected from these hosts:
Agropyron repens (L.) Beauv., Carex sp. (a sedge), Festuca rubra L., Holcus lanatus
L., Poa pratensis L., and Poa sp. None of these grasses ( Carex sp. is a doubtful host)
are obligatorily associated with the apparent restricted habitat of I. knighti. Collection
dates are from 15 May to 15 July. All localities are from sea level to 45 m. (150 feet)
elevation.

Specimens examined. 242 specimens were examined from the following localities (Map 8).
CANADA: BRITISH COLUMBIA: Jesse Island -Nanaimo (UBC, UCB, USMN); Mt. Doug-
las-Victoria (CNC); Royal Oak (CNC); Victoria (CAS, CNC, UCB, USNM). UNITED STATES:
CALIFORNIA: Humboldt Co., 5 mi. W Loleta-tidal mud flat (UALB). OREGON: Lane Co.,
12 mi. N Florence— Big Creek Rd. — Siuslaw Nat. For. (OSU). Tillamook Co., 2.3 mi. N
Neskowin on rt. 101 (OSU); 0.1 mi. N Rockway on rt. 101 (OSU); Sand Beach Cmpgrd. — 2
mi. W Sand Lk. (OSU); Sand Lk. (MSU); Tierra del Mar (OSU). WASHINGTON: King Co.,
Seattle (OSU). Pacific Co., Bay Center (OSU); Nahcotta, sweeping grasses just above tide line
on bay shore (AMNH, OSU); Wallapa Bay— near Nahcotta (WSU). Skagit Co., Mt. Vernon
(OSU).

Irbisia limata Bliven
Figs. 20a, b, 43, 66, 89; Map 5

Irbisia limata Bliven, 1963, pp. 78, 86, pi. 8, figs. 5, 5a (new species).

Diagnosis. I. limata differs from I. solani in these characters: polished, shining
black dorsum; virtually impunctate pronotum; longer rostrum (/. limata ranging
from 1 .88 to 2. 1 3 mm. for males and 1 .65 to 2.23 mm. for females; I. solani ranging
from 1.40 to 1.83 mm. for males and 1.60 to 1.80 mm. for females); flavescent first
antennal segment; left paramere and vesica of male. I. limata may be separated from

1. panda Bliven by the flavescent first antennal segment and legs of the former species.
Other distinguishing characters that separate these two species are found under the
diagnosis of I. panda.

Description. Macropterous male. Length 4.65-5.45, width 2.00-2.40, polished,
shining black, sparsely covered with recurved and appressed short fine testaceous to

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whitish setae. HEAD: width across eyes 1.07-1.20, vertex 0.48-0.55, dorsal width
of eye 0.28-0.31; triangular, polished; temporal areas indistinct; median depression
lacking; basal carina small, but broad in cross section, barely higher than collum,
declivous to level of vertex, straight; eyes broadly joined to frons, straight in dorsal
view, distance from tylus to ventral margin of eye 0.58-0.63, height of eye 0.50-
0.54, maximum interocular width 0.75-0.85. ROSTRUM: length 1.88-2.13, black
to fuscous, surpassing mesocoxae, usually surpassing metacoxae. ANTENNAE: fus-
cous or black; I, length 0.60-0.70, flavescent, base and anterior surface, fulvous,
fuscous or piceous, with appressed short black setae, and black bristles; II, 1.58-
2.01, sometimes testaceous on basal third; III, 1.00-1.38; IV, 0.88-1.05. PRONO-
TUM: length 0.94-1.08, anterior width 0.83-0.98, maximum width 1.55-1.88; sub-
conical, broadly convex, slightly rugulose, very slightly punctate, posterior margin
arcuate, lateral margins straight, rounded at junction with propleura, anterior angles
rounded, not prominent in dorsal view, with single black bristle, slightly sulcate in
lateral view; calli prominent, convex, polished, confluent anteriorly, reaching an-
terolateral margins of pronotum, collar widest medially; ventral third of propleura
and prostemum narrowly bordering coxae and attaining front margins, flavescent.
LEGS: flavescent; coxae, except for apices, piceous; trochanters, basal portion (in
variable degree) of femora, apices of tibiae, and tarsus sometimes piceous; femora
sometimes rufescent. VESTITURE: sparsely clothed with short, fine, testaceous to
white setae; recurved and appressed on hemelytra, suberect on head and pronotum;
venter with sparse suberect fulvous setae.

Macropterous female. More robust than male, otherwise not differing greatly in
structure, color and vestiture. Length 4.50-5.40, width 2.20-2.60. HEAD: width
across eyes 1 .10-1.20, vertex 0.53-0.60, dorsal width of eye 0.25-0.30, distance from
tylus to ventral margin of eye 0.63-0.68, height of eye 0.50-0.54, maximum inter-
ocular width 0.80-0.88. ROSTRUM: length 1.65-2.23, reaching and usually sur-
passing metacoxae. ANTENNAE: I, length 0.59-0.70; II, 1.54-1.90; III, 1.20-1.40;
IV, 1.00-1.13. PRONOTUM: length 1.00-1.20, anterior width 0.90-1.05, maximum
width 1.73-1.95.

Types. Holotype male. Fountain Springs, [Tulare Co.], Cal., Mar. 26. 1961; HO-
LOTYPE 6, Irbisia limata, 1963 B. P. Bliven [red]; CAS type no. 13856. I have
examined the holotype and the 39 paratypes mentioned in the original description;
all are conspecific and retained at the California Academy of Sciences, San Francisco.

Remarks. The distribution of /. limata is predominately contained within the blue
oak-digger pine vegetation association of the Sierran forest province. This species
also occurs, but to a lesser extent, in chaparral, California prairie, riparian forest and
valley oak savanna vegetation associations of both the Sierran and Californian chap-
arral provinces. Angles Camp, Calaveras County, is the northernmost locality of the
material on hand; the only record south of the Transverse Ranges is a single specimen
from Coachella Valley, Riverside County. Adult specimens have been collected from:
Aesculus californica (Spach) Nutt., Amsinckia sp., Avena sp., Bromus sp., Calochortus
luteus Dougl. ex Lindl., Clarkia cylindrica (Jeps.) Lewis & Lewis, C. mlliamsonii
(Dur. & Hilg.) Lewis & Lewis, Asteraceae, Poaceae, Sanicula tuberosa Torr. and
Vulpia sp. The period of occurrence is from 20 March to 14 July, and the altitudinal
range is from 240 to 1220 m. (800 to 4000 feet) elevation.

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Specimens examined. 246 specimens were examined from the following localities (Map 5).
UNITED STATES: CALIFORNIA: Calaveras Co., Angles Camp (CAS). Fresno Co., 12 mi.
SW Coalinga (UCB); Pine Flat (UCR); Squaw Valley (UCR). Kern Co., 28 mi. E Bakersfield
(UCB); Fountain Spgs. (OSU); Glenville (OSU); Havilah (UCR); Hobo Cmpgd. — Kern River
(UCB); 10-20 mi. S Isabella Lk. (AMNH); Miracle Spgs. (UCB); Taft (LACM); Tejon Pass-
Lebec (UCB); Woody (OSU, UCB). Madera Co., 2 mi. NE & 3 mi. W Bass Lks (UCB, CAS);
Coarsegold (UCB, UID); Northfork (CAS, USNM). Mariposa Co., 1 .9 mi. W Mt. Bullion (UCB);
Mariposa Co. (UCB). Riverside Co., Coachella Valley (CAS). Sta. Clara Co., Diablo Range
(CAS). Stanislaus Co., 1 mi. W LaGrange (UCB). Tulare Co., Ash Mt. HQ— N Kaweah (AMNH,
UCB); Badger (CAS); 11 mi. NW Calif Hot Spgs. (UCB); Coffee Cmp.-5 mi. E Springville
(UCB); Cold Spgs.— Sequoia N.F. (USNM); Fountain Spgs. & 8 mi. SE, (CAS & AMNH, UCR);
8 mi. NE Hammond (OSU); 5 mi. N Kaweah (UCB); Lemoncove & 4 mi. NE (UCB); Porterville
(F&A), 1 mi. NE Posey (UCB), Potwisha— Sequoia N.P. (CAS); Rose Crk.— 10 mi. N Columbia
(UCB); Sequoia N.P. (LACM); 8 mi. NE Three Rivers (USU); Tule River Cyn. (KU). Tuolumne
Co., Sonora (UCB).

Irbisia mollipes Van Duzee
Figs. 29a, b, 52, 75, 98; Map 14

Irbisia sericans var. mollipes Van Duzee, 1917b, p. 264 (new variation).

Irbisia mollipes: Van Duzee, 1921a, pp. 147-148, 152 (new status). Essig, 1926, p.

361. Carvalho, 1959, p. 105.

Irbisia umbratica: Bliven, 1963, pp. 69-70 (in part, not holotype).

Irbisia upupa Bliven, 1963, pp. 70-71, 85, pi. 7, figs. 8, 8a. New Synonymy.

Diagnosis. Irbisia mollipes is the only species in the genus in which the dorsum is
densely covered with suberect, long shining pale white to pale golden setae. Specimens
of I. bliveni from Mariposa County, California, to the northern limit of its range,
have vestiture similar to I. mollipes . I. mollipes differs from the former species in
these characters: the vertex is narrower (I. mollipes ranging from 0.40 to 0.50 mm.
males; from 0.48 to 0.55 mm. females; I. bliveni ranging from 0.56 to 0.64 mm.
males; from 0.60 to 0.70 mm. females), the legs are usually fulvous or rufescent, and
the sclerotized process is laterally inserted on the left side of the main trunk of the
vesica of the male.

Description. Macropterous male. Length 4.50-5.80, width 1.88-2.20, polished,
shining black clothed with dense, suberect long shining pale white to pale golden
setae. HEAD: width across eyes 1.07-1.23, vertex 0.40-0.50, dorsal width of eye
0.30-0.36; triangular, polished; temporal areas indistinct; median depression usually
lacking; basal carina straight, very small, but acute in cross section, scarcely higher
than collum, declivous to level of vertex; eyes broadly joined to frons, but produced
laterally, posterolateral margin arcuate posteriorly in dorsal view, distance from tylus
to ventral margin of eye 0.53-0.63, height of eye 0.46-0.56, maximum interocular
width 0.75-0.85. ROSTRUM: length 1.48-1.73, fulvous, base and apex infuscated,
reaching bases of mesocoxae. ANTENNAE: black; I, length 0.59-0.73, sometimes
slightly fuscous; II, 1.73-2.25; III, 0.91-1.30; IV, 0.93-1.20. PRONOTUM: length
0.91-1.15, anterior width 0.78-0.90, maximum width 1.58-1.84; conical, broadly
convex, distinctly punctate, posterior margin arcuate, lateral margins slightly concave,
broadly rounded at junction with propleura, anterior angles rounded, quite reduced
in dorsal view, not sulcate in lateral view; calli slightly elevated, polished, confluent

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anteriorly, broadly attaining anterolateral margin of pronotum and curving around
anterior angles. LEGS: fulvous, rufescent to piceous; apices of coxae and trochanters
sometimes testaceous; apices, bases and, in some specimens, extensive areas of tibiae
infuscated; tarsi infuscated or black. VESTITURE: densely clothed with long shining,
pale white to golden setae, erect on head, suberect on rest of dorsum; venter mod-
erately covered with suberect white to golden setae.

Macropterous female. More robust than male, but not differing greatly in structure,
color and vestiture. Length 4.70-5.80, width 2.00-2.90. HEAD: width across eyes
1.10-1.30, vertex 0.48-0.55, dorsal width of eye 0.29-0.38, distance from tylus to
ventral margin of eye 0.59-0.65, height of eye 0.48-0.55, maximum interocular width
0.85-0.93. ROSTRUM: length 1.65-1.85, barely surpassing base of mesocoxae. AN-
TENNAE: I, length 0.56-0.70; II, 1 .48-2. 1 3; III, 0.90-1 .38; IV, 1 . 1 3-1 .2 1 . PRONO-
TUM: length 0.98-1.28, anterior width 0.85-1.00, maximum width 1.83-2.10.

Types. Male holotype. Santa Cruz Co., Cal. no. 2B; W. M. Giffard, 7-VI-17; TYPE
mollipes [red]; CAS type no. 337. I have examined two paratypes and the holotype;
all are conspecfic and retained at the California Academy of Sciences, San Francisco.

Taxonomic consideration. I. upupa Bliven, the junior synonym, meets the habitus
of I. mollipes in structure, color and vestiture. Van Duzee (1921a) lists several
specimens which he determined as I. mollipes. All of this material is retained in the
collection of the California Academy of Sciences. I have examined these specimens
and have determined that they are not all entirely conspecific. The specimens are
correctly identified as: I. mollipes: Two males— SLObispo, Cal IV-24-19; EP Van
Duzee Collector. Male— Santa Cruz Isd. Cal. V-18-1010; EP Van Duzee Collector.
Male— Keen Camp Cal., Riverside Co., June 6-12, 1917; EP Van Duzee Collection.
Pair— Pasadena, Cal. 5-1-09, Grinnell; EP Van Duzee Collection (yellow). Female—
Pasadena, Cal. 4-9-09, Grinnell; EP Van Duzee Collection (yellow); Two females—
same data, retained at USNM. I. setosa: Female— Ross Cal, Marin Co., iii-3 1-18;
EP Van Duzee Collector. I. (sp.?), teneral: Female— Alpine San Diego Co., 3-11-
1914 Cal, EP Van Duzee; EP Van Duzee Collection (yellow). I. serrata: Pair— Top
of Las Vegas Range, VI.28’02 NM; EP Van Duzee Collection (yellow).

Remarks. I. mollipes is distributed wholly within the California chaparral province
from the San Francisco Bay area to San Diego County. Throughout this range the
majority of specimens were collected from three forest associations— mixed hard-
wood, blue oak-digger pine and southern oak. Adult specimens were found on the
following plants: Poaceae, Hordeum vulgare L., Juglans californica Wats., Lupinus
sp., Marah sp., Medicago sp., Mimulus longiflorus (Nutt.) Grant, Prunus ilicifolia
(Nutt.) Walp., Rubus sp. and Salix sp. The dates of occurrence range from 3 March
to 20 August. The preponderance of late summer dates suggests that this species may
occur later in the season than many other species of Irbisia. Specimens were collected
within the altitudinal range of 180 to 2300 m. (600 to 6800 feet).

Specimens examined. 320 specimens were examined from the following localities (Map 14).
UNITED STATES: CALIFORNIA: Alemada Co., Arroyo Mocho— 8 mi. SE Livermore (CAS);
Livermore (CAS); Niles Cyn. (CAS). Contra Costa Co., Moraga (AMNH). Kern Co., Frazier
Mt. Pk. (UCB); Mt. Pinos (UCB). Los Angeles Co., Arroyo Seco— nr. Pasadena (USNM);
Burbank (USNM); Chatsworth (LACM); Claremont (UCB); Covina (F&A); Elizabeth Lk. Cyn.
(LACM); Glendale (UCD); Green River Cmp.— Lwr. Sta. Ana Cyn. (CAS); Griffiths Prk. (USNM);

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Los Gatos (UCD); Pasadena (CAS, USNM), Pomona (UCR); Puente Hills (UCR); San Gabriel
Cyn.— nr. Dam no. 1 (LACM); Sierra Madre (USNM); Tanbark Flat (F&A, LACM, UCD,
UCR), Tapia Park (LACM); Topanga Cyn. (LACM); Whittier (LACM, UCR). Marin Co.,
Lagunitas (CAS). Merced Co., Pacheco Pass Summit (F&A). Monterey Co., 5 mi. NE Arroyo
Seco G. Sta. (UCB); Big Sur Crk. Trl. (CAS); Bixby Crk. (OSU); Bradley (CAS); Bryson (CAS);
Greenfield (F&A); Horse Bridge— 1.5 mi. SW Arroyo Seco G. Sta. (UCB); Indian Rd. — 2 mi.
S Arroyo Seco Grd. Stn. (UCB); Paloma Crk. — 5 mi. NE Arroyo Seco Grd. Stn. (UCB); Paraiso
Spgs. (CAS); Pleyto (CAS); Tassajara Hot Spgs. (CAS); Ventana Wldns. — 8 mi. SSE Hwy. 1 on
Palo Colorado Rd. (OSU). Riverside Co., Bautista Cyn. — 8 mi. SE Hemet (UCR); El Cariso
Rgr. Stn.— Hwy. 74, Clev. Nat. For. (UCR), Keen Cmp. (CAS); 2 mi. SE Poppet Flat— San
Jacinto Mts. (CAS, KU, UCB, UCR); Sandia Cyn. (T7S R4W Sec. 25) (UCR); Temecula Cyn.-
Sta. Margarita R. (UCR); Tenaga Cmpgd.— NW Murrieta (UCR). San Bernardino Co., 4 mi.
E Mentone (UCR); Wildwood Cyn. — 5 mi. E Calimesa (UCR). San Diego Co., Cuyamaca Lk.
(KU); Palomar (SDNH, UCD); Pine Valley (SDNH); Ramona (ASU); Warner Spgs. (SDNH).
San Mateo Co., Crystal Lks. (CAS); Crystal Spgs. (AMNH); Portola Valley (CAS). S.L.O. Co.,
Atascadero (CAS); 10 mi. SE Creston (UCB); Cuesta Cyn. Pk. (AMNH); La Panza Cmpgd.
(UCB); Morro Bay (CAS); 3 mi. W Paso Robles (UCB). Sta. Barbara Co., Beach— Water Cyn.
(UCD), Canada del Medio (SBMN, UCD), Central Valley (UCB, UCR), Coches Prietos (UCD),
Prisoner’s Harbor (UCB), Ridge btw. Central Valley & Islay Cyn. (UCB), Upper Valley (UCB),
Valley anchorage (UCB)— all Sta. Cruz Is.; 3 mi. SE Zaca Pk. (UCB). Sta. Clara Co., Alum
Rock Pk. (UALB, UCB); Mt. Hamilton (UCB); Page Mill Rd. (USNM). Independence (LACM).

Irbisia nigripes Knight
Figs. 21a, b, 44a, b, 67, 90; Map 19

Irbisia nigripes Knight, 1925a, pp. 94-95 (new species); 1925b, p. 182; 1968, p. 185.

Downes, 1927, p. 12. Carvalho, 1959, p. 105. Harling et al., 1977, p. 35. Kelton,

1980, pp. 84-85, map 23.

Diagnosis. This species is unique among the species of the genus in possessing an
entirely black pleural surface on most specimens. The pronotal disk is conspicuously
flattened and the anterior angles of pronotum are very prominent in dorsal and lateral
views. In addition to the previous characters, the structure of the sclerotized process
of the vesica of the male will serve to distinguish I. nigripes from I. serrata. The
process of I. nigripes is not preapically expanded nor does it possess a linear series
of strong spicules. The basal carina of the vertex of I. nigripes is considerably more
produced than on I. serrata.

Description. Macropterous male. Length 5.50-6.30, width 2.43-2.65, shining black,
clothed with woolly sericeous white setae. HEAD: width across eyes 1.18-1 .26, vertex
0.59-0.65, dorsal width of eye 0.28-0.31; triangular, smooth or slightly asperate;
temporal area shining; median depression broadly shallow; basal carina acutely prom-
inent in cross section, posteriorly arcuate mesad, declivous to level of vertex (declivate
area asperate); eyes broadly joined to frons, posterior margin of head slightly ante-
riorly arcuate at junction of eye and frons in dorsal view, distance from tylus to
ventral margin of eye 0.65-0.69, height of eye 0.45-0.50, maximum interocular width
0.88-0.39. ROSTRUM: length 1.70-1.89, black, just reaching apex of mesocoxae.
ANTENNAE: black; I, length 0.65-0.74; II, 1.76-2.03; III, 0.88-1.05; IV, 1.03-1.21.
PRONOTUM: length 1.00-1.13, anterior width 0.95-1.08, maximum width 1.83-
2.00, trapezoidal, disk somewhat flattened, rugulopunctate, posterior margin broadly

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REVISION OF IRBISIA

253

arcuate, lateral margins straight, rounded or broadly margined at junction with pro-
pleura, anterior angles very prominent in dorsal and lateral view, with explanate
flange directed somewhat anteriorly; calli weakly developed, broadly confluent an-
teriorly, not reaching anterolateral margins of pronotum; pro-, meso-, metapleura
and ostiolar peritreme entirely black except for narrow area bordering coxae; some
specimens with extreme posterior border of ostiolar peritreme white. LEGS: black;
coxae black. VESTITURE: dorsum covered with a mixture of recurved or recumbent
sericeous white setae and suberect fulvous to fuscous setae, disk of pronotum, es-
pecially anterior angles and lateral margins, with prominent, conspicuous, erect fus-
cous setae; venter with sparse, thin fulvous setae.

Macropterous female. More robust than male, but similar in color, structure and
vestiture. Length 5.60-6.40, width 2.65-2.90. HEAD: width across eyes 1.23-1.35,
vertex 0.68-0.75, dorsal width of eye 0.26-0.33, distance from tylus to ventral margin
of eye 0.65-0.76, height of eye 0.48-0.53, maximum interocular width 0.91-1.01.
ROSTRUM: length 1.83-2.06, just attaining to barely surpassing mesocoxae. AN-
TENNAE: I, length 0.65-0.73; II, 1.63-1.90; III, 0.88-1.08; IV, 0.98-1.28. PRONO-
TUM: length 1.08-1.30, anterior width 1.03-1.18, maximum width 2.00-2.28.

Types. Male holotype. Troy Ida, 3-31; HOLOTYPE By H. H. Knight Irbisia
nigripes [red]. Retained at Cornell University Collection, Ithaca, New York. I have
examined 1 3 paratypes and the holotype; they are all conspecfic. The paratypes are
deposited at CAS, CNC and USNM.

Taxonomic consideration. Male specimens from the northern portion of the range
of I. nigripes have sclerotized processes with fewer and less prominent spines (the
spines are not arranged in a linear series) as compared to the spines of the sclerotized
processes of more southern specimens. The process of individuals from Wasco Coun-
ty, Oregon and Klickitat County, Washington typify the southern condition with a
linear series of strong spicules. The single specimen from Crater Lake, Klamath
County, Oregon has a process that closely resembles the northern specimens.

Remarks. Although the number of specimens on hand of I. nigripes are few, its
distribution is similar to the northern and montane ranges of I. elongata and I.
fuscipubescens. I. nigripes appears to be the northern counterpart of I. shulli. The
latter is not known from Canada, whereas I. nigripes has a northern limit of 50° 15'
north latitude. Its range is bounded on the west by the Cascade Range where it occurs
in western ponderosa pine and sagebrush steppe vegetational associations. Through-
out the rest of its range both grassland and forest border situations provide suitable
habitats; fescue-wheatgrass, foothill prairie ( Agropyron-Festuca-Stipa ), sagebrush
steppe, western ponderosa pine and Douglas fir forests of the Rocky Mountain region
of Idaho and Montana are the predominate vegetational associations. Host data are
lacking, but adult specimens were collected from Balsamorhiza sp., Ceanothus sp.,
Chrysothamnus sp. and Dactylus glomerata L. Its elevational range is from 460 to
2130 m. (1500 to 7000 feet) and occurs from 22 March to 13 July.

Specimens examined. 167 specimens were examined from the following localities (Map 19).
CANADA: ALBERTA: Crows Nest (CNC); Kananskis-Hwy. 25, Rds. 20 & 27 (CNC); Twin
Butte (CNC); Waterton (CAS, CNC, UALB). BRITISH COLUMBIA: Cascade (CAS), Chase
(CNC); Christina Lk. (UCB); Creston (UCB); 5 mi. W Creston-Hwy. 3, 49° 5' 1 16° 37' (UBC);
10 mi. E Falkland (CNC); Fife, & 5 mi. E (CNC); Grand Forks, & 9 mi. E (CNC); Rock Crk.,

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& 3 mi. W (CNC); Trinity Valley (CNC); Vernon (CAS, CNC, UBC). UNITED STATES:
IDAHO: Bonner Co., 4 mi. S Elmira (UID); 2 & 7 mi. S Sandpoint (UID). Kootenai Co., 4 mi.
W Athol (UID); Coeur d’Alene (OSU). Latah Co., Deary (UID); Julietta (BYU, SHF, UNSM);
Laird Park (UID); Moscow Mt. (OSU); Troy (UID). MONTANA: Lake Co., 10 mi. S Bigfork
(MSU); east shore Flathead Lk. (MSU); “Lake Co.” (MSU). Missoula Co., Missoula (WSU).
Ravalli Co., Blodgtt Mt. (MSU); 15 mi. SE Hamilton (OSU); Montana Expt. Stn.— Florence
(MSU). Sanders Co., Camas Prairie (MSU). OREGON: Hood River Co., Hood R. (USNM).
Klamath Co., Crater Lk. Nat. Pk. (USNM). Wasco Co., 7 mi. N Simnasho (OSU). WASH-
INGTON: Chelan Co., Leavenworth (OSU, WWU). Klickitat Co., 2 mi. NW & 5 mi. NE Lyle
(OSU); Satus Pass (KU); Trout Lk. (OSU). Lincoln Co., 5 mi. SE Grand Coulee (OSU).

Irbisia oreas Bliven
Figs. 16a, b, 39, 62, 85; Map 6

Irbisia oreas Bliven, 1963, pp. 72-73 (new species).

Diagnosis. This species may be separated from other small species of the genus on
the basis of a combination of these characters: lack of light spot on bases of cuneus
(as in I. cuneomaculata ); long erect fuscous or fulvous setae which are densely dis-
tributed throughout the dorsum (not present on I. solani, I. sita and I. incomperta );
and densely distributed, long suberect testaceous or white setae on tibiae (longer and
more erect than I. setosa Van Duzee). The right paramere of the male of I. oreas is
diagnostic. It is elongate with a somewhat concealed small apical spine that gives the
paramere a truncated appearance.

Description. Macropterous male. Length 3.83^4.75, width 1.65-1.90, shining black,
densely clothed with a mixture of recurved, appressed, sericeous white setae and long
erect fuscous to fulvous setae. HEAD: width across eyes 0.98-1 .09, vertex 0.44-0.49,
dorsal width of eye 0.26-0.30; triangular smooth or shagreened; tylus rounded in
dorsal view; temporal areas indistinct and without vestiture; median depression
absent; basal carina small, straight declivous to level of vertex or abruptly raised
above the vertex; eyes broadly joined to frons, protruding laterally, posterior margin
straight in dorsal view; distance from tylus to ventral margin of eye 0.43-0.49, height
of eye 0.43-0.48, maximum interocular width 0.65-0.76; jugal-loral suture some-
times testaceous. ROSTRUM: length 1.33-1.53, black, fuscous medially, reaching
apex of mesocoxae. ANTENNAE: black or fuscous; I, length 0.45-0.55; II, 1.13-
1.48; III, 1.33-1.53; IV, 0.73-0.85. PRONOTUM: length 0.74-0.93, anterior width
0.70-0.83, maximum width 1 .40-1 .63; conical, broadly convex and slightly flattened,
rugulopunctate, posteriolateral margins slightly arcuate, lateral margins concave, very
broadly rounded at junction with propleura, anterior angles broadly rounded, indis-
tinct in dorsal view, not sulcate in lateral view; calli not protruding above disk,
without vestiture, asperate, broadly confluent anteriorly, not reaching anterolateral
margin. LEGS: black or piceous; apices of coxae and trochanters fulvous; tibiae
usually lighter shade of base color; tarsi black. VESTITURE: densely clothed with
a mixture of recurved sericeous white setae, and long fuscous or fulvous setae; se-
riceous setae woolly on head and pronotum, appressed on hemelytra; long setae are
erect on head, pronotum and scutellum, suberect on hemelytra; venter moderately
covered with suberect long testaceous or white setae; legs densely covered with long
suberect testaceous or white setae.

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REVISION OF IRBISIA

255

Macropterous female. More robust than male, otherwise not differing in structure,
color and vestiture. Length 4.50-5.10, width 2.00-2.40. HEAD: width across eyes
1.10-1.16, vertex 0.51-0.55, dorsal width of eye 0.26-0.30, distance from tylus to
ventral margin of eye 0.5 1-0.58, height of eye 0.33-0.35, maximum interocular width
0.77-0.83. ROSTRUM: length 1 .54-1.68, attaining and sometimes surpassing apices
of mesocoxae. ANTENNAE: I, length 0.47-0.55; II, 1.20-1.40; III, 0.60-0.85; IV,
0.80-0.95. PRONOTUM: length 0.90-1.00, anterior width 0.83-0.88, maximum
width 1.65-1.83.

Type. Holotype female. Congress [Yavapai Co.], ARIZ, IV-6-58; HOLOTYPE 2,
Irbisia oreas, 1963 B. P. Bliven [red]; CAS type no. 13858. No other specimens were
collected by Bliven. Retained at the California Academy of Sciences, San Francisco.

Remarks. The distribution of I. oreas includes the eastern and western boundaries
of the Mojave Desert. The south westernmost portion of the Rocky Mountain forest
province in Washington County, Utah, and southern border of the Upper Gila Moun-
tain forest province in Pinal and Yavapai Counties, Arizona are the eastern limits
of the range. Within Arizona, specimens are found in a transition zone between oak-
juniper woodland and mountain mahogany-oak scrub floristic association; in Utah
juniper-pinyon woodland floristic associations are occupied.

Bliven (1963) predicted that I. oreas would be collected in extreme southeastern
California; some specimens available to this study are from this area. Several floristic
associations provide habitat for I. oreas in California. In the Sierran forest province
of Kern County these associations are: California prairie, blue oak-digger pine forest
and Joshua tree scrub. In the American desert province of Los Angeles and San
Bernardino Counties Joshua tree scrub and Mojave creosote bush associations are
occupied. Chaparral, coastal sagebrush, southern oak and juniper-pinyon forests are
the associations within the range of I. oreas in the California chaparral province of
Riverside and San Diego Counties.

Adult specimens have been collected from the following nongraminoid plants:
Ceanothus sp., Cercocarpus betuloides Nutt, ex T. & G., Chenopodium murale L.,
Keckiella sp., Larrea sp., Lotus scoparius (Nutt, in T. & G.) Ottley, Lycium sp. nr.
panchii Grey, Plagiobothrys sp., Quercus agrifolia Nee, Q. chrysolepis Liebm., Salvia
apiana Jeps; Simmondisa chinensis (Link) C. K. Schneid [only the male plants],
Yucca brevifolia Engelm. in Wats, and Y. schidigera Roezl ex Ortgies. The only grass
record is Festuca sp. The period of occurrence ranges from 23 February to 23 May,
and the altitudinal range is from 550 to 1580 m. (1800 to 5200 feet).

Specimens examined. 1 80 specimens were examined from the following localities (Map 6).
MEXICO: BAJA CALIFORNIA NORTE: 3.4 & 6.3 mi. S El Condor (AMNH); 2 mi. W Las
Encinas (UCB). UNITED STATES: ARIZONA: Maricopa Co., Four Peaks Rd., mi. 6 (JTP);
Vulture Mine Rd.— 4 mi. S Wickenburg (AMNH). Pinal Co., Apache Jet. (AMNH). Yuma Co.,
Yuma (USNM). CALIFORNIA: Kern Co., 20 mi. N Isabella Lk. (AMNH); Last Chance Cyn.-
9 mi. N Ricardo (UCB); 2 mi. N Mojave (UALB); 3 mi. W Woffard Hts. (UCB). Los Angeles
Co., 3 mi. S Lancaster (OSU); 2 mi. S Palmdale (OSU). Riverside Co., Lk. Mathews (UCR);
Manifee Valley (hills on W end), 33° 39' N 1 17° 13' W (UCR); 2 mi. NE Moreno (UC); Murray
Cyn.— Palm Spgs. (CNC); Palm Cyn.— Agua Caliente Indian Rsv. (CAB, CNC); 3.5 mi. S Palm
Desert— 0.8 mi. N jet. Deep Crk. & Horsethief Crk. (T7S R6E Sec. 6) (UCR); Palm Sprgs.
(USNM); 2 mi. N Perris (UCD); Pinyon Rats— 16 mi. SW Palm Desert (CAS, UCB); Railroad

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Cyn.— 4 mi. E Elsinore (UCB); 1 mi. SE & 4 mi. N Redec (UCB & AMNH); Riverside (UCR);
5 mi. S & 5.5 mi. SE Sage (UCB & UCR); 5.6 mi. S Sage on R3 (T7S R1E Sec. 32) (UCR);
Snow Crk.— White Water (CNC); trail From Hwy. 74 (T7S R5E Sec. 1-6) (UCR); White Water
(CNC); Wilson Valley Rd. — 1 mi. N Hemet (OSU). San Bernardino Co., Devil Cyn. (UCR):
Granite Cove— Granite Mts.— 15.8 mi. SW Kelson (UCD); Granite Mts. — 3 mi. ESE Apple
Valley (UCB); 4 mi. E Mentone (UCR); Yucaipa (UCD). San Diego Co., 4 mi. E Campo (UCB);
Jacumba (F&A, UCB). Tulare Co., 2 mi. E Johnsondale (UCB). UTAH: Washington Co., Dixie
St. Pk. (USU).

Irbisia pacifica (Uhler)

Figs. 1, 3, 7a, 1 la, b, 34, 57, 80; Map 3

Rhopalotomus pacificus: Uhler, 1871, p. 471 (name not available).

Rhopalotomus pacificus Uhler, 1872, p. 415 (new species); 1875, p. 319.

Capsus pacificus: Uhler, 1886, p. 19 (new combination). Atkinson, 1890, p. 105.
Thyrillus pacificus: Uhler 1894, p. 267 (new combination). Gillette and Baker, 1895,
p. 39. Essig, 1915, p. 215; 1927, pp. 361-362. Van Duzee, 1916b, p. 38; 1917a,
p. 325; 1917b, p. 264. Downes, 1927, p. 12. Slater, 1950, pp. 36, 71, pi. 2, figs.
20, 21, 75, pi. 4, fig. 9. Knowlton, 1951, pp. 74-75.

Irbisia pacificus: Van Duzee, 1914, p. 24 (new combination). Carvalho, 1959, p. 105.
Kelton, 1959, p. 18, fig. 28.

Irbisia pacifica: Knight, 1968, p. 185. Haws, 1978, p. 100, fig. 33.

Diagnosis. I. pacifica is separated from all other species in the genus by its: strongly
pedunculate eyes, which project above the basal carina, elongated head structure (the
range of the distance between tylus and ventral margin of eye to height of eye ratio
is, for males 1:0.33 to 1:0.58, and for females 1:0.37 to 1:0.49; the range for I.
elongata is males 1:0.48 to 1:0.53 and females 1:0.48 to 1:0.50), very long antenna
II which is much longer than the width of pronotum, wide collar and somewhat fine,
transversely rugulose pronotal surface.

Description. Macropterous male. Length 5.75-8.20, width 2.20-3.10, black, sha-
greened, densely clothed with appressed, recurved, white sericeous setae. HEAD:
width across eyes 1.28-1.54, vertex 0.64-0.79, dorsal width of eye 0.30-0.38; tri-
angular, but very elongated below eyes, shagreened and asperate; temporal areas
asperate and glabrous; median depression indistinct beneath vestiture, a short lon-
gitudinal furrow; basal carina very broadly rounded and declivous, barely projecting
above collum, arcuate posteriorly; eyes pedunculate, projecting above basal carina
in lateral view; posterior margin of head straight in dorsal view; distance from tylus
to ventral margin of eye 0.85-1.13, height of eye 0.41-0.50, maximum interocular
width 0.88-1.10; apical half of clypeus and clypeal-jugal-loral sutures sometimes
fulvous or rufescent; antennal sockets below ventral margin of eyes. ROSTRUM:
length 2.33-2.75, fulvous or rufescent, apex fuscous or black, reaching apices of
mesocoxae, sometimes attaining apices of metacoxae. ANTENNAE: black; I, length
0.79-1.13, sometimes extensively rufescent; II, 2.20-3.35; III, 1.21-1.50; IV, 1.20-
1.55. PRONOTUM: length 0.91-1.25, anterior width 0.93-1.18, posterior width
1.50-2.10, subconical, broadly convex, somewhat flattened, transversely rugulose,
posterior margin straight, lateral margins slightly to mostly concave broadly rounded
in dorsal view, slightly to obviously sulcate dorsad of coxal cleft in lateral view; collar

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REVISION OF IRBISIA

257

wide; calli convex, rugulose, confluent anteriorly, broadly reaching anteriolateral
margin of pronotum, episterum sometimes with a fulvous spot. LEGS: testaceous,
fulvous or in some specimens rufescent; coxae fuscous preapically; trochanter with
fuscous spot; femora variously streaked and spotted with fuscous; apices and bases
of tibiae and tarsi black. VESTITURE: dorsum densely covered with appressed,
recurved, white sericeous setae; venter moderately covered with short white setae,
longer on midline.

Macropterous female. Similar in structure, color and vestiture, but with robust
stature; hemelytra strongly arcuate laterally. Length 6.20-8.20, width 2.63-3.20.
HEAD: width across eyes 1.38-1.56, vertex 0.70-0.85, dorsal width of eye 0.29-
0.34, distance from tylus to ventral margin of eye 0.93-1.15, height of eye 0.41-0.50,
maximum interocular width 0.98-1.18. ROSTRUM: length 2.30-2.85 reaching api-
ces of mesocoxae, sometimes attaining apices of metacoxae. ANTENNAE: I, length
0.80-1.08; II, 2.50-3.13; III, 1.3-1.58; IV, 1.28-1.55. PRONOTUM: length 0.95-
1.29, anterior width 0.98-1.28, maximum width 1.69-2.06.

Types. Originally described by Uhler from a series. The only specimen of this
series is a male with these label data: type M, P. R. Uhler Collection; Thyrillus
pacificus, Uhler, Snake Riv., Idaho; Thyrillus pacificus, Uhler, Snake Riv., Det. Uhler.;
USNM type no. 100950. Retained at the United States National Museum, Wash-
ington, D.C. I have examined this specimen (legs and antennal segment four are
absent) and as no other specimens of the syntype series has been located I have
designated it as the lectotype of Rhopalotomus pacificus.

Taxonomic consideration. Little variation in characters used to separate this species
from other species in the genus was noticed in this study. Both male and female
genitalic structures were uniform regardless of the collection locality of the specimens.

Remarks. Irbisia pacifica has the widest distributional range of any species in the
genus. It occurs at low to moderate elevations. The known northern limit of the
distribution is 50° 15' north latitude at Vernon, British Columbia, and the known
southern limit is 32° 45' north latitude at Lakeside, San Diego County, California.
The easternmost specimen was collected from Laramie County, Wyoming; this species
is extensively distributed in California. The majority of the distributional range is
occupied by the sagebrush steppe vegetational association. In British Columbia,
Idaho, Oregon and Washington the Palouse grassland associations (fescue- wheatgrass
and wheatgrass-bluegrass) provide the northernmost habitat. The grassland associ-
ations of the eastern portion of the range in Montana and Wyoming are foothills
prairie ( Agropyron-Festuca-Stipa ) and grama-needlegrass-wheatgrass respectively.
Within the Rocky Mountain forest ecoregion of Colorado and Utah, the juniper-
pinyon woodland and mountain mahogany-oak scrub vegetational associations pro-
vide I. pacifica habitats at slightly higher elevations. Oregon, in addition to the
previous habitats, possesses ponderosa shrub forest on the east side of the Cascade
Range and oak woodlands on the west side in which this species is found. Two
apparently isolated populations of I. pacifica are known from the Willamette Valley
of this state. In California, excluding the Mojave Desert, all the ecoregions have
yielded specimens of this species of Irbisia. The vegetational associations occupied
are: northern yellow pine forest, yellow pine-shrub forest, Oregon oak forest, mixed
evergreen forest, blue oak-digger pine forest, southern oak forest, chaparral, valley
oak, savanna and California prairie.

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I. pacifica has been collected in adult and immature stages on these grass hosts:
Agropyron cristatum (L.) Gaertn., A. desertorm (Fisch.) Schult., A. repens (L.) Beauv.,
A. smithii Rydb., A. spicatum (Pursh) Scribn. & Sm., A. trichophorum (Link) Richt.,
Avena sp., Bromus mollis L., Bromus sp., Elymus cinereus Scribn. & Merr., E.
condensatus Presl., E. triticoides Buckl., Festuca rubra L., and Hordeum sp. Adult
specimens have also been taken from these non-graminoid plants: Amsinckia sp.,
Cirsium cymosum (Greene) J. T. Howell, Medicago sativa L., Ranunculus sp., Rham-
nus sp., Rubus sp. and Wyethia sp. The altitudinal range is from 168 to 2377 m.
(550 to 11,000 feet) levation. The dates of occurrence are from 14 March to 11
August.

A single male specimen is known from Carnation, King County, Washington. I
have not collected material from this area, but would expect that the specimen (barring
a labeling error) was taken from a roadside, hedgerow or south facing hill situation.
This species is often pestiferous on forage grasses in the Intermountain states (Knowl-
ton, 1951; Haws, 1978).

Specimens examined. 3,512 specimens were examined from the following localities (Map
3). CANADA: BRITISH COLUMBIA: 5 mi. E Christina Lk. (UBC); Grand Forks (CNC);
Hedley, & 17 mi. W (UBD & CNC); Okanagan Falls (CNC); Oliver & Meyer Hat, Vaseaux
L., White R. (CNC, USNM); Osoyoos (CNC); Princeton (UBC); Summerland (CNC); Vernon
(CAS, UBC). UNITED STATES: CALIFORNIA: Alameda Co. (2 localities). Alpine Co. (2).
Amador Co. (2). Butte Co. (4). Cal\er as Co., Angels Cmp. (CAS). Colusa Co., Ramsey Cyn.
(UCD). Contra Costa Co. (6). El Dorado Co. (5). Fresno Co. (4). Inyo Co., 7 mi. N Parcher’s
Cmp. (CAS). Kern Co. (7). Lake Co. (1). Lassen Co. (8). Los Angeles Co. (6). Madera Co. (4).
Mariposa Co. Mariposa (UID). Mendocino Co. (7). Modoc Co. (8). Mono Co. (4). Monterey
Co. (7). Napa Co. (6). Nevada Co. (3). Orange Co. (2). Placer Co. (6). Plumas Co. (5). Riverside
Co. (9). Sacramento Co. (5). San Benito Co. (5). San Bernardino Co. (2). San Diego Co. (14).
San Francisco Co., Laguno Puerco (UCB). San Joaquin Co. (3). S.L.O. Co. (6). Sta. Barbara
Co. (2). Sta. Clara Co. (5). Shasta Co. (4). Sierra Co. (3). Siskiyou Co. (5). Solano Co. (2).
Sonomo Co. (3). Stanislaus Co. (2). Tehema Co. (2). Trinity Co. (6). Tulare Co. (5). Tuolumne
Co. (2). Ventura Co. (2). Yolo Co. (6). Yuba Co. (1). Most specimens are retained at CAS, F&A,
OSU, UCB, UCD & UCR. COLORADO: Chaffee Co., Buena Vista (CNC). Eagle Co., State
Bridge (CNC). Mesa Co., De Beque Cyn. (JTP). Moffat Co., Lay (CU); Maybell (KU). IDAHO:
Ada Co. (5). Adams Co. (1). Camas Co. (1). Canyon Co. (1). Caribou Co. (1). Cassia Co. (6).
Clearwater Co. (1). Elmore Co. (2). Franklin Co. (4). Fremont Co. (1). Idaho Co. (1). Jerome
Co. (1). Latah Co. (7). Lincoln Co. (3). Madison Co. (1). Nez Perce Co. (13). Oneida Co. (16).
Owyhee Co. (4). Payette Co. (1). Twin Falls Co. (5). Washington Co. (4). Most specimens are
retained at UID, USU & WSU. MONTANA: Gallatin Co., Bozeman (MSU). Jefferson Co.,
“Jefferson Co.” (MSU). Madison Co., Montana Expt. Stn. (MSU, USNM). Missoula Co., Mis-
soula (CAS); Montana Expt. Stn. (MSU). Park Co., Livingston; Park Co. (USNM). Sweet Grass
Co., Big Timber (MSU). Treasure Co., Hysham (MSU). Yellowstone Co., Huntley (MSU).
Beaver Crk. (KU). “Montana 46a” (USNM). NEVADA: Carson City, Carson City (CAS);
“Ormsby Co.” (CAS). Douglas Co., W Minden— Hwy. 19 (UCR). Elko Co., Carlin & 5, 10 mi.
W (KU, UCD); Lamorille Cyn. (UCD); Wells (USU). Eureka Co., Dumphy (UCD); Humboldt
R.-23 mi. W Carlin (UCB); Red House Rch. (UCD). Humboldt Co., Paradise Valley (UCD).
Lander Co., Austin (KU). Washoe Co., Reno (USNM); Sparks (CAS); Verdi, & 6 mi. E (UCB,
UCD, UCR). White Pine Co., Conners Pass (UCD); Ely, & 10 mi. S (KU & UCD); 17.5 mi.
N Hwy. 50— Steptoe Crk. Rd. (OSU). Nev. (USNM). OREGON: Baker Co. (3). Benton Co.,
Corvallis— Willamette Pk. (OSU). Deschutes Co. (3). Grant Co. (4). Harney Co. (7). Hood River
Co., Viente St. Pk. (OSU). Jackson Co. (5). Jefferson Co., 2.5 mi. NW Warm Spgs. (OSU).

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Josephine Co., 7 mi. S Grants Pass (OSU). Klamath Co. (13). Lake Co. (10). Linn Co., Lost
Prairie Cmpgd. (OSU). Malheur Co. (7). Sherman Co. (4). Umatilla Co. (3). Union Co. (3).
Union Co. (8). Wallowa Co. (4). Wasco Co. (3). Wheeler Co. (3). Yamhill Co., 6 mi. E Mc-
Minnville—Willamette R. (OSU). Most specimens are retained at OSU. UTAH: Beaver Co.
(1). Box Elder Co. (14). Cache Co. (18). Davis Co. (3). Iron Co. (2). Juab Co. (2). Kane Co.,
Long Valley (USNM, USU). Millard Co. (1). Rich Co. (1). Salt Lake Co. (2). Sanpete Co. (6).
Sevier Co. (2). Tooele Co. (4). Uintah Co. (1). Utah Co. (2). Wasatch Co. (1). Washington Co.,
Pinto (USNM). Weber Co. (3). Most specimens are retained at USU. WASHINGTON: Adams
Co. (1). Asotin Co. (3). Benton Co. (2). Garfield Co. (2). Grant Co. (3). King Co., Carnation
(OSU). Kittitas Co. (1). Klickitat Co. (1). Lincoln Co. (1). Okanogan Co. (2). Skamania Co. (1).
Spokane Co. (2). Walla Walla Co. (4). Whitman Co. (7). Yakima Co. (5). Most of the specimens
are retained at WSU. WYOMING: Albany Co., 23 mi. NE Bosler (KU). Carbon Co., Medicine
Bow Mts. (USNM). Laramie Co., 40 mi. NE Laramie (KU). Park Co., 27 mi. W Cody (KU).
Platte Co., Wheatland (UWY).

Irbisia panda Bliven
Figs. 30a, b, 53, 76, 99; Map 15

Irbisia panda Bliven, 1963, pp. 77-78, 86, pi. 8, figs. 4, 4a (new species).

Diagnosis. This species can be separated from other species of Irbisia by these
characters: impunctate pronotum; polished, shining black dorsum; light markings on
the ventral surface; sparse and short fuscous vestiture; relatively narrow vertex; black
antennae; mostly rufescent femora. I. panda and I. limata are superficially similar
but examination of the left paramere and vesica of the males reveals that they belong
in different species groups. In the former, the left paramere is very broadly curved
and the length of the shaft is small; the latter is more acutely curved with a longer
shaft. The sclerotized process of the vesica of I. panda is small, deflected to the left
of the trunk of the vesica, not expanded but preapically constricted, with the distal
portion sparsely covered with small spines. The process in I. limata is larger, expanded
on the left surface, not preapically constricted, and moderately covered with rows of
spines (which decrease in height basad). In addition the base of the clypeus of I.
panda is considerably more convex (projecting above the frons) than in I. limata.

Description. Macropterous male. Length 4.50-5.75, width 2.15-2.35, polished,
shining black, sparsely covered with short, fine recumbent fulvous or fuscous setae.
HEAD: width across eyes 1.04-1.23, vertex 0.43-0.50, dorsal width of eye 0.26-
0.31; triangular, polished; temporal areas indistinct; median depression a single,
slightly depressed line or lacking; basal carina very small, broadly rounded, in cross
section sometimes higher than collum, declivous to level of vertex, slightly posteriorly
arcuate mesad; eyes broadly joined to frons, straight in posterior view; distance from
tylus to ventral margin of eye 0.55-0.63, height of eye 0.48-0.55, maximum inter-
ocular width 0.70-0.81; jugal-loral junction testaceous to variable degree, clypeus
quite convex in lateral view, protruding above curvature of frons. ROSTRUM: length
1.83-2.05, piceous to fuscous, reaching apices of mesocoxae, sometimes surpassing
apices of metacoxae. ANTENNAE: fuscous to black; I, length 0.53-0.65, apex usually
testaceous; II, 1.50-1.79, base sometimes testaceous; III, 0.94-1.18; IV, 0.79-1.03.
PRONOTUM: length 1.03-1.20, anterior width 0.85-1.00, maximum width 1.70-
2.00; conical, slightly convex but with a flattened appearance, neither distinctly

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punctate nor rugulose, posterior margin arcuate, posterolateral margins greatly round-
ed, lateral margins straight, rounded at junction with propleura; anterior angles round-
ed, quite indistinct in dorsal view, not sulcate in lateral view; calli convex, polished,
broadly confluent anteriorly, reaching anterolateral margins of pronotum, with fov-
eate depression posteromedially of anterior angles; ventral third of propleura, pro-
sternum narrowly bordering coxae and attaining front margins, basalar plate and
posterior margin of epimeron ivory. LEGS: fuscous or piceous; coxae testaceous
apically; procoxae testaceous basally; femora rufescent medially, testaceous apically;
tibiae testaceous, piceous on bases and apices. VESTITURE: dorsum sparsely covered
with short, fine fulvous to fuscous setae, recumbent on hemelytra, erect and coarser
on head and lateral margins of pronotum; venter sparsely covered with suberect
fulvous setae.

Macropterous female. Stouter than male but similar in structure, color and ves-
titure. Length 5.10-6.15, width 2.40-2.75. HEAD: width across eyes 1.14-1.25,
vertex 0.50-0.56, dorsal width of eye 0.63-0.68, height of eye 0.50-0.54, maximum
interocular width 0.78-0.88. ROSTRUM: length 1.98-2.15, reaching bases of me-
tacoxae. ANTENNAE: I, length 0.60-0.70; II, 1.59-1.88; III, 1.00-1.13; IV, 0.81-
0.95. PRONOTUM: length 1.14-1.32, anterior width 0.90-1.08, maximum width
1.85-2.18.

Types. Male holotype. Rumsey, [Yolo Co.], Cal., V. 15.57; HOLOTYPE <5, Irbisia
panda, 1963 B. P. Bliven [red]; CAS type no. 13860. I have examined the holotype
and all twenty-four conspecific paratypes. These specimens are retained at the Cal-
ifornia Academy of Sciences, San Francisco.

Remarks. This species occupies the same vegetational association and provinces
as I. limata. Both species are sympatric from Calaveras County south, but the dis-
tribution of I. panda extends into Shasta County, the northern boundary of the blue
oak-digger pine forest. I. panda is also much more prevalent within the California
chapparal province than I. limata. Very few grass host records exist ( Avena sp.,
Bromus sp. and Vulpia sp.), perhaps indicating a diminished reliance on graminoids.
Adult specimens were collected from the following plants: Achillea sp., Amsinckia
sp., Asteraceae, Baccharis glutinosa Pers. Baeria sp., Brassica sp., Brodiaea laxa
(Benth. Wats.), Calothortus sp., Chaenactis sp., Clarkia Williamsonii (Dur. & Hilg.)
Lewis & Lewis, Cryptantha muricata (H. & A.) Nels. & Macbr., Delphinium sp.,
Eriodictyon californicum (H. & A.) Torr., Eriophyllum confertiflorum (DC.) Gray,
Fremontia calfornica Torr., Poaceae, Gilia capitata Sims, G. tricolor Benth., Lavia
sp., Lupinus densiflorus Benth., L. subvexus C. P. Sm. and Plagiobothrys sp. Adults
occur from 240 to 1220 m. (800 to 4000 feet) elevation and from 24 March to 30
May.

Specimens examined. 393 specimens were examined from the following localities (Map 15).
UNITED STATES: CALIFORNIA: Amador Co., Plymouth (F&A). Butte Co., 5 mi. NW
Oroville (UCB); Yankee Hill (UCB). Calaveras Co., Modelumne Hill (INHS, UCB); 4 mi. S
Railway Flat (UCB). Colusa Co., Rumsey Cyn. (UCD); Williams (UCD). Contra Costa Co.,
Mt. Diablo (UCB); Russelman Pk. (UCB). El Dorado Co., Greenwood (UCB); Nashville (UCD);
nr. Rescue (UCD); Riverton (UCB); Shingle (UCB); 6 mi. NW Shingle Spgs. (LACM). Fresno
Co., Coalinga, & 12 mi. W (CAS & UCB); Dunlap (LACM). Kern Co., Hobo Cmp.— Kern
River (UCB); Lebec (CAS); Miracle Spgs. (UCB); Tejon Pass (UCB); 1 mi. E Woody (UCB).
Lake Co., Clearlake Oaks, & 10 mi. E (F&A, UCD & UCB); N Frk. Cache Crk.-Hwy. 20

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(UCD). Madera Co., Coarsegold (UCB, UID); Oakhurst (UCB); Raymond (UCB); San Joaquin
Expt. Rng. (UCB). Mariposa Co., El Portal (F&A, UCB); 1.9 mi. W Mt. Mullion (UCB).
Monterey Co., 5 mi. NE Arroyo Seco Grd. Stn. (UCB); Horse Bridge— 1.5 mi. SW Arroyo Seco
Grd. Stn. (UCB). Napa Co., 4 mi. NW Benyessa (UCD); 10 mi. S & 20 mi. W Monticello Dam
(UCB); Samuel Spgs. (UCD). Nevada Co., Grass Valley (CAS); Nevada City. Placer Co., Auburn
& 10 mi. E (UCD & UCB); 5 mi. W Foresthill (UCD). Riverside Co., Riverside (UCD).
Sacramento Co., Folsom (F&A). San Benito Co., Limekiln Cyn.— SW Paicines (UCB); W entr.
Pinnacles Nat. Mon. (OSU). San Joaquin Co., Lone Tree Cyn. (F&A). Shasta Co., Redding
(WSU). Solano Co., nr. Dozier— 11 mi. S Dixon (UCD); 8 mi. W Winters (UCD). Sonoma
Co., Mark West Spgs. (CAS). S.L.O. Co., Adelaida— 15 mi. W Paso Robles (OSU); La Panza
Cmp.— 12 mi. NE Pozo (UCB, UCR); Nacimiento Dam (UCB). Sta. Barbara Co., Figueroa
Mt. (LACM). Stanislaus Co., Del Puerto Cyn. — Frank Raines Pk. (CAS, UCB, UCD). Sutter
Co., Maryville Butte (CAS, UCB). Tehema Co., 21 mi. NW Red Bluff— rt. 36 (OSU). Tulare
Co., Ash Mt. H.Q. (AMNH, UCB); 4 mi. N Kaweah (UCB); 10.3 mi. N & Lemoncove (AMNH
& CAS, UCB); 3 mi. NE Springville (UCR); 4.5 mi. S & 8 mi. NE Three Rivers (UCR, USU).
Tuolumne Co., Buck Mdws. (USNM); Jamestown (UCB); N. Frk. Tuolumne River— 3 mi. NE
Tuolumne (CAS). Yolo Co., Cache Crk. Cyn. (UCD); Putah Cyn. (UCD); 2 mi. NW Rumsey
(UCB).

Irbisia sericans (Stal)

Figs. 7c, 12a, b, 35, 58, 81; Map 10

Leptomerocoris sericans Stal, 1858, p. 188 (new species). Walker, 1873, p. 144.
Irbisia sericans: Reuter, 1875b, p. 548 (new combination); 1879a, p. 58; 1896, pp.
12-13, pi. 1, figs. 4a, 4b. Bergroth, 1885 p. 270. Sahlberg, 1885, pp. 68, 70.
Atkinson, 1890, p. 125. Ashmead, 1898, p. 341 (incorrectly cited as McAtee, 1899
in Carvalho, 1959). Heidemann, 1900, p. 504. Breddin, 1902, p. 540. Kirkaldy,
1906, p. 142. Oshanin, 1910, p. 760. Van Duzee, 1916b, p. 38; 1917a, p. 325;
1921a, pp. 145, 146, 151; 1921b, p. 193. Knight, 1921, p. 110. McAtee, 1923, p.
145. Essig, 1926, p. 361. Kiritshenko, 1926, p. 16, pi. 1, fig. 2. Downes, 1927, p.
12. Lindberg, 1927, p. 22. Slater, 1950, pp. 36, 71, pi. 2, figs. 16, 17, p. 75, pi. 4,
fig. 7. Carvalho, 1952, p. 89; 1959, pp. 105-106. Kelton, 1959, pp. 18, 60, figs.
27a-c. McKendrick and Bleicher, 1980, pp. 15-18, figs. 1-4.

Diagnosis. I. sericans, I. californica and I. sita can be separated from the remainder
of the species of Irbisia on the basis of the structure of the male genitalia. These
species are distinguished by the left paramere which is large, with a prominent sensory
lobe which projects above the surface of the arm, abruptly angled, without tubercles
on the outer surface, with a long preapically constricted shaft, and abruptly truncated
apex. I. sericans is allopatric with the other species, occurring as far south as Eugene,
Lane County, Oregon. /. sericans can be separated from the allied species by its dull
bronze luster, trapezoidal pronotum, with straight lateral margins, and transversely
rugose disk; black femora; densely distributed, shining, woolly, recurved vestiture;
series of minute spicules on the sclerotized process of the male vesica.

Description— large form. Macropterous male. Length 5.85-6.50, width 2.40-2.60,
dull black with shagreened luster, covered with shining gold, pale yellow or white
setae. HEAD: length 0.63-0.70, width across eyes 1.23-1.41, vertex 0.55-0.68, dorsal
width of eye 0.38; shagreened, impunctate; basal carina moderately developed, nar-
rowly rounded in cross section, and slightly posteriorly arcuate mesad; temporal areas

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distinct, glabrous, with transverse sulcus on posterior margin; distance from tylus to
ventral margin of eye 0.68-0.74, height of eye 0.54, interocular width maximum
0.88-0.95. ROSTRUM: length 2.09-2.23, black, reaching beyond apices of meso-
coxae approaching but not attaining metacoxae. ANTENNAE: black; I, length 0.73-
0.93; II, 1.85-2.23; III, 1.08-1.40; IV, 1.38-1.50. PRONOTUM: length 0.93-1.15;
anterior width 0.93-1.08; maximum width 1.73-1.96; disk strongly convex, trans-
versely rugose without distinct punctures, posterior margin broadly concave medially,
lateral margin broadly concave in dorsal view, rounded at junction with propleura;
calli slightly confluent anteriorly, asperate; anterior angles not greatly produced, but
evident in dorsal and lateral view. VESTITURE: with densely distributed suberect,
fine golden setae; setae mostly woolly on pronotal disk; venter with shining golden
setae. LEGS: black; tibiae fulvous with bases and apices black.

Brachypterous female. More robust than male, but very similar in color, structure
and vestiture. Length 5.20-6.20, width 2.60-3.15. HEAD: length 0.63-0.88, width
across eyes 1.33-1.53, vertex 0.65-0.73, dorsal width of eye 0.39, distance from tylus
to ventral margin of eye 0.75-0.88, height of eye 0.51-0.60; maximum interocular
width 0.95-1.05. ROSTRUM: length 2.20-2.41, fulvous, barely reaching apices of
metacoxae. ANTENNAE: I, 0.70-0.93; II, 1.80-2.40; III, 1.05-1.53; IV, 1.30-1.80.
PRONOTUM: length 0.90-1.11, anterior width 1.05-1.23, maximum width 1. 73-
LOO, membrane reduced, extending very slightly beyond cuneus, cells developed,
apex of abdomen exposed.

Description— small form. Brachypterous male. Measurements are less than or equal
to the diminutive values of the male large form, but very similar in color, structure
and vestiture. Length 4.70-5.00, width 2.20-2.30. HEAD: length 0.64-0.70, width
across eyes 1.19-1.28, vertex 0.55-0.58, dorsal width of eye 0.34, distance from tylus
to ventral margin of eye 0.63-0.68, height of eye 0.50, maximum interocular width
0.85. ROSTRUM: length 1.95-2.05, black, barely reaching apices of metacoxae.
ANTENNAE: I, 0.63-0.70; II, 1.65-1.78; III, 0.90-0.93; IV, 0.95. PRONOTUM:
length 0.83-0.88, anterior width 0.88, maximum width 1.51. Membrane reduced,
extending slightly beyond cuneus, cells developed, apex of abdomen exposed.

Brachypterous female. Measurements are less than or equal to the diminutive
values of the female of the large form, but very similar in color, structure and vestiture.
Length 4.80-5.55, width 2.43-2.63. HEAD: length 0.60-0.70, width across eyes 1.25-
1.28, vertex 0.60-0.68, dorsal width of eye 0.30-0.35; distance from tylus to ventral
margin of eye 0.70, height of eye 0.50-0.53, maximum interocular width 0.93-0.95.
ROSTRUM: length 1.98-2.18, infuscated fulvous, reaching or surpassing apices of
metacoxae. ANTENNAE: I, 0.65; II, 1.60-1.63; III, 0.85-0.88; IV, 1.00 PRONO-
TUM: length 0.80-0.91; anterior width 0.95, maximum width 1.63-1.65.

Types. This species was described from a pair of specimens from Sitka, on Baranof
Island, Alaska. These are specimens of the large form. I have not examined the
holotype but have before me a pair of topotypic specimens collected during the
Harriman Alaska Expedition of 1899 by T. Kincaid. These specimens are retained
in the collection of the United States National Museum, Washington, D.C. The St&l
types are likely to be in the collection of the Naturhistoriska Riksmuseet, Stockholm,
Sweden.

Remarks. The combined distribution of both forms of I. sericans is strictly humid
coastal Vancouveran. The Kamachatka Peninsula and Commander Islands of the

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Soviet Union delimit the western boundary of this species (and the genus). Hood
River and Eugene, Oregon are the eastern and southern boundary of the known
distribution respectively. Adult and immature specimens were collected from these
grass hosts: Bromus carinatus H. & A., B. mollis L., B. rigidus Roth, Calamagrostis
canadensis (Michr.) Beauv., Festuca arundinacea Schreb., F. rubra L., Holcus lanatus
L. and Poa pratensis L. Specimens were collected on Angelica sp. and Rubus sp.

The only qualitative study concerning utilization of grasses by this genus focuses
on /. sericans and C. canadensis (McKendrick and Bleicher, 1980). Preliminary
results indicate that the individual grass blades with the greatest feeding damage were
those blades with low levels of insoluble silica. Childs (1914) and Essig (1915) reported
the occurrence of this species from Santa Clara County, California. I have examined
these specimens and re-identified them as I. californica. Collection dates for speci-
mens from the westernmost portion of the range of the small form are from 1 1 July
to 3 September. Large form specimens from Oregon were collected from 29 April to
21 June. Specimens were collected within an altitudinal range from sea level to 150
m. (500 feet). The lack of specimens of this species (and all other Irbisia species)
from western Washington (particularly the Olympic penisula) probably indicates a
lack of collecting effort rather than a lack of appropriate habitat.

I. sericans has a puzzling distribution. Within the material from the Canadian
National Collection are four specimens of the small form from Quebec, Gatineau
Park— Harrington Lake. The specimens were collected on 28 May 1958. The habitat
surrounding the collecting site is coniferous and at an elevation of approximtely 152
m. (500 feet). Researchers at the Canadian National Collection, Ottawa who were
on that particular expedition relate that the locality, at the time of collection, was a
recently abandoned mink farm with a small clearing around it. Thirty years to the
day, with the permission of the Royal Canadian Mounted Police (the adjoining
property is the summer weekend residency for the Canadian Prime minister) L. A.
Kelton and I intensively swept the grasses on the now overgrown plot and did not
find any I. sericans ; specimens of Stenodemini were abundant.

The period of occurrence, vegetational association and elevation appears similar
to the conditions encountered in the western and main distribution of this species.
There are no known intervening locality records between Quebec and the Pacific
Northwest. If the collection record is accurate, the population at Harrington Lake is
disjunct from the nearest known locality by 3,800 km. (2,900 miles). A recent in-
troduction would explain the disjunction, as Gatineau Park is adjacent to Ottawa,
Ontario. However, the brachypterous form of I. sericans is most prevalent on the
Aleutian Peninsula and Islands which make an introduction a suspect scenario.
Additional collection records may reveal that the distribution of this species is trans-
continental at northern latitudes.

Specimens examined. 677 specimens of the large form were examined from the following
localities (Map 10). CANADA: BRITISH COLUMBIA: Cowichnan Bay (CNC); Hevenor In-
let—Pitt Is. (UBC); Kelsey Bay— Vancouver Is. (UBC); Miracle Beach— nr. Oyster R. (CNC);
Prince Rupert (AMNH, UBC) & Kaien (UBC); 0.7 mi. S Sayward— Vancouver Is. (OSU); Sugar
Lk. — 22 mi. SW Ocean Falls (UCD); Queen Charlotte Is.— Anthony Is., Burnaby Is., Graham
Is. [Jungle Beach (UBC), Massett (CNC, UBC), Tlell (CNC)], House Is., Kunghit Is., Louis Is.,
Moresby Is. [Sandpit (AMNH), Slim Inlet (UBC)] all UBC; Tofino (UBC); Ucluelet (CNC);
Vancouver (CAS, CU, USNM). UNITED STATES: ALASKA: Anton Larsen Bay-Kodiak

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Is. (USNM); Atica Is. (CAS); Belkofski (USNM); Douglas (USNM); Dutch Harbor (CAS, USNM);
Eagle River (USNM); Edna Bay— Dali Is. (CAS); Fox Point (USNM); Girdwook (USNM);
Glacier River— Unalaska Is. (CAS); 6 mi. NW Granite Crk. Cmpgd. — Kenai Pen. (KU); Homer
(OSU); Iliuliuk— Unalaska Is. (USNM); Johnson Pass— Kenai Pen. (KU); Juneau & 1 1 mi. N
(CAS, USNM & OSU); Katmai (TA&M, USNM); Ketchikan (UBC, UCD); King Salmon -
Naknek R. (CNC); Kodiak (USNM); Kukak Bay (USNM); Lk. Karluk (USNM); Little Sitkin
(USNM); McNeil River Cove & Moose Pass— Kenai Pen. (OSU & CNC); Naknek (CNC);
Nazan Bay— Atka Is. (CAS, F&A); Nunivak Is. (USNM); Ohison Mt. Rd. — Homer (USDA);
Olga Bay -Kodiak Is. (OSU); Seward (CAS, CNC); Sitka (UBC, USNM); Skilak Lk.- Kenai
Pen. (CNC); Thome Bay & 20 mi. W— P.O.W. Is. (OSU); Trap Bay— nr. Tenakee Inlet (Chkha-
gof Is.) (OSU); Valdez (USNM); Virgins Bay (USNM); Wrangell (CAS, USNM). OREGON:
Benton Co., Corvallis (CAS, OSU, TA&M, UBC, USNM); Findley Nat. Wldf. Rfg. (OSU);
Mary’s Peak (OSU); Mary’s River (OSU); McDonald St. For. (OSU); Monroe (OSDA); 4 mi.
NW Philomath (OSU). Clackamas Co., Barton (AMNH); 3 mi. SE Milwaukie (OSU). Hood
River Co., Hood River (CU, USNM). Lane Co., Good Pasture— Eugene (OSDA, OSU). Lincoln
Co., Silez (USNM). Linn Co., Shedd (USNM). Multnomah Co., Sauvie Is. (OSU). Siuslaw Co.,
10 mi. W Alsea (USNM). Tillamook Co., 0.1 mi. N Rockway (OSU). Washington Co., Dilley
(USNM). Yamhill Co., McMinnville (OSU); Muddy Valley (OSU); Yamhill River- Mc-
Minnville (OSU). WASHINGTON: Grays Harbor Co., Montesano (CNC, UCB, USNM).

327 specimens of the small form were examined from the following localities: CANADA:
BRITISH COLUMBIA: Tofino (UBC); Rankin Is. (UBC); Triangle Is. (UBC). UNION OF
SOVIET SOCIALIST REPUBLICS: Copper Is. [O. Mednyy] (USNM); Glenka [Glinka] -
Copper Is. (USNM); Karaginski [O. Karaginskiy] (USNM); Lissonkoraja [Lisinskaya Bukhta] —
Bering Is. (USNM); Nikolaski [Nikol’skoye] — Bering Is. (USNM); Petropaulski [Petropav-
lovsk-Kamchatsky]- Kamchatka Pen. (USNM). UNITED STATES: ALASKA: Adak Is. (CAS);
Andrew Lagoon— Adak Is. (USNM); Casco— Attu Is. (USNM); Cold Bay & Russel Crk.—
Aleutian Pen. (CNC, USNM); Finger Crk.— Adak Is. (USNM); Makushin Bay— Unalaska Is.
(CAS); Mount Tulik & Okmok-Umnak Is. (USNM & CNC); Popof Is. (USNM); Pyramid
Cove -Attu Is. (USNM); St. George Is. (CAS, OSU, USNM); St. Matthew Is. (CNC); St. Paul
Is. (AMNH, CAS); Terrible Mt.-Attu Is. (USNM); Thome Bay-P.O.W. Is. (OSU); Umuak
Is. (USNM); Unalakleet (CNC); Unga Is. (OSU).

Irbisia serrata Bliven
Figs. 23a, b, 46, 69, 92; Map 20

Capsus solani: Heidemann, 1910, pp. 200-201 (not holotype).

Irbisia brachycerus [<2]: Cockerell, 1910, p. 370. Vosler, 1913, p. 553 (in part). Van
Duzee, 1914, p. 24 (in part); 1921a, pp. 149, 152.

Irbisia mollipes: Van Duzee, 1921a, pp. 147-148 (in part).

Irbisia inurbana: Bliven, 1963, p. 81-82 (in part, not holotype).

Irbisia serrata Bliven, 1963, pp. 82-83, 86, pi. 8, figs. 8, 8a (new species).

Diagnosis. I. serrata is distinguished from allied species by a combination of
characters. It possesses a membranous sac which is medially attached to the left outer
surface of the sclerotized process of the vesica of the male. This character is unique
to the genus. The structure of the female genitalic characters is not diagnostic. The
following characters of I. serrata will separate it from I. sericans : pronotum with
straight lateral margins, confluent punctation, and prominent anterior angles, in
dorsal and lateral view; shorter antenna; macropterous female; vastly different male
parameres. I. serrata is distinguished from I. solani by: the prominent anterior angles

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265

of the pronotum, mostly shagreened dorsal surface of the head, generally more densely
distributed longer vestiture, strongly spined shaft of the left paramere of the male.
7. shulli Knight has a wider vertex and brachypterous female which distinguish this
species from 7. serrata. The preponderance of black color and flattened pronotal disk
of 7. nigripes serve to differentiate it from 7. serrata. I. serrata is distinguished from
7. fuscipubescens by the former’s densely distributed much longer vestiture, mostly
shagreened dorsal surface, and confluently punctate pronotum. 7. serrata is separated
from 7. cascadia by the former’s longer first antennal segment, rugulopunctate prono-
tum and longer vestiture.

Description. Macropterous male. Length 5. 10-5.80, width 2.30-2.65, shining black,
densely clothed with upright white to fulvous setae intermixed, in varying degree,
with recurved sericeous white setae. HEAD: width across eyes 1.10-1.24, vertex
0.45-0.59, dorsal width of eye 0.30-0.35; triangular, shagreened or slightly asperate;
temporal areas indistinct, basal carina produced, slightly declivous to level of vertex
posteriorly arcuate mesad; median depression indistinct, eyes broadly joined to frons;
posterior margin of head (including eyes) arcuate posterolaterally in dorsal view;
distance from tylus to ventral margin of eye 0.53-0.63, height of eye 0.48-0.55,
maximum interocular width 0.79-0.90. ROSTRUM: length 1.53-1.95, fuscous to
black, barely reaching to slightly surpassing mesocoxae. ANTENNAE: black; I, length
0.51-0.68, apex and base sometimes fulvous; II, 1.45-1.85; III, 0.73-1.05; IV, 1.00-
1.23. PRONOTUM: length 0.97-1.20; anterior width 0.88-1.03; maximum width
1.66-1.95; trapezoidal, disk broadly convex, scabrous and confluently punctate, pos-
terior margin broadly arcuate, lateral margins straight, margined at junction with
propleura, anterior angles, prominent, quite evident in lateral view, pronotal disk
“shelf-like” dorsad of coxal cleft; calli slightly flattened, evident but only slightly
produced, confluent anteriorly, smooth or asperate. LEGS: black; tibiae dark fuscous,
sometimes testaceous to fulvous, medially. VESTITURE: head, pronotum and scu-
tellum with upright shining white and/or fulvous setae, which become suberect or
recumbent on hemelytra; dorsum covered with recurved sericeous white setae; venter
with dense, thin light setae.

Macropterous female. More robust than male; however, similar in color, structure
and vestiture. Length 5.10-5.90, width 2.30-3.00. HEAD: width across eyes l.lS-
l^S, vertex 0.55-0.63, dorsal width of eye 0.30-0.35, distance from tylus to ventral
margin of eye 0.57-0.65, height of eye 0.49-0.56, maximum interocular width 0.88-
0.93. ROSTRUM: length 1.76-1.90, either reaching meso or metacoxae. ANTEN-
NAE: I, length 0.54-0.63; II, 1 .4 1-1 .73; III, 0.75-1 .00; IV, 1 . 1 0-1.23. PRONOTUM:
length 1.10-1.25, anterior width 0.98-1.08, maximum width 1.90-2.10. LEGS: fus-
cous or rufescent, coxae black, apices testaceous, trochanter testaceous, femora with
fuscous spots ventrally on some specimens, tibiae fulvous, bases and apices black,
tarsi black.

Types. Male holotype. Laribee Valley, Humboldt Co., Cal. IV.26.36; B. P. Bliven,
Collector, No. 615; HOLOTYPE 6 Irbisia serrata, 1963 [red]; CAS type no. 13863.
I have examined the holotype and all of the paratypes. Twelve paratypes are con-
specific with the holotype, four are misidentified and are specimens of 7. solani. All
of this material is retained at the California Academy of Sciences, San Francisco.

Taxonomic consideration. Heidemann ( 1 9 1 0), in the original description of Capsus

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solani, included within the distributional range of this species specimens from the
Wasatch Mountains, Utah. It is my opinion that these specimens are not conspecific
with 7. solani, but are misidentified specimens of I. serrata. No specimens, of the
material on hand, of 7. solani occur at or near this locality, whereas the locality is
within the distributional range of 7. serrata.

Van Duzee (1914) included several specimens of 7. serrata Bliven from Colorado,
New Mexico and Utah within his treatment of 7. brachycera. This misidentification
was apparently due to his confusion of the habitus of 7. brachycera. Van Duzee
(1921a) received specimens of 7. brachycera and the not yet described 7. serrata from
Sunset, Colorado. He mistakenly identified the specimens of 7. serrata as 7. brachycera
and named a new species 7. arcuata , for the true 7. brachycera specimens. Knight
(1941) made 7. arcuata the junior synonym of 7. brachycera after he examined the
type specimen of the latter species. Knight took this action after he had misidentified
many specimens of 7. brachycera as 7. arcuata, apparently following the example of
Van Duzee. I have examined specimens of 7. serrata identified as 7. brachycera by
both Van Duzee and Knight.

It is interesting that Knight did not describe those specimens of the not yet described
7. serrata as a new species, at the time he synonymized 7. arcuata, in light of the fact
that he had specimens of 7. serrata, with red marks on the labels (indicating new
species) in his collection.

Van Duzee (1921a) included specimens from the Las Vegas Range, New Mexico
in his treatment of 7. mollipes. These specimens are correctly identified as 7. serrata.
In all cases, I have added my determination labels to these critical specimens.

I have cited Cockerell (1910) in 7. serrata because he states that the identification
was made by Van Duzee. The specimens of concern were taken at Steamboat Springs,
Colorado. This locality is within the range of 7. serrata, although I have no topotypic
material. However, there is a good series of 7. shulli from Steamboat Springs. The
separation of the males of these two species can be difficult and the dissection and
examination of the genitalia are required for positive identification.

Remarks. I. serrata has caused great difficulty in the identification of the species
of Irbisia because of its very great distributional range, sympatry with nine other
species of Irbisia, and the relative obscurity of its original description (which did not
contain a key to the species).

7. serrata has one of the widest distributional ranges in the genus. It occurs as far
south as the Upper Gila Mountain forest province of Arizona inhabiting open sit-
uations in several floristic associations: Arizona pine forest and a transition zone
between oak-juniper woodland and mountain mahogany-oak scrub. Within the Rocky
Mountain forest province seven forest associations contain open areas providing
suitable habitat for 7. serrata', these associations are: western ponderosa, Douglas fir,
grand fir-Douglas fir, western spruce-fir, pine-Douglas fir, Arizona pine and south-
western spruce-fir. Many of these forest associations intergrade with sagebrush steppe
and juniper-pinyon woodland associations in Colorado, Idaho, New Mexico, Oregon
and Utah. East of the Cascade Range, ringing the Columbia Basin, in Oregon and
Washington fescue-wheatgrass and wheatgrass-bluegrass grasslands also provide 7.
serrata with habitat. West of the Cascade Range in the Pacific forest province openings
are found within cedar-hemlock-Douglas fir and silver fir-Douglas fir forests of
Oregon and Washington. The Sierran forest province in the Klamath Mountains of

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REVISION OF IRBISIA

267

northern California is essentially the southern limit of the distibution in this state;
a few specimens are known from the black oak-digger pine forests of Monterey and
Napa Counties.

Adult and immature specimens have been collected on these grass hosts: Agropyron
spicatum (Pursh) Scribn. & Sm., Dactylis glomerata L., Elymus glaucus Buckl., E.
triticoides Buckl., Festuca arundinacea Schreb., F. rubra L., Poa pratensis L. and
Sitanion hystrix (Nutt.) J. G. Sm. Both adults and immature stages were observed
feeding on these non-graminoid monocots: Allium cernuum Roth, A. crenulatum
Wieg., Brodiaea sp. and Iris tenax Dougl. Several other plants have had adults
collected from them: Abies procera Rehd., Alnus sp., Balsamorhiza sagittata (Pursh)
Nutt., Balsamorhiza sp., Ceanothus integerrimus H. & A., C. velutimus Dougl. ex
Hook, Cercocarpus ledifolius Nutt., Eriodictyon californica (H. & A.) Torr., Juniperus
sp., Lomatium dissectium (Nutt.) Math. & Const., Lupinus sp., Physocarpus capitatus
(Pursh) Kuntze, Picea sp., Pinus flexilis James, P. ponderosa Dougl. ex P. & C.
Lawson, Polygonum bistortoides (Pursh), Prunus pensylvanica L., Psuedotsuga men-
ziesii (Mirbel) Franco, Quercus breweri (Engelm. in Wats.) Jeps., Q. gambelii Nutt.,
Q. garryana Dougl., Q. turbinella Greene, Ribes cereum Dougl., R. roezlii Regel,
Salix sp., Spiraea sp. and Vicia sp. The altitudinal range of I. serrata is 15 m. (49
feet) at Puyallup, Washington and 3660 m. (12,000 feet) at Silverton, Colorado. Its
period of occurrence is 15 March at Globe, Arizona and 27 August at Blackwall,
British Columbia.

Specimens examined. 972 specimens were examined from the following localities: CANADA:
BRITISH COLUMBIA: Blackwell-Manning Pk. (UBC); Kushanook (CNC); Lytton (UBC);
North Bend (USNM); Madden L. & McIntyre Crk.- Oliver (CNC); 10 mi. W & Oliver (CNC);
Osoyoos— Anachrist Mt. (CNC); Salked Flats— nr. Inkitsaph (UBC); Skihist Cmp. — Fraser R.
(UCB); 10 mi. W Summerland (CNC). UNITED STATES: ARIZONA: Apache Co., Big Lk.-
Apache Nat. For. (CNC). Coconino Co., 9 mi. S Flagstaff (NAU); Yavapai Pt.— Grand Canyon
Nat. Pk. (USNM), Grand Cyn.- Grand Cyn. Nat. Pk. (CAS), Grand View Lookout (AMNH).
Gila Co., Globe (OSU); 1 mi. W Miami (UCR); Parker Crk. -Sierra Ancha Mts. (USNM); nr.
Roosevelt lk. (USNM). Maricopa Co., Reavis Rnch., trl. (JTP). Navajo Co., Lewis Cyn.— 2 mi.
S Pinedale (AMNH); 1 5-20 mi. SW Show Low (AMNH). CALIFORNIA: Del Norte Co., mp.
8.25 rt. 199— Middle Fort of Smith River. Humboldt Co., Blair’s Rch.— Redwood Crk; Dins-
mores (CAS); Iaqua (CAS); Laribee Valley (CAS). Lake Co., Blue Lakes (UCD). Lassen Co.,
Susanville (CNC). Marin Co., 4 mi. E Stinson Beach (AMNH). Mariposa Co., Clouds Rest—
Yosemite Nat. Pk. (LACM); Yosemite (UCB). Mendocino Co., 7 mi. W Eel River Rgr. Stn.
(UCB); Hopland (UCD); Latonville (CAS). Modoc Co., 1 mi. NW Ft. Bidwell (UCB). Monterey
Co., Jolon (UCD). Napa Co., Castle Crag (F&A); 5 mi. S Pope Valley (UCD). Placer Co., 5
mi. NE Auburn (WSU). Sacramento Co., 5 mi. N Folsom (F&A). Shasta Co., Castle Crag St.
Pk. (AMNH). Siskiyou Co., Ash Crk. Rgr. Stn. -9 mi. E McCloud (UCB); 1 mi. SE Bartle
(UCB); Caribou Mt. -Trinity Alps (UCB); Young’s Valley (PUC). Sta. Clara Co., 3 mi. W.
New Almenden (UCB). Trinity Co., Buttercreek Mdw. — 8 mi. W Hayfork (UBC); Carrville
(CAS); Coffee Crk. Rch. (UCB); Scott Mt. Mdw. -6 mi. E Calahan (OSU); Scott Mt. Peak
(UCB); South Fork Pass (UID). COLORADO: Boulder Co., Boulder, 4.5 mi. N & Flagstaff
Cyn. (CNC); Sunset (CAS). Clear Creek Co., Chicago Crk. (CNC). Conejos Co., Trujillo Mdw.
Cmp. — 3 mi. N Cumbres (AMNH). Custer Co., Davenport Cmp. 36 mi. S Florence (AMNH).
Dolores Co., Lizard Head Pass (USNM). Garfield Co., 2 mi. S Douglas Pass (AMNH); Glenwood
(USNM). Grand Co., Rabbit Ears Pass (USNM). Gunnison Co., Monarch Pass (USNM). Hins-
dale Co., Lk. City (KU). La Plata Co., La Plata (CNC); Rockwood (KU); 5 mi. N Rockwood
(OSU). Larimer Co., nr. Estes Pk. -Rocky Mt. Nat. Pk. (CNC, USNM); Ft. Collins (SHF). Las

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Animas Co., Cucharas Pass (CNC); Spanish Peaks (USNM). Mesa Co., East Fork Big Crk.—
10 mi. S Collbran (USNM); Mesa & 8 mi. S (USNM). Mineral Co., 16 mi. N Pagosa Spgs.
(TA&M). Montezuma Co., Mesa Verde Nat. Pk. (UCD). Montrose Co., 2 mi. S Columbine
Pass (JTP). Ouray Co., Summit Road— Ouray (AMNH); Ouray (AMNH). Routt Co., Meadows
(CNC). San Juan Co., Silverton (USNM). San Miguel Co., Comet Crk.— Telluride (AMNH);
Trout Lk. (USNM). Glen wood (USNM); Rices Spur (USNM). IDAHO: Adams Co., 7.5 & 8
mi. N Council (OSU & UID). Boise Co., 3 mi. N Idaho City (UID). Cassia Co., City of Rocks
(UID); Elba-Basin Pass (UID). Elmore Co., 13 mi. W Hill City (MSU). Nez Perce Co., Lenore
(USNM). Oneida Co., Black Pine Cyn. (USU); Holbrook (USU). NEVADA: Humboldt Co.,
Lye Crk.— Sta. Rosa Mts. (OSU). NEW MEXICO: Colfax Co., Cimarron Cyn. (USNM). Rio
Arriba Co., NE Truchas Pk. (USNM). Santa Fe Co., Aspen Ranch (USNM); Tesuque (USNM).
Sandoval Co., Jemez Spgs. (LACM, KU); Sandia Mts. (USNM). San Miguel Co., Holy Ghost
Cyn.— 14 mi. N Pecos (UALB); Mosimann Rch.— 15.7 mi. W Sapello (UALB); Panchuella
Cyn.— Cowles (USNM); “Top of Las Vegas Range,” (CAS). Taos Co., Hondo Cyn. — 7 mi. E
Valdez (USNM). Torrance Co., Estania (KU); Tajique (KU). OREGON: Baker Co., 5.5 mi.
SW Halfway (OSU), 14 mi. E Hereford (OSU). Benton Co., Alsea (USNM); Fish Hatchery,
Oak Crk. - both nr. Corvallis (OSU); Corvallis (OSU, UCB); MacDonald St. For. (OSU); Mary’s
Peak (T12S R7W Sec. 20 SE & SW 'A) (AMNH, OSU); Rock Crk. -5 mi. W Philomath (OSU).
Clatsop Co., Saddle Mt. St. Pk.— trail. (AMNH, OSDA, OSU). Crook Co., Mark’s Crk.— Crook
City (OSDA, OSU); 20 mi. E Prineville (OSU); 9.5 mi. E Post (OSU). Deschutes Co., Indian
Ford Crk. — 6 mi. W Sisters (OSU); Indian Ford Rd. — 2 mi. NE Sisters (T14S R10E Sec. 27)
(OSU); 4.2 & 14 mi. S Millican (AMNH, OSU); Pine Mt. Observatory (AMNH, OSU); RW
Sawyer St. Pk.— 1 mi. NW Bend (OSU). Douglas Co., Bradley Crk— nr. Glide (OSU); Grant
Co., 8 & 11 mi. N Seneca (OSU). Hood River Co., Dimmock St. Pk. — 2 mi. SW Parkdale
(OSU). Jackson Co., Applegate Rd. (OSU); Ashland (SOS); Colestin (OSDA); Medford (OSU);
Summit Green Spgs. Hwy. (OSU); 5 mi. W & Pinehurst (OSU); 3 mi. E Sams Valley (OSU);
Siskiyou Pass (CNC, OSU); 0.5 mi. S Siskiyou Smt. (old rt. 99) (AMNH, OSU); Tubb Spgs.—
6 mi. W Pinehurst (OSU). Josephine Co., Rough & Ready Botanical Wysd. (OSU); Woodcock
Crk. — 3 mi. SW Cave Jet. (T39S R5E Sec. 31) (OSU). Klamath Co., 11 mi. E & 15 mi. NW
Bly (OSU); Bly Mt. - 10 mi. W Beatty (OSU); 7 & 10 mi. W, 1 2 mi. SW Keno (OSU & AMNH);
Gerry Ranch — 1 2 mi. NW Klamath Falls (OSU); Spencer Crk. (OSU); Sprague River Pk. (OSU).
Lake Co., Chandler St. Pk. (OSU); Fremont Nat. For. (CAS); Quartz Crk. — 16 mi. E Bly (OSU).
Lincoln Co., Little Grass Mt. — 20 mi. N rt. 20 (OSU). Polk Co., Dalles (USNM); Grande Ronde
(OSU). Lane/Linn Cos., H. J. Andrews Expt. For.— 11 mi. N Blue River (T15S R5E Sec. 28
NW & SW y4) (All OSU); McKenzie Pass (USNM). Linn Co., Iron Mt. Trl. (OSU); Lost Prairie
(OSU); Peterson Butte (OSU); Roaring R. Fish Hatchery (USNM); Tombstone Prairie (OSU).
Umatilla Co., 2.2 mi. N Dale (OSU); Tollgate (OSU). Union Co., 12 mi. NE LaGrange (OSU).
Wallowa Co., Wallowa Lk. (CAS). Wasco Co., Bear Spgs. — 25 mi. W Maupin (OSU); Cherry
Hgts. Rd. — 5 mi. S The Dalles (OSU); 5.5 mi. E Mosier (OSU); 2 mi. S & 7 mi. N Simnasho
(OSU); The Dalles (CU); Wapinta (OSU). Wheeler Co., 4 mi. SW Fossil (OSU); 4.5 mi. S
Mitchell (OSU); Slide Mt.-9.5 mi. E rt. 26 on FS Rd. 2630 (OSU). Yamhill Co., Bald Mt.
(OSU); Forest Grove (OSU), 1 mi. S Pike (OSU). UTAH: Cache Co., Benson (USU); Blacksmith
Fork Cyn. (SHF, USU); East Cyn. (UCD, USU); Green Cyn. (USU); Logan (SHF, OSU, USU);
Logan Cyn. (USU); Providence Cyn. (USNM); Richmond (USU); Smithfield (USU); Wellsville
(KU, USU); Wellsville Cyn. (OSU, USNM, USU). Daggett Co., Elks Pk.- Ashley Nat. For.
(BYU); Grizzly Rdg. (USU); Skull Crk. (AMNH). Duchesne Co., Yellowstone Rgr. Stn. (BYU).
Grand Co., Lk. Oowah— La Sal Mts. (USU); Warner Rgr. Stn. — 28 mi. ESE Moab (AMNH).
Juab Co., Chicken Crk. Cyn. (USU); Little Valley (USU). Millard Co., Oak Crk. Cyn. (USU).
Salt Lake Co., Alta— Little Cottonwood Cyn. (CAS); Cowley Cyn. (OSU); Parleys Cyn. (UCD).
San Juan Co., Bears Ears-Elk Rdg. (BYU, USU); 3 mi. W, 10 mi. SW & Monticello (USU,
BYU); Navajo Mt. (CAS). Sevier Co., 20 mi. E Salina (UIS). Summit Co., 20 mi. E Salt Lk.

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REVISION OF IRBISIA

269

City (KU). Tooele Co., Johnson Pass (UCD). Utah Co., Provo (USNM); Springvile (BYU).
Wasatch Co., Upper Wolf Crk. Cyn. (USNM). Washington Co., Pintura (USNM); Santa Clara
(BYU); 15 mi. N St. George (USNM). Anderson Rch. (BYU). Copper Mts.— Tecoma Range
(BYU). Geyser Pass (USU). Warner (SHF). WASHINGTON: Chelan Co., Blewett (CNC).
Kittitas Co., Cle Elum (CAS); Easton (USNM); 20 mi. NNW Ellensburg (OSU). Klickitat Co.,
5.3 mi. E Bingen (OSU). Lincoln Co., 12 mi. N Davenport (OSU). Pierce Co., Ft. Lewis (UCD);
Graham (USNM); Mt. Rainier (USNM); Puyallup (UCB). Skamania Co., Dog Mt.— Pacific
Crst. Trl. (OSU). Spokane Co., 8 mi. S Spokane (OSU). Stevens Co., 10 mi. S Fruitland (OSU).
Thurston Co., Tenino (USNM); Yakima Co., Toppenish (USNM). Whitman Co., Kamiak Butte
St. Pk. (OSU); Pullman (USNM).

Irbisia setosa Van Duzee
Figs. 31a, b, 54, 77, 100; Map 16

Irbisia mollipes: Van Duzee, 1921a, pp. 147-148 (in part).

Irbisia setosa Van Duzee, 1921a, pp. 149-150, 152 (new species). Essig, 1926, p.

361. Carvalho, 1959, p. 106.

Irbisia ustricula Bliven, 1963, pp. 79-80, 86, pi. 8, figs. 6, 6a. New Synonymy.

Diagnosis. Recognition of this species is not difficult because of the unique vestiture.
I. setosa is the only species with moderately distributed uniformly long, erect bristle-
like fuscous setae on the dorsum of both sexes.

Description. Macropterous male. Length 4.70-5.40, width 1 .80-2. 1 5, shining black
densely clothed with sericeous woolly white setae and widely spaced long erect fuscous
bristle-like setae. HEAD: width across eyes 1 .03-1 . 15, vertex 0.46-0.50, dorsal width
of eye 0.25-0.31; triangular asperate; tylus acute in dorsal view; temporal areas
distinct, asperate; median depression present and distinctly smooth; basal carina
prominent, declivous to level of vertex, straight; eyes broadly joined to frons, pro-
truding laterally, posterior margin straight in dorsal view; distance from tylus to
ventral margin of eye 0.48-0.53, height of eye 0.4 1-0.49, maximum interocular width
0.70-0.78. ROSTRUM: length 1.43-1.60, fuscous to piceous apex always dark, at-
taining bases, and sometimes reaching apices of mesocoxae. ANTENNAE: piceous
or black; I, length 0.50-0.60; II, 1.38-1.74; III, 0.70-0.89; III, 0.83-1.00. PRONO-
TUM: length 0.85-0.97, anterior width 0.75-0.85, maximum width 1.50-1.73; con-
ical, broadly convex, confluently punctate or rugose, posterolateral margins arcuate,
lateral margins straight to concave, rounded or slightly margined at junction with
propleura, anterior angles broadly rounded in dorsal view, not sulcate in lateral view;
calli somewhat indistinct, asperate, narrowly confluent anteriorly, not reaching an-
terolateral margins of pronotum; extreme ventral apices of propleura, very narrowly
bordering mesocoxae and posterior half of ostiolar peritreme ivory. LEGS: casta-
neous, piceous or black; apices of coxae whitish; tibiae usually lighter shade of base
color, infuscated; apices of tibiae and tarsi black. VESTITURE: dorsum densely
clothed with sericeous woolly setae and moderately distributed, uniformly long, erect,
fuscous setae; suberect on posterior portion of hemelytra and cuneus; venter and legs
moderately covered with long suberect testaceous setae.

Macropterous female. Similar to male in most respects, but more robust. Length
4.70-5.45, width 2.13-2.35. HEAD: width across eyes 1.06-1.19, vertex 0.49-0.58,
dorsal width of eye 0.28-0.33, distance from tylus to ventral margin of eye 0.54-

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0.58, height of eye 0.45-0.49, maximum interocular width 0.75-0.85. ROSTRUM:
length 1.53-1.61 reaching bases and sometimes attaining apices of mesocoxae. AN-
TENNAE: I, length 0.50-0.60; II, 1.34-1.68; III, 0.78-0.93; IV, 0.80-1.09. PRONO-
TUM: length 0.88-1.00, anterior width 0.85-0.93, maximum width 1.66-1.90.

Types. Male holotype. Cazadero Cal, iv- 1 2- 1 8, EP Van Duzee; HOLOTYPE setosa
[red]; CAS type no. 803. 1 have examined the holotype and six conspecific paratypes;
these specimens are retained at the California Academy of Sciences, San Francisco
and the Canadian National Collection, Ottawa.

Taxonomic consideration. The junior synonym fits the species concept of I. setosa
in structure and vestiture.

Remarks. The distribution of this species occupies six floristic associations in which
oak is a major or minor component (Coast Range montane, Oregon oak, mixed
evergreen, mixed hardwood, blue oak-digger pine, and southern oak forests), as well
as southern jefFery pine forest and chaparral association. I. setosa occurs as far north
as Josephine County, Oregon within the Pacific forest province and follows this
province south into Californian chaparral province to the San Jacinto Mountains of
Riverside County. Specimens are known throughout the oak associations of the
Sierran forest province. A single specimen from Umatilla County, Oregon, near the
Columbia River, represents either a disjunct population or a labelling error.

Adult specimens have been collected from the following plants: Arctostaphylos sp.,
Ceanothus cuneatus (Hook.) Nutt., C. intergerrimus H. & A., Eriogonum nudum
(Dougl. ex Benth.) S. Stokes, Poaceae, Lupinus sp., Prunus subcordata Benth., Quercus
sp., Rhamnus sp., and Rhus diversibola T. & G. The dates of occurrence are 1 1 March
to 15 July, and elevational range is from 270 to 2260 m. (880 to 6500 feet).

Specimens examined. 164 specimens were examined from the following localities (Map 16).
UNITED STATES: CALIFORNIA: Alameda Co., hills back of Oakland (CAS). Butte Co.,
Butte Crk. Cyn. — nr. Chico (CAS); Oroville (CAS); Pentz (CAS). Colusa Co., Wilbur Hot Spgs.
(JTP). Contra Costa Co., Mt. Diablo— Russellman St. Pk. (CAS, UCB, UCD, UID); Orinda
(LACM). El Dorado Co., Placerville (UCB). Glenn Co., Black Butte (PUC). Humboldt Co., Blair
Rch.- Redwood Crk. (USNM). Kern Co., Fort Tejon (CAS); 3 mi. W Woffard Hts. (UCB).
Lake Co., Anderson Spr. (UCD); Blue Lks. (UCD); 4 mi. W Finley (UCB); Kelseyville (CAS);
Lakeport (CAS); 6 mi. S Middletown (UCB); 12 mLN Upper Lk. (UCB). Marin Co., Alpine
Lk. (UCB); Fairfax, & hills nr. (CAS & UCB); Mill Valley (CAS); Phoenix Lk. (CAS); Ross
(CAS). Mendocino Co., 5 mi. N Branscomb (UCB); 40.38 mp. on rt. 23 btw. Coptche & Ukiah
(OSU); 7 mi. NE & W Eel River Rgr. Stn. (UCB); 2 mi. NW Mendocino Pass (UCB); Ryan
Crk.— N Willits (UCB); U.C. Hopland Exp. Stn. -upper pond & H.Q. (UCB & CAS). Monterey
Co., Bryson (CAS); Carmel (CAS); Chew’s Rdg.— nr. White Oak Cmp. (UCB); Wiley Rch. — 6
mi. W Greenfield (UCB). Napa Co., Monticello Dam (UCD); 4 mi. S & 5 mi. SE Pope Valley
(UCD). Riverside Co., Pinyon Flat— 16 mi. SW Palm Desert (UCR). Sacramento Co., Car-
michael (F&A). San Benito Co., Pinnacles Nat. Mon.— W entr. (OSU). S.L.O. Co., 10 mi. S
Creston (UCB); La Panza Cmp. (UCB); Black Mt. — 6 mi. NE Pozo (UCB); 3 mi. W Paso Robles
(UCB). San Mateo Co., Huddart Pk. (UCB); King’s Mt. (UCB); Redwood City (UCD). Sta.
Barbara Co., Figueroa Cmp. — 13 mi. NE Los Olivos (UCB). Sta. Clara Co., Herbert Crk. — 3
mi. W New Almaden (UCB). Shasta Co., 6 mi. W Fall River Mills (UID, WSU); Harrison
Gulch Rgr. Stn.— rt. 36 nr. Platina (OSU); Redding (UID); Summit City (WSU). Solano Co.,
Green Valley (UCD). Stanislaus Co., W Adoke Crk. (UCB). Trinity Co., 6 mi. NE Hayford
(UCB). Tulare Co., Hospital Flat Cmpgd. — 5.7 mi. N Kemville (UCR); 2 mi. E Johnsondale

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(UCB). Tuolumne Co., Buck Mdws. (USNM). Ventura Co., 5 mi. SE Camarillo (OSU). OR-
EGON: Josephine Co., 3.6 mi. N Kerby (OSU). Umatilla Co., Umatilla (WSU).

Irbisia shulli Knight
Figs. 22a, b, 45, 68, 91; Map 21

Irbisia shulli Knight, 1941, pp. 75-76 (new species); 1968, p. 185. Slater, 1950, pp.

36, 71, pi. 2, figs. 11, 12, 13, p. 75, pi. 4, fig. 5. Carvalho, 1959, p. 106.

Diagnosis. This species is distinguished from allied species by several characters.
I. shulli and I. sericans both have brachypterous females, but the latter has a narrower
vertex, longer antennal segments and fulvous tibiae. The vertex to antennal segment
I ratio of I. shulli ranging from 1:0.84 to 1:1.00 for males and from 1:0.73 to 1:0.86
for females; for 7. sericans the range is from 1:1.04 to 1:43 for males, and from 1:
1 .08 to 1 : 1 .27 for females. 7. shulli can be separated from the closely related 7. nigripes
by the wider vertex and brachypterey of the female of the former species. The vertex
to antennal I ratio of 7. nigripes ranging from 1:1.02 to 1:1.19 for males, and from
1:0.91 to 1:1.11 for females. The sclerotized process of the vesica of the male of 7.
shulli is preapically expanded and slightly involuted as compared to 7. nigripes. I.
shulli, by virtue of its wide vertex and brachypterous female, is also distinct from 7.
serrata. The vertex to antennal I ratio of the latter species ranging from 1:1.05 to 1:
1.24 for males, and from 1:0.89 to 1:1.07 for females.

Description. Macropterous male. Length 5.20-6.15, width 2.05-2.95, moderately
shining black, covered with long erect fulvous setae. HEAD: width across eyes 1.20-
1.36, vertex 0.68-0.75, dorsal width of eye 0.26-0.33; triangular, smooth or slightly
asperate; temporal areas poorly differentiated, smooth, shining; median depression
usually not evident; basal carina blunt, moderately prominent, posteriorly broadly
arcuate mesad, declivous to level of vertex; eyes broadly joined to frons, posterolateral
margin arcuate posteriorly in dorsal view, distance from tylus to ventral margin of
eye 0.70-0.83, height of eye 0.42-0.49, maximum interocular width 0.90-1.03. ROS-
TRUM: length 1 .65-2. 1 8, black, approaching but not attaining mesocoxae to reaching
apex of metacoxae. ANTENNAE: black; I, length 0.55-0.73; II, 1.44-1.85; III, 0.78-
1 .25; IV, 1 .05-1 .30. PRONOTUM: length 1 .00-1 . 1 5, anterior width 0.96-1 .22, max-
imum width 1.73-2.09, trapezoidal, disk broadly convex, rugulopunctate, posterior
margin practically straight, lateral margins straight, broadly rounded at junction with
propleura, anterior angles prominent, rounded, slightly directed anteriorly in dorsal
view, quite evident in lateral view; calli, quite convex, confluent anteriorly, with or
without foveate impression on anteromedial angles, usually not reaching anterolateral
margins of pronotum. LEGS: black; tibiae with variably sized fuscous areas. VES-
TITURE: dorsum densely clothed with suberect fulvous to fuscous or erect white
setae, intermixed, in varying degrees, with recumbent sericeous setae; venter with
sparse thin fulvous setae.

Brachypterous female. Similar to male in color, vestiture and structure (except
membrane), but stature more robust. Length 4.30-5.4, width 2.45-2.88. HEAD:
width across eyes 1 .30-1 .45, vertex 0.73-0.85, dorsal width of eye 0.25-0.33, distance
from tylus to ventral margin of eye 0.75-0.90, height of eye 0.43-0.48, maximum
interocular width 1.03-1.23. ROSTRUM: length 1.80-2.03, approaching but not

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reaching mesocoxae to surpassing metacoxae. ANTENNAE: I, 0.53-0.66; II, 1.33-
1.75; III, 0.85-1.05; IV, 1.05-1.33. PRONOTUM: length 0.94-1.08, anterior width
1.05-1.30, maximum width 1.73-2.08; membrane reduced, not reaching beyond tip
of cuneus, cells not developed, tip of abdomen exposed.

Types. Male holotype. Lenore, Ida., May 7, 1938, Alt. 1000 ft., WE Shull. Holotype
retained at the United States National Museum, Washington, D.C. I have examined
the holotype and 74 paratypes; all are conspecific. The paratypes are deposited at
BYU, CNC, JFS and USNM.

Taxonomic consideration. Some specimens from populations collected from the
westernmost portion of the range of I. shulli are noticeably smaller in all body
measurements and possess erect thin white setae. This condition is obvious in spec-
imens collected on the west side of the Columbia River in Okanogan and Chelan
Counties, Washington, and the Osoyoos District of British Columbia. The seemingly
constant brachyptery of the female in this species suggests limited dispersal ability.
The location of these populations raises the possibility of isolation during pluvial
periods of the Pleistocene epoch. The smaller size may be a result of the drier
conditions of the Okanogan Valley.

A single macropterous female was (hind wings are still brachypterous) contained
in a large sample of 106 specimens of I. shulli collected from 5.5 miles SW of Halfway,
Baker County, Oregon. The entire series was collected from Balsamorhiza sp. This
occurrence suggests that the genes for macroptery are still present in the species but
the conditions necessary for its expression are conjectural.

Remarks. The distribution of I. shulli is rather wide, occupying woodland situations
from the west slopes of the Rocky Mountains in Colorado to the east slopes of the
Cascade Range in Washington. Several grassland, shrub, and forest vegetational
associations are contained within its range. In Colorado and Utah, I. shulli occurs
in mountain mahogany-oak scrub, sagebrush steppe and open situations within
Douglas fir forests associations. Foothill prairies are inhabited in Montana. The
vegetational associations for its range in Idaho, Oregon and Washington are sagebrush
steppe and wheatgrass-blue grasslands. The Palouse Prairie of Washington delimits
the northern border of this species. Adult specimens have been collected from: Agro-
pyron intermedium Beauv., Agropyron sp., Artemisia cana Pursh, Artemisia sp., Bal-
samorhiza sagittata (Pursh) Nutt., Carex sp., Ceanothus sp., Delphinium sp., Elymus
cinereus Scribn. & Merr., Lupinus sp., and Purshia tridentata (Pursh.) DC. Specimens
range in altitude from 305 to 2903 m. (1000 to 9524 feet). The high altitude record
is from Gore Pass, Colorado. Collection dates are from 23 April to 1 1 November.
The late record is from Franklin Basin, Utah.

Specimens examined. 1,935 specimens were examined from the following selected localities
(Map 21). CANADA: BRITISH COLUMBIA: Anarchist Mt.- Osoyoos (CNC); Kelowna
(USNM); Oliver & Madden L., McIntyre Crk., Meyer Flats, Vaseaux L., White Lk. (CNC,
USNM); 8 mi. E & E Osoyoos (CNC & UBC); Rock Crk. (CNC); Shingle Crk. - Penticton
(CNC); Summerland (CNC). UNITED STATES: COLORADO: Eagle Co., Muddy Pass Rd.-
10 mi. NE Wolcott (JTP). Grand Co., Gore Pass (CAS). Routt Co., Steamboat Spgs. (KU,
USNM). IDAHO: Adams Co., 7.5 mi. N Council (OSU). Blaine Co., 18 mi. NW Ketchum
(UID). Bonneville Co., 10 mi. NE Swan Valley. Caribou Co., 3 & 5 mi. E Wayan (UID). Custer
Co., Bear Crk. Cmpgd. (UID). Franklin Co., Cub River Cyn. (USU); Franklin Basin (USU);
Thomas Spg.— Cub River Cyn. (USU); Weston Cyn. (UID). Latah Co., Juliaetta (USNM);

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Moscow (CAS, OSU, UID); 3 mi. N Moscow (UID). Lemhi Co., 2 mi. E Baker (UID). Nez
Perce Co., Lenore (OSU, UID, USNM); Lewiston (USNM). Oneida Co., Holbrook (UDU).
Twin Falls Co., Twin Falls (UID). Valley Co., Donnelly (BSU). MONTANA: Gattatin Co.,
Bozeman (MSU, USNM). Powell Co., Garrison (UID). Sanders Co., Montana Expt. Stn. (MSU).
OREGON: Baker Co., 30 mi. ENE Baker (OSU); 5 mi. NW Ballards Landing (T5S R78E Sec.
31 SE >/4 SW lA) (OSU); 5.5 mi. SW & 9 mi. NE Halfway (OSU); 12 mi. ENE Keating (OSU);
6.5 mi. W & 7 mi. N Richland (OSU); Sparta (OSU). Grant Co., Blue Mt. Hot Spgs. (AMNH);
0.3 mi. S Fox (OSU); Fox Valley (JS); 5 mi. S John Day (OSU); Keerin’s Rch.— Izee (JS); 1 1
mi. N Seneca (OSU). Umatilla Co., Kamela (USNM); 1 mi. N Kamela (OSU); Tollgate (USNM);
nr. Ukiah (OSU); Weston (UID); 9 mi. E Weston (OSU); 0.4 mi. S jet. 244 & 395 (OSU). Union
Co., La Grande, 1 1 mi. SSE & 12 mi. NE (USNM & OSU); 25 mi. SE Pendleton (OSU); Phillips
Cyn.— 4.2 mi. W Elgin (OSU). Wallowa Co., 40 mi. N Enterprise (OSU). Wasco Co., 27 mi.
S Maupin (OSU); Mayer St. Pk. (OSU); 3.4 mi. S Shaniko (OSU). UTAH: Box Elder Co.,
Brigham City (UNSM, USU); Rosevere Crk.— Raft River Mts. (BYU). Cache Co., Ant & Upper
Ant Valey (USU); Blacksmith Fork Cyn. (OSU, USU); East Cyn. (USU); Elk Cyn. (USU);
Franklin Basin (OSU, USU); Hodges Cyn. (USU); Logan (OSU, USNM, USU); Porcupine Res.
(USU); Tony Grove Cyn. Cmpgd. (BYU, OSU, USNM, USU); Porcupine Res. (USU); Tony
Grove Cyn. Cmpgd. (BYU, OSU, USU); Wellsville (CNC, USU), Wellsville Cyn. (AMNH).
Daggett Co., naming Gorge Nat. Rec. A. (UCD); Skull Crk. Cmpgrd.— Ashley Nat. For. (AMNH).
Duchesne Co., Yellowstone Rgr. Stn. (BYU). Juab Co., Chicken Crk. Cmpgd. (USU). Morgan
Co., Morgan (USU); Peterson (USU); Porterville (USU). Rich Co., Allen Cyn. (SHF, USNM,
USU); Bear Lk. Overlook (USU); Garden Cty (SHF, USNM); Laketown (OSU); Monte Cristo
(USNM, USU); Randolph (USU); Woodruff (USNM). Sevier Co., Mattsson Rch.— Salina Cyn.
(USU). Wasatch Co., Current Crk. (KU); 20 mi. SE Heber (UID); Strawberry Res. (UCD,
USNM). Weber Co., Cousey Dam (USNM); 12, 18 mi. N & Huntsville (SHF, USNM, USU);
6, 10 mi. S & 7 mi. W Monte Cristo (USNM, USU). WASHINGTON: Asotin Co., 1.5 mi. S
Anatone (OSU). Benton Co., Snively Cyn., Rattlesnake Rdg.— Hanford A.E.C. Rsv. (WSU).
Chelan Co., 12 mi. SW, 13.4 mi. W Chelan (OSU). Douglas Co., 4 mi. W Waterville (OSU,
WWU). Ferry Co., 8 mi. SE Keller (OSU). Garfield Co., Pomeroy (WSU). Grant Co., 1 mi. E
Stratford (OSU). K ittatas Co., 10 mi. NW, 14 mi. N, 20.5 mi. NNW Ellensburg (OSU); 12 mi.
N Kittatas (OSU). Klickitat Co., 3 mi. E Bingen (OSU); 19 mi. NE Goldendale (OSU); 5 &
21.7 mi. NE Lyle (OSU); 3.7 mi. E Wahkiacus (OSU). Okanogan Co., 7 mi. NE Coulee City
(OSU); 6 mi. E Elmer City (OSU); Frye Hill top— 5 mi. SE Molson (AMNH); Molson (AMNH);
1 1 mi. W Okanogan (OSU); 3.7 mi. W Twisp (OSU). Spokane Co., Turnbull Wldf. Rfg. (OSU).
Stevens Co., 10 mi. S Kettle Falls (OSU). Yakima Co., 13.5 mi. W Naches (OSU); 27.6 mi. S
Toppenish (OSU); Wenas (OSU); Yakima (TA&M, USNM). WYOMING: Lincoln Co., Afton
(USU); Auburn (BYU); Commissary Rdg.— Salt River Range (UWY). Teton Co., Teton Cyn.—
West side Teton Mts. (UID).

Irbisia silvosa Bliven
Figs. 26a, b, 49, 72, 95; Map 4

Irbisia sita Van Duzee: 1921a, pp. 150, 152 (in part, not holotype).

Irbisia silvosa Bliven, 1961, pp. 48-49 (new species); 1963, p. 85, pi. 7, figs. 6, 6a.
Irbisia upupa: Bliven, 1963, pp. 70-71 (in part, not holotype).

Irbisia paulula Bliven, 1963, pp. 75-76, 86, pi. 8, figs. 2, 2a. New Synonymy.
Irbisia inurbana: Bliven, 1963, pp. 81-82 (in part, not holotype)

Diagnosis. Both I. silvosa and I. cuneomaculata are easily separated from all other
species of Irbisia by the presence of extensive light coloration on the ventral portion
of the head and pronotum of both sexes, the venter of the female and the apical one

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third of the left paramere of the male. I. silvosa is distinguished from I. cuneomaculata
by the lack of testaceous marking on the base of cuneus, and the markedly greater
stature and measurements of the former species.

Description. Macropterous male. Length 4.50-6.20, width 1.80-2.33, shining black,
densely covered with recurved, appressed sericeous white setae, sometimes inter-
mixed with long erect white or fuscous setae. HEAD: width across eyes 1.08-1.31,
vertex 0.48-0.63, dorsal width of eye 0.30-0.33, triangular, transversely asperate;
temporal areas poorly differentiated, smooth, without vestiture, with transverse sul-
cus on posterior margin; median depression well developed, wide; basal carina large,
wide and rounded, declivous to level of vertex, posteriorly arcuate mesad, projecting
above eyes in frontal view; eyes broadly joined to frons, protruding laterally; pos-
terolateral margins slightly arcuate posteriorly or straight in dorsal view; distance
from tylus to ventral margin of eye 0.50-0.68, height of eye 0.45-0.54, maximum
interocular width 0.73-0.95; variably marked with testaceous or ochraceous color-
ation on clypeal, jugal and loral sutures and gena; buccula and gula black. ROSTRUM:
length 0.60-2.05, ochraceous, base and apex infuscated with black, attaining apices
of mesocoxae, sometimes reaching bases of metacoxae. ANTENNAE: black; I, length
0.50-0.70; apical one to three quarters fulvous; II, 1.28-1.70, sometimes fulvous
basally; III, 0.73-0.91; IV, 0.80-1.20. PRONOTUM: length 0.90-1.13, anterior mar-
gin 0.83-1.06, maximum width 1.48-2.00; subconical, broadly convex or somewhat
flattened confluently punctured or rugose; posterolateral margins arcuate posteriorly,
posterior margin sometimes arcuate; lateral margins very slightly to moderately con-
cave, rounded at junction with propleura; anterior angles prominent, straight not
projecting anteriorly in dorsal view, sulcate in lateral view; calli prominent, asperate,
broadly confluent anteriorly, reaching anterolateral margins; ventral third of pro-
pleura, prostemum narrowly bordering coxae and attaining front margins, and vari-
able portion of basalar plate, epistemum, epimeron, and second abdominal stemite
testaceous or ivory. LEGS: black or fuscous; apices and perhaps bases of coxae, and
margins of trochanters testaceous or fulvous; femora sometimes extensively fulvous
or testaceous medially, with fuscous spots and streaks; tibiae fulvous or testaceous,
infuscated basally, apices fuscous; tarsi fuscous or black, in some specimens femora
extensively castaneous and tibiae rufescent. HEMELYTRA: insertion ochraceous or
testaceous. VESTITURE: dorsum densely covered with short recurved sericeous
white setae, which are woolly on head, pronotum and scutellum, appressed on he-
melytra; specimens sometimes densely covered with long white or fuscous setae,
which are erect on head, pronotum and scutellum, suberect on hemelytra; both types
of setae sometimes golden; venter moderately covered with thin suberect white to
fulvous setae, these setae are long on the midline of the stemites.

Macropterous female. More robust than male, with more extensive light coloration,
but similar in structure and vestiture. Length 5.00-6.30, width 2.10-2.70. HEAD:
width across eyes 1.18-1.41, vertex 0.56-0.70, dorsal width of eye 0.29-0.36, distance
from tylus to ventral margin of eye 0.58-0.73, height of eye 0.48-0.58, maximum
interocular width 0.83-1.05; testaceous or ochraceous coloration varies from a min-
imum on clypeal, jugal and loral sutures, preapical portion of clypeus, infuscated
juga and lora, and medial portion of gena, to a maximum on entire ventral portion
of head, ventrad of antennal insertions, except for the basal half of clypeus, buccula

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and gula. ROSTRUM: length 1.78-2.18, reaching apices of mesocoxae, sometimes
reaching apices of metacoxae. ANTENNAE: I, length 0.50-0.64, sometimes entirely
fulvous; II, 1.28-1.73; III, 0.79-1.08; IV, 1.00-1.25. PRONOTUM: length 1.00-
1.25, anterior width 0.91-1.13, maximum width 1.66-2.33. LEGS: fulvous, coxae
and trochanters with medial fuscous or black patch, femora with ventral fuscous
spots and streaks, tarsi black; in some specimens femora extensively castaneous and
tibiae rufescent. VENTER: testaceous or ochraceous, dorsolateral regions of stemites
sometimes somewhat infuscated or entirely black, posteriorly directed medial portion
of seventh stemite sometimes black. VESTITURE: both sericeous and long types of
setae sometimes golden.

Types. Male holotype. Woody, [Kern Co.], Cal., Mar. 5,61; HOLOTYPE <5, Irbisia
silvosa, 1963 B. P. Bliven [red]; CAS type no. 13864. I have examined the holotype
and twenty-four paratypes; all are conspecific and retained at the California Academy
of Sciences in San Francisco, California.

Taxonomic consideration. The following specimens of the paratypic series of I.
sita are I. silvosa : Atascadero, S.L.O. Co., California, 23 April 1919, V.D. (three
females); Pasadena, L.A. Co., California, 30 April 1909, F. Grinnell (two males);
Foster, San Diego Co., 22 April 1913, V.D. (one male); San Diego Co., 1 1 March
1914, V.D. (one male). I. paulula Bliven is the junior synonym of I. silvosa because
of identical genitalic structures of both sexes. Specimens that would be identified as
I. paulula differ from the typical I. silvosa by the former’s slightly smaller size and
lack of erect or suberect long white or fulvous setae. Of the material on hand,
specimens that do not have long setae are from Baja California Norte, and Fresno,
Kings, Los Angeles, Riverside, San Bernardino, San Diego, San Luis Obispo and Sta.
Barbara Counties, California. I have collected male and female specimens of I. silvosa
from south of El Condor, Baja California Norte, that do not have any of the light
coloration characteristic of this species. The femora (especially the metafemora) are
castaneous with black spots basad. The genitalia of both sexes do not differ from
more characteristically colored specimens.

Remarks. The distribution of I. silvosa is extensive in California, occupying many
floristic associations in four ecoregional provinces. Douglas County, Oregon is the
known northern limit of the distribution. This area and the northwestern counties
of California are in the Pacific forest province. Within this province specimens of
I. silvosa occupy openings and roadcuts in the following floristic associations: cedar-
hemlock-Douglas fir, redwood, and California mixed evergreen forests. The blue oak-
digger pine and southern oak forest associations of the eastern portion of the Sierran
forest province and the California chaparral province provide the majority of the
habitats of this species. Chaparral, juniper-piny on woodland, mixed hardwood, coastal
sagebrush and valley oak savanna floristic associations are also occupied in the
previously mentioned provinces. Specimens of I. silvosa have been collected in the
California grassland province from California prairie ( Stipa spp.) and riparian forest
floristic associations.

Adult specimens have been collected from these graminoid hosts: Agropyron sp.,
Avena fatua L., Avena sp., Bromus rigidus Roth, B. rubens L., Bromus sp., Elymus
sp., Festuca sp., Hordeum sp. and Vulpia sp. Other non-graminoid plants from which
adults have been collected are: Adenostoma sp., Amsinckia intermedia F. & M.,

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Amsinckia sp., Arctostaphylos sp., Artemisia californica Less., Astragalus sp., Baeria
chrysostoma F. & M., Baeria sp., Ceanothus crassifolius Torr., C. cuneatus (Hook.)
Nutt., C. intergerrimus H. & A., Cer cocarpus ledifolius Nutt., Chaeniactis sp., Cirsium
occidentale (Nutt.) Jeps., Crypthantha intermedia (Gray) Greene, Eriophyllum con-
fertiflorum (DC.) Gray, Lomatium dasycarpum (T. & G.) Coult. & Rose, Lotus
scoparius (Nutt, in T. & G.) Ottley, Lupinus sp., Malvastrum fasciculatus (Nutt.)
Greene, Nemophila sp., Penstemon antinhinoides Benth., Phacelia distans Benth., P.
tanacetifolia Benth., Plagiobothrys sp., Platystemon calif ornicus Benth., Prunus per-
sica Batch, Quercus chrysolepis Liebm., Q. dumosa Nutt., Q. lobata Nee, Q. wislizenii
A. DC., Rhamnus crocea Nutt, ex T. & G. (and ssp. ilicifolia (Kell.) C. B. Wolf, Ribes
sp., Salvia mellifera Greene, Salvia sp., Trichostema parishii Vasey, Trifolium vari-
egatum Nutt., and Yucca schidigera Roezl ex Ortgies. Adult specimens have been
colleted from 64 to 1622 m. (210 to 5322 feet) elevation and from 23 February to
27 May.

Specimens examined. 1,262 specimens were examined from the following localities (Map 4).
MEXICO: BAJA CALIFORNIA NORTE: 12.1 mi. NE Ensenada (UCR); 3.0, 3.4 & 6 mi. S
El Condor (AMNH); 3 mi. W Meling Rch. (UCB); 10 mi. S San Vicente (UCB). UNITED
STATES: CALIFORNIA: Alameda Co., Altamont (UCB); Arroyo Mocho, 22 mi. S Livermore
(UCB), 1 1 mi. E & 1 1-12 mi. SE Livermore (OSU & F&A). Amador Co., lone (UCD); Volcano,
& 3 mi. N (UCD). Butte Co., Chico (CAS); Oroville (CAS); Yankee Hill (CAS). Calaveras Co.,
Altaville (F&A); Angels Cmp. (CAS); Calaveras Res. (UCB); Mokelumne Hill (UCB); 4 mi. S
Railway Flat (UCB). Colusa Co., Wilbur Hot Spgs. (JTP). Contra Costa Co., Lafayette (UCB);
Mt. Diablo — Russelman St. Pk. (UCB, OSU). El Dorado Co., Cool (UCB); El Dorado (F&A);
Lotus (F&A); Marshall Gold Discovery St. Hist. Pk.— nr. Coloma (OSU); Nashville (UCD);
Pine Hill— W Rescue (UCD); Shingle Spgs. (F&A). Fresno Co., Ciervo Hills— 18 mi. SW
Mendota (UCB); 12 mi. W Coalinga (UCB); Dunlap (UCD); Jacalitos Cyn.— SW Coalinga
(CAS). Humboldt Co., Blocksburg (OSU); 2 mi. W Briceland (UCB); Bridgeville— behind cafe
(OSU); W Bridgeville -mp. 40.91 on rt. 36 (OSU); Dinsmores (CAS); Ft. Seward (UCB, OSU);
SW Garberville— mp. 0.28 on Co. rd. 6B (OSU); btw. Honeydew & Bullcreek mp. 0.46 (OSU);
Laribee Valley (CAS); Miranda— mp. 6.52 Ave. of the Giants (OSU); Red way— Redwood Dr.
(OSU). Kern Co., Antelope Cyn.— Tehachapi Mts. (UCR); 27 mi. SE Bakersfield (UCB, OSU);
Ft. Tejon (CAS); Glenville, & 2 mi. E on rt. 155 (UCB & OSU); Havilah, & 3 mi. NE (UCR
& UCB); Hobo Cmpgd.-Kem River (UCB); Lk. Isabella Cmpgd. (OSU); Lebec (CAS & UCB);
1 mi. NE McKittrick (UCB); 5 mi. NE Mojave (UCB); Quail Rsr. — 1 mi. E Keene School on
Hwy. 466 (UCB); Tejon Pass (UCB); Woody (CAS, UCB); 3 mi. W & 1 1 mi. NW Woffard
Hts. (UCB & OSU). Kings Co., 8 mi. SSW & Avenal (F&A & UCD); Kettleman Hills-3 mi.
E Avenal (UCB); McClure Valley (UCD). Lake Co., Blue Lks. (UCB); Butts Cyn. — 10 mi. SE
Middleton (F&A); Clear Oaks— mp. 29.16 on rt. 20 (OSU); Herendon Crk.— Lower Lk. (CAS);
Lk. Pillsburgh (UCB); Nice (UCB); 12 mi. N Upper Lk. (UCB); Walker Rdg. (UCD); rt. 20-
Lake Co. line (OSU); 0.2 mi. W intrsct. rts. 20 & 53 mp. 31.21 (OSU). Los Angeles Co., Bell
Glen Cyn. (LACM); 3 mi. SE Calamigos (UCB); Crater Cmp.— Sta. Monica Mts. (LACM);
Eagle Arch Hills (UCR); Elizabeth Lk. Cyn. (LACM); Frenchmans Flat (UCB); Ganesha Hills
(CU); 5 mi. SE Gorman (F&A); Irwindale (UR); Lk. Hughes (LACM); Newhall (CAS); Los
Angeles, Angeles Nat. For. (OSU, F&A, LACM, UCD, USNM); Pasadena (CAS, USNM); Pine
Cyn. Rd. — San Femado (LACM); San Gabrial Mts. (OSU); Solemint (CNC); Tanbark Flat
(UCB); Tapia Park (LACM); Westwood Hills (UCD); Woodland Hills (AMNH). Madera Co.,
Coarsegold (UCB); San Joaquin Expt. Rng. (OSU, UCB). Marin Co., Fairfax (CAS); Mill Valley
(CAS, SDNH); Phoenix Lk. (CAS); Pt. Reyes (CAS). Mariposa Co., 6 mi. NW Mariposa (UCB).

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277

Mendocino Co., Calif. Coast Range Prsv. 5 mi. N Branscomb (UCB); 1 1 mi. E Calpella (UCD);
btw. Coptche & Ukiah— mp. 40.68 on rt. 223 (OSU); 6 mi. N Hopland (OSU); Hopland, Univ.
Calif. Field Sta. (UCB); Longvale— mp. 0.24 on rt. 162 (OSU); 1 mi. N & 2 mi. SE Piercy
(UCB); Ryan Crk. (UCB); Tooby Mem. Co. Pk. (OSU); Ukiah (CAS, USNM); 8 mi. E Yorkville
(UCB); mp. 68.6 1 on Hwy. 101 (OSU). Merced Co., Pacheco Pass (F&A). Monterey Co., Arroyo
Seco Cmpgd. (UCD); Bryson (CAS); Carmel (CAS); 5 mi. N Escondido Cmpgd. — Sta. Lucia
Mts. (UCB); Jamesburg, & 1 mi. S— Sta. Lucia Mts. (UCB); King City (UCD); 8 mi. E Lucia
(UCB); Monterey (UCB); 10 mi. W Salinas— 0.5 mi. on cutoff to Carmel Valley (OSU); San
Ardo (CAS); Wiley Rch.— 6 mi. W Greenfield (UCB). Napa Co., Angwin (PUC); Bothe— Napa
Valley St. Pk. (OSU); Calistoga (WSU); nr. Conn Dam (AMNH); Lk. Berryessa (WSU); Mon-
ticello, & 1 1 mi. S (F&A & UCD); 3 mi. W Oakville (OSU); Pope Valley, & 5 mi. SE, 7 mi.
S (UCD); 7 mi. E Rutherford (UCD); Wooden Valley (UCD). Nevada Co., Alta Sierra (CAS);
8 mi. S Grass Valley (CAS); Vera Lk.— Nevada City (CAS). Placer Co., Auburn, & 5 mi. NE
(UCB & WSU); nr. Applegate (UCB); Newcastle (UCD). Riverside Co., Banning (USNM);
Cabazon (UCR); Cactus Spg. Trl.— btw. Hwy. 74 & Horsethief Crk. (T7S R5E Sec. 12) (UCR);
El Cariso Rgr. Stn.— Hwy. 74, Cleveland Nat. For. (UCR); Gavilan (UCR); Hemet (USNM);

H. James Rsr.— Lk. Fulmore (UCR); Lk. Mathews (UCR); Millard Cyn. (UCR); Moreno (USNM);

I. 5 mi. W & 2 mi. N Perris (UCR & UCD); Pinyon Flat - 16 mi. SW Palm Desert (UCB); 2
mi. N Poppet Flat on Hwy. 243 (UCR); Sage, & 5 mi. S (UCB); 5.6 mi. S Sage (T7S R1E Sec.
32) (UCR); Riverside (UCR); Wilson Valley Rd. — 1 mi. N Hemet (OSU); mp. 21.29 on rt. 243
(OSU). Sacramento Co., Arcade Crk. (F&A); Fair Oaks (F&A); 3 mi. N Folsom (UCD); Galt
(UCD). San Benito Co., Big Panoche Crk. (UCB); Hernandez (F&A); 2 mi. NE New Idria
(UCB); Paicines, & 12 mi. S (F&A & UCB); Panoche Pass (UCB); Pinnicles Nat. Mon. (F&A,
OSU); 10 mi. S Pinnicles Jet. on Hwy. 25 (UCB). San Bernardino Co., S Camp Angeles (UCR);

4 mi. E Mentone (AMNH, UCR); Redlands, & 10 mi. SW (CAS, UCD); Yucaipa (UCR). San
Diego Co., Alpine (UCB); Banner Grade (ASU); Boulevard (UCB); Descanso (CAS, UCB); 3
mi. SE El Cajon (UCB); 12 mi. N Escondido (F&A; Jacumba, & Round Mt. (F&A & SDNH);
Julian (UCR); Lk. Henshaw (UID); Pala (USNM); Pine Valley, & 2 mi. S (CAS, UCB); San
Diego (CAS); Sta. Ysabella (UID); 9.3 mi. NW Sissors Crossing (OSU); SE Sunshine Summit—
mp. 7.7 on rt. 79 (OSU); Warner Spgs.- Chihuahua Valley (SDNH); Witch Crk. (USNM). San
Joaquin Co., Lone Tree Cyn. (F&A). S.L.O. Co., Atascadero, & 7 mi. SW (CAS, UCB); 10 mi.
SE, 2.5, 4 & 10 mi. S, 4 mi. N Creston (CAS, UCB); 2 mi. NW Cuesta Pass— Sta. Lucia Rng.;
(UCB); Cuyama Valley (LACM); La Panza Cmpgd., & 12 mi. NE (UCB); Morro Bay (CAS);

5 mi. SE Nacimiento Dam (UCB); 2 mi. W Paso Robles (UCB); Pozo (UCB); Sta. Margarita,

6 5 mi. NE (UCB); Templeton (CAS, UCD). San Mateo Co., Palo Alto (CAS, CU). Sta. Barbara
Co., 1 mi. S Buellton (UCB); Los Prietos (UCB); Wons Crk.— below Lk. Cachuma (CAS). Sta.
Clara Co., Mt. Hamilton, & 1 1 mi. E (UCB); San Antonio Valley (UCB); San Jose (USNM);
Silver Crk. (UID); Singleton Rd. (UID). Sta. Cruz Co., Sta. Cruz (UCD). Shasta Co., Wiskey-
town St. Pk. on rt. 299 (OSU); Shasta Co. (F&A). Solano Co., Gates Cyn.— 6 mi. NW Vacaville
(UCD); 3 mi. SE Suisun (OSU); Vacaville (UCD). Sonoma Co., 1.5 mi. S Asti (UCD); Cazadero
(CU); Trinity Rd. (UCB). Stanislaus Co., Del Puerto Cyn.— Frank Raines Pk., 18 mi. W
Patterson (UCB, UCD). Tehama Co., 2 mi. N Red Bluff— jet. of rt. 36 & I 5 (OSU). Trinity
Co., 8 & 9 mi. E Forest Glen Summit on rt. 36 (OSU); E Mad River St. Pk.— mp. 4.63 on rt.
36 (OSU); 3.7 mi. S rt. 36 on Van Duzen Rd. (OSU). Tulare Co., Ash Mt. Rngr. Stn.— Sequioa
Nat. Pk. (AMNH); Fairview (UCB); 10.3 mi. N & Lemon Cove (AMNH & UCB). Tuolumne
Co., Buck Mdws. (USNM); Jamestown (UCB); N. Frk. Tuolumne River— 3 mi. NE Tuolumne
(CAS). Ventura Co., 2 mi. E Lk. Sherwood (UCB); Ojai (F&A); Quatal Cyn. (UCB). Yolo Co.,
Cache Crk. Cyn. Pk. — 7.2 mi. NW Rumsey (OSU, UCB, UCD); Davis (UCD); Putah Cyn.
(UCD). Yuba Co., Cherry Valley (USNM). OREGON: Curry Co., 1 mi. N Agness (T35S R1 1 W
Sec. 7) (OSU). Douglas Co., Whistler’s Bend- 12 mi. E Roseburg (AMNH).

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Irbisia sita Van Duzee
Figs. 14a, b, 37, 60, 83; Map 12

Irbisia sita Van Duzee, 1921a, pp. 150, 152 (new species). Carvalho, 1959, 106.
Irbisia neptis Bliven, 1963, p. 74. New Synonymy.

Diagnosis. I. sita is distinguished from other small (less than 5 mm. in length)
Irbisia on the basis of a combination of characters. The vestiture is a mixture of
dense, woolly sericeous white setae and moderately distributed long white or gray
setae. The anterior width of the pronotum is narrow, from 0.71 to 0.83 mm. in
males, and from 0.81 to 0.88 mm. in females, producing a ratio with the width of
the head across the eyes of from 1:0.66 to 1:0.73 in males, and from 1:0.70 to 1:
0.78 in females. The basalar plate is entirely black. The femora of both sexes are
mostly castaneous and may be rufescent. Positive identification of this species is
facilitated when the left paramere of the male is examined. This structure is abruptly
curved, with the apical half long, and the apex flattened at right angles to the axis of
the shaft. No other small species of Irbisia has a remotely similar paramere. The
structure of the sclerotized rings and dorsal labiate plate of the female are also
distinctive.

Description. Macropterous male. Length 4.25-5.00, width 1.15-1.80, polished
shining black, moderately clothed with a mixture of long, upright white or gray setae,
and dense, sericeous woolly white setae. HEAD: width across eyes 1.06-1.16, vertex
0.45-0.51, dorsal width of eye 0.28-0.33; triangular; tylus rounded in dorsal view,
smooth; temporal areas indistinct, polished; median depression indistinct or very
shallowly depressed; basal carina straight, moderately prominent, acute and abrupt,
or rounded or flattened and declivous to level of vertex; eyes broadly joined to frons,
protruding laterally, posteriolateral margin slightly arcuate posteriorly in dorsal view;
distance from tylus to ventral margin of eye 0.45-0.53, height of eye 0.45-0.51,
maximum interocular width 0.71-0.83. ROSTRUM: length 1.55-1.73, fulvous to
castaneous, attaining apices of mesocoxae. ANTENNAE: black; I, length 0.53-0.63;
II, 1.33-1.78; III, 0.85-1 .05; IV, 0.95-1 . 1 0. PRONOTUM: length 0.80-0.95, anterior
width 0.7 1-0.83, maximum width 1 .30-1 .63; conical, broadly convex, distinctly and
closely punctate, rugulose rear margins, posteriolateral margins arcuate, lateral mar-
gins slightly concave, rounded at junction with propleura anterior angles broadly
rounded in dorsal view, slightly sulcate in lateral view; calli slightly convex, smooth
or slightly asperate, confluent anteriorly, reaching anterolateral margins of pronotum.
LEGS: fulvous to castaneous; bases of coxae, apices of tibiae and tarsi black; femora
sometimes extensively black on basal half; tibia testaceous. VESTITURE: dorsum
moderately clothed with long white or gray setae, erect on head, pronotum and
scutellum, suberect and shorter on hemelytra; intermixed with dense, woolly, re-
curved sericeous white setae; venter sparsely clothed with thin erect white setae.

Macropterous female. More robust than male, otherwise similar in structure, color
and vestiture. Length 4.70-5.40, width 1.83-2.25. HEAD: width across eyes 1.12-
1.25, vertex 0.52-0.60, dorsal width of eye 0.28-0.31, distance from tylus to ventral
margin of eye 0.50-0.58, height of eye 0.46-0.53, maximum interocular width 0.80-
0.90. ROSTRUM: length 1.63-1.80, reaching apices of mesocoxae. ANTENNAE: I,
length 0.50-0.58, II, 1.30-1.58, sometimes fuscous; III, 0.85-1.00; IV, 0.91-1.05.

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REVISION OF IRBISIA

279

PRONOTUM: length 0.91-1.08, anterior width 0.81-0.98, maximum width 1.55-

1.88.

Types. Male holotype. San Diego Co., 4-11-14 Cal., EP Van Duzee; HOLOTYPE
sita [red]; CAS type no. 805. 1 have examined twelve paratypes which are conspecific
with the holotype. Seven paratypes were misidentified and are specimens of I. silvosa\
one paratype was misidentified and is a specimen of I. incomperta. All of the preceding
material (two conspecfic paratypes are at the Canadian National Collection, Ottawa)
is retained at the California Academy of Sciences in San Francisco.

Taxonomic consideration. The original description of this species clearly describes
the holotype and the paratypic series from Foster, San Diego County. These specimens
are all conspecific; several other paratypes were incorrectly identified. I. neptis Bliven,
the junior synonym of I. sita, fits within the habitus of this species in all aspects.

Remarks. The distribution of I. sita is strictly southern California. The northern-
most populations occur in Pinnacles National Monument, San Benito County; the
southern limit of the range is 30° north latitude, near El Rosario, in Baja California
Norte. Within this range, specimens occur in southern oak forest, chaparral and blue
oak-digger pine floristic associations. Adult specimens have been collected from these
grasses: Agropyron sp., Bromus sp., Elymus sp. and Hordeum vulgare L.; and non-
graminoids: Amsinckia intermedia F. & M., Astragulus pomonensis Jones, Ceanothus
crassifolius Torr., C. cuneatus (Hook.) Nutt., Ceanothus sp., Cercocarpus betuloides
Nutt, ex T. & G., Lotus scoparius (Nutt, in T. & G.) Ottley, Penstemon antrirrhinoides
Benth., Rhamnus crocea Nutt, in T. & G., Rhus trilobata Nutt, ex T. & G., Solanum
xanti Gray, Yucca whipplei Torr. Adult specimens have been collected from 6 March
to 29 April and from 30 to 750 m. (100 to 2460 feet) elevation.

Specimens examined. 80 specimens were examined from the following localities (Map 1 2).
MEXICO: BAJA CALIFORNIA NORTE: 12 mi. SE El Rosario (UCR); 4.3 mi. N Ensenada
(UCR); 7 mi. SE Maneadero (UCB); 1 1 mi. N San Vicente (UCB). UNITED STATES: CAL-
IFORNIA: Los Angeles Co., Burbank (USNM); Eve Cyn.— Pomona (LACM); Frenchman’s
Flat (UCB); Gorman (CAS); Pasadena (USNM); San Gabriel R.- mouth of N Frk. (UCR); Sta.
Monica Mts. (LACM); Tapia Park (LACM); Topanga Cyn. (LACM). Orange Co., Sta. Ana
(USNM). Riverside Co., Gavilan (UCR); Riverside (UCR). San Benito Co., Pinnacles Nat. Mon.
(UCB, UCD). San Bernardino Co., 4 mi. E Mentone (AMNH); Mountain Home— S. Bernardino
Mts. (AMNH). San Diego Co., Alpine (UCB); Fallbrook (UCR); 2 mi. E Lyons Valley (UCB);
Mission Dam (USNM); San Diego (AMNH, USNM); San Susana Mts. (LACM); SE Sunshine
Summit— mp. 7.7 on rt. 79 (OSU). S.L.O. Co., 10 mi. S Creston (UCB); La Panza Cmp. — 12
mi. NE Pozo (UCB); 5 mi. NE Sta. Margarita (UCB). Sta. Barbara Co., Los Prietos (UCB).
Ventura Co., N end of Casitas Res. (UCB).

Irbisia solani (Heidemann)

Figs. 6, 18a, b, 41, 64, 87; Map 9

Capsus solani Heidemann, 1910, pp. 200-201, fig. 3 (new species).

Irbisia brachycerus: Vosler, 1913, pp. 551-553, figs. 331, 332. Childs, 1914, p. 220
(misspelled Irbesia brachycerus ).

Irbisia brachyerus var. solani: Van Duzee, 1914, p. 24 (in part, new combination
and new status); 1917a, p. 325 (in part). Carvalho, 1959, pp. 104-105 (in part).
Irbisia sericans: Essig, 1915, pp. 213-214, fig. 189.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

Lygus brachycerus: Van Duzee, 1917b, pp. 265-266 (in part, new combination).
Lygus solani: Van Duzee, 1917b, p. 266 (new combination).

Irbisia solani: Van Duzee, 1916b, p. 38 (new combination); 1921a, pp. 145, 152.

Essig, 1926, p. 361. Essig and Hoskins, 1944, pp. 71-72. Herms, 1926, p. 269.

Tavemetti, 1933, p. 22. Lockwood, 1933, pp. 329-331. Carvalho, 1959, p. 106.

Hall, 1959, pp. 48-51, figs. 1-4. Bliven, 1963, p. 68.

Irbisia setosa: Van Duzee, 1921a, p. 149 (in part, not holotype).

Irbisia incomperta : Bliven, 1963, pp. 73-74, 86, pl.8, figs. 1, la (in part, not holotype).
Irbisia paulula: Bliven, 1963, pp. 75-76 (in part, not holotype).

Irbisia lacertosa Bliven, 1963, pp. 76-77, 84, pi. 8, figs. 3, 3a. New Synonymy.
Irbisia inurbana Bliven, 1963, pp. 81-82, 86, pi. 8, figs. 7, 7a. New Synonymy.
Irbisia serrata: Bliven, 1963, pp. 82-83 (in part, not holotype).

Diagnosis. I. solani is distinguished by: shining black, mostly smooth dorsum;
mixed vestiture of white to fulvous sericeous and short, thin setae; rounded and
indistinct anterior angles of pronotum; mostly smooth (with several slight spines)
outer surface of the left paramere, and non-expanded, linearly spiculate sclerotized
process of the male; narrow dorsal labiate plate, and sclerotized rings of the female.
In the Willamette Valley of Oregon, the above features will separate this species from
I. fuscipubescens, I. cascadia, I. sericans and I. serrata.

I. solani is distinguished from I. oreas by: the considerably less densely distributed
vestiture on the dorsum, tibiae lacking long fulvous setae, more convex calli, concave
lateral margins of pronotum, long shaft of left paramere and apically spined right
paramere of the male, and narrow sclerotized rings of the female. Separation of I.
solani and I. incomperta is difficult and requires examination of the sclerotized rings
of the female; those of the latter species are small and barely open centrally. I.
incomperta differs externally from I. solani by these characters: vertex narrower
(vertex to antennal segment I ratio ranging from 1 : 1 . 1 3 to 1 : 1 .35 in males, and from
1:1.00 to 1:1.04 in females of I. incomperta ; the ratio ranging from 1:0.82 to 1:1.06
in males, and from 1:0.79 to 1:0.92 in females of I. solani ); medial depression of the
head is obvious; lateral margins of the pronotum are concave; and the basalar plate
is ochraceous. I. solani differs from I. limata by: the punctate pronotal disk, shorter
rostrum, dark antennal segment I, and rounded membraneous lobes of the vesica of
the male. I. solani is separated from I. panda by the non-convex clypeal base of the
former species.

Description. Macropterous male. Length 3.80-5.20, width 1.70-2.50, shining black,
moderately or densely clothed with recumbent, sericeous, white to fulvous setae
intermixed with suberect, thin, fulvous setae. HEAD: width across eyes 1.00-1.28,
vertex 0.44-0.63, dorsal width of eye 0.25-0.33; triangular, smooth or slightly as-
perate; temporal areas distinct, bordered basally by transverse, short sulcus; median
depression indistinct or broad but shallow; basal carina straight, moderately prom-
inent, acute or slightly rounded in cross section, declivous to level of vertex (either
steep or gentle); eyes broadly joined to frons, moderately produced laterally, pos-
terolateral margins slightly arcuate posteriorly in dorsal view; distance from tylus to
ventral margin of eye 0.46-0.60, height of eye 0.44-0.53, maximum interocular width
0.70-0.88. ROSTRUM: length 1 .40-1 .83, black, piceous or fuscous, reaching bases
of mesocoxae, usually attaining but seldom surpassing, apices. ANTENNAE: black

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REVISION OF IRBISIA

281

or fuscous; I, length 0.40-0.58; II, 1.10-1.63; III, 0.55-0.88; IV, 0.73-1.05. PRONO-
TUM: length 0.80-1.05, anterior width 0.75-1.03, maximum width 1.43-1.80; sub-
conical, broadly convex, distinctly or confluently punctate, posterolateral margin
arcuate, lateral margins straight or very slightly concave, broadly rounded at junction
with propleura, anterior angles rounded, indistinct in dorsal view, slightly sulcate in
lateral view; calli moderately convex, smooth, confluent anteriorly, reaching antero-
lateral margins of pronotum with transverse depressions anterior of inner angles.
LEGS: black; coxae and femora testaceous apically; femora sometimes extensively
rufescent; tibiae testaceous, piceous on apices and bases; metatibiae sometimes ex-
tensively piceous. HEMELYTRA: fulvous on insertion. VESTITURE: dorsum mod-
erately or densely clothed with recumbent, sericeous white to fulvous setae intermixed
with suberect, thin, white to fulvous setae; head and pronotum predominantly clothed
with upright shining white setae; venter sparsely clothed with thin suberect fulvous
setae.

Macropterous female. Quite similar to male in structure, color and vestiture, but
with more robust stature. Length 4.60-5.50, width 2.10-2.55. HEAD: width across
eyes 1.13-1.29, vertex 0.55-0.65, dorsal width of eye 0.28-0.33, distance from tylus
to ventral margin of eye 0.53-0.65, height of eye 0.45-0.53, maximum interocular
width 0.8 1-0.93. ROSTRUM: length 1.60-1.80, reaching apices of mesocoxae, some-
times reaching apices of metacoxae. ANTENNAE: I, length 0.45-0.58; II, 1.10-
1.60; III, 0.70-1.10; IV, 0.83-1.05. PRONOTUM: length 0.95-1.15, anterior width
0.90-1.05, maximum width 1.70-2.00.

Types. The holotype mentioned in Heidemann’s original description (USNM No.
1 3227) has not been located and is presumed lost. I have examined eleven specimens
from the apparent paratype series and all are conspecific (see discussion under the
treatment of I. serrata ). One male specimen with these label data: Walnut Creek,
[Marin Co.], Cal.; feeding on potato 1910; injurious to Irish Potatoes; Type No.
13680 USNM; Capsus ( Rhopalotomus ) solani Heidm. O.H. is now designated as the
lectotype of C. solani. This specimen is retained at the United States National Mu-
seum, Washington, D.C.

Taxonomic consideration. Heidemann (1910) described Capsus solani. Van Duzee
(1914, 1917a) placed it in the genus Irbisia as a variety of I. brachycerus [a]. Van
Duzee, however, had misidentified the latter species; I. brachycera (by Van Duzee)
is conspecific with the then undescribed I. serrata. Van Duzee (1917b) subsequently
elevated solani and placed it in the genus Lygus. Van Duzee (1921a) finally replaced
solani in the genus Irbisia with species status. Carvalho (1959) cites I. solani as a
variety of I. brachycera. Knight (1941) examined the holotype of I. brachycera and
made I. arcuata its junior synonym. Knight did not recognize I. brachycera by Van
Duzee (1914, 1921) as a distinct species. However, Bliven (1963) gave I. brachycera
by Van Duzee species status as I. serrata. In the same paper Bliven agreed with Van
Duzee (1921a) (although he does not state this fact) in elevating I. solani. Herein I.
brachycera, I. serrata, and I. solani are treated as distinct species.

Vosler (1913) and Child (1914) in short economic papers followed the taxonomy
of Van Duzee (1914, 1917a) and placed I. solani as a variety of I. brachycera. Both
applied and taxonomic workers postdating 1941 except Carvalho (1959) refer to I.
solani as a distinct species. Many of the key specimens referred to in the above
literature were examined during the course of this study.

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Remarks. The distribution of I. solani is extensive in California, occupying ten
vegetational associations within the Pacific and Sierran forest, and California chap-
arral provinces. This species is also found in low lying situations in the Willamette
Valley of Oregon from Forest Grove to Eugene. Both sections of the range of I. solani
are dominated by oak woodlands. The vegetational associations occupied are: mixed
hardwood forest {Arbutus- Quercus), blue oak-digger pine forest {Pinus-Quercus),
southern oak forest {Quercus agrifolia), valley oak savanna {Quercus- Stipa) and Or-
egon oakwoods {Quercus garrayana). In Humboldt, Mendocino and Trinity counties
of northwestern California I. solani is distributed in mixed evergreen forests {Arbutus-
Chrysolepis-Lithocarpus-Pseudotsuga-Quercus-Rhodendrori) and coastal prairie-scrub
mosaic {Baccharis-Danthonia-Festuca) floristic associations.

Adult specimens were colllected from these grass hosts: Agropyron sp., Avena
barbata Broth., A. fatua L., Bromus mollis L., B. rigidus Roth, Bromus rubens L.,
Elymus glaucus Buckl., Festuca arundinacea Schreb., F. rubra L., Hordeum sp. and
Vulpia sp. Adult specimens have been taken on the following non-graminoid plants:
Achillea millefolium L., Amsinkia intermedia F. & M., Artemisia douglasiana Bess,
in Hook., Astragalus pomonensis Jones, Brassica sp., Carduus pycnocephalus L.,
Ceanothus cuneatus (Hook.) Nutt., Ceanothus sp., Conium maculatum L., Cryptantha
sp., Cytisus scoparius (L.) Link, Eriodictyon californicum (H. & A.) Torr., Fraxinus
latifolia Benth., Grindelia camporum Greene, Juglans californica Wats., Lupinus
albifrons Benth., L. micranthus Dougl. in Lindl., Lupinus sp., Penstemon antirrhi-
noides Benth., Plagiobothrus nothofulous (Gray) Gray, Prunus ilicifolia (Nutt.) Walp.
Quercus agrifolia Nee, Q. douglasii H. & A., Q. lobata Nee., Ranunculus sp., Rhamus
calif ornicus Esch., Salix sp., Solanum xantii Gray, Urtica holosericea Nutt, and Yucca
whipplei Torr. Several orchard and ornamental trees and agricultural crops have had
adult specimens collected on them as well: artichoke, citrus, lilac, nut, peach, pear
and potatoes. Specimens occur from 180 to 1420 m. (600 to 4650 feet) elevation
and from 7 March to 15 July.

Specimens examined. 2,837 specimens were examined from the following selected localities
(Map 9). UNITED STATES: ARIZONA: Yavapai Co., Prescott (CAS). CALIFORNIA: Ala-
meda Co. (17 localities). Butte Co. (3). Calaveras Co. (1). Colusa Co. (3). Contra Costa Co. (13).
El Dorado Co. (6). Fresno Co. (5). Humboldt Co. (14). Kern Co. (15). Lake Co. (7). Los Angeles
Co. (6). Madera Co. (4). Marin Co. (16). Mariposa Co. (1). Mendocino Co. (19). Merced Co.
(1). Monterey Co. (24). Napa Co. (17). Nevada Co. (2). Orange Co. (7). Placer Co. (7). Riverside
Co. (12). Sacramento Co. (5). San Benito Co. (11). San Bernardino Co. (8). San Diego Co. (12).
San Francisco Co. (3). San Joaquin Co. (2). S.L.O. Co. (15). Sta. Mateo Co. (9). Sta. Barbara
Co. (2). Sta. Clara Co. (14). Sta. Cruz Co. (2). Shasta Co. (6). Solano Co. (4). Sonomo Co. (15).
Stanislaus Co. (4). Sutter Co. (2). Tehema Co. (2). Trinity Co. (2). Tulare Co. (13). Tuolumne
Co. (4). Yolo Co. (3). Yuba Co. (10). All specimens are retained at CAS, F&A, OSU, UCB,
UCD & UCR. OREGON: Benton Co. (5 localites) (CAS, JS, OSU & USNM); 4.2 mi. N & 6
mi. SW Corvallis (OSU); Finley Nat. Wldlf. Rfg. (OSU); Misty Pt.- McDonald St. For. (OSU);
1 mi. W Philomath (OSU). Lane Co., Good Pasture Is.— Eugene (OSDA). Linn Co., Peterson
Butte (OSU). Marion Co., Salem (USNM). Washington Co., Blooming (OSDA); Cornelius
(AMNH); Dilley (USNM); Forest Grove (OSDA, USNM). Yamhill Co., McMinnville (OSU);
Muddy Valley (OSU); Yamhill R.- McMinnville (OSU). UTAH: Ut. (USNM). WASHING-
TON: Thurston Co., Tenino & Olympia (USNM).

1984

REVISION OF IRBISIA

283

METHODS OF CLASSIFICATION

Characters for each species were compiled subsequent to the completion of the
redescriptions. The characters of the male and the external morphology of the entire
body were analyzed to delineate species groups. The character set was subjected to
cladistic analysis using the PHYSYS program of Drs. J. S. Farris and M. F. Mickevich,
State University of New York at Stony Brook. General discussions of cladistic analysis
are found in Nelson and Platnick (1981) and Wiley (1981).

The options used in this algorithm produced two equally parsimonious trees of 44
steps when Capsus cinctus was chosen as the outgroup (see discussion of solani group).
These trees are the minimum total length (in terms of changes in character states)
arrangement of the data set. The relationships between the species are based on
synapomorphies or shared derived characters. Sister groups, regardless of the number
of furcations, originating from a node have a common hypothetical ancestor with
certain derived characters relative to all the groups to the left of that node on a tree
(Schuh and Polhemus, 1980). The PHYSYS program provided a list of the character
changes leading to a node (the changes are the length of the stem) and the number
of synapomorphies of each node as well as the tree itself. The presentation of the
characters is found in Appendix A, the character matrix is in Table 1, and the
arrangement of the characters is in Figure 101.

The data set used to determine the species groups contains characters from: the
left paramere (2 1 characters), right paramere (4 characters), vesica (6 characters), and
external morphology (3 characters). All the characters are coded as binary data.

With structures that appeared to be homologous in Capsus cinctus and the Irbisia
species, characters selected were entered into the data matrix in the following manner.
Where Capsus cinctus possessed a character it was assumed to be plesiomorphic (state
’O’), and those species of Irbisia with that character were also coded as the state ’O’.
Where C. cinctus did not possess the characters as identified in Irbisia, C. cinctus
was coded as state ’0’ as well. Those Irbisia species with that character (not found
in C. cinctus ) were coded as state ’1’, the apomorphic condition. These characters
only found in the Irbisia species, and not C. cinctus are therefore coded as being
present or absent. No assumptions were made as to transformation series or polarity
of these characters. For example, character 3, sensory lobe strongly projecting above
the surface of the arm is found in C. cinctus and coded state ’O’. Character 1, sensory
lobe not projecting above the surface of the arm, and character 2, sensory lobe slightly
projecting above the surface of the arm, do not occur in C. cinctus and C. cinctus is
again coded state ’0’ for these two characters. In another system of coding characters,
characters 1 and 2 could have been represented as two states of one multistate
character. This, however, would have required that one of the states be hypothesized
as plesiomorphic. I have thus choosen to code such characters, not present in, but
homologous to, the outgroup, as being present or absent to avoid an assumption of
directionality.

This method of coding was possible for characters from the parameres and head.
The structure of the male vesica is unique in both Capsus and Irbisia and therefore
did not permit coding its characters as homologous. It was not practical, given the

Table 1. Matrix of Characters for Irbisia species.

284

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

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1984

REVISION OF IRBISIA

285

scope of this revision, to develop homologies for the vesica considering the current
state of our knowledge of the tribe Mirini.

DISCUSSION OF SPECIES GROUPS

The monophyly of Irbisia in the tribe Mirini is substantiated by the morphological
characters presented in the generic and species descriptions. Several structures men-
tioned in the diagnosis of the genus Irbisia, pertaining to the genitalia of both sexes,
that separate Irbisia and the genus Capsus have not been included in the matrix for
analysis because of the autapomorphic nature of those characters. Cladistic analysis
reveals five species groups of Irbisia. The species and synapomorphies for each are
as found in Figure 10 1 . In the cladogram characters with no homoplasy are in regular
print, those with parallelism are in italics and those with reversal are in italics with
a degree symbol after the number. The following discussion presents the species
groups and their synapomorphies according to the characters in Table 1. The com-
ponent numbers refer to Figure 101, and the numbers in parentheses are the character
numbers with their states.

The brachycera group, component 27 —brachycera, elongata, pacifica and silvosa
uniquely possess: left paramere with sensory lobe not projecting above arm (1-1, 2-
0, 3-1), large ’L’ shaped angle (8-0), arm and shaft without spines (4-0, 17-0), arm
and shaft of equal length (9-1), shaft stout and of equal diameter throughout (16-0),
apex securiform with ventral and dorsal portions equally developed (18-1); right
paramere, unexpanded distally (22-1), with single terminal spine parallel to long axis
(24-0) and for all but silvosa, component 26, antennal socket with ventral margin
below ventral margin of eye (32-1).

The californica group, component 30— californica, cuneomaculata, sericans and
sit a uniquely possesses: left paramere with sensory lobe strongly projecting above
arm (1-0, 2-0, 3-0), sharp ’V’ shaped angle (5-1), arm and shaft without spines (4-
0, 17-0), shaft of greater length than arm (11-1), shaft slender and of equal diameter
throughout (13-1); right paramere, unexpanded distally (22-1), with single terminal
spine at right angle to long axis (23-1); vesica with sclerotized process tapering distally
(29-1).

The brachycera and californica groups show parallelism in the distally unexpanded
condition of the right paramere (22-1).

The remainder of the species of Irbisia, component 40 possess these synapomor-
phies: left paramere with sensory lobe slightly or moderately projecting above the
arm (1-1, 2-1, 3-1), arm with tubercles and spines (4-1), apex securiform with ventral
portion of blade larger than dorsal portion (19-1); right paramere with diameter
expanded distally (22-0), double terminal spine (25-1).

The solani group, component 33 — incomperta, knighti, limata, oreas and solani
uniquely possesses: left paramere with moderate ’C’ shaped angle (6-1) and shaft of
equal moderate thickness throughout (14-1). Two equally parsimonious trees were
derived in this analysis I have presented one (Fig. 101). In the other arrangement
oreas and limata reverse positions resulting in this character arrangement: component
32 possesses character 10-1; component 31 possesses character 29-1; oreas possesses
characters 11-1 and 29-1; limata posseses character 4-0.

Component 39 — bliveni, cascadia, castanipes, fuscipubescens, mollipes , nigripes.

286

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

Fig. 101. Cladistically derived relationships of the species of Irbisia based on 34 characters.

1984 REVISION OF IRBISIA 287

Table 2. Relationships of cladistically derived species groups to the distribution patterns
of the species of Irbisia.

WIDESPREAD

VANCOUVERAN

* brachycera, pacifica.

f knighti.

** serrata.

ft sericans.

NORTHERN ROCKY MTS.

CALIFORNIAN

* elongata.

f limata, incomperta,

** fuscipubescens.

oreas, solani.

nigripes.

* silvosa.

shulli.

ft californica, sita,
cuneomaculata.

SISKIYOU MTS.

castanipes, bliveni' .

** cascadia.

*** setosa, mollipes, panda.

* brachycera group, ** ni gripes group, *** setosa group

t solani group, ft calif or nica group 'Component 39

panda, serrata, setosa and shulli uniquely possesses: left paramere with broad ’U’
shaped angle (7-1), length of shaft shorter than length of arm (9-0), shaft diameter
constricted on apical two thirds (15-1), shaft with tubercles or spines (17-1). Within
this component are two species groups. The setosa group — mollipes, panda and setosa
uniquely possess: vesica with sclerotized process projecting sharply to the left (25-1)
and large accessory lobe in front of sclerotized process (26-1). The nigripes group,
component 38 —cascadia, fuscipubescens, nigripes, serrata and shulli uniquely pos-
sesses a shaft which is subapically constricted with a *T* shaped securiform apex (2 1 -
1). Within this species group component 37 uniquely possesses a cone shaped scler-
otized process with very large spines (30-1).

One or two species in each of these aforementioned species groups shows homo-
plasy in some characters. I. silvosa, in the brachycera group, lacks character 32-1. I.
cuneomaculata has parallelism with component 40 in character 19-1, and reversal
of character 24 to state ’O’ linking this species with the brachycera group. I. limata
has a reversal of character 4 to state ’O’ thus showing some similarity with both the
brachycera and californica species groups.

BIOGEOGRAPHY

The individual species ranges of Irbisia reveal several distributional patterns.
Table 2 lists the species of Irbisia within five distributional patterns. The species are
also arranged according to their membership in the cladistically derived species
groups. Each species group contains species which have ranges that cross distribu-
tional pattern boundaries. Within the californica species group the sister species I.
californica and I. sericans occur in the Californian and Vancouveran pattern re-
spectively. I. solani and I. knighti of the solani group display a similar relationship.
Sister species connect the Northern Rocky Mountain and Siskiyou Mountain patterns;
the California pattern is connected to the Northern Rocky Mountain pattern as well.

288

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

Distribution maps 1,2. 1.7. brachycera. 2. 7. elongata.

1984

REVISION OF IRBISIA

289

i!

Distribution maps 3-5. 3. I. pacifica. 4. I. silvosa. 5. I. limata.

290

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

Distribution maps 6-9. 6. 7. oreas. 7. 7. incomperta. 8. 7. knighti. 9. I. solani.

1984

REVISION OF IRBISIA

291

Distribution maps 10-13. 10. I. sericans. 11. I. californica. 12. /. sita. 13. I. cuneomaculata.

292

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

Distribution maps 14-18. 14. I. mollipes. 15. I. panda. 16. I. setosa. 17. I. bliveni. 18. I. castanipes.

1984

REVISION OF IRBISIA

293

Distribution maps 19, 20. 19. I. nigripes. 20. I. serrata.

294

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

Distribution maps 21-23. 21. I. shulli. 22. I. fuscipubescens. 23. I. cascadia.

1984

REVISION OF IRBISIA

295

Table 3. Distributional grouping of the species of Irbisia based on the Ecoregions of the
United States (Bailey 1978) with minor modification.

Domain

Division

Province

Species

Polar

1 species

Tundra

Bering Tundra

sericans

Subarctic

Alaska Range

sericans

Humid

Temperate

Warm Continental

Columbia Forest

elongata , fuscipubescens ,

23 species

4 species
Marine

Willamette-Puget Forest

cascadia, sericans, serrata.

14 species

Pacific Forest

solani

Northwestern

(bliveni), cascadia,
fuscipubescens, knighti,

sericans, serrata,

Northeastern

fuscipubescens, nigripes, shulli

Southern

bliveni, brachycera, californica,
cascadia, pacifica, serrata,
setosa, silvosa, solani.

Mediterranean

California Grassland

brachycera, californica,

17 species

(limata), pacifica, (silvosa),
solani

Sierran Forest

bliveni, brachycera, (cascadia),
castanipes, cuneomaculata
elongata, incomperta, limata,
oreas, pacifica, panda, serrata,
setosa, silvosa, solani

California Chaparral

brachycera, californica,
(castanipes), cuneomaculata,
incomperta, limata, mollipes,
oreas, pacifica, panda, setosa,
silvosa, sita, solani

Dry

10 species

Steppe

Great Plains Shortgrass

brachycera, pacifica

10 species

Prairie

Palouse Grassland

(brachycera), (fuscipubescens),
pacifica, serrata, shulli

Intermountain Sagebrush

brachycera, cascadia, elongata,
fuscipubescens, nigripes,
pacifica, serrata, shulli

Rocky Mountain Forest

brachycera, elongata, fusci-
pubescens, nigripes, pacifica,
serrata, shulli

Upper Gila Mountains

oreas, serrata, solani

Forest

Colorado Plateau

brachycera, serrata

Wyoming Basin

(brachycera), (pacifica)

( ) indicates that record within the province are few

296

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

The distribution patterns recognized in Irbisia roughly correspond to several dis-
tributional range types described by Monroe (1956). However, Monroe did not in-
clude heteropteran species in his scheme. Kelton (1975) presented the distribution
ranges for the North American species of the mirid genus Lygus. Six of the species
treated therein have distributional patterns which are similar to the widespread
pattern found in the Irbisia species.

Table 3 summarizes the distribution data presented in the species redescriptions.
The species are placed within the divisions and provinces of Bailey’s (1978) Ecore-
gions of the United States. I have subdivided the Pacific forest province into north-
western, northeastern and southern regions because many of the species have dis-
tinctly delineated ranges within this province. Species with few known or peripherally
arranged localities are placed in parentheses.

Vicariance biogeographic methods (Nelson and Platnick, 1 98 1) were attempted on
Irbisia with somewhat inconclusive results due to the extensive overlap of the dis-
tribution ranges of the species. However, I would like to point out several possible
areas of endemism that appear to me to be worthy of consideration by researchers
working with other taxa. These areas are: 1) coastal California including the western
and eastern flanks of the Coastal range (typified by Map 11); 2) the western foothills
and the pine areas of the Sierra Mountains (Map 5); 3) the northern Rocky
Mountains (Maps 19, 20); 4) the Klamath and Siskiyou Mountains (Map 23).

ACKNOWLEDGMENTS

Many people have assisted during all phases of this study and without their contributions
much of the research could not have been completed.

Type specimens were borrowed from: California Academy of Sciences, San Francisco, Dr.
Paul H. Amaud, Jr.; Cornell University, Ithaca, New York, Dr. Laveme L. Pechuman; and the
United States National Museum, Washington, D.C., Dr. Richard C. Froeschner and Mr. Thomas
J. Henry. The holotype of I. cuneomaculata Blatchley was examined by Mr. Arwin V. Pro-
vonsha, Purdue University, West Lafayette, Indiana.

In addition to the above, the following institutions or private collections and curators or
individuals kindly loaned specimens: Alaska Agricultural Experiment Station, Palmer, Mr.
David P. Bleicher; American Museum of Natural History, New York, Dr. Randall T. Schuh;
Arizona State University, Tempe, Dr. Frank F. Hasbrouck; Boise State University, Idaho, Dr.
Charles W. Baker; Brigham Young University, Provo, Utah, Dr. Stephen L. Wood; California
State Department of Food and Agriculture, Sacramento, Dr. Alan R. Hardy; Humboldt State
University, Areata, California, Dr. Richard L. Hurley; Illinois Natural History Survey, Urbana,
Dr. Donald W. Webb; James Entomological Collection, Pullman, Washington, Dr. William J.
Turner; Los Angeles County Museum of Natural History, California, Dr. Charles L. Hogue;
Montana State University, Bozeman, Ms. Sharon D. Rose; North Dakota State University,
Fargo, Dr. Edward U. Balsbaugh Jr.; Northern Arizona University, Flagstaff, Dr. Clarence D.
Johnson; Oregon State Department of Agriculture, Salem, Mr. Kenneth Goeden and Mr. Rich-
ard L. Westcott; Oregon State University, Corvallis, Dr. John D. Lattin, Dr. Paul W. Oman,
Dr. Loren K. Russell, Dr. Gary M. Stonedahl, Mr. Gary M. Cooper, Mr. Kenneth J. West;
Pacific Union College, Angwin, California, Dr. Lloyd E. Eighme; Polhemus Collection, Engle-
wood, Colorado, Dr. John T. Polhemus; San Diego Natural History Museum, California, Mr.
David K. Faulkner; Santa Barbara Museum of Natural History, California, Mr. Scott E. Miller;
Schuh Collection, Klamath Falls, Oregon, the late Mr. Joe Schuh; Snow Entomological Museum,
Lawrence, Kansas, Dr. Peter D. Ashlock and Dr. George W. Byers; South Dakota State Uni-
versity, Brookings, Dr. Burrus McDaniel; Southern Oregon College, Ashland, Dr. Marvin D.

1984

REVISION OF IRBISIA

297

Coffey; Texas A & M University, College Station, Dr. Joseph C. Schaffner; University of Alberta,
Edmonton, Canada, Dr. George E. Ball; University of Arizona, Tucson, Dr. Floyd G. Werner;
University of British Columbia, Vancouver, Canada, Dr. Geoffrey G. E. Scudder; University
of California, Berkeley, Dr. Jerry A. Powell; University of California, Davis, Dr. Robert O.
Schuster; University of California, Riverside, Mr. Saul I. Frommer and Dr. John D. Pinto;
University of Idaho, Moscow, Dr. William F. Barr; University of Nebraska, Lincoln, Dr. Brett
C. Ratcliffe; University of Wyoming, Laramie, Dr. Robert J. Lavigne; Utah State University,
Logan, Dr. Wilford J. Hanson; Western Washington State University, Bellingham, Dr. Gerald

F. Kraft.

Plant identifications were provided by Mrs. LaRae D. Johnston and Dr. Kenton L. Chambers,
Herbarium, Oregon State University; Ms. Pamela Ann Camp, Bureau of Land Management,
Spokane, Washington, and Dr. A1 H. Winward, United States Forest Service, Ogden, Utah.

Dr. J. M. Ferris, Purdue University, West Lafayette, Indiana, provided a CDC 6500 version
of the Wagner 78 maximum parsimony program of Dr. J. S. Farris, State University of New
York, Stony Brook. Dr. Charles Mitter, University of Maryland, College Park, provided access
to the PHYSYS program of Drs. Farris and M. F. Mickevich, State University of New York
at Stony Brook, on the University of Maryland computer facility. Dr. Norman I. Platnick,
American Museum of Natural History, New York, for discussing the phylogenetic relationships.

Ms. Bonnie B. Hall, Scientific Illustrator, Oregon State University, Systematic Entomology
Laboratory, Corvallis, illustrated the body of three species in dorsal view. Ms. Kathleen Schmidt,
Scientific Assistant, Americam Museum of Natural History, aided in the preparation of the
distribution maps and cladogram.

The majority of the work for this project was completed as partial requirement of a master
of science degree at Oregon State University. Financial support was provided by a research
assistantship through the Oregon State University Systematic Entomology Laboratory with
funds made available by the Science Technical Advisory Fund. Additional financial support
for publication of this manuscript was provided by Oregon State University Systematic Ento-
mology Laboratory.

I would like to express my sincere thanks to Dr. John D. Lattin, my major professor, for
suggesting this project. He allowed me to make use of his own preliminary notes on the genus,
aided in all facets of the research, and provided unerring guidance for its duration.

I wish to especially thank these individuals: Ms. Elyse Larsen Schwartz for the outstanding
preparation of the original manuscript, its subsequent revisions and typesetting the tables; Dr.
Loren K. Russell for assisting me in collecting specimens on several field trips in California
and Oregon; Dr. Paul W. Oman for providing me with numerous specimens from central
Washington; Dr. George F. Knowlton for bringing to my attention the unpublished research
of Mr. Earl J. Taylor, Yuma, Arizona, who investigated a complex of Irbisia species in Davis,
Yolo County, California during the early 1960’s; Mr. Jeff B. Knight, University of Nevada,
Reno, for graciously allowing me to incorporate his unpublished data on the biology of Irbisia
species; Mr. Gary M. Cooper, Dr. Paul W. Oman, Dr. Vincent de Paul Razafimahatratra and
Dr. Gary M. Stonedahl for their advice and sincere encouragement during this study; Dr. A1

G. Wheeler, Jr. for critically reading the manuscript; Dr. Randall T. Schuh for his help in
turning the thesis into a publication.

Museum Abbreviations

AKAS, Alaska Agricultural Experiment Station, Palmer

AMNH, American Museum of Natural History, New York

ASU, Arizona State University, Tempe

BSU, Boise State University, Idaho

BYU, Brigham Young University, Provo, Utah

CAS, California Academy of Sciences, San Francisco

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CNC, Canadian National Collection of Insects, Agriculture Canada, Ottawa

CU, Department of Entomology, Cornell University, Ithaca

F&A, California Department of Food and Agriculture, Sacramento

HSU, Humboldt State University, Areata, California

INHS, Illinois Natural History Survey, Urbana

JS, Schuh Collection (now at AMNH), Klamath Falls, Oregon

JTP, Polhemus Collection, Englewood, Colorado

KU, Snow Entomological Museum, University of Kansas, Lawrence

LACM, Los County Museum of Natural History

MSU, Montana State University, Bozeman

NAU, Northern Arizona University, Flagstaff

NDS, North Dakota State University, Fargo

OSDA, Oregon State Department of Agriculture, Salem

OSU, Oregon State University, Corvallis

PUC, Pacific Union College, Angwin, California

SBNM, Santa Barbara Museum of Natural History, California

SDNH, San Diego Natural History Museum, California

SDS, South Dakota State University, Brookings

SHF, Schaffner Collection, Texas A&M University, College Station

SOS, Southern Oregon College, Ashland

TA&M, Department of Entomology, Texas A&M University, College Station

UALB, University of Alberta, Edmonton, Canada

UAZ, University of Arizona, Tucson

UBC, University of British Columbia, Vancouver, Canada

UCB, California Insect Survey, University of California, Berkeley

UCD, Department of Entomology, University of California, Davis

UCR, Department of Entomology, University of California, Riverside

UID, University of Idaho, Moscow

UN, University of Nebraska, Lincoln

USNM, National Museum of Natural History, Washington, D.C.

USU, Department of Entomology, Utah State University, Logan.

UWY, University of Wyoming, Laramie

WSU, James Entomological Collection, Pullman, Washington

WWS, Western Washington State University, Bellingham

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APPENDIX A

Left paramere
Sensory lobe

1 . 1 — not projecting above surface of arm.

0— either strongly, slightly or moderately projecting above surface of arm.

2. 1 — slightly or moderately projecting above surface of arm.

0— either strongly or not projecting above surface of arm.

3. 1 — either slightly, moderately or not projecting above surface of arm.

0— strongly projecting above surface of arm.

Arm

4. 1 — with spines or tubercles.

0— without spines or tubercles.

Angle

5. 1— sharp ’V’ shaped.

0— either ’C\ *U* or ’L’ shaped.

6. 1 —moderate ’C’ shaped.

0— either ’V’, ’U’ or ’L’ shaped.

7. 1 — broad ’U’ shaped.

0— either ’V’, ’C’ or ’L’ shaped.

8. 1— either ’V’, ’C’ or ’U’ shaped.

0 — large ’L’ shaped.

Ratio of length of shaft to length of arm

9. 1 — length of shaft equal to or greater than lengthof arm.

0— length of shaft less than length of arm.

10. 1 — length of shaft equal to length of arm

0— length of shaft less than or greater than length of arm.

11. 1 — length of shaft greater than length of arm.

0— length of shaft less than or equal to length of arm.

Arm and shaft junction

12. 1 —junction with flange.

0— junction without flange.

Shape of shaft diameter

13. 1 — slender throughout.

0— either medium thickness, constricted on apical two thirds or thick throughout.

14. 1 —medium thickness.

0— either slender throughout, constricted on apical two thirds or thick throughout.

15. 1 —constructed on apical two thirds.

0— either slender throughout, medium thickness, or thick throughout.

16. 1— either slender throughout, medium thickness, constricted on apical two thirds or

thick throughout.

0— thick throughout.

Shaft

17. 1 —with spines.

0— without spines.

Apex of shaft

18. 1 — securiform with either ventral portion of securiform apex larger than dorsal portion

or ventral and dorsal portions equally developed.

0— at right angle to long axis of shaft.

19. 1 — securiform with ventral portion of securiform apex larger than dorsal portion.

0— either at right angle to long axis of shaft or securiform with ventral and dorsal

portions equally developed.

1984

REVISION OF IRBISIA

305

20. 1 — securiform with ventral and dorsal portions equally developed.

0— either securiform with ventral and dorsal portions equally developed or at right
angle to long axis of shaft.

21. 1— shaft constricted subapically; securiform apex somewhat T’ shaped.

0— shaft not constricted subapically; securiform apex not ’T’ shaped.

Right paramere
Diameter of shaft

22. 1 —unexpanded distally.

0— expanded distally.

Terminal spine

23. 1 — simple spine, at right angle to long axis of shaft.

0— either simple spine, parallel with long axis of shaft or double faceted spine.

24. 1 — either simple spine, at right angle to long axis of shaft or double faceted spine.

0— simple spine, parallel with long axis of shaft.

25. 1— double faceted spine.

0— simple spine either at right angle to long axis of shaft or parallel with long axis of
shaft.

Vesica

26. 1 — sclerotized process projecting to left side.

0— sclerotized process projecting ventrad.

27. 1 — accessory lobe in front of sclerotized process large.

0— accessory lobe in front of sclerotized process small.

28. 1— both membranous side lobes tapering distally.

0— both membranous side lobes broadly rounded distally.

29. 1 — sclerotized process tapering distally.

0— sclerotized process not tapering distally.

30. 1 — sclerotized process cone-shaped and with very large spines.

0— sclerotized process not cone-shaped with very large spines.

31. 1 — large spines in triangular arrangement.

0— without large spines in triangular arrangement.

Head structure

32. 1— antennal socket with ventral margin below ventral margin of eye.

0— antennal sockets with ventral margin in line or above ventral margin of eye.

33. 1 — ratio of width of vertex to length of antennal segment I equal to or less than one.
0— ratio of width of vertex to length of antennal segment I greater than one.

34. 1 — length of antennal segment I greater than width of head.

0— length of antennal segment I less than width of head.

306

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(3)

APPENDIX B

A. = Area

mp. = milepost

A.E.C. = Atomic Energy Commission

Mt.(s.) = Mountain(s)

Ave. = Avenue

Nat. = National

btw. = between

nr. = near

Clev. = Cleveland

Pen. = Peninsula

Cmp. = Camp

Pk. = Park

Cmpgd. = Campground

Prsv. = Preserve

Crk. = Creek

Pt. = Point

Co. = County

R. = River

Crst. = Crest

Rch. = Ranch

Cyn. = Canyon

Rd. = Road

Dr. = Drive

Rdg. = Ridge

Entr. = Entrance

Rec. = Recreational

Exp. = Experiment

Res. = Reservoir

Expt. = Experimental

Rfg. = Refuge

For. = Forest

Rgr. = Ranger

Frk. = Fork

Rng. = Range

Ft. = Fort

Rsr. = Reserve

Grd. = Guard

Rsrh. = Research

Hist. = Historic

Rsv. = Reservation

HQ. = Headquarters

rt. = route

Hts. = Heights

Sec. = section

Hwy. = Highway

Spg.(s.) = Spring(s)

intrsct. = intersection

S.L.O. = San Luis Obispo

Is. = Island

Smt. = Summit

Jet. = Junction

St. = State

L.A. = Los Angeles

Sta. = Santa

Lk. = Lake

Stn. = Station

LWr. = Lower

Trl. = Trail

Mdw. = Meadow

Wldf. = Wildlife

Mem. = Memorial

Wldns. = Wilderness

Mon. = Monument

Wysd. = Wayside

.

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Journal of the

New York Entomological Society

VOLUME 92 JULY 1984 NO. 3

CONTENTS

A Revision of the Black Grass Bug Genus Irbisia Reuter (Heteroptera: Miridae)

Michael D. Schwartz 193-306

OCTOBER 1984

No. 4

70673

Vol. 92

Journal

of the

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Entomological Society

(ISSN 0028-7199)

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NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 307-315

ORMENARIA RUFIFASCIA
(HOMOPTERA: FULGOROIDEA: FLATIDAE):
DESCRIPTIONS OF NYMPHAL INSTARS AND
NOTES ON FIELD BIOLOGY1

Stephen W. Wilson and James H. Tsai

Department of Biology, Central Missouri State University,
Warrensburg, Missouri 64093 and
Ft. Lauderdale Research and Education Center,

University of Florida, IFAS, Ft. Lauderdale, Florida 33314

Abstract.— The five nymphal instars of Ormenaria rufifascia (Walker) are described and
illustrated. A key to instars and comparisons of the morphology of 5 genera of U.S. flatid
nymphs are provided. Features useful in separating nymphal instars include spination of me-
tatibiae and metatarsomeres, numbers of metatarsomeres and abdominal waxpads, and differ-
ences in body and wingpad sizes. One generation per year was observed in south Florida.
Average developmental times (x ± SD) for the first-fifth instars are 23 ± 2.2, 20 ± 2.0, 19 ±
2.3, 24 ± 1.9, and 8 ± 1.1 days, respectively. Mean adult longevity is 12 ± 1.6 days. Of 13
palm species and one non-palm plant studied, only Sabal palmetto (Walt.) Lodd. and Latania
lontaroides (Gaertn.) H. E. Moore served as breeding hosts for O. rufifascia.

Ormenaria rufifascia (Walker) has been recorded from Florida, Georgia, and Cuba
(Metcalf and Bruner, 1948). It has been reported feeding on 9 species of palms and
3 species of other plants (Metcalf and Bruner, 1948; Mead, 1965; Table 1). Although
occasionally common on palms, injury is apparently slight (Mead, 1965). Other than
records of host plants, little information is available on the biology of this flatid.
Mead (1965) noted that nymphs were observed feeding in conspecific groups and
are present from late April-May, and adults from mid-May-July, in Florida. Moore
(1961) reported that adults produce tymbal vibrations.

Information on the morphology of this flatid is very limited. Metcalf (1923) pro-
vided a color illustration of an adult’s head and thorax and Metcalf and Bruner ( 1 948)
described the adult and illustrated the male genitalia. Mead (1965) briefly described
the nymphal color patterns. Of the 33 species of U.S. flatids (Metcalf, 1957), only
the immatures of Anormenis septentrionalis (Spinola), Metcalf a pruinosa (Say), Or-
menoides venusta (Melichar) (Wilson and McPherson, 1981) and Cyarda sp. near
acutissima Metcalf and Bruner (Wheeler and Hoebeke, 1982) have been described.

This paper presents descriptions of the five nymphal instars and notes on the field
biology of O. rufifascia in south Florida.

MATERIALS AND METHODS

Description of immatures. Specimens to be described were obtained from field
cages (see below) and preserved in 70% ethyl alcohol. The first instar is described in

Fla. Agric. Exp. Stn. Journal Series No. 5443.

308

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

detail, but only major differences are described for subsequent instars. Comparative
statements refer to earlier instars (e.g., more numerous). Measurements are given in
mm as mean ± SE. Length was measured from apex of vertex to apex of abdomen,
width across the widest part of the body, and thoracic length along the midline from
anterior margin of the pronotum to posterior margin of the metanotum. Several
specimens of each instar were cleared in 1 0% KOH to observe obscure features such
as body pits and waxpads. Twenty specimens of each instar were measured.

Field study. The development of O. rufifascia was studied from January to June
1983 in a small palm planting at Ft. Lauderdale, Broward County, Florida. Cabbage
palm [Sabal palmetto (Walt.) Lodd] was used as test plant. Sleeve cages made of
clear butyrate tubing measuring 22 cm long (dia: 5 cm) enclosed portions of pinnae
for nymph release. Newly hatched nymphs were collected in the field and released
singly into the sleeve cages. Specimens were observed and moltings recorded every
1-2 days. Daily temperature was recorded throughout the 6-month study period for
comparing the nymphal developing rate.

A group of 13 species of palms including Carpentaria acuminata (H. Wendl. &
Drude) Becc., Veitchia merrilli (Becc.) H. E. Moore, Pritchardia eriostachya Becc.,
Veitchia merrilli (Becc.) H. E. Moore, Pritchardia eriostachya Becc., Ptychosperma
nicolai (Sand, ex Andre) Burret, Livistonia chinensis (Jacq.) R. Br. ex Mart., Wash-
ingtonia robusta H. Wendl., Heterospathe elata Scheff, Cocos nucifera L., Phoenix
dactylifera L., Phoenix roebelenii O’Brien, Caryota mitis Lour, Dictyosperma album
(Bory) H. Wendl. and Drude, and Latania lontaroides (Gaertn.) H. E. Moore; and
one non-palm, screw pine (Pandanus utilis Bory), were examined bi-weekly in the
same study area for the presence of O. rufifascia.

RESULTS AND DISCUSSION

Descriptions of Nymphs

Nymphs are green with faint, longitudinal orange stripes and posterior filaments
of white wax. Specimens turn white to stramineous when preserved in alcohol.

First instar (Fig. 1). Length 1.41 ± 0.018; thoracic length 0.60 ± 0.003; width
0.58 ± 0.006.

Form elongate, depressed, widest across mesothorax; body white (when preserved
in alcohol), some specimens with posterior tergites tinged light brown.

Vertex rounded anteriorly, obscure pits in anterolateral comers, overlapped by
pronotum posteriorly. Frons ovoid, length subequal to width; anterior margin convex,
frontoclypeal juncture concave; lateral margins carinate (outer carinae), outwardly
convex, and paralleled by pair of inner carinae ca. % distance from midline to outer
carinae; row of pits between each inner and outer carina. Clypeus narrowing distally;
consisting of subconical basal postclypeus and beaklike cylindrical distal anteclypeus.
Beak 3 -segmented, extending to base of metacoxae; segment 1 covered by anteclypeus,
segments 2 and 3 subequal. Eyes red. Antennae 3-segmented; scape ringlike; pedicel
subcylindrical, ca. 2 x length of scape; flagellum whiplike distally with bulbous base
slightly longer than pedicel.

Thoracic nota divided by longitudinal middorsal line into 3 pairs of plates. Prono-
tum rounded anteriorly, overlapping vertex, posterior margin sinuate; each plate with

1984

NYMPHS OF ORMENARIA

309

Figs. 1-5. O. rufifascia nymphs. 1. First instar. 2. Second instar. 3. Third instar. 4. Fourth
instar. 5. Fifth instar. Vertical bars = 0.5 mm.

weak, outwardly curved carina forming anterior margin then curving posterolaterally
and bordered medially by row of ca. 9 obscure pits. Mesonotum with median length
ca. 1 V2 x that of pronotum, posterior margin slightly sinuate; each plate with 2 obscure
pits in lateral V2. Metanotum with median length subequal to that of mesonotum,
posterior margin slightly curved. Pro- and mesocoxae elongate, posteromedially di-
rected; metacoxae globose, fused to metastemum giving appearance of being sub-
rectangular and transverse. Metatrochanters each with row of small teeth on medial
aspect. Remaining segments of legs with very fine setae. Metatibiae with transverse
row of 4 black-tipped spines at apex on ventral aspect. Each tarsus with 2 tarsomeres;
pro- and mesotarsomere 1 wedgeshaped; metatarsomere 1 cylindrical with transverse
row of 4 black-tipped spines at apex on ventral aspect; tarsomere 2 of all tarsi
subconical, curved, with pair of small claws and median pulvillus at apex.

Abdomen 9-segmented, segments 1-7 visible dorsally, segments 8-9 telescoped
anteriorly. Tergites 3-7 curving around lateral margins to ventral aspect. Very obscure
pits present on some tergites. Tergite 7 notched medially on posterior margin. Each
segment with the following number of waxpads on either side of midline: segment
6 with 1 small, obscure, whitish-yellowish oval waxpad on tergite, segments 7-8 each
with 1 elongate, oval, caudal, whitish waxpad. Segment 9 without waxpads, elongate

310

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

vertically, surrounding anus, with small, ventral, fingerlike process on either side of
midline.

Second instar (Fig. 2). Length 1.79 ± 0.035; thoracic length 0.83 ± 0.007; width
0.79 ± 0.007.

Frons with length ca. % x its width. Antennae with bulbous portion of flagellum
ca. V2X length of pedicel.

Pronotal plates each with 2 rows of obscure pits bordering carina; ca. 1 5-20 total
pits on plate. Mesonotal median length ca. 2 ¥2 x that of pronotum; each plate with
group of ca. 5 obscure pits midway between midline and lateral margin and ca. 2
pits near lateral margin. Metatibiae with 1 black-tipped spine in distal % of lateral
aspect of shaft and transverse row of 5 black-tipped spines at apex on ventral aspect.
Metarsomere 1 with transverse row of 5 black-tipped spines at apex on ventral aspect.

Abdominal tergite 6 apparently with 2 small, very obscure, whitish, oval waxpads
on each side in lateral ¥2.

Third instar (Fig. 3). Length 2.54 ± 0.059; thoracic length 1.22 ± 0.006; width
1.20 ± 0.010.

Frons length ca. V4 x its width.

Pronotal plates each with 2-3 irregular rows of obscure pits bordering carina; ca.
25-30 total pits on plate. Mesonotal plates each with group of ca. 10 obscure pits
midway between midline and lateral margin and ca. 4 pits near lateral margin.
Metanotal median length ca. % that of mesonotum, with ca. 3 pits near lateral margin.
Metatibiae with 2 black-tipped spines in distal ¥2 of lateral aspect of shaft and
transverse row of 6 (rarely 5) black-tipped spines at apex on ventral aspect.

Abdominal tergites 3-6 each with ca. 5 pits laterally on each side; tergite 6 with
3 small, obscure, whitish, oval waxpads on each side in lateral ¥2.

Fourth instar { Fig. 4). Length 3.89 ± 0.076; thoracic length 1.75 ± 0.013; width
1.88 ± 0.020.

Antennae with bulbous portion of flagellum ca. ¥3 x length of pedicel.

Pronotal plates each with 3 irregular rows of obscure pits bordering carina; ca. 35-
40 total pits on plate. Mesonotal plates each with group of ca. 12-15 obscure pits
midway between midline and lateral margin; pits near margin apparently absent;
wingpad distinctly lobate and covering ca. V2 of metanotum laterally. Metatibiae with
3 (rarely 2) black-tipped spines on lateral aspect of shaft. Metatarsi with 3 tarsomeres;
tarsomere 1 with transverse row of 7 black-tipped spines at apex on ventral aspect;
tarsomere 2 with 1 black-tipped spine on each side at apex on ventral aspect; tar-
somere 3 similar to terminal tarsomere of previous instars.

Abdominal tergite 6 with 4 small, obscure, whitish, oval waxpads on each side in
lateral ¥2.

Fifth instar (Fig. 5). Length 4.68 ± 0.117; thoracic length 2.36 ± 0.017; width
2.88 ± 0.061.

Antennae with bulbous portion of flagellum ca. ¥4 x length of pedicel.

Pronotal plates each with 3-4 irregular rows of obscure pits bordering carinae; ca.
40-45 total pits on plate. Mesonotal plates each with 3 obscure pits on lateral ¥3;
wingpad extending to apex of metanotal wingpad and to third abdominal tergite.

Abdominal tergites 3-6 with pits generally more numerous. Tergite 6 with 5 small,
obscure, whitish, oval waxpads on each side in lateral ¥2.

1984

NYMPHS OF ORMENARIA

311

Fig. 6. Nymphs in life illustrating waxy exudate.

Key to Nymphal Instars*

1. Metatarsi with 2 tarsomeres (Figs. 1-3) 2

- Metatarsi with 3 tarsomeres (Figs. 4, 5) 4

2. Metatibiae lacking spines on shaft, with an apical row of 4 spines; pronotal carinae
bordered by 1 row of pits; abdominal tergite 6 with 1 oval waxpad on each side (Fig.

1) First instar

- Metatibiae with 1 or more spines on shaft and an apical row of 5-6 spines; pronotal

carinae bordered by 2 irregular rows of pits; abdominal tergite 6 with 2-3 waxpads on
each side (Figs. 2, 3) 3

3. Metatibiae with 1 spine on shaft; abdominal tergite 6 with 2 waxpads on each side

(Fig. 2) Second instar

- Metatibiae with more than 1 spine on shaft; abdominal tergite 6 with 3 waxpads on

each side (Fig. 3) Third instar

4. Mesonotal wingpads covering ca. Vi of metanotum laterally; abdominal tergite 6 with

4 waxpads on each side (Fig. 4) Fourth instar

- Mesonotal wingpads extending to apex of metanotum; abdominal tergite 6 with 5

waxpads on each side (Fig. 5) Fifth instar

This key will also work for nymphs of A. septentrionalis, M. pruinosa, and O. venusta.

3 1 2 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Table 1. Plants associated with Ormenaria rufifascia and used in rearing experiments.

Taxon

Common name

Reference

Adults

col-

lected

Support

nymphal

devel-

opment

Arecaceae

Carpentaria acuminata

Carpenter palm

This study

No

No

(H. Wendl. & Drude) Becc.

Caryota mitis Lour

Fish tail palm

This study

No

No

Cocos nucifera L.

Coconut palm

This study

Yes

No

Co/pothrinax wrightii
Griseb & H. Wendl.

Barrel palm

Metcalf and Bruner
(1948)

Yes

—

Dictyosperma album (Bory)

Hurricane palm

This study

No

No

H. Wendl. & Drude

Hcterospathe data Scheff.

Sagisi palm

This study

No

No

Latania loddigesii Mart.

Blue latan palm

Mead (1965)

Yes

—

Latania lontaroides (Gaertn.)

Red latania palm

This study

Yes

Yes

H. E. Moore

Livistonia chinensis (Jacq.)

Chinese fan palm

Mead (1965)

Yes

—

R. Br. ex Mart.

Phoenix canaricnsis Hort.

Canary Islands
date palm

Mead (1965)

Yes

—

Phoenix dactylifera L.

Date palm

This study

No

No

Phoenix roebelenii O’Brien

Pigmy date palm

This study

No

No

Pritchardia eriostachya Becc.

Fan palm

This study

No

No

Pritchardia sp.

Fan palm

Mead (1965)

Yes

—

Ptychosperma nicolai

This study

No

No

(Sand, ex Andre) Burret

Sabal etonia Swingle

Scrub palmetto

Mead (1965)

Yes

—

Saba/ palmetto (Walt.) Lodd.

Cabbage palm

Mead (1965)

Yes

Yes

Serenoa repens (Bartr.) Small

Saw palmetto

Mead (1965)

Yes

—

Veitchia merrilli (Becc.)

Christmas palm

This study

Yes

No

H. E. Moore

Washingtonia robust a H. Wendl.

Washington palm

Mead (1965)

Yes

-

Orchidaceae

Cattleya sp.a

Cattleya orchid

Mead (1965)

Yes

-

Pandanaceae

Pandanus utilis Bory

Screw pine

This study

No

No

Moraceae

Ficus sp.a

Fig

Mead (1965)

Yes

-

Rosaceae

Rosa sp.a

Rose

Mead (1965)

Yes

-

Unconfirmed host record (Mead, 1965).

1984

NYMPHS OF ORMENARIA

313

Table 2. Duration (in days) of the 5 nymphal instars and adults of Ormenaria rufifascia on
Sabal palmetto in south Florida from January to June, 1983.

Instar

Range

Mean ± SD

# observed

1

18-28

23 ± 2.2

36

2

17-25

20 ± 2.0

28

3

15-26

19 ± 2.3

24

4

19-28

24 ± 1.9

21

5

5-10

8 ± 1.1

20

Adult

8-16

12 ± 1.6

20

Comparisons of Species of Flatid Nymphs

O. rufifascia is the fifth U.S. flatid in which the immatures have been described
and illustrated. Immatures of the five species can be separated by color (in life; all
turn white when preserved in alcohol) and external morphology. Identification by
association with adults when collecting is risky since at least 3 species feed in mixed
species feeding assemblages (Wilson and McPherson, 1980).

Live specimens of O. rufifascia and O. venusta are both green but O. rufifascia has
orange longitudinal stripes whereas O. venusta has white stripes; the other 3 species
are white in life. Preserved specimens can be separated by the following features.
Late instar A. septentrionalis and Cyarda sp. (near acutissima) generally bear dark
markings on the mesonotal wingpads, A. septentrionalis has a j -shaped waxpad on
each side of abdominal segment 6 and c-shaped waxpads on abdominal segment 7.
The waxpads on segments 6 and 7 are all ovoid on Cyarda sp. and the other species.
O. venusta bears small, dark marks laterally on abdominal tergites 6-7, which are
absent in the remaining species. M. pruinosa and O. rufifascia are the most mor-
phologically similar. M. pruinosa is smaller, the vertex is truncate anteriorly, bears
a weak median carina on the frons, and has ca. 6-8 weakly developed teeth on the
inner margin of each metatrochanter. O. rufifascia is larger, has a broadly rounded
vertex, lacks a median carina on the frons, and bears ca. 1 0 strongly developed teeth
on the inner margin of each metatrochanter.

Field study. Ormenaria rufifascia adults have been collected or observed on 12
species of palms and non-palms (Metcalf and Bruner, 1948; Mead 1965; Table 1),
but tests have never been conducted to determine if these or any other palms are
breeding and/or feeding hosts. During a 2-year period (1982-1983), 14 species of
palms and 1 non-palm species (Table 1) were examined in Fort Lauderdale and only
cabbage palm, S. palmetto and red latania palm, L. lontaroides, supported nymphal
development (Table 1). Cabbage palm is a native North American plant distributed
throughout Florida and is a common host for O. rufifascia. Red latania palm is an
uncommon introduced ornamental palm and is not an important host for O. rufi-
fascia.

Field observations and cage rearing studies indicate that O. rufifascia is univoltine
with five nymphal instars (Table 2). Numerous dissections of palm leaves were made

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in an effort to discover oviposition sites. Eggs were never found. However, females
probably insert eggs in palms because they have a sawlike ovipositor similar to those
of flatids known to deposit eggs in plant tissues (Wilson and McPherson, 1981). The
oviposition host is probably cabbage palms, because first instars were always found
on this plant. Females were dissected in order to obtain eggs however none was found.
As with other North American flatids (Wilson and McPherson, 1981), eggs are prob-
ably laid in summer, and hatch in January. Neither immatures nor adults have ever
been found on palms from July to December in south Florida. The average instars
(1-5) durations ( x ± SD) in the field were 23 ± 2.2, 20 ± 2.0, 19 ± 2.3, 24 ± 1.9,
and 8 ± 1.1 days, respectively (Table 2). First instars were detected the second week
of January. Adults emerged during the second week of May; mean adult longevity
was 12 ± 1.6 days, with the last adults found in late June. Average monthly tem-
peratures for January through June were 62.3, 65.0, 65.5, 65.8, 68.8, 75.1, and 77.9°F,
respectively. The relatively rapid development of fifth instars may have resulted from
the higher May temperatures.

Early instar nymphs exhibit gregarious and sedentary behavior near the basal
portions of pinnae. Nymphs were never observed on the upper surfaces of palm
leaves. The numbers of nymphs per aggregation ranged from 5 to 2 1 . A white wax
excreted by nymphs and presumed to serve a protective function (Hepburn, 1967)
was always associated with nymphal aggregations. As nymphal development pro-
gressed, later instars tended to disperse and were often found singly on the undersides
of leaves. Adults were often found near nymphal aggregations, but were rarely in
groups of more than 3 individuals.

Occasionally large nymphs were observed with drops of honeydew attached to the
posterior end. Nymphal aggregations were tended by 1 ant species, Camponotus
floridanus Buckley. When disturbed, nymphs jumped up to 80 cm and away from
aggregations, adults jumped and flew erratically while producing audible vibrations.

ACKNOWLEDGMENTS

We express our appreciation to D. G. Burch and H. M. Donselman, University of Florida,
AREC, Fort Lauderdale, for their assistance in identification of palm species. We thank J. C.
Nickerson at Florida Department of Agriculture and Consumer Services, Plant Industry Di-
vision, Gainesville and C. R. Thompson, University of Florida, AREC, Fort Lauderdale, for
identifying Camponotus floridanus Buckley. We also thank Mr. Willey Durden, Aquatic Plant
Laboratory, ARS, Southern Region, USDA for producing Figure 6.

LITERATURE CITED

Hepburn, H. R. 1967. Notes on the genus Epiptera (Homoptera: Achilidae). J. Georgia
Entomol. Soc. 2:78-80.

Mead, F. W. 1965. Ormenaria rufifascia (Walker), a planthopper pest of palms (Homoptera:
Flatidae). Ha. Dep. Agric. Entomol. Circ. 37, 2 pp.

Metcalf, Z. P. 1923. A key to the Fulgoridae of eastern North America with descriptions of
new species. J. Elisha Mitchell Sci. Soc. 38:139-230.

Metcalf, Z. P. 1957. General catalogue of the Homoptera. Fasc. IV. Fulgoroidea, Part 13.
Hatidae and Hypochthonellidae, 574 pp.

Metcalf, Z. P. and S. C. Bruner. 1 948. Cuban Hatidae with new species from adjacent regions.
Ann. Entomol. Soc. Amer. 41:63-1 18.

1984

NYMPHS OF ORMENARIA

315

Moore, T. E. 1961. Audiospectrographic analysis of sounds of Hemiptera and Homoptera.
Ann. Entomol. Soc. Amer. 54:273-291.

Wheeler, A. G. and E. R. Hoebeke. 1 982. Host plants and nymphal descriptions of Acanalonia
pumila and Cyarda sp. near acutissima (Homoptera, Fulgoroidea: Acanaloniidae and
Flatidae). Fla. Entomol. 65:341-349.

Wilson, S. W. and J. E. McPherson. 1980. Mixed species feeding assemblages of planthopper
nymphs (Homoptera: Fulgoroidea). Great Lakes Ent. 13:185-187.

Wilson, S. W. and J. E. McPherson. 1981. Life histories of Anormenis septentrionalis, Metcalfa
pruinosa, and Ormenoides venusta with descriptions of immature stages. Ann. Entomol.
Soc. Amer. 74:299-3 1 1 .

Received March 14, 1984; accepted May 24, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 316-341

A REVISION OF THE GENUS

HORRIDIPAMERA MALIPATIL (HEMIPTERA: LYGAEIDAE)

James A. Slater and Le-Yi Zheng

Section of Systematic and Evolutionary Biology,

University of Connecticut, Storrs, Connecticut 06268 and
Department of Biology, Nankai University,

Tianjin, People’s Republic of China

Abstract.— The genus Horridipamera Malipatil is revised. Two new species H. medleri from
West Africa and H. compacta from South India are described. A key to species, a cladogram,
and discussion of phylogenetic relationships are included. All species are diagnosed. Extensive
distributional data are included. A dorsal view drawing of H. medleri and 33 drawings with
details of genitalia are given. There is a discussion of the relationships of Horridipamera to
other genera.

The genus Horridipamera was erected by Malipatil (1978) for three species from
Australia and the Orient. The erection of the genus was part of an effort on Malipatil’s
part to recognize and segregate monophyletic units from the old omnibus genus
Pachybrachius Hahn. He selected the widespread Plociomerus nietneri Dohm as type
species and included two new species {robusta— eastern Australia, New Zealand and
cantrelli — Queensland).

Harrington (1980) included eight additional species, several of which are African.
The genus is thus represented throughout the tropics and subtropics of the Eastern
Hemisphere.

Several of the species are extremely common, almost ubiquitous in tropical areas
yet there is no modern work available to enable a student to readily identify species,
nor is there a discussion of the intra-generic relationships.

We accept, for the present, Harrington’s conclusion that Horridipamera is a mono-
phyletic group although her statement that the broad base of the right conjunctival
spine is a synapomorphy that distinguishes Horridipamera from Paraparomius, Togo
and Eucosmetus is not true for some of the species that she places in the genus
although it is true of nietneri, the type species.

Harrington (1980) erected the genus Stalaria with Pamera ferruginosa Stal as type
species. She included Pachybrachius kisseis Linnavuori and Pachybrachius nysias
Linnavuori in Stalaria. We have examined the lectotype of Stal’s Pamera ferruginosa
from the Stockholm Museum. It is not the species considered to be ferruginosus by
Harrington. It is a species of Horridipamera closely related to H. pullata (Hesse) (see
species discussion.) Stalaria is thus based upon a misidentified type species. The
International Rules (Article 70a) state that such cases are to be referred to the com-
mission asking either that the nominal species be designated as the type species or
that the species named by the designator be selected. In the present case the specimens
believed by Harrington to be ferruginosus are conspecific with Pachybrachius kisseis
Linnavuori. To avoid adding an additional name to the literature (since placing

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REVISION OF HORRIDIPAMERA

317

Stalaria as a junior synonym of Horridipamera would leave kisseis and nysias without
a generic name) we are revising Stalaria and will be asking the commission to set
aside Pamera ferruginosa as the type species of Stalaria and replacing it with
Pachybrachius kisseis Linnavuori. The status of Stalaria thus will at that time become
sub judice and it should be used in the sense of kisseis Linnavuori at least until the
commission has acted.

Diagnosis. Horridipamera can be recognized on the basis of the following char-
acteristics. Generally black and white or brown and white; phallic type II (see Har-
rington, 1980); anterior pronotal lobe somewhat globose, essentially impunctate; a
deeply impressed line separating anterior pronotal collar from remainder of anterior
lobe; male fore tibia usually spined; head slightly prolonged behind eyes; phallus
lacking appendages or outgrowths except for one pair of conjunctival spines and two
vesical marginal processes.

Harrington’s cladogram separates Horridipamera and Paraparomius from such
genera as Togo, Eucosmetus and Paraeucomstus on the most tenuous synapomor-
phies. The latter are “held together” in a clade by having the head broader than the
transverse pronotal impression. However, several species of Horridipamera have this
condition. We believe that the species we have included in Horridipamera do, on
the basis of their derived phallic features (lack of appendages or outgrowths on phallus
other than a pair of conjunctival spines and two vesical mariginal processes), form
a monophyletic unit. We suggest that all members of the genera noted above need
investigation to see if some may also be a part of this clade.

The abbreviations used for collectors are as follows: SS = J. and S. Slater; SSS =
J. and S. Slater, Schuh; SSSS = J. and S. Slater, Schuh, Sweet; BAC = Brink, An-
derson, Cederholm; HDKB = Hevel, Dietz, Karunaratne, Balasooriya; CDHHS =
Cederholm, Danielsson, Hammarstedt, Hedqvist, Samuelsson. Locality: K.N.P. =
Kruger National Park.

All measurements are in millimeters.

PHYLOGENETIC RELATIONSHIPS

We consider the following conditions to be derived in Horridipamera.

1. Asymmetrical conjunctival spines.

2. Bifid left conjunctival spine.

3. T-shaped outgrowth near proximal end of seminal duct.

4. Separation of distal and proximal vesical processes.

5. Loss of serrations on distal vesical processes.

6. Reduction of proximal vesical process.

7. Possession of long hairs on the dorsal surface.

8. Loss of spine near middle of male fore tibiae.

9. Elongation of the legs.

10. Narrow body shape.

1 1 . Strongly produced eyes.

12. Strongly swollen anterior pronotal lobe.

13. Striped scutellum.

14. Possession of four evenly spaced spines near distal end of male fore tibia.

15. Apical vesical process saw-shaped.

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16. Proximal portion of proximal vesical process broadened and plate-like.

17. Loss of distal vesical process.

In the cladogram (Fig. 1) we have relied heavily upon features of the phallus.
Unfortunately many of what appear to be synapomorphies are reduction phenomena.
Most of the reduction and loss is in the vesical processes. Where reduction and loss
occur, the remaining portions of the vesical processes have a very similar appearance
which we consider unlikely to have been the case if these reductions were homoplasies.
Where the apical vesical process is completely lost homoplasy is as likely as syn-
apomorphy.

Species of Horridipamera belong to Harrington’s (1980) “genitalia type II.” In this
clade the plesiomorphic condition of the vesical processes is one in which there was
a continuous tape-like sclerotization of the vesicular membrane (Figs. 10, 12, 13).
A polarity sequence appears to be present in the following manner. First, separation
of the tape-like vesicular sclerotization into two separate, but still elongate and
twisted, processes (and subsequently into three as in species of Pseudopachybrachius).
The next stage in the sequence, that is observable in Horridipamera at least, is the
reduction of the distal process into a slender non-serrated strip (Fig. 8) and the
subsequent complete loss of the distal process. The proximal process becomes reduced
to form first an elongate slender twisted structure, then is reduced to a short somewhat
“C-shaped” but still serrate-margined condition (Fig. 9).

Our other derived conditions are based largely on out-group comparison and
include such features as the long hairs on the dorsal body surface, loss of the male
fore tibial spines, striped scutellum, etc.

The more difficult areas to separate homoplasies from synapomorphies are in the
elongation and narrowing of the body, elongation of the legs, increase in the size and
globosity of the anterior pronotal lobe, and shape of the head. It is reasonably clear
from out-group comparison that these are derived features but again and again in
the Myodochini one sees a tendency to develop degrees of ant mimicry with which
the above features are frequently associated. Thus in the cladogram we have used
these chiefly as autapomorphies.

key to species of Horridipamera

1 . Male fore tibia lacking a distinct median spine 2

la. Male fore tibia with one or more conspicuous spines present midway along shaft or

on distal half 3

2. No erect long hairs present on pronotum and scutellum; explanate lateral area of

corium uniformly pale except for dark apex; antennal segments I, II, and III black
or very dark castaneous; head width as great as or greater than width of anterior
pronotal lobe; body relatively slender (Ceylon) emersoni

2a. Pronotum and scutellum with numerous conspicuous long, erect hairs present; ex-
planate corial margin with a dark spot that reaches lateral margin at level just caudad
of end of claval commissure; antennal segments I, II, and III chiefly pale yellow,
only distal ends of segments II and III dark; width of anterior pronotal lobe greater

than width of head across eyes; body comparatively stout (S. India) compacta

3. Male fore tibia with 3-4 small equidistantly placed spines on distal V2 cantrelli

3a. Male fore tibia armed near middle with 1 or 2 large spines 4

4. Hemelytra pilose, with long and erect hairs (view laterally) 5

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REVISION OF HORRIDIPAMERA

319

Fig. 1 . Cladogram of species of Horridipamera.

4a. Hemelytra not pilose, without conspicuous long and erect hairs (but sometimes with

elongate hairs present on pronotum and scutellum) 7

5. Anterior lobe of pronotum huge, in males at least twice as long as posterior lobe;

fourth antennal segment with a pale basal annulus (Australia) robusta

5a. Anterior lobe of pronotum smaller, in males less than twice as long as posterior lobe;

fourth antennal segment with or without a pale basal annulus 6

6. Hairs on antennal segments II and III twice as long as diameter of segments; de-

cumbent hairs on hemelytra slender, silky not flattened nor somewhat scale-like;
anterior lobe of pronotum more swollen and globular frequently higher than posterior
lobe; fourth antennal segment uniformly dark; vesica with a reduced ridge-like distal
process and an elongate proximal process fading gradually at one end (Figs. 5, 8)
(Africa) bergrothi

6a. Hairs on antennal segments II and III shorter, subequal to diameter of segment;
decumbent sericeous hairs on hemelytra shorter and broader, somewhat scale-like;
anterior lobe of pronotum relatively narrow, rarely higher than posterior lobe; fourth
antennal segment often with a pale basal annulus; vesica only with proximal process,
this semi-oval, flap-like, margin inconspicuously toothed (Fig. 9) (Oriental and Aus-
tralian) nietneri

7. Scutellum with a pair of oblique brown stripes. A somewhat angulate sclerotized
outgrowth present on proximal part of ejaculatory duct (Fig. 7); punctures on pale
macula before dark apex of corium dark, of same color as other punctures on corium

(Asia and Africa) inconspicua

7a. Scutellum uniformly black or chocolate brown; ejaculatory duct without a distinct

outgrowth proximally; punctures on pale corial macula either pale or dark 8

8. Fourth antennal segment with a conspicuous pale basal annulus perlonga

8a. Fourth antennal segment uniformly dark 9

9. Body relatively broad and stout, total body length less than four times width of
pronotum across humeral angles; eyes not strongly protruding; width of head across
eyes not greater than maximum width across anterior pronotal lobe; middle and hind

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femora darkened only on distal V3 to V2; hind tibiae largely pale, infuscated only near

proximal and occasionally extreme distal ends perlonga

9a. Body more elongate, total body length at least four times as great as width across
pronotal humeral angles; eyes more prominently produced, width across eyes some-
time greater than maximum width across anterior pronotal lobe, if not anterior
pronotal lobe strongly swollen ( medleri ); degree of dark coloration on hind femora
variable; hind femora usually entirely dark, occasionally only darkened on proximal
V2 to %; hind tibiae usually infuscated (dark brown) for entire length 10

1 0. Width across eyes less than, or subequal to, maximum width across anterior pronotal

lobe; latter strongly swollen and appearing globose; ground color of hemelytra pale
with infuscated dark stripes and dashes medleri

1 0a. Width across eyes greater than maximum width across anterior pronotal lobe; anterior
pronotal lobe relatively slender and linear, not appearing globose; ground color of
hemelytra dark brown with a strongly contrasting white subapical corial macula ... 11

1 1. Head, pronotum, and scutellum chocolate or very dark red brown; white subapical

corial macula strongly contrasting with ground color of corium pullata

1 la. Head, pronotum, and scutellum light reddish; subapical corial macula only weakly

differentiated from ground color of corium ferruginosa

Horridipamera inconspicua (Dallas)

Rhyparochromus inconspicuus Dallas, 1852, p. 547.

Diplonotus rusticus Scott, 1874, pp. 430-431. New Synonymy.

Pamera spinicrus Reuter, 1882, pp. 17-18. New Synonymy.

Pamera ebenaui Reuter, 1887, pp. 96-97 . New Synonymy.

Pachybrachius inconspicuus Slater, 1964a, p. 1 127.

Horridipamera inconspicuus Slater, 1979, p. 22.

Diagnosis. Body relatively short and stout. Scutellum dark red-brown with a con-
spicuous pair of light orange to light reddish brown oblique streaks on basal half.
Hemelytra chiefly pale yellow with punctures dark brown. Pale macula at inner corial
angle large, very conspicuous by virtue of dark streaks extending anteriorly and
posteriorly from the pale macula. Usually a dark apical corial macula and a small
dark spot at edge of explanate lateral margin (but not reaching margin) at level of
inner pale corial macula. Thus corium appearing largely pale and subapical pale
macula not strikingly differentiated. Pronotum reddish brown with humeral angles,
and often extreme posterior margin, pale yellow. Fore femora dark reddish brown,
with pale distal ends. Usually middle femora pale and hind femora with a narrow
subdistal dark macula. Antennae variable in color, segments II and III usually pale
yellow; segment IV dark sometimes with a pale subbasal annulus present. Second
labial segment white or pale yellow. Membrane pale with irregular darker spots and
streaks scattered throughout.

Male fore tibiae each with a distinct spine midway along shaft. Dorsal surface
lacking numerous elongate upstanding hairs but with decumbent sericeous pubes-
cence. Width of anterior pronotal lobe slightly greater than width of head across eyes
and conspicuously swollen. Labium reaching at least to middle of mesostemum.

Sperm reservoir with wedge-shaped bulb, very broad proximally, tapered distally;
wings strongly bent ventrad and produced into a narrow elongate blunt projection
(Fig. 7). Vesical process of phallus interconnected, elongate, complexly twisted, evenly

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REVISION OF HORRIDIPAMERA

321

serrated (Fig. 13). Conjunctival spines as in Figures 28, 29. Paramere with elongate
acute inner projection, inner margin proximal of projection produced into a broad
flap-like flange, outer projection short, broad, blade thick (Fig. 35). Ejaculatory duct
with a peculiar bent, sclerotized projecting outgrowth near proximal end (Fig. 7).

As might be expected in a species with such a wide range there is considerable
variation in color. The scutellar streaks while usually diagnostic may be much reduced
in dark specimens and in unusually pale specimens may not be appreciably differ-
entiated from the color of the remainder of the scutellar surface. However, the
combination of pale corium in which the distal light macula is little differentiated,
relatively pale membrane, stout body, etc., make this a generally readily recognizable
species without recourse to the definitive differentiating characters of the male gen-
italia.

The correct name to apply to this species has been a matter of difficulty. We (Zheng
and Slater, 1984) have selected a specimen of this species from West Africa as the
neotype of Rhyparochromus inconspicuus Dallas (see discussion in that paper) which
is the oldest available name.

First instar nymph. South Africa: Pretoria, Transvaal, 18.XII.1967 (SSS). (Reared
from eggs laid by identified female.) Head and pronotum dull orange. Meso- and
metanota gray with reddish borders. Abdomen dull orange red shading to red pos-
teriorly along intersegmental sutures and laterally. Sclerotized areas about scent gland
orifices on terga 3-4, 4-5, 5-6 gray with larger pigmented area posterior to suture
than anterior to it. Legs uniformly dull white. Antennal segments I, II, III white
tinged with reddish. Fourth segment a strongly contrasting orange-red. Length head
0.37, width 0.34, interocular space 0.24. Length pronotum 0.15, width 0.34. Length
mesonotum 0.10, width 0.37. Length metanotum 0.07, width 0.39. Length abdomen
0.61. Length labium 0.73, slightly exceeding posterior margin of metacoxae. Length
antennal segments I 0.10, II 0.17, III 0.17, IV 0.32. Total body length 1.20.

Second instar nymph. (As above.) Head and anterior lh of pronotum dark brown.
Ground color of remainder of body dull yellow with a narrow dull brown longitudinal
stripe on either side of midline and an oblique irregular streak extending dorso-
laterad near middle of segment on each side.

Abdominal terga 1 , 2 and anterior Vi of 3 dull gray brown. Remainder of abdomen
with dark central markings similar to those of instar 5. Abdomen with red striping
along intersegmental sutures. Dark sclerotized areas about scent gland orifices as well
developed anterior to orifices as posterior to them; sclerotization between terga 3-4
somewhat wider than that between terga 4-5 or 5-6. Femora conspicuously tinged
with red on distal halves. Antennal segment I and proximal Vi of II white, former
with a narrow, dark, basal annulus. Distal lh of antennal segment II and all of segments
III and IV red.

Length head 0.5 1 , width 0.49, interocular space 0.29. Length pronotum 0.24, width
0.49. Length mesonotum 0.17, width 0.54. Length metanotum 0.10, width 0.63.
Length abdomen 0.98. Length labium 0.98, reaching well between metacoxae. Length
antennal segments I 0.15, II 0.24, III 0.22, IV 0.43. Total body length 1.95.

Fifth instar nymph. South Africa: 1 7 mi NE Pretoria, Pienaars River Dam 1 .XI. 1 967
(SSS). General coloration bright tan, or light brown interspersed with light yellow as
follows: a longitudinal stripe running completely through pronotum on either side
of midline and a short longitudinal vitta on posterior pronotal lobe slightly mesad

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of humeral angles; a series of oblique stripes on scutellum and mesothoracic wing
pads giving a striped appearance to these areas; distal ends of femora, tibiae, first
tarsal segments and second and third antennal segments pale yellow.

Color of abdomen complexly variegated: ground color pale yellow, an elliptical
brown macula present around dorsal scent gland openings between terga 3-4, 4-5
and 5-6, that between 3-4 broader than succeeding. Tergum 2 dark brown; tergum
3 mesally chiefly brown but this coloration extending posteriorly to scent gland
macula only as a pair of narrow bars and a mesal convex extension from the main
darkened central area; tergum 4 with an ovoid central dark patch, enclosed on either
side by a pincer-like dark band that coalesces posteriorly with macula surrounding
terga 4-5 scent gland orifices; terga 5, 6 and 7 each with a dark central macula, that
on tergum 5 especially large. Oblique dark rays extending cephalo-laterad from dark
central areas of each segment, sutures between segments also dark. Explanate lateral
margins of pronotum and wing pads pale yellow with narrow dark brown outer edges.
First antennal segment pale yellow with a strongly contrasting dark brown basal
annulus. Fourth antennal segment uniformly reddish brown. Femora and second
tarsal segment dull brown.

Head slightly declivent. Tylus attaining middle of first antennal segment. Vertex
moderately convex. Epicranial suture with stem present but very short. Length head
0.81, width 0.90, interocular space 0.56. Lateral pronotal margins narrowing evenly
but moderately from humeral angles to anterior margin; both anterior collar and
transverse impression conspicuous. Anterior pronotal lobe much larger and more
convex than posterior. Length pronotum 0.79 (length anterior pronotal lobe 0.59,
length posterior pronotal lobe 0.20), width pronotum 1.07. Mesothoracic wing pads
broad, reaching to or almost to suture between abdominal terga 3-4. Length meso-
thoracic wing pads 1.10. Length abdomen 1.76. Fore femur generally with 2 or 3
spines on inner rank and a single spine on outer. Labium extending posteriorly to
or between mesocoxae. Length labial segments I 0.29, II 0.37, III 0.56, IV 0.59.
Length antennal segments I 0.32, II 0.73, III 0.61, IV 0.88. Total body length 4.07.

We have examined 3rd and 4th instar nymphs from Meintjies Kop, Pretoria, South
Africa 19.III.1968 (SSS). These closely resemble instar 5 in color.

Biology. This species is abundant in early succession stages of old fields in South
Africa. It is one of the most common of the lygaeid species that comes to lights in
Pretoria where it presumably lives in gardens and weed grown lots. We were unable
to establish any definite breeding host, but on several occasions took adults and
nymphs along roadsides below a variety of weeds and grasses. It apparently is not
entirely, if at all, a grass feeder, since it often occurs on recently burned over veld
with much bare ground and where only scattered forbs produce a seed crop.

Distribution. H. inconspicua is distributed throughout much of the Ethiopian Re-
gion, occurs on at least some of the islands of the Indian Ocean and through the
Oriental Region to China and Japan.

It is considerably less common than perlonga in West Africa. By contrast incon-
spicua is an abundant, almost ubiquitous, species in South Africa. The distributions
of inconspicua and perlonga in fact closely resemble those of Pseudopachybrachius
reductus (Walker) and P. capicola (Stal) where the former is abundant in West Africa,
but scarce in South Africa and capicola the reverse. Cases like these suggest former
allopatry for these disturbed habitat or early succession stage insects. If so the sub-

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REVISION OF HORRIDIPAMERA

323

sequent sympatry is possibly due to disturbance by man that has led to the elimination
of former ecological barriers.

H. inconspicua has previously been reported from: Senegal (Slater and Wilcox,
1973) and (Scudder, 1982-as spinicrus and ebenaui)-, Sierra Leone (Dallas, 1852);
“Guinea” (Lethierry and Severin, 1894— as spinicrus)-, Ghana (Reuter, 1882— as
spinicrus ); Ruanda (Schouteden, 1957); Zaire (Slater, 1972); Sudan (Linnavuori, 1978 —
as spinicrus)-, Tanzania (Scudder, 1962a); Southwest Africa (Schumacher, 1913— as
ebenaui?)-, Cape Verde Is. (Lindberg, 1958— as ebenaui); South Africa (Slater, 1964b);
Madagascar (Reuter, 1887— as ebenaui)-, Rodriguez 1. (China, 1924— as ebenaui)-,
Sri Lanka (Slater, 1979); Japan (Scott, 1874— as rusticus)’, China (Horvath, 1879 —
as nietneri).

Additional material examined. SENEGAL: 4, off Senegal at sea, 25.x. 1957 (Cape-
ner). 1 , Guede, 1 .iii. 1 946 (Risbec). GUINEA: 9, off coast, xii. 1957 (Capener). GHANA:
1, Accra, 14.xi.1969 (Campbell). NIGERIA: 2, Ile-Ife, xi.1969 (Medler). 5, same,

27. 111. 1969. 18, same, 10. iii. 1969. 1, same, 25.iii. 1969. l,Keffi, B.P. State, 23. xii. 1968
(Medler). 7, Bedeggi, NW State, 14.xii. 1974 (Medler). 1, Lagos, Univ. Campus,
ii. 1 975 (Hamid). ST. THOMAS IS.: 1, Neves, Cote N 25 km S Tome, 2 1.x. 1973
(Schmitz). CAPE VERDE IS.: 3, Tiago, Sierra Pico Antonia, 10. ii. 1945 (Panelius).
SUDAN: 2, Blue Nile, Wadi Medari, 1 1-12. xi. 1962 (Linnavuori). 2, Upper Nile,
Malakal, 5-20. i. 1963 (Linnavuori).TANZANIA: 2, Mlingano, 5.xi. 1963 (Robert-
son). 1, same, 13.vi.1963. 2, Ukiriguru, 8.xi.l960 (Robertson). 1, same, 29.iv.1960.

1, Ilonga, 29.ix.1963 (Robertson). 2, same, 3.x. 1963. 7, Tabora, 27.xi.1963 (Rob-
ertson). 2, same, 28. xii. 1963. 1, same, l.i. 1964. BOTSWANA: 2, 2 mi N Gaberones,

21. x. 1974 (Samuel Slater). SOUTH AFRICA: Transvaal: 1, 20 mi SW Rustenburg,
20.x. 1974 (Samuel Slater). 1, Rustenburg, 4.xii.l950 (Capener). 2, same, 4.xii.l951.
3, 10 mi N turnoff, 19 mi E Groot Marico, 22.x. 1974 (Samuel Slater). 18, Pretoria,

22. x. 1967 (SSS). 79, same, 23.x. 1967. 2, same, 27.x. 1967. 21, same, 2.xi.l967. 2,
same, 3.xi.l967. 2, same, 5.xi. 1967. 1, same, 6.xi. 1967. 1, same, 7.xi.l967. 1, same,
22.xi.1967. 1, same, 24.xi.1967. 3, same, 25.xi.1967. 6, same, 27.xi.1967. 3, same,
xi.1967. 1, same, 4.xii.l967. 9, same, 5.xii.l967. 6, same, 9.xii.l967. 3, same, 3.L1968.

2, Nat. Botanical Garden, 24.xi. 1967 (SSS). 3, Fountains, 28.x. 1968 (SSS). 2, Irene,
Smut’s Farm, 1 0.i. 1 968 (SSSS). 6, Meintjies Kop, Pretoria, 22.x. 1967 (SSS). 1, same,

18. 111. 1968. 1, same, 19.iii.1968. 1, same, 29.x. 1967. 1, Lyttleton, 26.xii.1967 (SS).
1, same, 17.xii.1967. 2, same, 18.xii.1967. 1, same, 12.i.l968. 1, same, 19.L1968.
1, same, 26.ii.1968. 2, same, 29. ii. 1968. 1, same, 20.xi.1968. 1, 10 mi N Mookeetsi,
13.xii.1967 (SSS). 1, Dendron, xi. 197— (van Ark). 5, Hartebeesport Dam, 20 mi W
Pretoria, 30.x. 1967 (SSS). 1, 14 mi NE Potgietersrus, 26.x. 1967 (SSS). 3, Mariepskop
nr. Klaserie, 6,300 ft, 30.xi.1967 (SSS). 1, 10 mi SSW Skukuza (K.N.P.), 26.iv.1968
(SSSS). 1, 10 mi N Satara Camp (K.N.P.), 28.iv.1968 (SSSS). 1, 3 mi E Satara Camp,
Nwanedzi River (K.N.P.), 29.iv. 1968 (SSSS). 2, 9 mi N Pietersburg, 26.x. 1967 (SSS).
9, 15 mi SSW Pietersburg, 25.x. 1967 (SSS). 1, Pietersburg, 26.x. 1967 (SSS). 6, base
Magoebaskopf 4,000 ft, 1 2.xii. 1 967 (SSS, Munting). 1 , 6 mi N Warmbaths, 7. xii. 1 967
(SSS). 1, Zoutpansberg, 4,500 ft, 5 mi N Louis Trichardt, 8.V.1968 (SSSS). Cape
Province: 1, Van Zylsrust, Kuruman River, 2.xi.l967 (Coaton). 6, Kimberley, 17-
1 8.i. 1 968 (SSSS). 2, 31 mi NE Calvinia, 10.x. 1974 (S. Slater & Ecker). Orange Free
State: 10 mi Petrus Steyn to Reitz, 27.xii.1967 (SSS). 1, Lindly, xii. 1951 (Weber).
Natal: 10 mi W Paulpietersberg, 7.xi.l967 (SSS). CEYLON: 2, Anu. Dist. Wildlife

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Bungalow, Hunuwilagama, Wipattu, 200 ft, 1 0-1 9.iii. 1970 (Davis and Rowe). 1,
Kan. Dist. 5 mi NW Maheyangana, 30.iii-l.v. 1971 (Spangler). 2, Tri. Dist., China
Bay, 200 ft, 9-1 l.ix. 1976 (HDKB). INDIA: 1, Pulney Hills, Kodiakanal, 6,500 ft,
v.1953 (Nathan). PHILIPPINES: 1, Mt. Maquiling, Luzon, 15.xi.1947 (Gines). JA-
PAN: 2, Nishinomiya, P.R. Settsu, 27.xi.1952 (Nakanishi).

We have dissected specimens from South Africa, Nigeria, Tanzania, Japan and
Ceylon. No significant differences have been found.

Horridipamera cantrelli Malipatil
Horridipamera cantrelli Malipatil, 1978a, pp. 94-96.

Diagnosis. It is readily distinguishable from all species of Horridipamera except
robusta by the presence of four small equidistantly spaced spines on the distal half
of each male fore tibia.

H. cantrelli is closely related to nietneri as is indicated by the lack of the distal
vesical process and the reduction of the proximal process, the quadrate sperm res-
ervior wings that only curve moderately posteriorly and the similarly shaped para-
meres.

H. cantrelli has a pale basal annulus on the fourth antennal segment, pale pronotal
humeri, spotted fore femora and chiefly pale hemelytra. The coloration of the hem-
elytra is very similar to that of inconspicua. The known specimens range in length
from 5.5 to 6.2 mm.

Malipatil’s (1978) description and figures are excellent. The spotted fore femora
lack of a greatly swollen anterior pronotal lobe and the characteristic fore tibial (male)
spines should enable the species to be readily recognized.

Distribution. AUSTRALIA: Known only from the type localities in Queensland.

Horridipamera perlonga (Scudder)

Pachybrachius pullatus Slater, 1964b ( nec Hesse), p. 217.

Pachybrachius perlongus Scudder, 1969, p. 173.

Horridipamera perlongus Harrington, 1980, p. 99.

Diagnosis. Elongate, slender. Head, pronotum and scutellum piceus to reddish
brown. Scutellum uniformly dark, lacking oblique reddish brown stripes. Anterior
pronotal lobe impunctate. Clavus and corium lacking upright hairs. Fore femora
except proximal and distal ends chocolate brown. Middle and hind femora pale
yellow with a broad subdistal dark annulus. Antennal color variable from nearly
uniformly dark to chiefly pale yellow on segments I, II, and III. Fourth antennal
segment frequently with a yellow basal annulus. Males with a spine present near
middle of each anterior tibia. Hemelytra usually with a conspicuous subdistal white
corial macula, lateral explanate corial margin pale. Labium at least attaining middle
of mesostemum, first segment remote from base of head, second segment usually
white or pale yellow. Membrane usually dark with veins white and with a narrow
white apical median stripe.

Sperm reservoir bulb wedge-shaped similar to that of inconspicua but slightly less
narrowed distally. Wings strongly bent downward but evenly narrowed (Fig. 6) not
produced into an elongate narrow projection. Vesical processes of phallus more or

1984

REVISION OF HORRIDIPAMERA

325

less connected, complexly twisted, proximal process saw-like (Fig. 10). Left con-
junctival spine broad and blade-like throughout most of length, slightly bifid and
terminating in an acute spine at distal end (Fig. 20). Right conjunctival spine evenly
tapering (Fig. 21). Paramere with expanded margin proximad of acute inner projec-
tion subtruncate with proximal margin concave (Fig. 32).

H. perlonga is a variable species. The fourth antennal segment usually has a pale
proximal annulus, but many specimens, from West Africa particularly, have the
fourth segment entirely dark. The head, pronotum, and scutellum are always dark
(though varying from almost black to reddish brown) whereas the hemelytra are
variable in color. In some specimens the clavus and corium are almost completely
pale with only the large apical corial macula, an area at the inner corial angle and
some of the punctures retaining the dark brown coloration. More commonly the
corium has a broad irregular dark transverse fascia (frequently incomplete), a dark
patch near the base and the clavus strongly infuscated with brown so that only the
proximal Vi of the cubital vein, an irregular basal area and small “dash” opposite
the claval commissure is pale.

There is, as in many myodochines, considerable variation in the relative length
and degree of convexity of the anterior pronotal lobe. The difference in the length
of the antennae is remarkable. Thus when one compares an elongate slender, pale
specimen with a short robust dark one it is difficult to believe that only a single
species is involved. However, all intermediate conditions are present in the study
series, and there are no differences in the male genitalia. We conclude that a single
species is represented.

Biology. This is an uncommon species in South Africa where much of the African
field work of the senior author has been concentrated. On May 1, 1968 we took
adults and nymphs of perlonga under a creeping forb with many seeds present that
was growing below a large Natal mahogany tree ( Trichilia emetica Vahl.). This was
a disturbed habitat, much trampled by hippopotami, near Letaba Camp, Kruger
National Park, South Africa.

Fifth instar nymph. South Africa: Kruger National Park 4 mi E Letaba Camp,
Letaba River, l.V. 1968 (SSSS). Very unlike inconspicua in general coloration. Head
and pronotum dark chocolate brown, the latter nearly unicolorous, lacking any in-
dication of pale longitudinal stripes near midline; scutellum and wing pads dark the
latter very obscurely marked; a pair of faint diagonal paler streaks on scutellum and
a small pale yellow macula anteriorly near lateral margin of each wing pad and a
second adjacent on explanate flange of wing pad. Ground color of abdomen dull red.
Dark areas in central portion of terga 3-7 similar to inconspicua but tending to form
a broad median dull brown stripe; tergum 4 mesally with only a spot in center but
with a broad dark transverse fascia that covers almost entire tergum laterad of scent
gland orifices. All terga darkened laterally. Abdomen below dull reddish with a
strongly contrasting pale yellow oblique band on sterna 4 and 5 running antero-
laterad from meson of sternum 5. Antennae and femora dull reddish; tibiae brown;
first tarsal segment pale yellow, second tarsal segment somewhat infuscated.

Head slightly more formicoid than is that of inconspicua. Tylus reaching middle
of 1st antennal segment. Stem of epicranial suture relatively elongate. Length head
1.20, width 0.90, interocular space 0.54.

Explanate pronotal margins much less strongly developed than are those of in-

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conspicua. Length pronotum 0.85 (length anterior pronotal lobe 0.66, length posterior
pronotal lobe 0.20), width 1.02. Length mesothoracic wing pads 1.36. Length ab-
domen 2.32. Fore femora with 2-3 spines on inner rank, one spine on outer rank.
Labium apparently attaining mesocoxae. Length labial segments I 0.71, II 0.68, III

0. 49, IV 0.37. Length antennal segments I 0.43, II 0.90, III 0.78, IV 0.90. Total body
length 5.18.

The nymphs of H. perlonga and H. inconspicua are extremely different in ap-
pearance. H. perlonga has a much more ant-like facies with its dark coloration, white
banded abdominal venter and long head and legs. H. inconspicua on the other hand
gives no suggestion of ant mimicry and the general appearance is of a pale streaked
and striped insect. The median coloration of the abdominal terga is similar but the
other striking differences in the nymphs suggest that these two species are not closely
related.

Egg. South Africa: K.N.P., 4 mi E Letaba Camp, Letaba River, l.V. 1968 (SSSS).
Egg elongate elliptical, truncate at anterior end (“typical” rhyparochromine shape);
surface lacking numerous hairs. Six short closely set micropylar processes at anterior
pole. Length 1.17, maximum width 0.39.

There does not appear to be a significant difference between the eggs of inconspicua
and perlonga.

Distribution. H. perlonga is a widespread, ubiquitous species in West Africa. It
occurs over much of tropical Africa and on Madagascar as well. In South Africa it
is known only from the tropical and subtropical areas of the eastern Transvaal and
Natal.

H. perlonga was originally described from Guinea (Scudder, 1969) and has been
subsequently reported from Senegal by Scudder (1971, 1982) and Slater and Wilcox
(1972) and from the Sudan by Linnavuori (1978). Slater’s (1964b) record of Pachy-
brachius pullatus from Eranchi, Swaziland belongs here as do his (1972) records of
“ Pachybrachius nr. pullatus ” from Zaire.

Additional material examined. SENEGAL: 7, Npak, 1 1 km S Ziguichor, 8.xi.l977
(CDHHS). 1, in forest 1 km NE Djibelor about 7.5 km W Ziguichor, 8.xi.l977
(CDHHS). 1, Cap Skiring, 10.xi.1977 (CDHHS). GAMBIA: 2, Tendeba Camp near
River Gambia, 14.xi.1977 (CDHHS). 1, River Tanji, 3 km SW Brufut, 28.xi.1977
(CDHHS). 1, outside Abuko Nature Reserve at waterworks, 26.xi.1977 (CDHHS).

1 , Abuko Nature Reserve, 1 8.xi. 1977 (CDHHS). 1 , 2 km S Kitty, 7 km SSW Brikama
Road Junction, 27.xi.1977 (CDHHS). GUINEA: 2, off coast wind NNW, xii.1957
(Capener). GHANA: 181, Tafo, 4-9.X.1967 (SSS). 1, Tafo, 28.xi.1965 (Leston). 1,
same, 29.xi.1965. 1, same, 9.xii.l965. 1, same, 17.xii.1965. 1, same, l.vi.1967. 1,
same, 7.viii.l967. 1, Kade, 28. v. 1966 (Leston). 2, Goaso, 2 1 .iii. 1 969 (Leston). 1,
Kpandu, 1 .xi. 1 969 (Leston). 1 no abd., Osiem, 24.vii. 1 967 (Leston). 2, Wiawso W.R.,
31. iii. 1969 (Leston). 5, Accra, 15.xi.1969 (Campbell). 2, same, 14.xi. 1969. 1, same,
22. xi. 1969. 2, same, 16.xi.1969. l,same, l.xii.1969. l,Mt. Atewa, 2,000 ft, 6.x. 1967
(SSS). NIGERIA: 4, Oban RH, SE State, 7.iv.l975 (Medler). 2, Ile-Ife, lO.iii. 1969
(Medler). 4, same, iv. 1 969. 6, same, 27. iii. 1 969. 1 , same, ix. 1 974. 1 , same, 25. iv. 1 969.
1, same, 5.V.1969. 2, same, 10. iii. 1969. 1, same, xi.1969. 1, same, 27.X.1969. 2, U.
Ora, M.W. State, xi.1974 (Medler). 1, Sopoba FR, M.W. State, viii.1973. 9, Umuahia
CRIN, EC State, 9.iv.l975 (Medler). 2, Ikom CRIN, SE State, 4.iv.l975 (Medler).
1, Obudu CR, SE State, 27.ix.1973. 6, Udo FR, M.W. State, ll.iv.1975 (Medler).

1984

REVISION OF HORRIDIPAMERA

327

1, Ibadan, W. State, 15.iii. 1969 (Medler). 1 Benin, NIFCR, M.W. State, l.iv.1975
(Medler). CAMEROON: 1, (Carayon). UGANDA: 1, Lake George, 27. v. 1940 (Ste-
phenson). 1, Kawanda, 19.vi.1958 (Odhiambo). 1, Kassesse Busongoro, 20.V.1940
(Stephenson). TANZANIA: 3, Mlingano, 13.vi.1963 (Robertson). 1, same,
29.viii. 1963. l,same, 22.i.l966. l,Ilonga, 14.iv. 1965 (Robertson). 6, same, 8. i. 1965.
4, same, 26. ii. 1964. 1, same, 18.iv. 1965. 4, same, 29.iv.1963. SOUTH AFRICA:
Natal: 5, Eshowe, 15.xi.1967 (SSS). 1, St. Lucia Park, Zululand, 26.i.l968 (Brink-
man). 1, Lake St. Lucia, Charters Creek, 12.xi.1967 (SSS). Transvaal: 1, Letsitele
Valley, Gravelotte District, 18.xi. 1958 (Capener). 2, base Magoebaskloof, 4,000 ft,
12.xii. 1967 (SSS). MADAGASCAR: 1, E District Maroantsetra, v. 19 — (J. Vadon).
1, Sambirano, Nossi-Be, Foret de Lokobe, i. 1 960 (Andria Robinson). 1, Est. Dct.
Sambava R. N. xii-Marojejy, Beondroka 1,200 m, vi. 1960 (P. Soga). 1, Est. Anka-
lampona 130 m, Navana-Marpamtsetra, iii.1958 (Soga-Raharizonina).

Horridipamera pullata (Hesse)

Pamera pullata Hesse, 1925, p. 79.

Pachybrachius pullatus Slater, 1964a, p. 1140 (Pt.).

Horridipamera pullatus Harrington, 1980, p. 99.

Diagnosis. Very elongate, slender with unusually long legs and antennae. Head,
pronotum and scutellum black to dark red-brown. All femora chiefly dark chocolate
brown, pale only on proximal lh to lA and occasionally at extreme distal ends of fore
femora. Antennae chocolate brown to reddish brown nearly uniformly colored; fourth
segment never with a proximal pale annulus. Scutellum lacking oblique reddish vittae,
uniformly dark. Second labial segment not pale, similarly colored to other segments.
Hemelytra strongly suffused with chocolate (as in dark specimens of perlonga), a
large subapical elliptical white corial macula present. Membrane dark with veins
narrowly pale and a short apical median pale vitta present.

Eyes prominent, strongly elevated above vertex and produced. Width across eyes
noticeably broader than width of anterior pronotal lobe.

Male genital structures very similar to those of perlonga. Bulb of sperm reservoir
more quadrate, not appreciably narrowing distally. Left conjunctival spine relatively
shorter much more deeply bifid at distal end (Fig. 16). Right conjunctival spine is
very elongate and sharply acute (Fig. 1 7). Flange-like area proximad of inner pro-
jection of paramere narrow, more elongate and tapered than in perlonga.

Although similar to perlonga in size, and to darker specimens of the latter in color,
pullata is a more slender and long legged species with more “uptilted” and protruding
eyes. Occasional specimens of perlonga may have the head slightly broader than the
anterior pronotal lobe and when this condition occurs in specimens with dark fourth
antennal segments identification may be difficult without comparative material. We
have found two differences in color that appear to be diagnostic. In pullata the middle
and hind femora are completely dark chocolate brown on the distal % to 3A and the
second labial segment is brown and concolorous with the other labial segments. In
perlonga the middle and hind femora have only a small subdistal dark annulus and
the second labial segment is white or dull yellow. The majority of specimens of
perlonga may be separated from pullata by the pale basal annulus on the fourth
antennal segment.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Distribution. H. pullata has previously been thought to be a South African species.
This is not true. Hesse (1925) originally described it from Southwest Africa and Slater
(1964b) reported it from Swaziland. Hesse’s record was from Ovamboland in north-
ern Southwest Africa and Slater’s Swaziland record pertains to perlonga. H. pullata
is a tropical African species. It is sympatric with perlonga in West Africa from where
most of our study material has come. Scudder (1969) reported it from Guinea. It
also occurs in Tanzania and has been reported by Linnavuori (1978) from the Sudan
and Chad. Thus it will probably prove to be widespread in tropical Africa. It is
appreciably less common than perlonga in collections from areas where the two have
been taken together. See discussion under H. ferruginosa.

Material examined. IVORY COAST: 1, Lamto (Toumodi), 17.iii.1964. 1, same,
10.iv.1962. 1, same, 1 0.iii. 1 964. 1, same, 4.xii.l962. GHANA: 1, Accra, l.xii.1969
(G. W. Campbell). 8, Tafo, 7.x. 1967 (SSS). 3, same, 8.x. 1967; 2, same, 4-9. x. 1967.
1, Tafo, 7.vii.l965 (Leston). 1, same, 2.viii. 1965. 1, same, 11.x. 1965. 1, same,
22.vii.1967. l,same, 14.x. 1967. 1, Salt Pond, 1 7. ii. 1966 (Leston). 1, Kade, 2.ix. 1965
(Leston). 2, Obuasi, 7.ii. 1966 (Leston). 1, Prestea W.R., 13. ii. 1966 (Leston). l,Legon,
29.viii. 1 968 (Leston). 1 , Osiem, 2 1 .x. 1 966 (Leston). NIGERIA: 1 1 , Umuahia CRIN,
EC State, 9.iv.l975 (Medler). 2, Ikom CRIN, SE State, 4.iv.l975 (Medler). 7, Oban
RH, SE State, 7.iv.l975 (Medler). 1, Ile-Ife, 5.V.1969 (Medler). TANZANIA: 1,
Mlingano, 5.xi.l963 (Robertson).

Horridipamera ferruginosa (Stal), New Combination
Pamera ferruginosa Stal, 1874, p. 151.

As previously noted Harrington (1 980) cited Pamera ferruginosa as the type species
of her new genus Stalaria. Harrington, however, had misidentihed ferruginosa, con-
fusing it with Pachybrachius kisseis Linnavuori. As noted in the generic discussion
we are revising “ Stalaria ” and will be appealing to the International Commission to
have ferruginosa set aside as the type species of Stalaria and asking that P. kisseis
Linnavuori be named the type species.

P. ferruginosa Stal is actually a species of Horridipamera. Stal (1874) in his original
description keyed ferruginosa into a couplet with species whose males have a spine
midway along the shaft of the tibia. Harrington (1980) notes the lack of such a spine
as one of the diagnostic features of Stalaria. Harrington’s misidentihcation apparently
resulted from her use of specimens in the senior author’s collection from Upemba
National Park, Zaire. These specimens had been reported by Slater (1972) as “ Pachy-
brachius nr. ferruginosus (Stal)” with the statement: “The series reported above dilfers
from the type of ferruginosus in several important characteristics and probably rep-
resents an undescribed species.” The specimens are in fact Stalaria kissei (Linnavuori)
(1978).

The status of ferruginosa within Horridipamera is a matter of some concern. The
lectotype is very closely related to, if not conspecific with, pullata Hesse. However,
it is light reddish brown, whereas all of the West African specimens that we have
examined are chiefly dark chocolate brown. Scudder (1977) when selecting the lec-
totype noted that it was “slightly teneral.” This may well be true. We are reluctant
to synonymize the two species when such a striking color difference is present. We

1984

REVISION OF HORRIDIPAMERA

329

feel that study of a series from Principe Island is needed before formal synonymy is
warranted.

The following measurements are from the male lectotype bearing labels as indicated
by Scudder (1977). Head length 1.10, width 1.00; interocular space 0.50. Pronotum
length 0.78, width 1.28. Scutellar length 0.80, width 0.70. Length claval commissure
0.52. Midline distance apex clavus-apex corium 1.14. Midline distance apex corium-
apex membrane 0.90. Length labial segments I 0.64, II 0.74, III 0.44, IV 0.40. Length
antennal segments I 0.50, II 1.02, III 0.94, IV 1.10. Total body length 5.36.

H. ferruginosa is not included in the preceding cladogram but has all of the derived
conditions that pertain to pullata.

Horridipamera nietneri (Dohm)

Plociomerus nietneri Dohm, 1860, p. 404.

Pachybrachius nietneri Slater, 1964a, p. 1 133.

Horridipamera nietneri Malipatil, 1978, p. 90.

Diagnosis. Very similar in color and structure to bergrothi differing chiefly in the
much shorter hairs, especially those on the antennae which for the most part are
scarcely longer than the diameter of the segment, the more flattened sericeous hem-
elytral pilosity, the (usually) less globose anterior pronotal lobe and the shorter head.

Bulb of sperm reservoir not strongly tapered distally, but bluntly rounded at distal
end. Reservoir wings curving posteriorly, but less extremely so than in perlonga and
conspicua, tapering evenly, blunt at distal ends. No distal vesical process present.
Proximal process small, consisting of a crescentic serrated process (Fig. 9) lying on
right side of ejaculatory duct immediately distad of right conjunctival spine. Right
conjunctival spine much longer than left and expanded into a point distally before
acute apical tooth. Both spines terminating in acute apical tooth-like points (Figs.
22, 23). Paramere with blade and prominently produced outer projection short and
rounded; inner projection strongly recurved (Fig. 33).

H. nietneri and H. bergrothi are allopatric and probably derived from a common
ancestor. The external differences as well as those of the male genitalia appear to be
constant.

Malipatil (1978a) discusses the variability of nietneri in detail pointing out that
specimens from New Caledonia, New Hebrides, Samoa, and some other Pacific
islands are smaller than mainland specimens and that there is considerable variation
in the conjunctival spine. This is true of the limited material we have examined from
these islands and from Fiji as well. Specimens from these islands also have a very
conspicuous white annulus on the fourth antennal segment and relatively short up-
right hemelytral hairs.

The color pattern in some of the Pacific island populations is remarkably different
from those of the Asiatic mainland. In Pacific specimens the hemelytra tend to be
black with broad white or pale yellow lateral margins. A very similar coloration
occurs in emersoni on Ceylon, Paraeucosmetus pacificus Malipatil (Fiji, Western
Samoa, New Hebrides, Tonga, Niue) and in eastern Pacific populations of Pseudo-
pachybrachius guttus (Dallas). The reason for this apparent color convergence is
unknown.

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We have accepted the synonymy as given by Malipatil (but have not examined
types) except that of Pamera emersoni Distant which is a quite different species.

Fifth instar nymph. Mt. Coot-Tha, Brisbane, Queensland, Australia 10.1.1972 (J.
A. Slater). General shape and color pattern similar to perlonga but reddish brown
rather than black or chocolate brown on head and pronotum. Median dark areas on
abdominal terga as in perlonga but much lighter than dark elliptical areas around
scent gland orifices. Coloration mesally on tergum 3 projecting posteriorly to scler-
otization about scent gland orifices between terga 3-4 suture in the form of three
“prongs” as in inconspicua.

Malipatil (1978b) has described and figured the 5th instar nymph in detail (from
Queensland). Neither his figure nor description shows the small dark spot with a
parenthesis-like mark surrounding it which is present mesally on the 4th abdominal
tergum.

Egg. Described by Malipatil (1978b). Said to have 5 chorionic processes, an average
total length of 1.14 and average maximum width of 0.34.

Distribution. H. nietneri is found almost throughout the Oriental Region. It also
occurs in northern and eastern Australia and east into the Pacific Islands at least as
far as the Carolines and Samoa. It was originally described from Ceylon. Slater ( 1 964a)
lists records from Burma, India, China, Malaya, Cambodia and Thailand on the
mainland and from the following islands: Bonins, Carolines, Guam, Christmas, New
Caledonia, Fiji, Japan, Ryukyus, Taiwan, Samoa, Celebes, Java, Sumatra, Philip-
pines, Moluccas and Laccadives and Australia. Malipatil (1978a) adds New Guinea,
New Hebrides and the Tonga Islands. He gives detailed information on the Australian
distribution where it is chiefly confined to Queensland and Northern Territory but
he states that it also occurs in New South Wales. Tomokuni (1982) reports it from
Minami-Iwojima (San Agustino) Island of the Volcano Islands.

Additional material examined. PHILIPPINES: 3, Manila, (Luzon), iv.1947 (Enns).
1, Mt. Maquiling (Luzon), 18.viii.1950. 1, same, 18.i.l947 (Santas). 4, same, 1 9.ii. 1 947
(Agbozala). 3, same, 2.xi. 1949, 50 ft (Uichanco). 1, same, 10.x. 1949 (Hosillos). 1,
same, 20. ix. 1951 (Bowagan). 1, Manila (domestic airport waiting room), 29.x. 1963
(McFarland). 1, IRRI Farm, Luzon, ll.x.1972 (Pawar). 1, same, 8.V.1972. 1, same,
v. 1972. 1, same, iv. 1972. 1, same, iii.1972. 1, same, vi. 1972. 2, same, 12.x. 1972. 1,
same, 23.xii.1972. 2, same, 3.iii.l972. 2, same, x. 1972. 1, same, i. 1972. 2, same,
7.xi.l972. 1, Sagada (Luzon), Mt. P., 9.iv. 1972 (Pawar). 1, Ilagan (Luzon), Isabela,
25.V.1972 (Pawar). 1, Munoz, NE, 23. v. 1972 (Pawar). 1, Vigan Island (S. Luzon),
28. v. 1972 (Pawar). 1, Ladoy Island (N. Luzon), 27. v. 1972 (Pawar). 1, St. Rosa,
Laguna, Luzon, 6. iii.1972 (Pawar). 1, Malolos, Buula (Luzon), 12.iv. 1972 (Pawar).
1, Apari, Cagayan (Luzon), 26.V.1972 (Pawar). 1, Legaspi, Albay, 16.i. 1964 (Mc-
Farland). BORNEO: 1, Kota Kinbulu, 29.xii.1967 (Roche). 1, same, 25.xii.1967.
OKINAWA (RYUKYUS): 3, Kin, 18.ix.1945 (Slater). 1, same, 7.ix.l945. 1, same,
14.vii.1945. INDIA: 5, Anamalai Hills, Cinchona, 3,500 ft, iv.1959 (Nathan). 2,
same, v. 1959. 1, Coimbatore, x.1951 (Nathan). 4, same, x.1953. 1, same, xi.1964.
l,same, 22.xii.1951. l,same, ix. 1951. 1, Orissa, C.E. India, Jeypore, 1,775 ft, x.1958
(Nathan). 1, Kurumbagarum, Karikal Territory, viii. 1955 (Nathan). 1, same, xi. 1951.
1, Nilgiri Hills, Sinpare, 3,400 ft, v.1954 (Nathan). SRI LANKA: 2, N. Centr. Prov.,
Polonnaruwa, 1 0.ii. 1 962 (BAC). 2, N. Centr. Prov., Kandurukanda, 20 mi NE Ha-
barana, 8.ii. 1 962 (BAC). 4, Anu. Dist., Wildlife Soc. Bungalow, Hunuwilagama,

1984

REVISION OF HORRIDIPAMERA

331

Wilpattu, 1 0-1 9.iii. 1970 (Davis, Rowe). 2, “Rat. Dist.,” Uggalkaltota 350 ft, Irri-
gation Bungalow, 31.i-8.ii. 1970 (Davis, Rowe). 1, Kandy District, 5 mi NW Ma-
hiyangana, 30.iii-9.iv.1971 (Spangler). 1, “Ham. Dist.,” Wirawila Wewa, 85 ft,
26.x. 1970 (Flint). 2, “Kal. Dist.,” Agalawatta, 13-14.X.1976 (HDKB). NEW CAL-
EDONIA: 6, Yate (under tall grasses), 9.vii. 1977 (Alex Slater). FIJI: 1, Viti Levu,
Nadarivatu, 3 1 .i. 1 968 (Gross). 2, Nausori Highlands 16 mi E Nandi, 16.vii.1977
(Alex Slater). 2, same, 20 mi E Nandi. NEW HEBRIDES: 3, Efate Isl., Port Villa,
10-13.vii.1977 (Alex Slater).

Horridipamera bergrothi (Horvath)

Pamera bergrothi Horvath, 1892, p. 261.

Pachybrachius bergrothi Slater, 1964a, p. 1113.

Horridipamera bergrothi Harrington, 1980, p. 99.

Diagnosis. Robust, heavy bodied. Dorsal surface including hemelytra and ap-
pendages thickly clothed with elongate upstanding hairs. Elongate hairs intermixed
with decumbent, silvery, somewhat tomentose, pubescence. Head, pronotum and
scutellum uniformly dark chocolate brown, scutellum lacking oblique light vittae.
Hemelytra chiefly dark brown (similar to nietneri). Clavus either completely dark or
with anterior half of cubital vein pale. Corium with explanate margin chiefly pale
but this area invaded by a dark brown macula at level of distal end of claval com-
missure and at distal end of corium. Subdistal “white macula” often reduced to two
or even three distinct spots. Corium frequently with a pale area on anterior half, but
this not reaching claval suture. Membrane dark with pale veins and an apical median
vitta. Antennae brown; fourth segment lacking a pale basal annulus; first segment
often pale on distal half. Fore femora dark. Middle and hind femora with distal
halves dark brown, strongly contrasting with pale proximal halves. Second labial
segment pale.

Anterior pronotal lobe strongly globose, broader than width of head across eyes.
Labium reaching nearly to mesocoxae.

Bulb of sperm reservoir subquadrate, not narrowing distally. Reservoir wings as
in nietneri. Distal vesical process long slender non-serrated (Fig. 8). Proximal vesical
process with a serrated crescent-shaped proximal portion (similar to that of nietneri)
and an additional slender distally twisting portion (Fig. 5). Conjunctival spines short,
broad, nonbifid and trianguloid (Figs. 24, 25). Paramere with a very large inner
distally bifid flange; outer projection broad and rounded (Fig. 34).

This large dark species is readily recognizable from other African members of the
genus by the numerous upright hemelytral hairs. The chiefly dark hemelytral surface
is also distinctive. There is some variation in color. For example, the second and
third antennal segments may be dull yellow. Occasionally the fourth segment is paler
than the distal end of the third, but there is never a pale proximal annulus.

H. bergrothi is in general habitus and color as well as many morphological features
more closely related to the Oriental nietneri than to any of the African species.

Distribution. Reported by Slater (1964a) from Cameroon, Ethiopia and Ghana; by
Scudder (1969) from Guinea; by Scudder (1982) from Senegal; by Linnavuori (1978)
from Sudan and as “nr bergrothi” by Slater (1972) from Zaire. It probably is dis-
tributed throughout tropical Africa.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Material examined. IVORY COAST: 1, Lamto (Toumodi), 30. i. 1962. 2, same,

17.111.1964. 1, same, 22. v. 1964. 3, same, 20.ii.1962. 1, same, ll.ii.1964. 1, same,

18.11.1964. 1, same, 6.iii. 1962. GHANA: l,Tafo, 7.x. 1967 (SSS). 1, Tafo, 21.vi. 1965
(Leston). 1, same, 2.ii. 1966. 1, same, 8.iii.l966. 1, Sunyani, 12.xii.1965 (Leston). 1,
same, 13.xii.1965. 1, Legon, 28.viii.1968 (Leston). 1, same, 30.ix.1968. 1, same,

1. x. 1968. 1, Techiman, 9.xii.l965 (Leston). 1, Tokos, 6.xi.l967 (Leston). NIGERIA:

2, Umuahi CRIN, EC State, 9.iv. 1975 (Medler). 1, Ile-Ife, 1 l.viii. 1969. 1, same,
27.iii. 1969. 1, same, W State, 9.iii. 1975. TANZANIA: Ilonga, 29. ix. 1963 (Robert-
son). 2, same, 3.X.1963. 2, same, 14.iv.1965. 1, same, 10. ii. 1965. 1, same, 2.ii.l964.
UGANDA: 1, Kassesse Busoyfora, 5. v. 1940 (Stephenson).

Horridipamera robust a Malipatil

Horridipamera robusta Malipatil, 1978a, p. 92.

Diagnosis. A large dark strongly hirsute species. Coloration much as in nietneri.
Fore femora completely black. Antennae brown, contrasting with velvety black col-
oration of head and pronotum. Fourth antennal segment with a prominent sub-basal
pale annulus. Anterior pronotal lobe very large and swollen (twice as long as posterior
lobe in males, 1 x/i times as long in females). Fore tibia armed with a large spine near
middle and 2 or 3 smaller spines distally.

Malipatil (1978a) gives a good detailed description and figures of anatomical details,
including those of the male genitalia.

Immature stages. Malipatil (1978b) has described the 4th and 5th instar nymphs
and the egg.

The 5th instar nymph is said to be separable from that of nietneri by having the
anterior pronotal lobe about 4 times as long as the posterior lobe. In nietneri the
anterior lobe is approximately 3 times as long as the posterior.

Egg. The egg of robusta is said by Malipatil {ibid.) to have 4-6 chorionic processes,
and average length of 1.23 and an average width of 0.36.

Distribution. Originally described from Queensland and northern New South Wales
in Australia and from North Island, New Zealand. Malipatil (1978a) notes that it is
sympatric with nietneri in southeastern Queensland.

Horridipamera emersoni (Distant), New Status
Pamera emersoni Distant, 1909, p. 491.

Pachybrachius nietneri Scudder, (nee Dohm), 1962b, p. 770 (part).

Horridipamera nietneri Slater, (nee Dohrn), 1979, p. 22 (part).

Diagnosis. Elongate, slender, parallel-sided. Head, pronotum, scutellum, antennal
segments I, II, III, and distal Vi of IV black. Humeral pronotal angles pale yellow.
Basal half of fourth antennal segment with a broad white annulus. Hemelytra with
much of clavus and inner portion of corium heavily infuscated with dark brown
giving outer portion appearance of a pale yellow stripe which blends distally with
what in many other species is a distinct subapical pale macula. Apex of corium and
apical corial margin dark. Membrane chocolate brown with a broad strongly con-
trasting median white vitta. Second labial segment white or pale yellow. Fore femora

1984

REVISION OF HORRIDIPAMERA

333

(except distal ends) and distal half of middle and hind femora black or dark casta-
neous.

Pronotum and scutellum lacking numerous upright elongate hairs, but with nu-
merous short decumbent sericeous hairs present.

Eyes relatively large and somewhat protruding. Head broader than narrow, rela-
tively unswollen, anterior pronotal lobe. Labium extending posteriorly onto meso-
sternum. Both sexes lacking a prominent spine midway along shafts of fore tibiae.

Although synonymized with nietneri by Scudder ( 1 962) this is a distinct species.
The male genitalia of the two species are completely different. The lack of a spine
on the fore tibia of the male and the lack of elongate upstanding hairs on the hemelytra
will also readily separate it from nietneri. In addition nietneri is a much heavier
bodied, more robust species. While, as previously discussed, nietneri is quite variable
in color, on Ceylon where the two species are sympatric nietneri has the dark areas
of the corium more extensively developed, often covering most of the corium except
for the explanate margin, and, of course, the subapical pale macula. In nietneri the
head is narrower than the anterior pronotal lobe, the latter being more swollen and
globose than in emersoni.

Bulb of sperm reservoir moderately tapering distally, distal end rounded. Reservoir
wings moderately curving posteriorly (similar to nietneri ), irregularly tapering. Distal
vesical process elongate and slender, relatively straight. Proximal vesical process with
a broadened serrated plate-like area (Fig. 12). Conjunctival spines large, elongate,
acutely pointed, nonbifid; right spine (Fig. 27) longer than left (Fig. 26). Paramere
with inner projection small but with an enormous distally bifid flange projecting
from it (Fig. 30).

Scudder may have considered the white margined Pacific island populations of
nietneri to represent emersoni leading him to synonymize the two.

In general body shape emersoni most closely resembles the African species pullata
but the latter lacks a white annulus on the fourth antennal segment, its males have
a mid fore tibial spine, the subapical pale corial spot is distinct, the eyes even more
strongly protruding and the genitalia are distinctive (see discussion above).

H. emersoni appears to be a member of the perlonga-pullata group despite the
absence of the fore tibial spine.

Distribution. Known only from Sri Lanka. Slater’s (1979) series of nietneri was
mixed. Specimens he reported as nietneri from “Kal. District, Morapatiya near
Agalawatta” and “Rat. Dist., Bultota Pass 3000 ft.” are emersoni.

Horridipamera medleri, new species
Fig. 2

Description. Elongate, parallel-sided, with head broad and appendages elongate,
slender. Body somewhat ant-mimetic. General coloration black to chocolate brown
including entire head, pronotum, scutellum, antennae, fore femora (except extreme
proximal and distal ends), and distal % of middle and hind femora. Hemelytra chiefly
dull white but marked with dark chocolate brown as follows: an irregular band through
center of clavus; a large elongate corial macula between R+M and CU at level of
claval commissure; entire apical corial margin; a large apical corial macula; majority

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Fig. 2. Horridipamera medleri new species. Holotype, dorsal view.

1984

REVISION OF HORRIDIPAMERA

335

of hemelytral punctures; membrane, except for a large apical white patch and bases
of veins.

Head elongate, moderately declivent anteriorly. Tylus attaining distal xh of first
antennal segment. Eyes protruding. Width across head subequal to that of anterior
pronotal lobe. Head considerably narrowed behind eyes, with fine, dense punctures
and clothed with dense, decumbent sericeous hairs, except for a long ‘S’-shaped area
in front of each ocellus. Vertex moderately convex. Head length 0.92, width 1.12,
interocular width 0.68, eye length 0.38, postocular distance 0.20.

Dorsal surface lacking numerous elongate upstanding hairs but clothed (including
clavus and corium) with conspicuous decumbent sericeous pubescence.

Pronotum subshining. A deep transverse constriction present. Collar with distinct
rows of punctures. Anterior pronotal lobe swollen, lateral margins forming a smooth
evenly rounded curve. Posterior pronotal lobe with shallow, evenly separated punc-
tures. Length anterior pronotal lobe 1.20, maximum width 1.20; length posterior
lobe 0.60, width 1.48. Scutellum with a low inconspicuous median elevation. Length
scutellum 0.92, width 0.84. Length claval commissure 0.56. Midline distance apex
clavus-apex corium 1.20. Midline distance apex corium-apex membrane 0.88. An-
terior tibiae only slightly curved proximally and each tibia with a prominent median
spine. Labium extending well onto mesostemum. Length labial segments I 0.80, II
0.80, III 0.54, IV 0.42. Antennae slender, terete. Length antennal segments I 0.66,
II 1.32, III 1.14, IV 0.32. Total body length 6.40.

Sperm reservoir bulb moderately narrowing distally. Reservoir wings strongly curv-
ing posteriorly and tapered (as in perlonga) but not produced into an elongate pro-
jection as in inconspicua. Vesical marginal processes more or less interconnected,
elongate, twisted almost identical to conditions found in perlonga and pullata. Right
conjunctival spine large, elongate and tapering to an extremely slender, acute point
(Fig. 15). Left conjunctival spine with a broadened area on outer margin but not
distinctly bifid and ending in a large acute spine (Fig. 14). Paramere similar to that
of perlonga but inner flange not truncate and outer projection evident only as an
evenly rounded area.

H. medleri most closely resembles pullata (Hesse). It is a relatively heavier, less
slender species than is pullata. The anterior pronotal lobe is distinctly broader and
more swollen. This is especially evident in lateral view. The corium is paler in color
with the area of the corium laterad of the furrow (exocorium) either entirely white
or at most with only a small, incomplete submedian dark macula. The fore femur
has a large spine on the outer row proximad of the middle (lacking in pullata ). The
conjunctival spines are distinctive in shape.

Holotype. Male, NIGERIA: Oban RH SE State, 7.IV.1975. (J. T. Medler). In
American Museum of Natural History.

Paratypes. 1 male, 2 females, same data as holotype. In J. A. Slater collection.

This species is named in honor of Dr. John Medler for his generosity and in
recognition of his contributions to our knowledge of West African Hemiptera.

Horridipamera compacta, new species

Description. Relatively stout, parallel-sided. Head, anterior pronotal lobe and scu-
tellum black. Posterior pronotal lobe, except for a yellow spot before each humeral

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Figs. 3-13. Horridipamera spp. 3. H. inconspicua, apex of proximal vesical process. 4. H.
compacta, apex of proximal vesical process. 5. H. bergrothi, proximal vesical process, dorsal
view, right side. 6. H. perlonga, phallus, showing sperm reservoir conjunctival spines and vesical
processes. 7. H. inconspicua, phallus, semilateral view, showing sperm reservoir and proximal
portion of ejaculatory duct. 8. H. bergrothi, distal vesical process. 9. H. nietneri, proximal
vesical process. Dorsal view, right side. 10. H. perlonga, vesical process. 11 . H. compacta,
vesical processes. 12.//. emersoni, proximal vesical process. 13. H. inconspicua, vesical process.

angle, castaneous. Hemelytra variegated, pale yellow and dark red brown. Dark
coloration as follows: a complete but irregular transverse vitta across middle of
hemelytra which broadly attains lateral corial margins; a large apical macula; streaks
on distal portion of clavus; and membrane with exception of paler veins. All hem-

1984

REVISION OF HORRIDIPAMERA

337

elytral punctures dark brown. Antennae chiefly ochraceous to very light brown. Base
of segment I, distal ends of segments II and III and apical half of segment IV dark.
Fourth segment with a prominent pale annulus. Fore femora castaneous with pale
distal ends. Middle femora pale with a series of brown spots distally. Hind femora
pale with a conspicuous dark annulus on distal xh. Head, pronotum and scutellum
(but not hemelytra) thickly clothed with elongate upstanding hairs.

Head shining, moderately declivent anteriorly with dense, fine, shallow punctures.
Tylus reaching to distal xh of first antennal segment. Vertex only moderately convex.
Eyes not strongly produced. Head length 1.08, width 1.12, interocular width 0.64,
eye length 0.33, postocular length 0.19.

Pronotum subshining. Anterior lobe strongly swollen. Length pronotum 0.94, width
1.34; length posterior lobe 0.68, width 1.68. Collar with deep, irregularly arranged,
punctures. Scutellar length 1.0, width 0.50. Length claval commissure 0.60. Midline
distance apex clavus-apex corium 1.18. Midline distance apex corium-apex mem-
brane 0.80. Fore femora strongly swollen, width fore femur 0.64. Front tibiae dis-
tinctly curved without a prominent spine near middle of shaft. Labium attaining
middle of mesosternum, first segment remote from base of head. Length labial seg-
ments I 0.84, II 0.84, III 0.50, IV 0.44. Length antennal segments I 0.52, II 1.18,
III 1.06, IV 1.20. Total body length 6.40.

Sperm reservoir bulb distinctly tapering (similar to that of inconspicua ). Reservoir
wings relatively little curved posteriorly, projecting strongly laterad, distal margin
strongly convex, proximal margin nearly straight, ending in a nipple-like end.

H. compacta is a relatively stout species that most closely resembles perlonga in
general habitus. It differs from perlonga and related species by the numerous elongate
hairs on the pronotum and scutellum. The absence of a median spine on the fore
tibia of the male suggests relationship with emersoni (as shown in Fig. 1), but this
could be the result of independent loss. The male genitalia are distinctive. The widely
separated vesical proximal and distal marginal processes (Fig. 11) will immediately
distinguish compacta from either inconspicua or perlonga. The strongly swollen fore
femora and pale antennae with the darkened distal ends of segments 2 and 3 are also
useful recognition features.

Holotype. Male, INDIA (South), Shevaroy Hills, Yercaud 4,500 ft, 9. XII. 1954 (P.
S. Nathan). In American Museum of Natural History.

Paratypes. 1 female, INDIA (South), Annamalai Hills, Cinchona, 3,500 ft, X.1955
(P. S. Nathan). 1 female, Annamalai Hills, Cinchona, X.1955 (P. S. Nathan). In J.
A. Slater collection.

Incertae sedis

Horridipamera subsericea (Breddin)

Pamera ( Entisberus ) subsericea Breddin, 1907, p. 205.

Entisberus subsericea Distant, 1910, p. 59.

Pamera subsericea Bergroth, 1918, p. 105.

Pachybrachius subsericea Slater, 1964a, p. 1 142.

Horridipamera subsericea Harrington, 1980, p. 99.

This species was originally described from Ceylon and has had a checkered no-

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Figs. 14-35. Horridipamera spp. 14. H. medleri, left conjunctival spine. 15. H. medleri,
right conjunctival spine. 16. H. pullata, left conjunctival spine. 17. H. pullata, right conjunctival

1984

REVISION OF HORRIDIPAMERA

339

menclatural history (see Slater, 1 964a). Harrington (1980) placed it in Horridipamera
but with an indication that she had not seen material and was unsure of its correct
generic position.

We do not believe it will prove to be a species of Horridipamera. The body length
of “3% mm” is very much less than that of any known species of the genus. In fact,
of the species placed in the old “omnibus” genus Pamera (later Pachybrachius ), the
description would seem to better “fit” Pseudopachybrachius guttus than any species
of Horridipamera.

Plociomerus undulatus Dohrn
Plociomerus undulatus Dohrn, 1860, p. 404.

We have examined the type which is in Institute Zoology, Polish Academy of
Sciences, Warsaw, Poland. It consists of a scutellum, meso- and metapleuron, one
hind leg and one middle leg. It is not Pseudopachybrachius gutta as had been stated
by Zheng and Slater (1984) but is a larger species of myodochine. It probably is a
species of Horridipamera or Paraeucosmetus but it is impossible to know what species
it is. The hind leg has the femur darkened distally but not with so sharp a contrast
as have several species of both genera.

ACKNOWLEDGMENTS

We are indebted to the following for the loan or gift of valuable material: Dr. Adam Kedziorek
(Polish Academy of Sciences); Dr. Karl Krombein (National Museum of Natural History,
USNM); the late Dr. Dennis Leston; Dr. Rauno Linnavuori (Raisio, Finland); the members of
the Lund University Senegal-Gambia Expedition; Dr. M. B. Malipatil (Northern Territory
Museum of Arts and Sciences, Darwin); Dr. John Medler (University of Wisconsin); Dr. A. D.
Pawar (International Rice Institute, Philippines); Dr. I. A. D. Robertson (formerly Cotton
Research Program, Tanzania) and Dr. T. E. Woodward (Queensland Museum).

We wish to thank the late Dr. W. H. Coaton and the staff of the Plant Protection Institute
of Pretoria, South Africa for assistance to the senior author in field work and facilities; Dr.
Randall Schuh (American Museum of Natural History) for discussions on distribution and aid
in field work; Dr. M. H. Sweet (Texas A&M University) and Mr. Samuel Slater for aid in field
work in Africa; Ms. Mary Jane Spring and Mrs. Elizabeth Slater (University of Connecticut),
respectively, for preparation of the illustrations and extensive help in the preparation of the
manuscript.

This work was supported in part by a grant from the National Science Foundation.

spine. 18. H. compacta, left conjunctival spine. 19. H. compacta, right conjunctival spine. 20.
H. perlonga, left conjunctival spine. 21. H. perlonga, right conjunctival spine. 22. H. nietneri,
left conjunctival spine. 23. H. nietneri, right conjunctival spine. 24. H. bergrothi, left conjunc-
tival spine. 25. H. bergrothi, right conjunctival spine. 26. H. emersoni, left conjunctival spine.
27. H. emersoni, right conjunctival spine. 28. H. inconspicua, left conjunctival spine. 29. H.
inconspicua, right conjunctival spine. 30. H. emersoni, paramere. 31. H. compacta, paramere.
32. H. perlonga, paramere. 33. H. nietneri, paramere. 34. H. bergrothi, paramere. 35. H.
inconspicua, paramere.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

LITERATURE CITED

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Breddin, G. 1907. Berytiden & Myodochiden von Ceylon aus der Sammelausbeute von Dr.
W. Horn (Rhynch. het.) II. Deutsch ent. Zeit. 1907:203-220.

China, W. E. 1924. The Hemiptera-Heteroptera of Rodriguez together with the description
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Dallas, W. S. 1982. List of the specimens of Hemipterous insects in the collection of the
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Distant, W. L. 1909. LXV. Oriental Rhynchota Heteroptera. Ann. Mag. Nat. Hist. (8):3:491-
507.

Distant, W. L. 1910. The Fauna of British India, Including Ceylon & Burma. Rhynchota,
Vol. 5. Heteroptera: Appendix, 362 pp.

Dohm, A. 1860. Zur Heteropteren-Fauna Ceylon’s. Stettin ent. Ztg. 21:399-409.

Harrington, B. J. 1980. A Generic Level Revision and Cladistic Analysis of the Myodochini
of the World (Hemiptera, Lygaeidae, Rhyparochrominae). Bull. Am. Mus. Nat. Hist.
167:45-116.

Hesse, A. J. 1 925. A list of the Heteropterous & Homopterous Hemiptera of Southwest Africa.
Ann. S. Afr. Mus. 23:1-178.

Horvath, G. 1879. Hemiptera-Heteroptera a. Dom. Joanne Xantus in China & in Japonia
collecta. Term Tud. Fuz. 3:141-152.

Horvath, G. 1892. Hemiptera nova Africana. Term Tud. Fuz. 15:254-267.

Lethierry, L. and G. Severin. 1894. General Catalogue of the Hemiptera, Vol. II. Heteroptera.
F. Hayez, Imprimeur de l’Academie Royale de Belgique, Brussels.

Lindberg, H. 1958. Hemiptera Insularum Caboverdensium Systematik, Okologie & Ver-
breitung der Heteropteren & Cicadinen der Kapverdischen Inseln. Comment. Biol. 19:
1:246 pp.

Linnavuori, R. 1978. Hemiptera of the Sudan, with remarks on some species of the adjacent
countries 6. Aradidae, Meziridae, Aneuridae, Pyrrhocoridae, Stenocephalidae, Coreidae,
Alydidae, Rhopalidae, Lygaeidae. Act. Zool. Fenn. 153:1-108.

Malipatil, M. B. 1978a. Revision of the Myodochini (Hemiptera: Lygaeidae: Rhyparochromi-
nae) of the Australian Region. Aust. J. Zool. Suppl. Ser. no. 56:1-178.

Malipatil, M. B. 1978b. Immature Stages of Some Myodochini of the Australian Region
(Hemiptera: Lygaeidae: Rhyparochrominae). Aust. J. Zool. 26:555-584.

Reuter, O. M. 1882. Ad cognitionem Heteropterorum Africae occidentalis. Ofv. Finska Vet.-
Soc. Forh. 25:1-43.

Reuter, O. M. 1887. Ad cognitionem Heteropterorum Madagascariensium. Ent. Tidskr. 8:
77-109.

Schouteden, H. 1957. Contributions a l’etude de la faune entomologique du Ruanda-Urundi
(Mission P. Basilewsky 1953) CXXV. Heteroptera Lygaeidae, Pyrrocoridae et Berytidae.
Ann. Mus. Congo Beige. 8:58:247-268.

Schumacher, F. 1913. Ein Beitrag zur Kenntnis der Rhynchoten-Fauna Sudafrikas. In: Schultze
zool. und anthrop. Ergebnisse and Forschungreise in Sudafrika. Denkschr. med.-naturw.
Ges., Jena 17:49-88.

Scott, J. 1874. On a collection of Hemiptera-Heteroptera from Japan, descriptions of various
new genera and species. Ann. Mag. Nat. Hist. (4): 14:289-304, 426-452.

Scudder, G. G. E. 1962a. The World Rhyparochrominae (Hemiptera: Lygaeidae) I. New
Synonymy and Generic Changes. Canad. Ent. 94:764-773.

Scudder, G. G. E. 1962b. Mission Zoologique de l’LR.S.A.C. en Afrique orientale (P. Basi-
lewsky et N. Leleup, 1957). Ann. Mus. Roy. Afr. Centr., Sci. Zool., 1 10, pp. 400-453.

Scudder, G. G. E. 1969. The Lygaeidae (Hemiptera) collected by Dr. M. Mrazek and Mrs. I.
Korecka in the Republic of China. Act. Mus. Morav. 54:169-182.

1984

REVISION OF HORRIDIPAMERA

341

Scudder, G. G. E. 1971. New Lygaeidae (Hemiptera) from the Niokolo-Koba National Park,
Senegal. Bull, de l’LF.A.N. 33:A:3:7 18-736.

Scudder, G. G. E. 1977. The World Rhyparochrominae types (Hemiptera: Lygaeidae) XIII.
The Stcil types. Ent. Scand. 8:29-35.

Scudder, G. G. E. 1982. Recherches Scientifiques dans les Parcs Nationaux du Senegal IX.

Hemiptera Lygaeidae du Parc National Niokolo-Koba. Mem. de l’LF.A.N. no. 92:150.
Slater, J. A. 1964a. A Catalogue of the Lygaeidae of the World. 2 Vol. University of Con-
necticut, Storrs.

Slater, J. A. 1964b. Hemiptera (Heteroptera): Lygaeidae. S. Afr. Anim. Life 10:15-228.
Slater, J. A. 1972. The Lygaeidae of Upemba National Park, Mission G. F. deWitte (He-
miptera: Heteroptera). Parc National de L’Upemba 72:2:17-77.

Slater, J. A. 1979. Hemiptera, Heteroptera: Lygaeidae from Sri Lanka (Ceylon). Report no.

45 of Lund University Ceylon Expedition in 1962, pp. 1-27.

Slater, J. A. and D. B. Wilcox. 1972 (1973). Contribution a l’etude biologique du Senegal
septentrional XXII. Hemiptera Lygaeidae. Bull, de l’LF.A.N. 34:A:4:943-965.

St&l, C. 1874. Enumeratio Hemipterorum, Pt. 4. K. Svenska Vetensk. Acad. Handl. 12:1:1 —
186.

Tomokuni, M. 1982. Heteroptera of Minami-Iwojima Island. Conservation Reports of the
Minami-Iwojima Wilderness Area, Tokyo, Japan, pp. 335-341.

Zheng, L. and J. A. Slater. 1984. A revision of the lygaeid genus Pseudopachybrachius. Syst.
Entomol. 9:20:95-115.

Received February 21, 1984; accepted July 2, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 342-343

NEW SYNONYMY AND A NEW SPECIES IN THE GENUS
MORMIDEA (HEMIPTERA: PENTATOMIDAE)

L. H. Rolston

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University Agricultural Center,

Baton Rouge, Louisiana 70803

Abstract.— The name Mormidea kirkaldyi is proposed for a species described previously but
misdetermined. Mormidea bridarolli Piran, 1963, is placed in the synonymy of M. montandoni
Kirkaldy, 1902.

At the time I revised Mormidea, the type series of M. bridarolli Piran was un-
available, and the location of the syntypes of M. montandoni Kirkaldy was unknown
to me (Rolston, 1978). Recently the private collection of the late A. A. Piran, con-
taining the type series of M. bridarolli, was acquired by the Museo Argentina de
Ciencias Naturales “Bernardino Rivadavia,” Buenos Aires, and by chance the syn-
types of M. montandoni were located in the Institut Royal des Sciences Naturelles
de Belgique, Brussels.

Dr. Jocelia Grazia, at the Universidade Federal do Rio Grande do Sul, kindly
compared a specimen that I had determined as M. bridarolli with the holotype of
that species and found the specimens to be conspecific. Through the courtesy of Dr.
G. Schmitz, at the Musee Royal de TAfrique Central, the syntypes of M. montandoni
were loaned to me.

In the revision of Mormidea I misdetermined M. montandoni Kirkaldy, 1902,
which proves to be a senior synonym of M. bridarolli Piran, 1963. The species which
I believed to be M. montandoni is a new species for which I propose the name
Mormidea kirkaldyi. The synonymy of these two species is:

Mormidea kirkaldyi, new species

Mormidea montandoni: Rolston, 1978, p. 174, figs. 6-8 (misdetermination).

Mormidea montandoni Kirkaldy, 1 902
Mormidea montandoni Kirkaldy, 1902, p. 165.

Mormidea bridarolli Piran, 1963, pp. 108-109; Rolston, 1978, p. 175, fig. 9.

The type series of Mormidea kirkaldyi new species consists of 755 and 729. The 6
holotype is labeled: (a) “Divisoria, Loreto, Peru, November 1947, Alt. 1500 ft.” (b)
“W. Weyrauch Coll. Donor Wm. Procter”. Deposited in the American Museum of
Natural History. Paratypes: labeled as holotype (355, 422 AMNH)1; labeled as ho-

1 Paratype deposition: American Museum of Natural History (AMNH); author’s collection
(LHR); National Museum of Natural History (NMNH).

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MORMIDEA

343

lotype except “Doner” misprinted as “Coner” (9 AMNH); (a) “Divisoria, Loreta,
Peru, May 17, 194, Alt. 4500 ft.” (b) “J. C. Pallister, Coll. Doner Frank Johnson”
(9 AMNH); (a) “El Partidero, Ecuador, XI. 14. 1935” (b) “W. Macintyre, Collector”
(9 LHR); (a) “Upper Rio Huallaga, Peru, 1.V.26, F6123” (b) “H. Bassler Collection,
Acc. 33591” (3 LHR); “Brazil-Ecuador border, Davis” (8 LHR); “Puyo, ECUADOR,
Napo-Pastaza Prov. 15 April 1958, W. R. Hodges, Elev. 2,500 feet” (8 NMNH).

The type series of Mormidea montandoni Kirkaldy consists of 188 and 399. The 8
lectotype, here designated, is labeled: (a) “Coll. R. I. Sc. N. B., Equateur, Ambato,
ex. Coll. Schouteden” (purple label) (b) “Mormidea montandoni Kirkaldy. T” (c)
“syntype” Paralectotypes: 5<$<3, 299 labeled as lectotype except (b) “Mormidea mon-
tandoni Kirkaldy. co-type”; 8 labeled as lectotype except white label with locality
data is pasted on purple label between “Coll. R. I. Sc. N. B.” and “ex Coll. Schou-
teden”; 9 labeled as above except “(Mission)” follows “ambato.” The entire type
series is in the Institut Royal des Sciences Naturelles de Belgique.

In the revision of Mormidea, a drawing intended as a front piece appears without
a caption on page 212. The location suggests that this drawing pertains to M. lunara
Rolston, but the specimen illustrated is M. collaris Distant.

LITERATURE CITED

Kirkaldy, G. 1902. Miscellanea rhynchotalia. No. 4. (Heteroptera) Entomol. 35:164-167.
Piran, A. A. 1963. Hemiptera neotropica VIII. Especies nuevas o poco conocidas de las faunas
de Colombia, Peru, Brazil, Bolivia y Paraguay. Physis 24(67): 107-1 12 (“Brasil” in error
for Ecuador).

Rolston, L. H. 1978. A revision of the genus Mormidea (Hemiptera: Pentatomidae). J. New
York Entomol. Soc. 86(3): 16 1-2 19.

Received November 22, 1983; accepted April 10, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 344-348

PELLAEA SANTAROSENSIS (HEMIPTERA: PENTATOMIDAE),

A NEW SPECIES FROM COSTA RICA

L. H. Rolston

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University Agricultural Center,

Baton Rouge, Louisiana 70803

Abstract.— Pellaea candens (Distant, 1890) is placed in the synonymy of Pellaea stictica
(Dallas, 1851), and Pellaea santarosensis, a new species from Costa Rica, is described. A
lectotype is designated for Rhaphigaster sticticus Dallas, 1851.

The genus Pellaea Stal, 1872, has until recently contained three nominal species:
P. stictica (Dallas, 1851), with the synonym Pentatoma aspera Walker, 1867, and
an unavailable synonym, Nezara nebulosa Distant, 1891, whose interesting history
is discussed by Froeschner (1981); P. candens (Distant, 1890); and P. panamensis
(Distant, 1890), a species transferred by Rolston (1983) to Acrosternum Fieber sub-
genus Chinavia Orian.

Dallas (1 85 1) described P. stictica, as Rhaphigaster sticticus, from three specimens,
but only one of these can now be found in the British Museum (Natural History).
This male, here designated lectotype, bears the following labels: (a) “Type” on red-
edged disk, (b) “44/85” on one side of white disk and “B. Guiana” on other side,
and (c) “ Rhaphigaster sticticus ” cut from Dallas’s publication. The specimen lacks
antennal segment 5 on the left, segments 4 and 5 on the right, left anterior and
posterior tarsi, and right anterior leg.

After examining the crushed and broken female holotype of P. candens, described
by Distant (1890) as Nezara candens, I conclude that this specimen is a callow
example of P. stictica. The features by which Distant characterized P. candens, insofar
as the condition of the holotype permits comparisons, fall within the range of variation
exhibited by female specimens of P. stictica.

There is, however, a second species of Pellaea, a description of which follows.

Pellaea santarosensis, new species
(Figs. 1-10)

Description. Castaneous to light castaneous dorsally, often with rufous suffusion
along anterolateral margins of pronotum and on head; connexiva sometimes rufous,
usually with 2 subcircular black to metallic green macules on each segment, 1 in each
lateral angle. Ventral surfaces light castaneous, usually with irregular rufous suffusion,
or entirely rufous.

Punctation on head dark castaneous to black; most punctures in 2 longitudinal
bands on each jugum, these bands continuing onto vertex, the lateral band on each
side hooking laterad around ocellus. Width of head across eyes 3. 0-3. 3 mm, length
2.4-2. 7 mm; interocular width 1.8-1. 9 mm. Superior surface of antennifers with

1984

NEW SPECIES OF PELLAEA

345

Figs. 1-6. P. santarosensis. 1. Genital cup; paramere (p). 2. Left paramere, lateral face. 3.
Distal part of aedeagus, dorsal view; median penial lobe (mpl). 4. Same, lateral view; conjunctiva
(c). 5. Genital plates, caudo ventral view; basal plate (bp); 8th paratergite (pt8); 9th paratergite
(pt9); triangulum (t). 6. Distal part of spermatheca; spermathecal bulb (sb); proximal flange (pf);
sclerotized rod (sr); sclerotized cap (sc).

black to metallic green macule. Basal 2 segments of antennae black laterally, this
mark enveloping segment 2 apically to varying extent; segment 3 black, a narrow
pale ring covering basal joint; distal 2 segments fuscous to light brown, paler basally;
length of segments 0.6-0. 7, 1.2-1. 3, 1.4-1. 6, 1.4-1. 5, 1.3-1. 5 mm. Rostral segments
2-4 about 1.6-1. 8, 1.2-1. 5, 1. 0-1.1 mm long; last segment fuscous, apex between
metacoxae.

Most punctures on pronotum arranged in transverse, vermiform, dark castaneous
lines separated by polished interstices, most of these calloused, rarely in part ivory,
and contrasting strongly with remaining interstices. Humeri obtusely rounded, not
produced; anterolateral margins of pronotum straight to slightly convex; densely
punctate depressed line along anterior submargin interrupted mesially; pronotal width
at humeri 7. 0-7. 8 mm, mesial length 2. 7-3.0 mm.

Punctation of scutellum and coria somewhat irregularly disposed, a few subcal-
loused impunctate lacunae often present on coria. Scutellar width at base 4. 7-5. 2
mm; length 5. 1-5.8 mm, 6-12% greater than basal width. Hemelytral membranes
fumose.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Figs. 7-10. P. stictica. 7. Genital cup; paramere (p). 8. Left paramere, lateral face. 9. Distal
part of aedeagus; median penial lobe (mpl); conjunctiva (c). 10. Distal part of spermatheca;
spermathecal bulb (sb); spermathecal duct (sd); sclerotized rod (sr).

Ventral surface of head virtually impunctate; punctation of pleura moderate, con-
sisting in part of minutely darkened punctures; punctation of abdominal venter fine,
undifferentiated by color. Most setae on distal part of femora and along each tibial
sulcus arise in black dot; some dots on superior surface of femora may be enlarged
and confluent, most often on hind femora; superior femoral surface not prolonged
distally into spine; each tibial sulcus usually with black macule basally, occasionally
much of sulcus black. Spiracles narrowly ringed with black.

Basal plates posteriorly bent dorsad at nearly right angle; greatest dimension from
bend to posterior margin subequal to Vi length of remainder of plate; mesial margin
concave apically, exposing triangulum (Fig. 5). Both 8th and 9th paratergites concave
basally. Spermathecal bulb bearing 3 slender diverticula of unequal lengths (Fig. 6).
Spermathecal duct enlarged proximad of proximal flange; membranous dilation sur-
rounding sclerotized rod attached proximally to sclerotized cup.

Pygophore deeply and widely emarginate ventrally; ventral margins black; inferior
ridge large, exposed fully from caudal view. Parameres in form of stout hook with
large apical tooth curving ventrad, their apices hanging over inferior ridge and visible
from caudal view; lateral surface of each with field of stout pegs (Figs. 1 , 2), invested

1984

NEW SPECIES OF PELLAEA

347

generously with long hairs. Median penial lobes forming cup around ejaculatory duct;
conjunctiva reduced to ventral remnant (Figs. 3, 4).

Holotype. 6 labeled “Santa Rosa National Park, Guanacaste Prov. COSTA RICA.
2-4 May, 1980. D. H. Janzen & W. Hailwacks” Deposited in the American Museum
of Natural History.

Paratypes. 2529, 1 466: 1 labeled as holotype (222, 6 in each of following institutions:
AMNH, BMNH, CAS, FSCA, NR, NMNH; 2 RNH, 2 2 66 LHR); labeled as holotype
except date “7-8 May 1980” (222 LHR); labeled as holotype except date “5-7 June
1980” (2 UP); labeled as holotype except date “16-18 Jul 1980” (2 RNH); labeled
as holotype except date 9-17 Mar 1981 (222 DBT); labeled as holotype except date
1-15 Jun 1982 (222, 5 UFRGS, 6 RNH, 2 LHR, 366 UP, 5 DBT); labeled as holotype
except date “ 1 5-3 1 Jul 1 982 (2 UP); “Santa Rosa Nat. Pk. Prov. Guanacesta COSTA
RICA 12-14 Dec 1979, D. H. Janzen (2 UP).

Comments. This species differs notably from P. stictica in the form of the pygo-
phore, parameres, genital plates, spermatheca, absence of a spine at the apex of the
femora, and ventral markings. In P. stictica the concavity on each side of the genital
cup is interrupted by an obtuse, diagonal carina (Fig. 7); the parameres are elaborate
(Fig. 8); the 8th paratergite is slightly convex basally, the 9th paratergite nearly flat
basally, and the margin of the basal plates usually straight with the plates contiguous,
although sometimes the medial angle of each is diagonally truncated; the diverticula
of the spermathecal bulb are subequal in length and the duct between the proximal
flange and sclerotized rod is at least Vi longer than in P. santarosensis (Fig. 10); the
superior surface of the femora is prolonged distally into a small, angulate tooth; and
the venter has many large, black macules, with each spiracle in a large spot. The
aedeagus is similar but distinctive in the 2 species (Figs. 4, 9).

In the key to genera by Rolston and McDonald (1981), couplet 14 should be
replaced by the following:

14. Most punctures on pronotum arranged in transverse, vermiform lines separated by

polished, calloused interstices Pellaea St&l

Pronotal punctures rather evenly disposed, or if not interstices not calloused 1 5

ACKNOWLEDGMENTS

I am indebted to Daniel H. Janzen, University of Pennsylvania, for the specimens of Pellaea
santarosensis and to W. R. Dolling, British Museum (Natural History), for the privilege of
examining relevant type material.

LITERATURE CITED

Dallas, W. S. 1851. List of the specimens of Hemipterous insects in the collection of the
British Museum.

1 Paratype Depositories: American Museum of Natural History (AMNH); British Museum
(Natural History) (BMNH); California Academy of Sciences (CAS); Donald B. Thomas (DBT);
Florida State Collection of Arthropods (FSCA); author’s collection (LHR); National Museum
of Natural History (NMNH); Naturhistoriska Riksmuseum, Stockholm (NR); Rijksmuseum
van Natuurlijke Historic (RNH); Universidade Federal do Rio Grande do Sul (UFRGS); Uni-
versity of Pennsylvania (UP).

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Distant, W. L. 1880-1893. Insecta. Rhynchota, Hemiptera-Heteroptera. In: F. D. Godman
and O. Salvin, Biologia Centrali-Americana. London, Vol. 1.

Froeschner, Richard C. 1981. Heteroptera or true bugs of Ecuador: a partial catalog. Smith-
sonian Contributions to Zoology, no. 322.

Rolston, L. H. 1983. A revision of the genus Acrosternum Fieber, subgenus Chinavia Orian,
in the Western Hemisphere (Hemiptera: Pentatomidae). J. New York Ent. Soc. 91(2):
97-176.

Rolston, L. H. and F. J. D. McDonald. 1981. Conspectus of Pentatomini genera of the Western
Hemisphere— Part 2 (Hemiptera: Pentatomidae). J. New York Ent. Soc. 88(4), 1980:
257-272.

Received November 22, 1983; accepted April 10, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 349-351

KARYOTYPE OF CONOMYRMA FLAVA (MCCOOK)
(HYMENOPTERA: FORMICIDAE) 1

James C. Cokendolpher and Oscar F. Francke

Departments of Entomology and Biological Sciences,

Texas Tech University, Lubbock, Texas 79409

Abstract.— The karyotype (2N — 26) of the ant Conomyrma flava is reported from material
collected in central Texas. The chromosome numbers and morphology match those of Cono-
myrma bicolor from the western U.S.A. more closely than those of Conomyrma spp. from Peru
and Brasil. A lactic acid dissociation, air-drying technique with Giemsa staining for ant chro-
mosomes is described.

Conomyrma Forel and all its species were considered to belong in Dorymyrmex
Santschi, until Kusnezov (1952) separated the two genera, Dorymyrmex and Cono-
myrma. He further divided Conomyrma into two subgenera, Biconomyrma Kus-
nezov and Conomyrma. Although these subgenera subsequently were elevated to
generic status by Kusnezov (1959), Biconomyrma was later synonymized with Cono-
myrma by Snelling (1973).

The taxonomy of the North American Conomyrma species is uncertain, and the
genus is in need of revision. Snelling (1973) synonymized all but three of the nominal
taxa from the U.S.A. One species, C. insana (Buckley), cannot be recognized with
certainty because the type material is lost (J. C. E. Nickerson and J. C. Trager, pers.
comm.) and the original description (Buckley, 1866) is vague. Conomyrma flava
(McCook) was synonymized with C. insana by Snelling (1973), but has since been
determined to be a valid species by the late William F. Buren (J. C. Trager, pers.
comm.). We use the name C. flava for the specimens reported here and have deposited
voucher specimens, as indicated below, for later study.

Apparently unaware of the taxonomic changes proposed by Kusnezov (1952),
Crozier (1968, 1970) reported the karyotypes of the following three species of Cono-
myrma: Dorymyrmex bicolor (Wheeler), Dorymyrmex Ithoracius (Santschi), and Do-
rymyrmex ‘Ipulchellus (Santschi) ( =Dorymyrmex sp. in 1968 paper). Two of the
species reported by Crozier are from South America, whereas C. bicolor is from the
U.S.A. We herein report the karyotype of a second Conomyrma sp. from North
America.

MATERIALS AND METHODS

Workers and brood of Conomyrma flava (McCook) collected at Camp Verde, Kerr
Co., Texas, were maintained in the laboratory until suitable material (last instar
larvae) became available. Slides were prepared from five larvae and scored for diploid
number and centromeric position.

1 Supported by the Texas Department of Agriculture Interagency Agreement IAC (83-84)-
0853. Contribution No. T-10-154, College of Agricultural Sciences, Texas Tech University.

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

The larval heads were removed and opened in hypotonic solution. A 1% sodium
citrate and 0.075 M KC1 hypotonic solutions were used for times from 5 to 30
minutes, respectively. Although all preparations were similar in chromosome spacing
among individual spreads, the best results were obtained using 0.075 M KC1 solution
for 15 to 20 minutes. The heads were fixed in Camoy’s fixative (3:1 absolute meth-
anol : glacial acetic acid) for 30 minutes and then placed in a drop of dissociate solution
(3: 1 glacial acetic acid : 85% lactic acid) on the middle of a clean dry microscope slide.
Maceration of tissues with a pin and forceps aided dissociation of cells within the
dissociate solution. The dissociate solution will destroy the preparation if left in
contact with the cells for more than a couple of minutes. The moment the cells
became transparent to the unaided eye, three or four drops of fixative were dropped
onto the dissociated cell solution, and the slides were tilted back and forth several
times to spread the solution. After that, any remaining solution was poured off and
the slides were air dried for 24 hours, and then stained for 10 minutes in 6% Giemsa
stock solution in 15 M Sorenson’s buffer (pH 6.8). The above procedure is similar
to that proposed by Crozier (1968), differing mainly by the use of lactic acid disso-
ciation and Giemsa staining. The cells were not treated with colcemid or colchicine
as we, like Mehlhop and Gardner (1982), found this step unnecessary and we were
concerned with possible alterations of the karyotype by these agents as indicated by
Smith (1965).

Centromere classification follows that of Levan et al. (1964) as modified by Crozier
(1970).

The slides are not coverslipped and are numbered TTU Prep. #32-42. A voucher
series of the preserved workers and brood are deposited in the Entomological Col-
lection, The Museum, Texas Tech University (cat. no. 6476).

RESULTS AND DISCUSSION

The normal diploid chromosome number of the somatic head cells (presumably
cerebral ganglia) of five worker larvae was 2N = 26. A total of seventeen cells from
the five specimens were examined with no variation in counts. The karyotype (Fig.
1 ) consists of a pair of large subacrocentric, two pairs of medium metacentric, and
1 0 pairs of submetacentrics-to-subacrocentrics ranging in relative size from medium
to small.

The chromosome number of 2N = 26 for C. flava is identical to that of C. bicolor
reported by Crozier (1 970, Fig. 1 D), both species being from western North America.
By contrast, the two species from Peru and Brasil, C. ‘Ithoracica and C. Ipulchella,
respectively, have 2N = 18 (Crozier, 1970, Fig. IE, F). The karyotypes of both South
American species consist of a single pair of large metacentrics or submetacentrics
(almost subacrocentrics) and eight medium-sized metacentric chromosome pairs. In
contrast, the karyotypes of the two North American species consist of a pair of large
acrocentric-to-subacrocentric, two or five pairs of medium-sized metacentrics, and
10 or seven pairs of acrocentrics-to-submetacentrics ranging in size from small to
medium.

The differences noted here and in karyotypes by Crozier (1970) suggest two separate
groupings; however, these groups do not correspond to the genera/subgenera proposed
by Kusnezov (1952, 1959). Many more karyotypes and a thorough taxonomic re-

1984

KARYOTYPE OF CONOMYRMA FLA VA

351

Fig. 1 . Karyotype of Conomyrma flam (McCook), 2N = 26.

vision of Conomyrma spp. will be necessary to determine trends of karyotypic evo-
lution in this genus.

ACKNOWLEDGMENTS

We would like to thank Drs. J. C. Everett Nickerson and James C. Trager for information
on the taxonomy of Conomyrma and for reading the manuscript. Drs. M. Kent Rylander, James
K. Wangberg, and Mr. Frederick B. Stangl, Jr. also commented on the manuscript. We would
like to thank Dr. Rylander for allowing us the use of a microscope and camera in his care. Dr.
Nobuo Tsurusaki (Sapporo, Japan) kindly shared his knowledge of chromosomal preparation
techniques with the senior author.

LITERATURE CITED

Buckley, S. B. 1866. Descriptions of new species of North American Formicidae. Proc. Ent.
Soc. Phila. 6:152-172, 335-350.

Crozier, R. H. 1 968. An acetic acid dissociation, air-drying technique for insect chromosomes,
with aceto-lactic orcein staining. Stain Tech. 43(3): 17 1-1 73.

Crozier, R. H. 1970. Karyotypes of twenty-one ant species (Hymenoptera; Formicidae), with
reviews of the known ant karyotypes. Can. J. Genet. Cytol. 12:109-128.

Kusnezov, N. 1952. El estado real del grupo Dorymyrmex Mayr (Hymenoptera, Formicidae).
Acta Zool. Lilloana 10:427-448.

Kusnezov, N. 1959. Die Dolichoderinen-Gattungen von Siid-Amerika (Hymenoptera, For-
micidae). Zool. Anz. 162:38-51.

Levan, A., K. Fredga and A. A. Sandberg. 1964. Nomenclature for centromeric position on
chromosomes. Hereditas 52:201-220.

Mehlhop, P. and A. L. Gardner. 1 982. A rapid field technique for preparing ant chromosomes
for karyotypic analysis. Stain Tech. 57(2):99— 1 0 1 .

Smith, S. G. 1965. Heterochromatin, colchicine, and karyotype. Chromosoma (Berlin) 16:
162-165.

Snelling, R. R. 1973. The ant genus Conomyrma in the United States (Hymenoptera: For-
micidae). Los Angeles Co. Mus., Contrib. Sci. 238:1-6.

Received May 7, 1984; accepted June 6, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 352-364

KEY TO THE MALES OF THE NOMINATE SUBGENUS OF
EUSCHISTUS IN SOUTH AMERICA, WITH
DESCRIPTIONS OF THREE NEW SPECIES
(HEMIPTERA: PENTATOMIDAE)

L. H. Rolston

Department of Entomology,

Louisiana Agricultural Experimental Station,

Louisiana State University Agricultural Center,

Baton Rouge, Louisiana 70803

Abstract.— Keys to subgenera of the genus Euschistus Dallas and to the South American
species and one subspecies of the nominate subgenus are provided for males. Three new species
of the nominate subgenus are described: E. carbonerus, E. incus, and E. rohus. Euschistus
bifibulus var. guayaquilinus Kuhlgatz, 1903, is placed in the synonymy of E. taurulus Berg,
1878. E. lizerianus Pennington, 1922, is transferred to the genus Agroecus Dallas.

Most of the South American species of the genus Euschistus Dallas, 1851, are
contained in the subgenera Lycipta Stal, 1862, Euschistomorphus Jensen-Haarup,
1922, and Mitripus Rolston, 1978. Species of these subgenera may be identified from
the literature.

Recent papers on Lycipta are a revision by Rolston (1982), a nomenclatural cor-
rection by Thomas (1983), and a description of an additional species by Hildebrand
and Grazia (1983). Grazia and Hildebrand (1982) transferred Berecynthus monrosi
Piran, known only from females, to Euschistus. Although they did not place this
species in a subgenus, their drawings of the head and spermatheca indicate that E.
monrosi very likely belongs in the subgenus Lycipta. If this is so, there are currently
12 species in this subgenus.

Euschistomorphus, whose salient feature is the apically contiguous juga, contains
the species E. longiceps Berg, 1891, and E. albidus Jensen-Haarup, 1922, both quite
distinctive.

Rolston (1978) proposed the subgenus Mitripus to contain seven species, but he
misidentified two of these species, one of them previously undescribed, for want of
examining the relevant types. Grazia (pers. comm., 1983) proposes to correct these
errors, raising to eight the number of species in this subgenus.

In the nominate subgenus there are 14 species and one subspecies known from
South America, and another species, E. nicaraguensis, may occur there since its range
includes Panama. Rolston (1974) described or redescribed nine of these taxa, viz. E.
agudus, E. bifibulus, E. cremator cremator, E. cremator orbiculator, E. emoorei, E.
herns, E. nicaraguensis and E. rufimanus.

Here keys are provided for the males of the subgenera and species-group South
American taxa of the nominate subgenus. One uncommon species, E. quadripunc-
tatus, is redescribed, and three new species described.

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SYSTEMATIC CHANGES

Euschistus taurulus Berg, 1878

Euschistus taurulus Berg, 1878, Anal. Soc. Cient. Arg. 5:305-306.

Euschistus bifibulus var. guayaquilinus Kuhlgatz in Bayern, 1903, Berliner Entomol.

Zeit. 47 (1902):247, 254-256, fig. 2, 2a. New Synonymy.

Kuhlgatz (1903) included two drawings in his description of E. bifibulus var.
guayaquilinus, one a ventral view of a male. This drawing shows quite clearly a
median, obtuse tooth on the posterior pygophoral margin, a character which in
combination with the form of the humeri and widely spaced, posteriorly inclined
pronotal denticles is sufficient to identify the insect as E. taurulus, a common species
throughout South America except in the extreme northwest.

Agroecus lizerianus (Pennington), New Combination
Euschistus lizerianus Pennington, 1922, Physis 6:316-317.

The female type in the Drake collection, National Museum of Natural History,
Washington, D.C., was examined. The form of the genital plates and double row of
setae on the interior femoral surfaces, with each seta arising from a small tubercle,
place this species in the genus Agroecus Dallas, 1851. The species is apparently distinct
from those already in Agroecus.

KEY TO SUBGENERA FOR MALES

1 . Juga contiguous or nearly so before tylus for distance subequal to or greater than

diameter of eye Euschistomorphus

- Juga often convergent but usually well separated apically, rarely briefly contiguous
2

2(1). Genital cup with pale membranous cushion on each lateral wall, this cushion
running just beneath anterolateral portion of rim then bending ventrad near an-
terior wall of genital cup Lycipta

Genital cup with dark rigid carinae or denticles on lateral walls 3

3(2). Pair of proctigeral tubercles located about midway between apex and base of
proctiger or nearer base than apex; rim of genital cup interrupted on each side of
superior ridge by diagonal depression (excepting E. convergens ) Mitripus

- Proctigeral tubercles located on distal half of proctiger; anterolateral and anterior

portions of genital cup rim continuous Euschistus

KEY TO SPECIES AND SUBSPECIES FOR

MALES OF NOMINATE SUBGENUS

1 . Evaporative areas with dark punctures 2

- Evaporative areas lacking dark punctures, occasionally with small reddish spots
5

2(1). Denticles on pronotum widely spaced (Fig. 1); tylus usually acute apically

rufimanus St&l

- Denticles on pronotum closely spaced, most separated by distance no greater than
length of denticle (Fig. 22); tylus rounded apically

3

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Figs. 1-1 3. 1-3. Anterolateral pronotal margin and humerus. 1 . E. rufimanus. 2. E. quickua.

3. E. taurulus. 4, 5. Posterior margin of pygophore and inferior ridge, caudal view. 4. E. taurulus.
5. E. bifibulus. 6, 7. Scutellar apex. 6. E. heros. 7. E. quadripunctatus. 8, 9. Posterior pygophoral
margin, caudal view. 8. E. atrox. 9. E. nicaraguensis. 10, 11. Variation in posterior pygophoral
margin of E. emoorei, caudoventral view. 12. Connexival margin of E. quickua. 13. Paramere
of E. quickua. Dimensional lines equal 0.5 mm.

3(2). Scutellar apex with pale macule or broad border (Fig. 6); sutures between stemites

black except laterad of spiracles hews (Fabricus)

- Scutellar apex usually unmarked, occasionally with narrow pale border; sutures

between stemites usually concolorous with stemites 4

4(3). Anterior disk of pronotum convex in profile; posterior margin of pygophore from
caudal view with moderately deep emargination slightly sinuous at bottom (Fig.

9) nicaraguensis Rolston

- Anterior disk of pronotum an inclined plane in profile; posterior margin of py-
gophore trisinuately emarginate from caudal view (Fig. 8) atrox (Westwood)

5(1). Pair of brown calluses on apex of scutellum separated by small pale area (Fig. 7)

quadripunctatus St<U

6

Apex of scutellum not so calloused

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355

Figs. 14-21. 14, 15. Posterior pygophoral margin, caudoventral view. 14. E. agudus. 15.

E. backhauseni. 16, 17. Parameres. 16. E. agudus. 17. E. backhauseni. 18, 19. Variation in
posterior pygophoral margin of E. crenator cremator, caudoventral view. 20, 21. Variation in
posterior pygophoral margin of E. crenator orbiculator, caudoventral view. Dimensional lines
equal 0.5 mm.

6(5). Humeri produced into short spine usually directed anterolaterad (Fig. 25)

incus new species

Humeri produced laterad, obtuse to spinose 7

7(6). Most denticles on pronotum separated from each other by distance equal to or

greater than length of denticle (Figs. 2, 3) 8

Most denticles on pronotum closely spaced, adjacent denticles usually contiguous

basally or separated by distance less than length of denticle 10

8(7). Lateral margins of abdomen strongly serrate (Fig. 12); tylus surpassing juga, sep-
arated apically from each jugum by small incision; parameres with long stout

process (Fig. 13) quickua Piran

- Lateral margins of abdomen weakly serrate; tylus and juga subequal in length, not

separated apically by distinct incisions; parameres lacking process 9

9(8). Posterior margin of pygophore smoothly concave from caudal view (Fig. 5); rim
of genital cup on each side of superior ridge smoothly rounded; serrated carina

within genital cup on each lateral wall bifibulus (Palisot de Beauvois)

Posterior margin of pygophore with small protuberance mesially (Fig. 4); rim of

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genital cup on each side of superior ridge with thin black carina which bends into
genital cup near its termination about midway between superior and inferior ridges;
carina within genital cup on each lateral wall entire, with one denticle anteriorly

taurulus Berg

10(7). Slight medial protuberance on posterior margin of pygophore minutely notched

from caudoventral view (Fig. 34) rohus, new species

Posterior margin of pygophore without median tubercle 11

11(10). Conspicuous ivory macule on apex of scutellum; humeri stout, spinose, almost
entirely black laterad of hemelytra; width across humeri usually 7.0 mm or more,
rarely less 12

- Any ivory mark on apex of scutellum confined to narrow border; humeral shape
varies, black usually marginal or absent; width across humeri rarely as great as

7.0 mm 14

12(1 1). Antennae uniformly sordid yellow to rufous agudus Rolston

Each of last 2 antennal segments dark brown to black with broad pale band basally

13

13(12). Pronotal denticles black carbonerus, new species

- Pronotal denticles brownish yellow backhauseni Berg

14(1 1). Dorsal punctation on pronotum nearly uniform in density; black punctures if any

not concentrated along denticular margin; emargination in posterior margin of
pygophore broad, shallow, sinuous to subquadrate from caudoventral view (Figs.

10, 11) emoorei Rolston

Punctures along denticular margin usually crowded, black; posterior margin of
pygophore either sinuous and mesially concave, or convex with shallow concavity

mesially 15

15(14). Posterior margin of pygophore sinuous, concave mesially from caudoventral view

(Figs. 18, 19) cremator cremator (Fabricius)

- Posterior margin of pygophore convex with narrow shallow concavity mesially

from caudoventral view (Fig. 20); rarely sinuously convex with broad shallow
concavity mesially cremator orbiculator Rolston

Euschistus quadripunctatus Stal, 1860
Figs. 7, 22-24

Euschistus quadripunctatus Stal, 1860, p. 20

Description. Head as long as wide across eyes, 2.3-2. 5 mm. Margins of head
obtusely angular before eyes, briefly subparallel before apex of antenniferous tuber-
cles, 1.1 -1.2 mm wide here, slightly incised at apex, dorsally bronze black blending
to dark brown on disk (Fig. 22). Slight submarginal impression before eyes, base of
tylus somewhat elevated, disk otherwise rather flat; punctation even (excepting im-
punctate area mesad of each eye) deep, coarse, black or fuscous. Distance between
outer margins of ocelli 1.2-1. 3 mm. Antenniferous tubercles largely visible from
above. Basal segment of antennae not quite reaching apex of head, sordid yellow
with fuscous streak laterally; segments 2-5 fuscous, superior surface of 2 sometimes
pale, 3-4 narrowly and 5 broadly annulated at base with sordid yellow; length of
segments 0.6-0. 8, 1. 1-1.2, 1. 1-1.4, 1.6-1. 7, 1.8 mm. Head sordid yellow beneath
with black streak from eyes along suture above antenniferous tubercles; punctation
concolorous with sclerites, moderately coarse along buccalae, finer elsewhere. Buc-
culae obtusely lobed anteriorly, evanscent at base of head, subequal in length to basal

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357

Figs. 22-24. E. quadripuntatus, holotype. 22. Head and pronotum in part. 23. Posterior
pygophoral margin and inferior ridge (ir) caudal view. 24. Pygophore, caudoventral view.
Dimensional lines equal 0.5 mm.

segment of rostrum; remaining rostral segments 1.8-1. 9, 1.2-1. 4, 0. 9-1.0 mm in
length.

Pronotum 7. 7-8. 4 mm wide at humeri, 2.4-2. 7 mm long at meson; punctation on
disk moderately dense and strong, becoming fine, dense, confused along anterolateral
margins and on humeri. Humeral angles produced laterad, elevated, ivory at very
apex and on posterolateral margin. Denticles small, closely spaced, black. Ivory spot
toward meson on posterior border of cicatrices, disk otherwise yellowish brown
deepening to black where punctation confused. Scutellum longer than wide at base,
3. 8-4.0 mm wide basally, 4. 1-4.5 mm long; frena reaching about % distance from
base to apex; disk somewhat rugose; punctation uniform; fovea in each basal angle
small, black; pair of finely rugose dark brown calluses on apex of scutellum separated
by small pale area. Punctation on coria less dense than that on scutellum; white callus
at end of radial vein; membrane brown, veins simple or branched. Connexiva partially
exposed; punctation fine, shallow, dense; each segment sometimes with narrow trans-
verse ivory line at base, otherwise not conspicuously alternated.

Venter sordid yellow, concolorously punctate. Thorax with 4 black spots, one on
each subcoxae, one on each mesopleuron near distal end of supracoxal cleft. Evap-
orative areas matte, rugose, reaching nearly half way from ostiole to lateral thoracic
margins. Mesocoxae and metacoxae each with single dark spot on anterior surface;
femora and tibiae with many black spots; basal tarsal segments, and sometimes others,
darkest apically. Spiracles black; elongated black spot at anterolateral angles of all
sternites bearing spiracles.

Posterior margin of pygophore broadly emarginate, emargination containing 4
concavities (Figs. 23, 24). Inferior ridge largely visible from caudal view.

Length 11.5-12.5 mm.

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Distribution. Brazil (Guanabara, before 1960 the Federal District coextensive with
Rio de Janeiro).

Comment. Redescribed from 2 males, one the holotype in the Naturhistoriska
Riksmuseet, Stockholm.

Euschistus incus, new species
Figs. 25-30

Description. Dorsum light brown to light castaneous or gray; punctation light
castaneous to black, somewhat fine, often in aggregations on pronotum. Brownish
yellow beneath, sometimes finely flecked, tinted or blotched with orange or red;
punctation almost entirely concolorous, very fine on aciculate abdominal venter.
Length 9.7-10.9 mm.

Width and length of head subequal, 2. 1-2.3 mm wide across eyes; lateral jugal
margins briefly parallel or subparallel above apex of antennifers; tylus as long as or
slightly longer than juga (Fig. 25). Basal 3 segments of antennae nearly concolorous
with ventral surface of head, last 2 segments darker, usually basal segment and
sometimes basal 3 segments fuscous dotted; basal segment not reaching apex of head;
length of segments 0.5-0. 6, 0.7-0. 9, 1. 1-1.3, 1. 1-1.3, 1.1-1. 2 mm. Rostral segments
2-4 about 1.6-1. 8, 0.7-1. 0, 0.8-0.9 mm.

Pronotum 2. 2-2. 5 mm long at meson, 2. 6-2. 8 times as wide across humeri. Den-
ticles on anterolateral margins of pronotum concolorous with adjacent disk, some-
times with black punctures on base, nearly always discrete and sometimes widely
separated, occasionally as few as 5 denticles on a side. Humeri moderately produced;
apex acute, usually spinose, directed anterolaterad or rarely laterad.

Scutellum usually a little longer than wide at base, 3. 3-3. 9 mm across base; fovea
in basal angles small, shallow, blackish; disk rather smooth, moderately convex;
narrow margin of apex impunctate, occasionally conspicuously paler than disk. Mem-
branes of hemelytra hyaline; veins simple or branched, brown. Connexiva moderately
exposed, dark with pale marginal spot in middle of each segment; punctation close,
coarse, pale.

Evaporative areas unicolorous. Small fuscous dots on femora and tibiae least
numerous on hind legs. Basal angles of sternites each with black macule; apical angles
immaculate. Spiracles concolorous with supporting sclerite.

Broad emargination in posterior margin of pygophore sloping on each side to low
median protuberance of varying width (Figs. 27, 28); median protuberance sometimes
inapparent from caudal view, rarely absent; posterior pygophoral margin sinuously
truncate from dorsal view (Fig. 26). Anterolateral angles of inferior ridge each bearing
compressed black denticle oriented diagonally, its anterior limit laterad of posterior
limit. Lateral rims of genital cup roughened by low black carinae, these variable in
number, directed diagonally cephalad from within cup toward rim; larger carina,
mesially concave in profile, located entad and cephalad of row of lesser carinae along
each lateral rim. Parameres strongly cupped; oval area of fine denticles on lateral
face at apex (Fig. 29). Apex of conjunctiva on each side with or without lightly
pigmented spot; lateral diverticulum on each side entirely hyaline (Fig. 30). Large
sclerotized cap present at base of penisfilum. Thecal processes short; median thecal
lobe weakly developed, lateral thecal lobes absent.

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Figs. 25-30. E. incus. 25. Head and pronotum in part. 26. Posterior pygophoral margin,
dorsal view. 27. Posterior pygophoral margin and inferior ridge, caudal view. 28. Posterior
pygophoral margin, caudoventral view. 29. Paramere. 30. Aedeagus, lateral view; conjunctiva
(c); diverticulum (d); sclerotized cap (sc); thecal process (tp). Dimensional lines equal 0.5 mm.

Types. Holotype: male, labeled (a) “No. 53, Lima, Peru, IV- 1965” (b) “F. Asneros,
M. Tello.” Deposited in National Museum of Natural History, Washington, D.C.
Type No. 72132. Paratypes: 11 males, 10 females. “Cuenca, Ecuador, June 1939,
Lucio Vivar, preying on lep. larva” (9 LHR); “Peru, Chira Valley, V- 1 953, on cotton”
(6 NMNH); “Paila, Peru” remainder illegible (6 NMNH); “Canete, Mch. 4, 1941, E.
J. Hambleton” (3 LHR); same data except date “Mch. 13, 1941” (9 NMNH); same
data except date “Mch. 10, 1942” (6 UNLP); same data except date “Mch. 17, 1942”
(9 UNLP); “Canete, Feb. 11, 1941, E. J. Hambleton, on cotton (9 NMNH); same
data except date “Mch. 17, 1941” (9 NMNH); same data except date “Mch. 6, 1941”
(S NR); “No. 23-39, Piura, 3 1 -V- 1939, Schaefer Coll,” (266, 9 NMNH: 9 NR): “Piura,
Peru, May 1, 1941, E. J. Hambleton, on cotton” (6 LHR); “Huacachima Yea, E.
Escomel” (6 AMNH); (a) “Lobitos, H. P. Manton” (b) “Brit. Mus., 1939, 489” (399,
266 BMNH)

Distribution. Western Peru and southern Ecuador.

Comment. The peculiar humeri will immediately identify most specimens. The
male genitalia suggest an affinity with E. crenator (F.).

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Euschistus rohus, new species
Figs. 31-37

Description. Light brown dorsum usually tinted with red, becoming light castaneous
on head and anterior portion of pronotum; head thinly edged in black, anterolateral
margins of pronotum and apex of humeri black bordered. Brownish yellow beneath,
concolorously punctate. Length 7. 9-9. 4 mm.

Head 1.8-1. 9 mm across eyes, subequal in length; lateral jugal margins sigmoid,
tapering to apex; tylus slightly exceeding juga (Fig. 31). Antennae light brown with
tiny dark dots on 3 basal segments; each of last 2 segments usually castaneous with
pale but not clearly annulated base; basal segment reaching apex of head; length of
segments 0.5-0.6, 0.9-1.0, 1.0-1. 1, 1.3-1. 5, 1.4-1. 5 mm.

Pronotum 2. 2-2. 4 mm long at meson, 2. 8-3. 2 times as wide at humeri. Antero-
lateral margins evenly concave, closely and rugosely denticulate; narrow black border
toward apex consisting of black punctures, toward humeri involving entire margin.
Humeri acutely produced laterad, neither turned forward nor much elevated above
disk.

Scutellum 3. 0-3. 4 mm wide at base, subequal in length; each basal angle with 2
or 3 confluent punctures, scarcely foveate; color and punctation of apex as on disk.
Punctation of hemelytra darker and stronger on costal margins at base than on disks;
impunctate lacuna at distal end of radial vein neither calloused nor differentially
colored; membranes lightly fumose, veins simple or branched. Connexiva little or
not at all exposed, darker than coria, at least along margin, with small pale marginal
spot in middle of each segment; punctation close, coarse, pale.

Pleural surfaces of thorax with 5 small black dots as usual for genus; evaporative
areas unicolorous. Small fuscous dots scattered on femora and tibiae. Spiracles con-
colorous with sternites; basal angle of stemites black spotted, apical angles immac-
ulate.

Posterior margin of pygophore convexly arcuate from dorsal view (Fig. 32), sin-
uously concave from caudal view and sinuously truncate from caudoventral view, a
low median minutely notched truncate protuberance apparent from both caudal and
caudoventral views (Figs. 33, 34). Small black denticle on each lateral wall of genital
cup just visible above paramere. Parameres quite thin apically with reticulated area
on lateral surface (Figs. 35, 36). Conjunctiva on each side terminating in hook with
acute black apex (Fig. 37); lateral lobes of theca strongly developed, extending pos-
teriorly well beyond median lobe.

Type. Holotype: male, labeled (a) “Paramba, Ecuador” (b) “Collection Rosenburg.”
Deposited in National Museum of Natural History, Washington, D.C. Type no.
75557. Missing last antennal segment on right, last 2 on left, mesothoracic and
metathoracic legs on left, tarsi of these legs on right. Paratypes: 4 males, 4 females.
Same data as holotype (9 LHR); “Cachabe, low c, XII-96 (Rosenburg) (69 BMNH,
S UNLP); same data plus second label “Ecuador, Rosenburg, 99-104” (6 LHR);
“Paramba, 3500', II 97, dry season, Rosenburg” (9 BMNH); “60 K NW Cali, Co-
lombia, 29-VI-1972” (9 HDE); “Pipe Line Rd, Canal Zone, 22-X-1972, L. H. Rol-
ston” (6 LHR).

Distribution. Ecuador, Colombia, Panama (Canal Zone).

Comments. This species closely resembles E. vetus Rolston from Costa Rica. The

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361

Figs. 31-37. E. rohus. 31. Head and pronotum in part. 32. Posterior phygophoral margin,
dorsal view. 33. Same, caudal view. 34. Same, caudoventral view. 35. Paramere. 36. Same,
rotated 90 degrees. 37. Aedeagus, lateral view, conjunctiva (c). Dimensional lines equal 0.5
mm.

males are readily identified by the genitalia, but the females of these 2 species cannot
be separated with confidence. The ranges of these species are not known to overlap,
but they may do so in western Panama.

Euschistus carboneurs, new species
Figs. 38-44

Description. Dorsum dark brown to fuscous becoming black on humeri, with apex
of scutellum ivory; punctures black, evenly distributed excepting scattered subcal-
loused spots. Venter brownish yellow, concolorously punctate. Length 9.5-10.7 mm.

Width of head across eyes 1. 9-2.0 mm, length subequal. Lateral margins of head
tapering sinuously from eyes to rather narrowly rounded apex; tylus a little longer
than juga (Fig. 38). Antennae brownish-yellow to fuscous with darker dots on basal
or basal 3 segments and pale ring on proximal fourth of last two segments; basal
segment reaching apex of head; length of segments 0.6-0. 7, 0. 9-1.1, 1.3-1. 8, 1.4-
1.8, 1.4-1. 6 mm.

Pronotum 2. 3-2.6 mm long at meson, 2.9-3. 3 times as wide at humeri. Antero-
lateral margins concave from dorsal view, denticulate nearly to base of humeri;
denticles closely spaced, black. Humeri acutely produced laterad, apically elevated

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Figs. 38-44. E. carbonerus. 38. Head and pronotum in part. 39. Posterior pygophoral
margin, dorsal view. 40. Posterior pygophoral margin and inferior ridge, caudal view. 41.
Posterior pygophoral margin, caudoventral view. 42. Paramere. 43. Aedeagus, lateral view;
conjunctiva (c). 44. Genital plates; basal plate (bp); 2nd gonocoxae (gx2); paratergite 9 (pt9).
Dimensional lines equal 0.5 mm.

above pronotal disk. Small subcalloused spot near posteromesial margins of each
cicatrice little or no paler than cicatrice.

Scutellum as wide as or slightly wider than long, 3. 3-3. 8 mm across base; a few
black punctures in each basal angle forming shallow fovea. Membranes of hemelytra
fuscous; veins simple or branched. Connexiva narrowly exposed; punctation dense,
pale; small pale spot usually present submarginally in middle of each segment.

Evaporative areas unicolorous. Femora black dotted, more sparsely so than tibiae.
Anterolateral and posterolateral angles of abdominal sternites minutely marked in
black. Spiracles concolorous with surrounding surface.

Broad emargination in posterior margin of pygophore concave to moderate depth
from caudal view (Fig. 40), shallowly concave from caudoventral view (Fig. 41),
truncate and slightly sinuous from dorsal view (Fig. 39). Small black denticle on each

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lateral wall of genital cup obscured by parameres. Conjunctiva on each side termi-
nating in apically pigmented hook (Fig. 43). Lateral lobes of theca large, extending
posterolaterad beyond median lobe. Parameres as in Figure 42.

Basal plates overlapping along basal portion of medial margin; posterior margin
sinuous, convexity at lateral angle appressed to basal concavity of 9th paratergites
(Fig. 44). Second gonocoxae longitudinally grooved mesially.

Types. Holotype: male, labeled “Brazil, Para, Jacareacanga, Dec. 1968, M. Al-
varenga.” Deposited in American Museum of Natural History. Paratypes: 6 males,
2 females. Same data as holotype (9 AMNH, <3 LHR, <3 NMNH); same data as holotype
except date and collector “Feb. 1969, F. R. Barbosa” (6 AMNH); “Santarem, Acc.
No. 2966” (<3 CMNH, <3 UNLP); “Rio de Jan. Brazil. Acc. No. 2966” (9 LHR);
“Brazil, Rondonia” on white paper pasted on purple museum label (<3 IRSNB). The
last paratype listed and both female paratypes have the binomen hand lettered on
the paratype label; it is printed on the other paratype labels.

Distribution. Apparently widely distributed in Brazil, although all but one specimen
came from along the Amazon River and one of its tributaries, the Guapore River,
which marks part of the eastern Bolivian border.

Comments. This species belongs in a group of six species that are similar in size,
appearance and genitalia: E. atrox, E. heros, E. nicaraguensis, E. agudus, E. back-
hauseni and E. carbonerus. The first three species have dark punctures in the evap-
orative areas and the last three species have unicolorous evaporative areas. The color
of the pronotal denticles, black in E. carbonerus, separate this species from E. agudus
and E. backhauseni, both of which have ivory colored denticles. Each of these three
species differ somewhat in the shape of the posterior pygophoral margin and para-
meres (Figs. 14-17, 40, 42).

ACKNOWLEDGMENTS

I am indebted to the following individuals and institutions for the loan of specimens and
other assistance relevant to this study: W. R. Dolling (British Museum Natural History); H.
Dodge Engleman, Coco Solo, Panama; Richard C. Froeschem (National Museum of Natural
History, Washington, D.C.); Per Lindskog and Per Inge Persson (Naturhistoriska Riksmuseet,
Stockholm); Ricardo A. Donderos and Luis de Santis (Universidad Nacional de La Plata, La
Plata, Argentina); G. Schmitz (Musee Royal de l’Afrique Central and Institut Royal des Sciences
Naturelles de Belgique, Tervuren); Randall T. Schuh (American Museum of Natural History,
New York); and G. Wallace (Carnegie Museum of Natural History, Pittsburg).

The following abbreviations are used in the text for paratype depositories: American Museum
of Natural History (AMNH); British Museum (Natural History) (BMNH); Carnegie Museum
of Natural History (CMNH); H. Dodge Engleman collection (HDE); Institut Royal des Sciences
Naturelles de Belgique (IRSNB); Author’s collection (LHR); National Museum of Natural
History (NMNH); Naturhistoriska Riksmuseet (Stockholm) (NR); Universidad Nacional de La
Plata (UNLP).

LITERATURE CITED

Berg, C. 1878. Hemiptera Argentina: essayo de una monographia de los Hemipteros Heter-
opteros y Homopteros de la Republica Argentina. An. Soc. Cient. Arg. 5(6):297-314.
Grazia, J. and R. Hildebrand. 1982. Revisao do genero Berecynthus St&l, 1862 (Heteroptera,
Pentatomidae, Pentatomini). Rev. Bras. Entomol. 26(2): 173-1 82.

364

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Hildebrand, R. and J. Grazia. 1983. Uma nova especie de Euschistus Dallas, 1851 (Heter-
optera, Pentatomidae) Iheringia, ser. Zool., Porto Alegre 62:81-88.

Kuhlgatz, T. in T. von Bayern. 1903. VI Rhynchoten. Berliner Entomologische Zeitschrift
47(1902):243-278, pis. 4, 5.

Pennington, M. S. 1922. Hemipteros nuevos para la Republica Argentina (primera parte).
Physis (Buenos Aires) 6:315-319.

Rolston, L. H. 1974. Revision of the genus Euschistus in Middle America (Hemiptera, Pen-
tatomidae, Pentatomini). Entomol. Am. 48(1): 1-102.

Rolston, L. H. 1978. A new subgenus of Euschistus (Hemiptera: Pentatomidae). J. New York
Entomol. Soc. 86(2): 102-120.

Rolston, L. H. 1982. A revision of Euschistus Dallas subgenus Lycipta St&l (Hemiptera:
Pentatomidae). Proc. Entomol. Soc. Wash. 84(2):28 1-296.

St&l, C. 1 860. Bidrag till Rio Janeiro-traktens Hemipter-fauna. Kongliga Svenska Vetenskaps
Akademien 2(7): 1-84.

Thomas, D. B., Jr. 1983. On the homonymy of Euschistus luridus Dallas, 1851 (Hemiptera:
Pentatomidae) Proc. Entomol. Soc. Wash. 85(1): 182.

Received November 22, 1983; accepted April 10, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 365-370

CHEMICAL ANALYSIS OF THE MALE AEDEAGAL
BLADDER IN THE FIRE ANT, SOLENOPSIS INVICTA BUREN1

David E. Ball, H. J. Williams and S. B. Vinson

Department of Entomology, Texas A&M University,

College Station, Texas 77843

Abstract.— The aedeagal bladder in the male reproductive system of the fire ant Solenopsis
invicta Buren lies within the external genitalia. The lumen is lined by a single layer of squamous
epithelium surrounded by heavy musculature. In mature, sterile and mated males it contains
a milky substance but no fluid is found in newly eclosed males. The fluid in the bladder consists
of fatty acid esters (glycerol monodecanoate and glycerol monododecanoate) in a proteinaceous
matrix. The function of the secretion remains unknown.

The male reproductive system in the imported fire ant, Solenopsis invicta Buren
consists of the testes, vas deferens, seminal vesicles, accessory gland, ejaculatory duct,
wedge, and external genitalia (Ball and Vinson, 1984). An additional organ, the
aedeagal bladder, has been described in many species of male ants but its function
is unknown and no chemical analyses of its contents have been reported.

The aedeagal bladder was first described by Clausen (1938). Forbes ( 1 954) discussed
the anatomy and histology of the aedeagal bladder in Camponotus pennsylvanica
DeGeer in considerable detail. Forbes (1958) also described the aedeagal bladder in
the army ant, Eciton hamatum F. Later, Forbes and Do-Van-Quy (1965) investigated
the histology of the bladder in the legionary ant, Neivamyrmex harrisi (Haldeman).
Ford and Forbes (1980) showed that the bladder is present in several species of
doryline ants and Hagopian (1963) has described an aedeagal bladder in the ponerine
ant, Rhytidoponera metallica F. Smith. In each case, the aedeagal bladder was found
to be an ovoid muscular sac lined with squamous epithelium, filled with a basophilic
granular mass and emptying posteriorly into the region of the ejaculatory duct between
the inner genitalic valves.

Trakimas (1967) described an aedeagal bladder in Myrmica rubra L. but she stated
that the granular material in the lumen stains acidophilicly and is found in a greater
amount in mature males with degenerated testes. Tice (1967) did an extensive study
on the male reproductive system of the black imported fire ant, Solenopsis saevissima
richteri Forel, and also reported a muscular aedeagal bladder with a granular aci-
dophilic substance in the lumen. Ball and Vinson (1984) found an aedeagal bladder
lined with a single layer of squamous epithelium in the male reproductive system of
the red imported fire ant, S. invicta Buren, and determined that the material in the
lumen contains a small amount of protein. They also determined that the bladder
material is found in approximately equal amounts in mature, mated and sterile males.

1 This manuscript is approved as TA 19307 by the Director of the Texas Agricultural Ex-
periment Station in cooperation with A.R.S./U.S.D.A. This research was supported by the
Texas Department of Agriculture Interagency agreement IAC (84-85) 1018.

366

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

The purpose of this investigation was to determine the chemical composition of
the contents of the aedeagal bladder in the fire ant, S. invicta.

MATERIALS AND METHODS

Dissection and collection procedures. Newly eclosed (one day old) males and sterile
males (Hung et al., 1974) at least two weeks old were collected from laboratory
colonies. Sterile males lack testes and are, therefore, incapable of sperm production.
Additional males, collected as they flew from the surface of the mound prior to
mating (preflight) and after they returned (postflight) were also examined.

Males were pinned through the ventral surface of the thorax, the terminal gastral
segments were grasped with microforceps and pulled apart exposing the external
genitalia, the claspers were carefully pulled and the accessory glands were severed
from the claspers. The claspers were then quickly positioned with the posterior end
of the genital valves pointing upward and the muscular contractions within the
claspers caused the expulsion of the aedeagal bladder material which was collected
by capillary action using a flame-drawn pipette.

Chemical analysis. A variety of chemicals was used to test the solubility of the
aedeagal bladder material. Samples were collected from 5 mature males and placed
into sterilized glass tubes containing either acetone, ethanol, pentane, 0.1 N NaOH,
0.1 N HCL or distilled water (pH 7).

During dissection of the aedeagal bladder, the expelled material appeared as a
white particulate material resembling deposits of uric acid. Therefore, the material
was collected from 20 mature males, placed into 0. 1 N NaOH and tested concurrently
with a standard uric acid sample on thin layer chromatography (TLC) on cellulose
using 0. 1 N NaOH as the solvent. The plate was then examined under ultraviolet
light. The material also appeared to have droplets of suspended oil indicating lipid-
like substances. To determine the presence of free lipids, the aedeagal bladder material
was collected from 40 males and placed into ethyl ether. Standard samples (Sigma
Chem. Corp., St. Louis, Mo. 63178) of glycerides (distearin, triolein, and monopal-
mitin), a fatty acid (oleic acid), a steroid (stigmasterol) and an ester (palmitic acid
ethyl ester) were mixed with chloroform to obtain final concentrations of 1 jug/jul.
Five and 1 0 /i\ drops of the sample and standards were spotted onto a silica gel TLC
plate and air dried. The plate was then placed in a chromatography tank with a
hexane: ethyl ether solvent (50:50). The solvent front was allowed to migrate to
within 10 mm of the top and the plate was then transferred to an iodine developing
tank. Aedeagal bladder material and standard spots were then compared and RF
values computed.

As a further analysis, the material, collected from 40 mature males, was dissolved
in pentane as previously determined and injected onto a column gas chromatograph
(GC). The samples were analyzed on a Tracor 550 GC with a flame ionization detector
modified with an all glass variable split ratio collection system. Separations were
obtained on a 1.83 M x 4 mm i.d. column of 3% OV-101 on Chromosorb 750, 100-
120 mesh at a nitrogen flow rate of 60 ml/min programmed from 50 to 270°C at
10°C/min. Standard monoglycerides were analyzed under the same conditions. Mass
spectral (MS) analyses were performed on a Finnigan 1020 OWA quadrupole GC-

1984

CHEMICAL ANALYSIS IN SOLENOPSIS

367

Table 1 . RF values for TLC lipid analyses of standard and gland extract samples. Solvent-
ethyl ether/hexane (50:50), iodine vapor visualization on silica gel F254 plate.

Sample

RF value

Monoplatmitin

0.05

Distearin

0.39

Triolein

0.61

Oleic acid

0.45

Stigmasterol

0.67

Palmitic acid ethyl ester

0.67

Gland extract

0.05

MS unit using a 12 M x 0.33 mm i.d. BPI column, 6 psi, temperature programmed
from 60 to 220°C at 20°C/min. Monoglyceride standards (Sigma Chem. Corp., St.
Louis, Mo. 63178) were used, with RT 9.41 min for C10 and 10.65 min for C12 under
the above conditions. Hexane gland extracts were subjected to GC-MS analysis
without preliminary purification.

The polyacrylamide disc gel electrophoresis procedure described by Davis (1964)
was used for verification of the presence of lipoproteins. A 7.5% separating gel was
allowed to polymerize in glass tubes (i.d. 4.5 mm, length 75 mm) precleaned in
chromic acid followed by the addition of a 3.3% stacking gel to the top of the
separating gel. The aedeagal bladder material from 20 males was collected and placed
in 100 /i\ of an aqueous 1% mercaptoethanol/ 1 % SDS disruption buffer solution.
Standards (Sigma Chem. Corp., St. Louis, Mo. 63178) consisting of 100 ng each of
phosphorylase A (330 K), IgG (150 K), bovine serum albumin (67 K), ovalbumen
(43.5 K) and cytochrome C (12.5 K) were also placed in 100 ng of disruption buffer.
Standards and the aedeagal bladder sample were heated for 1 5 min at 60°C before
applying 30 yul aliquots per tube. Electrophoresis proceeded at 8 mA per gel tube for
ca. 4 hr.

A Cahn gram electrobalance was used to determine the percent composition of
proteins and/or lipids in the aedeagal bladder material. The contents of the bladder
from 30 mature males were extracted, placed on aluminum foil pans, weighed and
moved to a desiccator. Once a constant weight was obtained, the material was im-
mersed 3 times in hexane and returned to the desiccator. After reaching a constant
weight again, the material was placed 3 times in chloroform : methanol (50:50) for
5 min each. Two samples and a control (empty pan) were analyzed concurrently.

Transmission electron microscopy was used to observe the ultrastructure of the
bladder material. The claspers with the intact aedeagal bladder were removed from
several mature males and fixed in 2% glutaraldehyde/2% acrolein/ 1.5% DMSO in
0.133 M sodium cacodylate buffer (pH 7.4) at room temperature for 6 hr. The
specimens were then rinsed 3 times in buffer and post-fixed in 2% osmium tetroxide
in 0.1 M buffer and 0.2 M sucrose for 3 hr. The samples were rinsed 3 times in 0.1
M buffer, 2 times in distilled water and transferred to 1% aqueous uranyl acetate
overnight. After dehydration in an ethanol series, the specimens were embedded in

368

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Fig. 1. Transmission electron micrograph of the aedeagal bladder material surrounding a
lysed cell. Insert: Magnification of the fibrillar material.

Epon araldite resin, vacuum aspirated and allowed to polymerize for 24 hr at 40°C
and then 48 hr at 60°C. The blocks were sectioned at 500 A, stained with Toluidine
blue and viewed on a Ziess 10C/CR transmission electron microscope at 80 kV.

RESULTS AND DISCUSSION

Chemical analysis. Upon dissection in distilled water, the aedeagal bladder material
appeared as a white, granular substance and was insoluble in water, although a few

1984

CHEMICAL ANALYSIS IN SOLENOPSIS

369

suspended oil droplets were observed. Only slight dissolution occurred when the
extract was placed in acetone and ethanol. However, the material completely dis-
solved in 0.1 N NaOH but not in 0.1 N HCL.

TLC analysis for uric acid was negative for the gland extract. Additional TLC
analysis for the presence of free lipids showed that the material contained mono-
glycerides as shown by equal RF values (Table 1).

GC analysis of several samples showed gland contents to be variable perhaps due
to contamination during dissection but most samples (4 out of 6) showed the presence
of peaks with retention times equal to those of monodecanoyl and/or monododec-
anoyl glycerol. These samples were subjected to GC-MS analysis. Materials with
retention times equal to those of the monoglyceride standards had mass spectra
identical to those of the standards. Monododecanoyl glycerol was present in 4 of the
6 samples; monodecanoyl glycerol was in 1 sample in a 1:10 ratio. Other unidentified
highly variable materials made up from 10 to 80% of the samples containing mon-
oglycerides. Monoglyceride spectra were in excellent agreement with those given in
the EPA/NIH Mass Spectral Data Base (Heller and Milne, 1978).

Polyacrylamide gel electrophoresis showed the separation of one large band with
a molecular weight of ca. 40-90 K indicating the presence of lipoprotein(s).

The results using the gram electrobalance showed that nearly all of the aedeagal
bladder material was composed of polar lipids since the material was insoluble in
hexane but completely dissolved in chloroform : methanol.

Transmission electron microscopy showed the presence of several lysed cells sur-
rounded by slender fibrillar material in the bladder substance (Fig. 1). Closer ex-
amination revealed miniscule branching with dark, round globules adhering to the
surface of the fibrils (Fig. 1, insert).

This study has shown that the aedeagal bladder in the male fire ant, S. invicta,
consisted of two monoglycerides in a protein matrix. The source of this material is
unknown, and does not appear in the bladder until 5 to 7 days post emergence. No
change in the epithelial cells of the aedeagal bladder in preflight or postflight males
was observed nor did the cells appear to be secretory. The function of the gland also
remains unknown. Since the aedeagal bladder material has been found in approxi-
mately equal amounts in preflight and mated, postflight males, it seems unlikely that
it is used primarily during mating.

Several studies have shown that lipids play a major role in the biology of the fire
ant. Vinson et al. (1967) demonstrated that the triglyceride of linoleic acid was the
most active phagostimulant for workers. Vander Meer et al. (1982) analyzed several
glands in mated queens of various ages and found large amounts of triacylglycerols
in the crop and oesophagus which declined rapidly until 25 days following mating
coinciding with the first brood production. Nelson et al. (1980) reanalyzed the cu-
ticular methylalkanes in the cuticle of S. invicta and S. richteri Forel and determined
that the triglycerides originally found by Lok et al. (1975) had different structures
than were reported, although no function was demonstrated. However, the fatty acids
listed in these reports were not the same as we found in the aedeagal bladder material.

ACKNOWLEDGMENTS

We wish to thank Terry Coghlan for his assistance with the polyacrylamide disc gel electro-
phoresis procedure and Becky Matheson for the transmission electron microscopy.

370

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

LITERATURE CITED

Ball, D. E. and S. B. Vinson. 1984. Anatomy and histology of the male reproductive system
of the fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae). Int. J. Morph, and
Embryol. 13:283-294.

Clausen, R. 1938. Untersuchungen fiber den mannlichen Copulationsappart der Ameisen,
Speciell der Formicinae. Mitteil. Schweizer. Entomol. Gesells. 17:233-346.

Davis, B. J. 1964. Disc electrophoresis. II. Method and application to human serum proteins.
Ann. New York Acad. Sci. 121:404-427.

Forbes, J. 1954. The anatomy and histology of the male reproductive system of Camponotus
pennsylvanicus DeGeer (Formicidae, Hymenoptera). J. Morph. 95:523-556.

Forbes, J. 1958. The male reproductive system of the army ant, Eciton hamatum Fabricius.
Proc. Tenth Intemat. Congr. Entomol. 1:593-596.

Forbes, J. and D. Do-Van-Quy. 1965. The anatomy and histology of the male reproductive
system of the legionary ant, Neivamyrmex harrisi (Haldeman) (Hymenoptera, Formici-
dae). J. New York Entomol. Soc. 73:95-1 1 1.

Ford, F. C. and J. Forbes. 1980. Anatomy of the male reproductive systems of the adults and
pupae of two Doryline ants, Dorylus (Anomma) wilverthi Emery and D. (A.) nigricans
Illiger. J. New York Entomol. Soc. 88:133-142.

Hagopian, M. 1963. An anatomical and histological study of the male ponerine ant, Rhyti-
doponera metallica F. Smith (Formicidae, Hymenoptera). Ph.D. dissertation, Fordham
Univ., Univ. Microfilms, Ann Arbor, Michigan, 124 pp.

Heller, S. R. and G. W. A. Milne (eds.). 1978. “EPA/NIH Mass Spectral Data Base.” U.S.
Gov. Printing Office, Washington, D.C.

Hung, A. C. F., S. B. Vinson and J. W. Summerlin. 1974. Male sterility in the red imported
fire ant, Solenopsis invicta Buren. Ann. Entomol. Soc. Amer. 67:909-912.

Lok, J. B., E. W. Cupp and G. J. Blomquist. 1975. Cuticular lipids of the imported fire ants
Solenopsis invicta and richteri. Insect Biochem. 5:821-829.

Nelson, D. R., C. L. Fatland, R. W. Howard, C. A. McDaniel and G. J. Blomquist. 1980. Re-
analysis of the cuticular methylalkanes of Solenopsis invicta and S. richteri. Insect Bio-
chem. 10:409-418.

Tice, M. C. 1967. The anatomy and histology of some of the systems of the male of the
imported fire ant, Solenopsis saevissima richteri Forel (Hymenoptera: Formicidae). Ph.D.
dissertation, Fordham Univ., Univ. Microfilms, Ann Arbor, Michigan, 122 pp.

Trakimas, W. B. 1967. An anatomical and histological study of the male myrmicine ant,
Myrmica rubra L. (Hymenoptera: Formicidae). Ph.D. dissertation, Fordham Univ.,
Univ. Microfilms, Ann Arbor, Michigan, 125 pp.

Vander Meer, R. K., B. M. Glancey, and C. S. Lofgren. 1982. Biochemical changes in the
crop, esophagus and post-pharyngeal gland of colony-founding red imported fire ant
queens ( Solenopsis invicta). Insect Biochem. 12:123-127.

Vinson, S. B., J. L. Thompson and H. B. Green. 1967. Phagostimulants for the imported fire
ant, Solenopsis saevissima var. richteri. J. Insect Physiol. 13:1729-1736.

Received April 10, 1984; accepted October 1, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 371-402

REVISION OF THE HARVESTMAN GENUS LEPTOBUNUS
AND DISMANTLEMENT OF THE LEPTOBUNIDAE
(ARACHNIDA: OPILIONES: PALPATORES)

James C. Cokendolpher

Department of Entomology, Texas Tech University,

Lubbock, Texas 79409

Abstract.— The harvestman genus Leptobunus Banks is revised. Leptobunus calif ornicus Banks
and Leptobunus borealis Banks are redescribed, and Leuronychus parvulus Banks is transferred
to Leptobunus and redescribed. Lectotypes for L. borealis and L. parvulus are designated.
Leptobunus aureus n. sp. and Leptobunus pallidus n. sp. are described from California, U.S.A.
and Bering Island, U.S.S.R., respectively. Leptobunus atavus Cockerell, Leptobunus koreanus
Roewer, Leptobunus mexicanus Goodnight and Goodnight, and Leptobunus spinulatus Banks
are all excluded from the genus Leptobunus. The North American Tertiary fossil species is
maintained as Amauropilio atavus (Cockerell), whereas all Recent species excluded from Lep-
tobunus are reported in the following new combinations: Mitopus koreanus (Roewer), Para-
nelima mexicana (Goodnight and Goodnight), and Metopilio spinulatus (Banks). In conjunc-
tion, the Mexican species Diguetinus raptator Roewer is removed from synonymy with
“ Diguetinus” spinulatus (Banks). All the species are illustrated, including the male and female
genitalia, for which new anatomical terminology is presented. A dichotomous key to the species
and many new distributional records from western North America and northeast Asia are
provided.

The history of the family-group name Leptobunidae is discussed and the name is attributed
to Banks (1901). The family is abandoned and morphological characters traditionally used for
the classification of “Leptobunidae” are discussed. The type genus of the Leptobunidae, Lep-
tobunus Banks, is transferred to the Phalangiidae, subfamily Phalangiinae. The genera Cos-
mobunus Simon, Leuronychus Banks, and Homolophus Banks are transferred to the Gagrellidae,
subfamily Leiobuninae, and Trachyrhinus Weed is maintained in the Gagrellidae, subfamily
Gagrellinae. The North American genus Protolophus Banks is tentatively placed in the Gagrelli-
dae, subfamily Sclerosomatinae. The South American genus Thrasychirus Simon is retained in
the Neopilionidae, subfamily Enantiobuninae. The North American genus Globipes Banks is
maintained in an assemblage that is as yet unnamed. The New World members of the genera
Homolophus and Cosmobunus are not congeneric with Old World members referred to those
genera.

Attempts to establish phylogenetically meaningful classifications of the Phalan-
gioidea have been hindered by the confusion regarding the Leptobunidae. Until the
present, apparently no worker has studied examples of all genera assigned to this
family. Nathan Banks described several Nearctic species and genera but apparently
never studied any Old World members of the group. Carl-Friedrich Roewer examined
and described examples of all the genera assigned to the Leptobunidae except for
Leptobunus Banks, the type genus. Without knowing the true nature of Leptobunus,
Roewer could not have arrived at a reasonable classification for the Leptobunidae.
The lack of comparative material of Leptobunus spp. in European museums has also
hindered the study of the family. Most classifications by European authors involving

372

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

the Leptobunidae have been based on the North African, southern European genus
Cosmobunus Simon, which is not closely related to Leptobunus. To clear the con-
fusion regarding this family, I examined examples of all nominal species ever referred
to the Leptobunidae, and this is one of a series of papers dealing with those genera.
In the present contribution Leptobunus, the type genus, is revised. Furthermore, the
polyphyletic Leptobunidae is dismantled and placed in synonymy of the Phalangiidae.

MATERIALS AND METHODS

All anatomical measurements are in millimeters and were obtained using a bin-
ocular dissecting microscope equipped with an ocular micrometer (Cokendolpher,
1981a). Descriptions are based on all available specimens, but measurements were
taken only on a maximum of 10 specimens per sex. Measurements are reported as
ranges when more than one specimen was available. No measurements were taken
of the relatively unsclerotized genital operculum. The chelicera, palpus, and penis
were removed from the body and immersed in 100% glycerol for study, and then
placed in microvials within the specimen vials. A tuft of cotton placed in the glycerol
was used to support the structure when lateral views were desired. The female genitalia
were first dehydrated with absolute ethyl alcohol and then examined in 100% clove
oil. Ovipositors were soaked in absolute ethyl alcohol (to remove all traces of clove
oil) prior to rehydration and placement in microvials. All illustrations were prepared
with the aid of a camera lucida. The morphological terminology and description
format essentially follows Cokendolpher (1981b).

The following terminology relating to the anatomy of the genitalia is being adopted.

Penis. In situ, the penis lies in a dorsoventral plane, thus the orientation of the
truncus corresponds to that of the abdomen. As the penis truncus is attached to the
gonads and related structures on the end opposite the glans, this attachment region
becomes the basal or proximal portion of the truncus, and the basal portion of the
glans is attached to the truncus. As the glans is not rigidly fixed to the truncus, it can
be bent at any angle from 0° to about 160° to the plane of the truncus. This lack of
rigidity results in the need to define the glans orientation surfaces. As the glans
generally is at or near 135° to the truncus in preserved material, the glans surface
that is more anterior (distal) is hereby considered the dorsal surface. The dorsal
surface of the glans is generally straight, whereas the ventral surface is rounded. The
paired primary setae are located laterad or laterally on the glans. The stylus, like the
glans, is not rigidly attached (see Figs. 8, 9), but as no structures of taxonomic
importance have been discovered on the stylus, the use of basal, or proximal, and
distal (tip) will suffice.

Ovipositor. In situ, the ovipositor lies in a dorsoventral plane, and its orientation
follows that of the abdomen. The distal end of the ovipositor consists of a three
segmented furca. In the present descriptions, the furcal joints are numbered from
the basal attachment, the distal joint being the 3rd. The setae bearing lobes on the
3rd furcal joint are the apical sensilla. The slit sensilla are located distally on the
dorso- and ventromedial surfaces of the 2nd furcal joint. When specific sensilla
numbers (#) are reported, they are given in the following formula: dorsal left # —
dorsal right #/ventral left # — ventral right #. The 1st furcal joint lacks sensilla. The

1984

REVISION OF LEPTOBUNUS

373

ovipositor segments or rings are numbered from the distal end. The seminal recep-
tacles are located within the 2nd and 5th ovipositor rings, with the basal portions
generally located within the 2nd or 3rd ring, and the receptacles extend to about the
4th or 5th ring. Thus, the distal and proximal portions of the ovipositors and seminal
receptacles are opposite. Each seminal receptacle consists of a large distal loop (^pri-
mary loop) and one to many basal loops or coils on the basilateral margin; the basal
loops or coils are actually the extreme basilateral portion of the primary loop. The
basimesal margin often has a small tube in which the termination is indistinct. This
small tube is used as a marker for the region in which the basal loops should occur
when the loops are less convoluted (see Figs. 31-33).

AMNH

ARM

BMM

BMNH

CAS

CNC

EP

JCC

LACM

MCZ

NHR

NMB

osu

PDB

RGH

ROM

SMF

UCB

UCR

USNM

ZIZM

ACRONYMS FOR COLLECTIONS, LOCATIONS, AND CURATORS

American Museum of Natural History, New York, N. I. Platnick
A. R. Moldenke, Corvallis

Thomas Burke Memorial Washington State Museum, Seattle, R. L.
Crawford

British Museum (Natural History), London, F. R. Wanless and P. D.
Hillyard

California Academy of Sciences, San Francisco, D. H. Kavanaugh, W.
J. Pulawski, and V. F. Lee

Canadian National Collections of Insects, Arachnids and Nematodes,
Ottawa, C. D. Dondale and J. H. Redner
Exline-Peck Collection, Warrensburg, W. B. Peck
J. C. Cokendolpher, Lubbock

Los Angeles County Museum of Natural History, Los Angeles, C. L.
Hogue

Museum of Comparative Zoology, Cambridge, H. W. Levi and J. M.
Hunter

Naturhistoriska Riksmuseet, Stockholm, T. Kronestedt
Naturhistorisches Museum, Basel, E. Sutter

Oregon State University, Systematic Entomology Laboratory, B. B. Frost

and G. M. Stonedahl

P. D. Bragg, Vancouver

R. G. Holmberg, Edmonton

Royal Ontario Museum, Toronto, D. Barr

Senckenberg Natur-Museum und Forschungsinstitut, Frankfurt, M.
Grasshoff

University of California Berkeley, Essig Museum, Berkeley, C. E. Gris-
wold

University of California Riverside Entomological Teaching and Re-
search Collection, Riverside, S. E. Frommer

National Museum of Natural History (U.S. National Museum), Wash-
ington, R. E. Crabill, Jr.

Zoologisches Institut und Zoologisches Museum, Hamburg, G. Rack

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SYSTEMATICS

PHALANGIIDAE SIMON

PHALANGIINAE SIMON

Leptobunus Banks

Liobunum: Banks, 1894a: 160 (in part).

Leptobunus Banks, 1893:209 (in part), 1894a: 163, 1894b:40 (misidentification), 1894c:
146 (misidentification), 1898: 182 (misidentification), 1899:350, 1900a: 199, 1900b:
484, 1901:674, 1904:362, 1911:420, 1923:238; Simon, 1902:45; Cockerell, 1907a:
605 (in part), 1907b:620 (misidentification); Roewer, 1910:270, 1923:877, 1952:
268, 1957:355 (misidentification); Kishida, 1930:55; Redikorzev, 1936:33; Com-
stock, 1940:71; Goodnight and Goodnight, 1942b: 13 (in part); Star^ga, 1965:5
(misidentification), 1978:209; Silhavy, 1976:296 (misidentification); Cokendol-
pher, 1980:134, 1981b:309; Cokendolpher and Cokendolpher, 1982:1216.
Leptobrunus: Myers, 1921:19 ( lapsus calami).

Leuronychus : Banks, 1901:675 (in part), 1911:420 (in part); Roewer, 1910:271 (in
part), 1923:878 (in part), 1957:356 (in part); Comstock, 1940:73 (in part); Holm-
berg et al., 1981:19 (in part).

Liomitopus Schenkel, 1951:49; Forcart, 1961:53.

Type species. Leptobunus californicus Banks, by original designation.

Diagnosis and comparisons. Leptobunus is similar to Mitopus Thorell, Tchapinius
Roewer, and Liopilio Schenkel. These four genera differ from other phalangiid genera
by lacking large spines or tubercles on the preocular area and the ventral surfaces of
the palpal femora, by having the supracheliceral lamellae short and smooth, palpi
without apophyses (juveniles included), and generally by having a tooth on the basal
segment of the chelicerae. Leptobunus and Liopilio differ from Mitopus and Tcha-
pinius by having the palpal claws denticulate, penis glans convex beneath, glans
primary setae very large and often bifurcate, and by lacking strong spines or tubercles
anywhere on the body (except occasionally on appendages). Leptobunus differs from
Liopilio by having the palpal tarsi of males globular proximally and devoid of den-
ticles ventrally. The palpal femora and patellae of Liopilio are expanded distally and
densely covered with setae medially. The paired primary setae of the penis glans are
located near the stylus junction in Liopilio, whereas in Leptobunus they are centrally
located. The stylus-glans junction membrane is dorsally expanded in Liopilio, ven-
trally expanded in Leptobunus. The 2nd furcal joint of the ovipositor of Liopilio has
one or two slit sensilla per side, whereas Leptobunus has one to six (rarely one)
sensilla per side.

Description. Medium to small sized phalangiids with soft smooth bodies (Figs. 1-
5); dorsum and venter with only scattered setae; last abdominal stemite indented.
Ocular tubercle low, canaliculate, round to slightly wider than long, with scattered
setae. Chelicerae not enlarged, with tooth on ventral surface of basal joint; moveable
finger with or without apophysis (Figs. 40-50). Supracheliceral lamellae short and
smooth. Scent gland pores visible from above, elongate. Palpi (Figs. 36-39) covered
only with setae, except for one or two tubercles on distodorsal surface of female
femora; distomesal margins of femora and patellae not expanded (not even in ju-
veniles); distal ends of femora with lyriform organs on dorsolateral margins, dor-
somedial campaniform sensilla lacking; male tibiae and tarsi modified, proximal

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REVISION OF LEPTOBUNUS

375

portion of tibiae extended on mesal margin, proximal end of tarsi globular, expanded
slightly on mesal margin, ventral rows of denticles lacking; claws with small pointed
teeth at base. Legs relatively short; all articles round in cross-section. Penis truncus
not grooved on distodorsal margin, glans round with large primary setae medially,
stylus-glans junction membrane ventrally expanded (Figs. 6-21). Ovipositor often
darkly pigmented with 19-35 rings plus 3 segmented furca; 1-6 slit sensilla per side
on 2nd furca joint, seminal receptacles consisting of paired loops (Figs. 22-35).

INCLUDED SPECIES

Leptobunus consists of five species in two species groups. The californicus group
contains Leptobunus californicus Banks, Leptobunus borealis Banks, and Leptobunus
pallidus, new species. The parvulus group contains: Leptobunus parvulus (Banks),
new combination, and Leptobunus aureus, new species.

Members of the californicus group can be easily recognized by the details of the
genitalia and lack of a distinct apophysis on the moveable finger of the male chelicerae.
The primary setae of the penis glans in members of the californicus group are often
pointed or round and always much shorter in length than the stylus. Californicus
group members have the primary loops of the seminal receptacles more than four
times as long as wide. Members of the parvulus group often have tibiae IV with one
or two pseudosegments. The male chelicerae in parvulus group members have a
distinct apophysis on the moveable finger. The primary setae of the penis glans in
members of the parvulus group are bilobed and equal to or slightly longer than the
stylus. The primary loops of the seminal receptacles in members of the parvulus
group are less than three times as long as wide.

EXCLUDED SPECIES

Amauropilio atavus (Cockerell)

Leptobunus atavus Cockerell, 1907a:605, fig. 2; 1907b:620.

Amauropilio atavus: Cokendolpher and Cokendolpher, 1982:1216-1217, fig. 1.

This species is known from one Tertiary fossil. The holotype female from the
Florissant Formation, Teller Co., Colorado, U.S.A. (AMNH cat. no. FI-49 18834
Paleontology coll.) was not reexamined for the present study.

Mitopus koreanus (Roewer), New Combination

Leptobunus koreanus Roewer, 1957:355, pi. 26, fig. 23; Starfga, 1965:5; Silhavy,

1976:296.

Female holotype from Korea (date, collector, and specific locality unknown), SMF
cat. no. RII/3694/8, examined. This species conforms to Mitopus Thorell and is
hereby transferred to that genus. A redescription is in preparation.

Paranelima mexicana (Goodnight and Goodnight),

New Combination

Leptobunus mexicanus Goodnight and Goodnight, 1942b: 13, fig. 31.

Female holotype from mountain meadow at Cerro Tancitaro, Michoacan, Mexico,

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Figs. 1-5. Dorsal views of Leptobunus spp. 1. L. californicus male. 2. L. borealis male. 3.
L. pallidus female. 4. L. aureus male. 5. L. parvulus male. Scale line = 2.0 mm.

16 July 1941 (H. Hoogstraal), AMNH, examined. This species is related to other
North American species of Paranelima di Caporiacco and Mexican species referred
to Nelima Roewer. Until the mesoamerican species of these genera are revised, the
affinities of P. mexicana will remain unresolved. Further material of both sexes of
several species needs to be collected before such a revision can be realized.

Metopilio spinulatus (Banks), New Combination

Leptobunus spinulatus Banks, 1898:182; Pickard-Cambridge, 1905:585; Roewer, 1910:
257.

Hadrobunus spinulatus: Roewer, 1923:920.

Diguetinus spinulatus: Goodnight and Goodnight, 1942b: 11 (in part), 1945:13 (in
part), 1947:45 (in part); Roewer, 1956:252 (misidentification).

Female holotype from Tepic, Nayarit, Mexico (date and collector unknown), MCZ,
examined. Comparisons of the holotype of L. spinulatus and paratypes of Diguetinus

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377

raptator Roewer (2 males, 3 females, Guadalajara, Jalisco, Mexico, SMF cat. no. RI/
623) reveal they are not conspecific with one another as Goodnight and Goodnight
(1942b) had indicated. Leptobunus spinulatus is properly a member of Metopilio
Roewer. Metopilio and Diguetinus Roewer are quite similar, but are currently con-
sidered separate genera (Cokendolpher, 1984a).

KEY TO THE SPECIES OF THE GENUS Leptobunus

la. Male chelicera with distinct apophysis on moveable finger (Figs. 46-50); stylus of
penis equal to or shorter than primary setae of glans (Figs. 16, 18); primary loop of
seminal receptacle short, less than three times as long as wide (Figs. 31-34) (parvulus

group) 2

lb. Chelicera without apophysis on moveable finger (Figs. 40-45); stylus of penis much
longer than primary setae of glans (Figs. 8, 12); primary loop of seminal receptacle

long, more than four times as long as wide (Figs. 23, 29) ( californicus group) 3

2a. Body color golden, legs speckled; primary setae of glans separated more than the length

of the stylus (Fig. 16) L. aureus, new species

2b. Body color brown, gray or bronze, legs banded; primary setae of glans closely spaced

(Fig. 18) L. parvulus (Banks)

3a. Tibiae II with 1-4 pseudosegments; male palpal tibiae expanded on ventral and mesal

surfaces (Figs. 57-60); dorsum of abdomen with a well defined pattern (Figs. 1,2)... 4
3b. Tibiae II without pseudosegments; male palpal tibiae not expanded or enlarged (Figs.

51, 52); dorsum with faint pattern at most (Fig. 3) L. pallidus, new species

4a. Male palpal tibiae and tarsi noticeably expanded on mesal margins (Figs. 59, 60);
dorsum of penis glans slightly depressed (Figs. 8, 9); basal loops of seminal receptacles

consisting of three coils or loops (Figs. 23, 24) L. californicus Banks

4b. Male palpal tibiae and tarsi only slightly expanded (Figs. 57, 58); dorsum of penis
glans straight (Fig. 1 2); basal loops of seminal receptacles consisting of two coils or
loops (Figs. 26, 27) L. borealis Banks

Distribution. Aleutian and Bering Sea Islands, southern Alaska, and south along
coastal states to central California and western Nevada in western North America.

Leptobunus californicus Banks
Figs. 1, 6-10, 22-24, 36-41, 59, 60, 63

Leptobunus californicus Banks, 1893:210, 1901:674, 1904:362, 1911:420; Roewer,
1910:270, 1923:877; Comstock, 1940:72; Goodnight and Goodnight, 1942b:14;
Cokendolpher, 1980:134.

Leptobrunus californicus: Myers, 1921:19 ( lapsus calami).

Liomitopus laevis Schenkel, 1951:49, fig. 47; Forcart, 1961:53.

Types. Female holotype of Leptobunus californicus in Nathan Banks Coll., MCZ,
examined. Banks (1893) stated that the specimen was collected by Davidson in
southern California and later (1904) stated that Davidson collected the specimen but
restricted the locality to Los Angeles. As no further material of this species has been
collected in southern California, and because the type vial is simply labeled “Cal.,”
I believe the type locality was erroneously designated and restricted. As Liomitopus
laevis is the only synonym, I choose its type locality as that of L. californicus. Male

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Figs. 6-14. Male genitalia of Leptobunus spp. 6-1 1. L. californicus. 6. Lateral view penis.
7-9. Distal ends of penes, lateral views. 10, 1 1. Gians setae. 12-14. L. borealis. 12. Distal end
of penis, lateral view. 13. Gians seta. 14. Lateral view penis. Scale line = 0.50 mm for Figs. 6,
7, 14; 0.12 mm for Figs. 8, 9, 12; 0.03 mm for Figs. 10, 1 1, 13.

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REVISION OF LEPTOBUNUS

379

lectotype and female paralectotype of Liomitopus laevis from Yosemite, Mariposa
Co., California, 29 Dec. 1938 (H. Schenkel-Rudin), NMB no. 81 -a, examined.

Diagnosis and comparisons. Leptobunus californicus can be separated from L.
borealis and L. pallidus, the other members of its group, on the basis of differences
in genitalia and male palpi and number of tibiae II pseudosegments.

Distribution. Eastern portion of central California and adjoining Nevada (Fig. 63).

Description. Males: Total length 3.34-5.41, greatest width 2.75-3.82, maximum
height 1.80-2.50; body creamy white to light yellowish-brown (rarely silvery white)
with brown central pattern; central pattern with considerable spotting and splotches
(Fig. 1). Ocular tubercle length 0.43-0. 51, width 0.46-0. 55, height 0.20-0. 24, distance
from anterior edge of cephalothorax 0.32-0.40; concolorous with dorsum of ceph-
alothorax, except anterior border shaded dark brown; black ring encircling eyes, ring
extended slightly anteriorly. Venter creamy white, always lighter than dorsum; with
some brown mottlings on lateral margins posteriorly; anal operculum brown with
white splotches. Chelicerae (Figs. 40, 41) yellow brown with no marking, teeth and
tips of fingers dark brown to black; moveable fingers lacking apophyses. Palpi light
yellow brown with light brown shading on distal ends of femora; faint longitudinal
brown line on dorsum and brown splotches (often lacking on lateral surfaces of
patellae); tibiae with (at most) a dark spot on dorsoproximal tip; tarsi darkened on
distal tip; tibiae and tarsi noticeably enlarged ventrally and ventromesally (Figs. 36,
37, 59, 60). Palpal segment lengths: femora 0.96-1.22, patellae 0.48-0.65, tibiae
0.70-0.81, tarsi 1.24-1.35. Legs yellowish-brown to creamy light brown. Coxae un-
marked. Femora with extreme basal end, dorsal and lateral distal tips, and subdistal
region white; area between subdistal and distal white regions dark brown; basal 3A
shaded brown distally. Patellae subdistally shaded dark brown, extreme distal end
white, basal end yellowish-brown. Tibiae I, III, IV with subbasal and subdistal dark
brown bands; II with subdistal dark brown band and brown shading on proximal
sides of pseudosegment junctions. Tarsi shaded brown at pseudosegment junctions.
Metatarsi I with single pseudosegment; tibiae II with 2-3 (commonly 3) pseudo-
segments; tibiae IV lacking pseudosegments. Femora I-IV lengths (respectively):
3.77-4.06, 7.90-8.20, 4.30-4.46, 5.64-6.20; tibiae I-IV lengths (respectively): 3.27-
3.52, 6.03-6.41, 3.40-3.50, 4.09-4.70. Penis (Figs. 6-1 1) with glans slightly concave
dorsally; glans primary setae simple or bifurcate, spaced approximately one setal
length apart or slightly more, setae much shorter than stylus length.

Females: Total length 5.3 1-7.3 1, greatest width 3.50-4.53, maximum height 2.00-
4.18; generally larger and more robust than males, with more distinct and darker
color patterns, except basal bands on leg femora and basal bands on tibiae II absent
or greatly reduced. Leg coxae often with mottlings, lacking distal band; palpal tibiae
rarely with brown line (faint when present). Ocular tubercle length 0.38-0.50, width
0.45-0.60, height 0. 18-0.26, distance from anterior edge of cephalothorax 0.30-0.42.
Palpal segment lengths: femora 0.90-1.02, patellae 0.51-0.65, tibiae 0.69-0.84, tarsi
1.21-1.50. Metatarsi I with 0-2 (rarely 0) and tibiae II with 2-4 (averaging 3) pseu-
dosegments, tibiae IV lacking pseudosegments. Femora I-IV lengths (respectively):
2.72-4.19, 6.60-8.50, 3.00-4.53, 4.53-6.67; tibiae I-IV lengths (respectively): 2.45-
3.48, 4.65-6.80, 2.49-4.51, 3.19-4.42. Ovipositor (Fig. 22) with considerable vari-
ation in ring segment counts: the five examined with 19, 23, 30, 32, and 35 rings

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Figs. 15-21. Male genitalia of Leptobunus spp. 15-17. L. aureus. 15. Lateral view of penis.
16. Distal end of penis, lateral view. 17. Gians seta. 18-21. L. parvulus. 18. Distal end of penis,
lateral view. 19, 20. Gians setae. 21. Lateral view penis. Scale line = 0.50 mm for Figs. 15, 21;
0.12 mm for Figs. 16, 18; 0.03 mm for Figs. 17, 19, 20.

each; furca 3 segmented (one female from Norden, California, with 4 segmented
furca; distal two joints normal, first joint split in two on both sides). Slit sensilla
generally 2-2 (range 1-2 to 4-4) per side on furca segment II. Seminal receptacles
with long, often convoluted, primary loops and three basal loops (Figs. 23, 24).

Immatures: As adult females, except leg femora II and IV lacking proximal brown
bands; palpi often white to light yellowish-brown, generally without markings.

Natural history. Adults are known from late June to late September, and a single
collection on 29 December. The majority of collections are from August. Juveniles
(later instars) are known from late June to late September. Several of the females
collected during August are filled with eggs. A dozen eggs from two different indi-
viduals average 0.54 mm in size (range 0.40-0.68 mm). Specimens have been col-
lected at altitudes from 1,850 to 3,350 m, almost all from 2,350-2,450 and 2,900-
3,100 m. In general, the more northern localities are at lower elevations. The few
collections that were labeled as to habitat indicated that adults and immatures were
beneath rocks, from crevices in large granite rocks, and were on snow at night (during

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REVISION OF LEPTOBUNUS

381

June). This species appears to be restricted to the Sierra District (Schick, 1965) of
the Sierra Nevadan Biotic Province.

Specimens examined. U.S.A. CALIFORNIA: Placer Co., Norden, 5 Aug. 1951 (J.
F. Gustafson), 3 females (AMNH); Tuolumne Co., Sonora Pass, 26 Aug. 1965 (V.
Lee), 2 immatures (CAS), Sonora Pass Summit, 26 Aug. 1965 (V. Lee), 1 female
(CAS), 4.8 km W Sonora Pass, 14-16 Aug. 1966 (R. R. Snelling), 1 immature (LACM),
Tioga Pass, Yosemite Park, 8-10-1931 (W. Ivie), 3 females (AMNFI); Madera Co.,
Lake Ediza, 2 Aug. 1951 (P. Raven), 1 female (CAS); Inyo Co., South Lake, Bishop
Creek, 17 Aug. 1941 (W. M. Pearce), 3 females (AMNH), Mirror Lake, 14 Aug. 1953
(W. McDonald), 1 female (AMNH); Mono Co., E slope White Mountain, 27 June
1979 (D. H. Kavanaugh), 1 female, 3 immatures (CAS), E shore Silver Lake, 8 Aug.
1957 (R. Casebeer), 1 female (LACM), Sawmill Campground, 6.5 km N of Tioga
Pass, 30 July 1980 (C. Griswold), 1 immature (UCB), Big Bend Campground, 8 km
W Lee Vining, 21 Sept. 196 1 (W. Ivie and W. J. Gertsch), 1 female (AMNH); Wildyrie,
Mammoth Lakes, 18 Aug. 1941 (W. M. Pearce), 2 males, 2 females (UCR); Mariposa
Co., Yosemite, 29 Dec. 1938 (H. Schenkel-Rudin), lectotype male, paralectotype
female (NMB), Yosemite Creek Camp, Yosemite Park, 18 Sept. 1941 (W. Ivie), 1
immature (AMNH); Nevada Co., Sagehen Creek, 12.9 km N Truckee, 21 July 1966
(G. B. Wiggins), 1 female (ROM); Alpine Co., Silver Creek, 14.5 km SW Markleeville,
26 July 1980 (T. C. Meikle and C. Griswold), 1 male, 2 females (UCB); Fresno Co.,
Inyo National Forest, John Muir Wilderness, Cascade Valley, 7 Aug. 1972 (V. F.
Lee), 1 male, 1 female (CAS); Fresno Co., Graveyard Meadows, 12 Aug. 1959 (B.
Firstman), 13 males, 3 females, 4 immatures (AMNH), Graveyard Lakes, 13 Aug.
1959 (B. Firstman), 1 male, 1 female, 2 immatures (AMNH), Anne Lake, 15 Aug.
1959 (B. Firstman), 1 female (AMNH), Olive Lake, 16 Aug. 1959 (B. Firstman), 3
females, 2 immatures (AMNH), locality uncertain— “Southern California,” pre-1893
collection (Davidson), holotype female (MCZ). NEVADA: Washoe Co., Mt. Rose,
2.4 km W Pass Summit, 27 Aug. 1969 (H. B. Leech), 3 males, 2 females (CAS).

Leptobunus borealis Banks
Figs. 2, 12-14, 25-27, 42, 43, 57, 58, 61

Leptobunus borealis Banks, 1899:350, pi. a, hg. 7, 1900b:484, 1901:674, 1911:420,
1923:238; Roewer, 1910:271, 1923:877, hg. 1,039; Redikorzev, 1936:33; Com-
stock, 1940:72; Star^ga, 1978:209; Strand, 1906:473.

Types. Banks (1899) based his description on “several specimens” collected on
Bering Island and Copper Island. In the description he attributes measurements to
a male and female, but from his description of the palpi and his illustration there is
no doubt he examined only females and immatures. Only a female and three im-
matures of the original type series are still available. This series collected on Bering
Island, July-Aug. 1897, by Barrett-Hamilton is clearly labeled in Banks’ handwriting
“ Leptobunus borealis Bks Type.” The vial also contains a label “Type No. 4060
U.S.N.M.” which corresponds to the data given by Banks. As no single specimen
was designated as the holotype, I hereby designate the adult female as the lectotype
and three immautres as paralectotypes.

Diagnosis and comparisons. In addition to differences in the genitalia and male

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palpi noted in the key, L. borealis can be distinguished from L. californicus by its
shorter legs.

Distribution. Komandorskiye Ostrova of U.S.S.R., Aleutian Islands, Pribilof Is-
lands and southern Alaska of U.S.A. (Fig. 61).

Description. Males: Total length 3.56-4.12, greatest width 2.89-3.50, maximum
height 2.07 -2.27 ; dorsum white to creamy white with extensive dark brown markings,
those on cephalothorax appearing velvety; abdomen with pattern of cross with many
white spots and splotches, area surrounding cross mottled with brown splotches and
rows of brown dots (Fig. 2). Ocular tubercle length 0.35-0.40, width 0.37-0.42, height
0. 1 8-0. 1 9, distance from anterior edge of cephalothorax 0.30-0.3 1 ; brown with dark
ring encircling eyes, paler half ring on areas dorsal to dark eye ring. Venter white to
light tan, with two dark brown spots at base and posterior to base of genital operculum;
row of brown spots on anterior areas of stemite junctions (less distinct on the most
posterior stemites) and lateral margins of all stemites shaded brown (lighter in color
than spots). Leg coxae yellowish-brown, mottled with brown, distally shaded to
indistinct brown band. Chelicerae (Figs. 42, 43) with basal segment creamy yellowish-
white; distal segment light yellow brown; both segments with light brown mottlings
on lateral surfaces; teeth and tips of fingers black; moveable fingers lacking apophyses,
rarely with slight “bump” (Fig. 43). Palpi yellow brown with darker brown markings
on distodorsal surfaces of femora, lateral and mesal surfaces of patellae and tibiae,
and semidistinct line down patellae and proximal % to 3A of tibiae dorsally; basal
portion of tarsi enlarged (Figs. 57, 58). Palpal segment lengths: femora 0.90-0.94,
patellae 0.56, tibiae 0.68-0.73, tarsi 1.18-1.23. Legs yellowish-brown to tan; femora
with distal dark brown leg bands, extreme distal ends white dorsally on lateral regions,
often with indistinct subdistal white band; patellae mottled with brown except for
thin light line dorsally; all tibiae with brown bands (often indistinct) distally, I, III,
IV with a second brown band on proximal lh, extreme proximal end tan to yellowish-
white; dorsa of tibiae I, III, IV (rarely II) with faint light brown lines; tarsi and
metatarsi pseudosegment junctions shaded brown; metatarsi I and tibiae II both with
single pseudosegments, tibiae IV lacking pseudosegments. Femora I-IV lengths (re-
spectively): 2.63-2.67, 4.62-5.18, 2.94-3.10, 4.25-4.57; tibiae I-IV lengths (respec-
tively): 2.30-2.42, 3.92-4.16, 2.42-2.48, 3.15-3.25. Penis with glans dorsum straight
to slightly convex (Figs. 12-14); primary setae of glans simple, approximately one
setal length apart; setae much shorter than stylus.

Females: Total length 5.27-6.02, greatest width 3.20-4.28, maximum height 2.60-
3.27; similar to males except ocular tubercle lighter in color, creamy white; leg femora
and tibiae have more brown splotches; leg coxae and trochanters of some females
with considerable amount of brown splotches and mottlings. Ocular tubercle length
0.40-0.46, width 0.40-0.41, height 0.19-0.23, distance from anterior edge of ceph-
alothorax 0.28-0.41. Palpal segment lengths: femora 0.82-1.00, patellae 0.51-0.60,
tibiae 0.59-0.75, tarsi 1.1 1-1.28. Metatarsi I with single pseudosegment (one spec-
imen had 2), tibiae II with one pseudosegment (rarely 2 or 3), tibiae IV lacking
pseudosegments. Femora I-IV lengths (respectively): 1.92-2.48, 3.80-4.46, 2.30-
2.80, 3.21-4.1 1; tibiae I-IV lengths (respectively): 1.90-2.33, 3.19-4.00, 1.98-2.54,
2.58-3.08. Ovipositor (Fig. 25) with considerable variation in furca II slit sensilla
counts; range 2-6 per side, counts of 3-3/6-4 and 5-4/5-4 not uncommon; the single

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REVISION OF LEPTOBUNUS

383

Figs. 22-27 . Female genitalia of Leptobunus spp. 22-24. L. californicus. 22. Distal end
ovipositor. 23, 24. Seminal receptacles. 25-27. L. borealis. 25. Distal end of ovipositor. 26,
27. Seminal receptacles. Scale line = 0.22 mm for Figs. 22, 25; 0.05 mm for Figs. 23, 24, 26,
27.

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ovipositor examined with 29 rings, 3 segmented furca. Seminal receptacles (Figs. 26,
27) with long primary loop and two basal loops.

Immatures: Essentially as in adult females, except body more gray than brown;
leg femora with distinct dark band on distal end and single dark band at midpoint;
chelicerae uniformly grayish-yellow brown; leg coxae and chelicerae lacking splotches
and mottlings.

Natural history. Specimens from the present study were all collected during July
and August. Banks (1923) reported that this species was common on the Pribilof
Islands from the first of June until “fall.” Five females collected during the later part
of July were filled with large eggs. Collections from Homer, Alaska, were from “drift”
on beach at high tide along with juveniles of the harvestman Sabacon sp. Leptobunus
borealis is apparently restricted to the maritime subarctic, occurring from sea level
to approximately 300 m in elevation.

Specimens examined. U.S.S.R. KOMANDORSKI YE OSTROVA: Bering Island
(=Behrings om), 14-19 August 1879 (Vega Expedition no. 1079), 1 female (NHR),
July-August 1897 (Barrett-Hamilton), lectotype female, 3 immature paralectotypes
(USNM type no. 4060). U.S.A. ALASKA: Andreanof Islands, Atka Island, 28 July
1 907 (E. C. VanDyke), 3 males, 5 females (CAS); Adak Island, 4 July 1 948 (I. Newell),
1 penultimate male (AMNH); Pribilof Islands, St. Paul Island, 1899 (T. Kincaid), 2
subadults (BMM), 10 August 1919 (G. D. Hanna), 3 females (CAS); Homer, 20-25
July 1945 (J. C. Chamberlin), 3 females, 1 immature (AMNH).

Leptobunus pallidus, new species
Figs. 3, 28, 29, 44, 45, 55, 56, 61

Types. Female holotype and two immature (female? and male) paratypes from
Bering Island, Komandorskiye Ostrova, U.S.S.R., 14-19 August 1879 (Vega Expe-
dition), NHR.

Etymology. The specific epithet from Latin, referring to the pale color of the body.

Diagnosis and comparisons. Leptobunus pallidus is a member of the californicus
group. In addition to the differences in the genitalia noted in the key, L. pallidus
differs from other Leptobunus spp. by the increased number of setae on the 1st furcal
joint of the ovipositor. The lack of pseudosegments on metatarsi I and tibia II will
also serve to separate L. pallidus from other members of the genus, but the lack of
pseudosegments on metatarsi I is also characteristic of many immatures of other
species in the genus.

Distribution. Known only from Komandorskiye Ostrova of U.S.S.R. (Fig. 61),
where L. borealis also occurs.

Description. Female: Total length 5.35, greatest width 2.86, maximum height 2.90;
body creamy white to yellow white; dorsum with light brown pattern, interrupted
with scattered white spots (Fig. 3). Ocular tubercle length 0.40, width 0.45, height
0.15, distance from anterior edge of cephalothorax 1 .27; white with light brown rings
around eyes. Coxae, genital operculum, and abdominal sternites creamy white. Che-
licerae light yellow brown with dark brown teeth. Palpi whitish-yellow to light yellow
brown, without markings. Palpal segment lengths: femur 0.80, patella 0.51, tibia
0.69, tarsus 1 .26. Legs whitish-yellow to light yellow brown with faint bands of brown
on distal ends of femora, dorsum of patellae, and subdistal portions of tibiae; metatar-

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385

Figs. 28-35. Female genitalia of Leptobunus spp. 28, 29. L. pallidus. 28. Distal end of
ovipositor. 29. Seminal receptacle. 30-33. L. parvulus. 30. Distal end of ovipositor. 31-33.
Seminal receptacles. 34, 35. L. aureus. 34. Distal end of ovipositor. 35. Seminal receptacle.
Scale line = 0.22 mm for Figs. 28, 30, 35; 0.05 mm for Figs. 29, 31-34.

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si I and tibiae II lacking pseudosegments. Femora I-IV lengths (respectively): 2.31,
3.90, 2.51, 3.69; tibiae I-IV lengths (respectively): 2.00, 3.85, 2.18, 2.69. Ovipositor
consisting of 25 rings; furca 3 segmented, first furcal joint with greatly increased
number of setae (Fig. 28), second joint with 3-5 slit sensilla per side. Seminal re-
ceptacles with three basal loops (Fig. 29).

Immatures: Similar to adult female, except dorsal pattern faint or lacking. Leg
bands more distinct than in adult female. Subadult male (total length 3.60) with palpi
only slightly modified (Figs. 55, 56). Moveable finger of chelicerae without apophysis
(Figs. 44, 45).

Natural history. The only known specimens were collected during the middle of
August on Bering Island. According to Danks (1981) the Commander Islands are
steep mountainous islands of volcanic origin with peaks reaching nearly 1,000 m.
The islands are very wet, and due to marine influences the climate is relatively mild
for the latitude (55°N). The islands are primarily covered by forest-tundra vegetation.
Specimens examined. Only the type series.

Leptobunus parvulus (Banks), New Combination
Figs. 5, 18-21, 30-33, 46-48, 53, 54, 62

Liobunum parvulum Banks, 1894a: 163.

Leuronychus parvulus: Banks, 1901:675, 1911:420; Roewer, 1910:273, 1923:879,
1957:356; Comstock, 1940:73; Holmberg et al., 1981:19.

Leuronychus pacificus: Roewer, 1910:273 (misidentification in part), 1923:879 (mis-
identification in part). Of the three male and two female L. pacificus (Banks)
reported from Puget Sound (Tacoma, Washington) by Roewer, one is a male L.
parvulus (Banks).

Types. The two male and one female syntypes from Olympia, Washington, de-
scribed by Banks (1894a) in the Nathan Banks Coll., MCZ, examined. I designate
the larger male (total length 3.72) as the lectotype and the smaller male (total length
3.45) and female as paralectotypes. The lectotype has been placed in a separate vial
and labeled accordingly. The lectotype is missing both first legs whereas the male
paralectotype is missing legs II and IV. I do not regard the female from Puget Sound
(SMF no. 174), reported by Roewer (1923, 1957) as a cotype, as part of the original
type series. When Banks described this species he mentioned only two males and
one female from Olympia, none from Puget Sound. The vial with the female from
the SMF bears the label: “ Leuronychus parvulus 1 Expl. Banks Puget Sound Roewer
det. 1910 No. 174.”

Diagnosis and comparisons. Leptobunus parvulus is similar to L. aureus\ these two
form the parvulus group. In addition to the differences in the genitalia, L. parvulus
can be distinguished from L. aureus by its larger size.

Distribution. Southeastern Alaska, western British Columbia, Washington, Oregon,
and northern California in western North America (Fig. 62).

Description. Males: Total length 3.35-4.78, greatest width 2.43-3.18, maximum
height 1 .85-2.60; body generally whitish-yellow, but ranges from golden tan to opales-
cent white; often with many tiny white specks and larger white splotches. Abdomen
with faint cross pattern of brown spots and splotches; pattern most distinct posteriorly

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387

with some opalescent spotting (Fig. 5). Some individuals with faint line indicating
posterior edge of cross. Areas on each side of line are essentially the same, small
brown mottlings with some opalescent spots. Posterior and lateral areas of abdomen
often shaded dark brown with many white opalescent spots. Cephalothorax with
many brown maculations. Scent gland pore often dark blue, encircled by black. Ocular
tubercle length 0.37-0.41, width 0.40-0.43, height 0.13-0.22, distance from anterior
edge of cephalothorax 0.22-0.32; whitish-yellow to opalescent white with black rings
around eyes. Venter of abdomen yellowish- white to opalescent white with some light
brown spots; few specimens with spots fused, forming line across stemite. Anal
operculum generally brown without white spot. Coxae ranging from yellowish-white
with no marks to yellowish-brown with many white reticulations, and broad subdistal
brown bands, covering approximately Vs of coxae length; bands sometimes replaced
by faint brown splotches on anterior and posterior margins distally. Chelicerae (Figs.
46-48) yellowish-brown with black teeth, noticeable spur on moveable finger. Palpi
yellowish-brown; femora with either two subdistal brown splotches on sides or with
subdistal dark brown band with distal white patches; patellae often with distodorsal
brown splotches and two small brown specks on sides, occasionally with faint brown
median line dorsally and less distinct brown lines on dorsolateral and dorsomesal
margins; tibiae dark brown at base, shaded to yellow brown distally; tarsi unmarked,
only slightly swollen basally (Figs. 53, 54). Palpal segment lengths: femora 0.88-1.16,
patellae 0.47-0.62, tibiae 0.68-0.80, tarsi 1.13-1.39. Legs yellowish-white to light
yellowish-brown with white rings on distal ends of all femora and tibiae, subdistally
on patellae and a light stripe on dorsal surfaces of patellae; femora with broad subdistal
rings of dark brown (varies from Vs to almost % length), when brown rings are short
followed by second white ring and lighter brown band (approximately Vs to Vi length
from basal end); tibiae (except for II) banded as on femora, II with subdistal brown
band V* to its length and dark rings on basal half of each pseudosegment; patellae
dark brown on median portion, yellowish-brown basally. Tarsi with brown rings on
basal half of each pseudosegment; metatarsi I with 1 or 2 (rarely 2) pseudosegments,
tibiae II with 2-6 (see comments under variation) pseudosegments, tibiae IV with
0-2 (generally lacking) pseudosegments. Femora I-IV lengths (respectively): 3.02-
4.93, 5.78-9.88, 3.00-5.90, 4.36-8.30; tibiae I-IV lengths (respectively): 2.33-4.62,
4.63-6.80, 2.63-5.00, 3.18-6.62. Truncus of penis not noticeably enlarged basally
(Figs. 18-21); primary setae of glans bilobed, approximately one setal length apart,
setae subequal in length to stylus; distal end of glans constricted.

Females: Total length 4.59-6.82, greatest width 2.79-4.62, maximum height 2.50-
3.84; body similar to male, more robust with darker colors and more distinct dorsal
pattern. Some specimens bronze to silvery gold with dark brown patterns, and specks
of gold on dorsum. Ocular tubercle length 0.40-0.46, width 0.39-0.50, height 0.1 1-
0.20, distance from anterior edge of cephalothorax 0.24-0.40. Generally, brown
dorsal lines distinct on palpal patellae and tibiae. Palpal segment lengths: femora
0.80-0.97, patellae 0.40-0.58, tibiae 0.59-1.00, tarsi 1.19-1.37. Legs as on males
except basal portion of femora often light yellowish-brown and characteristic bands
of males mottled with brown splotches; tarsi and metatarsi speckled with brown,
often lacking bands. Metatarsi I with 1-3 (rarely lacking) pseudosegments, tibiae II
with 3-7 (see comments under variation), tibiae IV without pseudosegments (rarely

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Figs. 36-39. Leptobunus californicus palpi. 36. Male, lateral view. 37. Male, mesal view.
38. Female, lateral view. 39. Female, mesal view. Scale line = 1.0 mm.

one present). Femora I-IV lengths (respectively): 2.81-3.94, 6.18-9.40, 3.18-4.24,
4.89-6.80; tibiae I-IV lengths (respectively): 2.52-3.55, 5.20-6.42, 2.60-4.06, 3.68-
4.64. Ovipositor with 24-30 darkly pigmented rings (N = 3); furca 3 segmented, 2nd
joint with 1-4 (averaging 2, N = 14) slit sensilla per side (Fig. 30). Seminal receptacles
(Figs. 31-33) with very short primary loop; basal loops very irregular (see comments
under variation).

Immatures: Similar to adult females, except with body ranging from opalescent
white with few marks to pale yellowish-white with dark brown patterns. The dis-
tinctive abdominal cross pattern is present on juveniles as small as 1.0 mm total
length.

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389

Variation. Throughout its range this species exhibits considerable variation in color
and size. Leg pseudosegment counts and male chelicerae also differ between popu-
lations. Specimens from northern California, particularly Siskiyou Co., generally have
slightly higher tibiae II pseudosegment counts: range 3-7, averaging 5. Specimens
from central California, Oregon, and northwards average 3, range 2-5, tibiae II
pseudosegments. The moveable finger of the chelicera is equipped with a large apoph-
ysis on males from Washington and Canada (Figs. 46-47), whereas males from
Oregon and California generally have smaller apophyses (Fig. 48). The female seminal
receptacles also vary considerably, but no geographical trends were noted except for
the more convoluted basal loops on some females from central California (Fig. 33).
It must be stressed, though, that these are only trends and cannot be relied upon for
specific locality identifications. Specimens from Siskiyou County are particularly
problematic and may represent a relict population.

Natural history. The majority of the collections of juveniles (all stages) and adults
were made during August and September. Latitude does not seem to affect signifi-
cantly the seasonal abundance of L. parvulus. A few collections are known from
April to July, and October to December. Some females collected during September
(California and British Columbia) were filled with eggs. A female collected and caged
on 28 September (by PDB at Vancouver) laid 21 eggs (letter dated 8 June 1982 from
Robert G. Holmberg, Athabasca University, Edmonton, Alberta).

Leptobunus parvulus appears to be restricted to the Sierra Nevadan Biotic Province
of California (Schick, 1965) and the Vancouveran Province of Van Dyke (see Leech,
1972). Specimens have been collected from sea level (Alaska, Oregon, and British
Columbia) to ca. 2,700 m elevation (central California). Throughout the range in
California L. parvulus occurs under 2,700 m, Oregon under about 1,800 m, and
under about 1 00 m further north. These are only general trends, though, as exceptions
are known (i.e., Mt. Blunt, Alaska at 760-1,020 m).

Holmberg et al. (1981) reported that this species has been taken from the tops of
29 m tall cedar, fir, and hemlock trees. Those specimens were all juveniles and were
collected during June, July, and August in British Columbia. The juveniles were
collected at 20, 23, 26, and 29 m heights in Western Red Cedar (Thuja plicata Donn),
Douglas Fir ( Pseudotsuga menziesii (Mirb.) Franco), and Western Hemlock ( Tsuga
heterophylla (Raf.) Sarg.), and additional juveniles were also obtained at 17 m height
in cedar (letter dated 8 June 1982 from Robert G. Holmberg, Athabasca University,
Edmonton, Alberta). This species is commonly encountered on low branches and
shrubs and on Alder trunks in forests of southern British Columbia (letter dated 8
June 1982 from Robert G. Holmberg, Athabasca University, Edmonton, Alberta).
Specimen labels indicate that adults and immatures were also collected from the
leaves or branches of Abies grandis (Dougl.) Lindl., Picea sp., Pinus contorta Dough,
Quercus garryana Dough, Gault heria shallon Pursh, and Vaccinium occidentale Gray.
Two immatures were also collected from the fern Polystichum, and one of these had
mature sporangia attached to two different leg tarsi.

Labels accompanying museum specimens indicate this species is active at night
and is attracted by UV light. Specimens from Haines, Alaska, were found under
“drift” on beach, whereas specimens in the Trinity Alps, California, were at snowfield
edges.

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Figs. 40-50. Leptobunus spp. male chelicerae. 40, 42, 44, 46, 49. Mesal views. 41, 43, 45,
47, 48, 50. Anterior views of moveable fingers. 40, 41. L. californicus. 42, 43. L. borealis. 44,
45. L. pallidus, subadult. 46-48. L. parvulus. 49, 50. L. aureus. Scale line = 1.0 mm.

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391

Specimens examined. U.S.A. ALASKA: Glacier Bay Natl. Mon., Lituya Bay, Mt.
Blunt, 8-9 Aug. 1979 (D. H. Kavanaugh), 1 immature (CAS); Haines, 20-24 Aug.
1945 (J. C. Chamberlin), 8 males, 3 females, 34 immatures (AMNH). WASHING-
TON: Thurston Co., Olympia, pre-1 894 collection (T. Kincaid), lectotype male, male
and female paralectotypes (MCZ), Tacoma, Puget Sound, 9 Sept. 1906 (Kpt. R.
Paessler), 1 male (ZIZM); Jefferson Co., Port Ludlow, Puget Sound, 12 Oct. 1907
(Kpt. R. Paessler), 1 female (ZIZM), Puget Sound, pre-1 9 10 collection (Kpt. R.
Paessler?), 1 female (SMF); Grays Harbor Co., Moclips, 25 Aug. 1981 (A. Moldenke),
1 immature (JCC). OREGON: Polk Co., 12.8 km W Grande Ronde, 27 Nov. 1940
(W. Ivie), 1 female (AMNH); Hood River Co., Mt. Hood, Still Creek, Still Creek
Forest Camp, 28-29 Sept. 1966 (Wiggins, Yamamoto, Odum), 1 1 males, 4 females
(ROM); Benton Co., Parker Creek, Mary’s Peak, 24 Sept. 1966 (Wiggins, Yamamoto,
Odum), 5 males, 6 females, 1 immature (ROM), McDonald Experimental Forest,
13 Sept. 198?, 29 July 1981, 1 Sept. 1981, 16 Oct. 1981, 25 June 1982, 15 Sept.
1 982, 24 Nov. 1 982 (B. Fichter and A. Moldenke), 1 4 males, 9 females, 1 7 immatures
(JCC), Lewisberg Saddle, 30 May 1981, 29 July 1981, 27 Aug. 1981, 16 Oct. 1981,

1 Nov. 1981, 25 June 1982 (B. Fichter and A. Moldenke), 13 males, 3 females, 61
immatures (ARM), Ballards Bay State Park, 24 Nov. 1981 (B. Fichter and A. Mol-
denke), 5 males, 3 females (JCC), Corvallis Chip Ross Park, 25 June 1982, 16 Aug.
1982 (A. Moldenke and B. Fichter), 4 immatures (ARM), Mary’s Peak summit, 2
Oct. 1981 (B. Fichter and A. Moldenke), 1 female (ARM), Mary’s Peak subsummit,

2 Sept. 1981, 2 Nov. 1981, 6 Aug. 1982, 2 Sept. 1982 (B. Fichter and A. Moldenke),

3 males, 8 females, 20 immatures (ARM); Jefferson Co., head Metolius River, Riv-
erside Forest Camp, 19-23 Sept. 1966 (Wiggins, Yamamoto, Odum), 4 males, 1
female (ROM), Spring Creek, 19-20 April 1952 (V. Roth), 1 female (CAS); Lane
Co., Honeyman State Park, 20 June 1966 (T. Briggs and V. Lee), 1 immature (CAS);
Fairview Peak summit, 28 May 1981 (A. Moldenke), 17 immatures (ARM), Mutton
Meadow, SE of Rigdon, 14 July 1981 (A. Moldenke), 2 immatures (ARM), Rujada
Campground, 26 May 1 98 1 (A. Moldenke), 1 immature (ARM), Neptune State Beach,
5 Aug. 1982, 10 Oct. 1982, 24 Nov. 1982 (B. Fichter and A. Moldenke), 2 males, 2
females, 9 immatures (JCC), Willamette National Forest, Sacandaga Campground,
15 July 1981, 21 Aug. 1982 (B. Fichter and A. Moldenke), 5 immatures (ARM),
West Lake, 23 June 1982, 10 Oct. 1982 (B. Fichter and A. Moldenke), 7 males, 8
females, 12 immatures (ARM), Washburn State Park, 24 Oct. 1981, 24 June 1982,
5 Aug. 1982, 10 Oct. 1982 (B. Fichter and A. Moldenke), 6 females, 19 immatures
(JCC), Andrews Experimental Forest, 11 July 1981, 2 Aug. 1981, 3-5 Sept. 1981,
13 Oct. 1981, 10-12 July 1982, 12-18 Aug. 1982 (B. Fichter and A. Moldenke), 11
males, 18 females, 62 immatures (JCC); Deschutes Co., South Sister, 31 Aug. 1941
(B. Malkin), 1 female (AMNH), 32 km S of Sisters, 11 Sept. 1948 (Roth), 1 female
(AMNH), Sisters Mtns., Green Lake, 28-30 Aug. 1952 (V. Roth), 1 male (CAS),
Crescent Lake, 22 Aug. 1982 (A. Moldenke), 1 male (JCC); Jackson Co., 32 km NE
of Ashland, 1 Sept. 1959 (W. J. Gertsch and V. Roth), 1 immature (AMNH); Klamath
Co., Odell Lake, 30 Aug. 1981 (B. Fichter and A. Moldenke), 6 immatures (JCC);
Tillamook Co., Cascade Head, 5 Aug. 1982 (B. Fichter and A. Moldenke), 2 im-
matures (JCC); Linn Co., Rooster R. Rock, 24 Oct. 198 1 (B. Fichter), 1 female (JCC);
Coos Co., Brandon, 24 Nov. 1981 (B. Fichter and A. Moldenke), 1 male (JCC);
Lincoln Co., Gleneden Beach, 13 Sept. 1981, 24 June 1982, 5 Aug. 1982 (B. Fichter

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and A. Moldenke), 6 males, 5 females, 25 immatures (JCC), Cascade Head, N of
Lincoln City, 18 June 1981, 13 Sept. 1982 (B. Fichter and A. Moldenke), 1 male, 2
females, 6 immatures (ARM); Marion Co., Silver Creek Falls, July 1941 (J. C.
Chamberlin), 1 immature (AMNH), Ochoco National Forest, Summit Campground,

1 1 June 1982 (B. Fichter and A. Moldenke), 1 immature (JCC). CALIFORNIA: Del
Norte Co., Middle Fork Smith River, 4 Sept. 1963 (J. and W. Ivie), 1 immature
(AMNH); Siskiyou Co., Trinity Alps, Caribou Basin (south rim), at snowfield edges,

12 Aug. 1980 (D. H. Kavanaugh), 1 immature (CAS), Panther Meadows, Mt. Shasta
Ski Bowl, 2 Sept. 1959 (V. Roth and W. J. Gertsch), 5 males, 5 females (AMNH),
Tule Lake, 14.4 km SW Tulelake, 15 Sept. 1965 (J. and W. Ivie), 6 immatures
(AMNH), Paynes Creek, 8 km by road SE of Medicine Lake, 22 Aug. 1966 (H. B.
Leech), 3 males, 1 female, 1 immature (CAS); Plumas Co., S side Lake Almanor, 5
Sept. 1959 (V. Roth and W. J. Gertsch), 3 females, 2 immatures (AMNH); Butte
Co., Little Butte Creek, Toadtown, 4 April 1977 (C. L. Hogue), 1 female (LACM);
Sierra Co., 3.2 km N Calpine, 6 Sept. 1959 (W. J. Gertsch and V. Roth), 1 male, 1
female, 2 immatures (AMNH); Placer Co., Squaw Valley, 20 Sept. 1961 (W. Ivie
and W. J. Gertsch), 2 females (AMNH); Eldorado Co., Fallen Leaf Lake, 9 Sept.
1959 (W. J. Gertsch and V. Roth), 1 male, 1 female (AMNH); Tuolumne Co.,
Pinecrest, 10 Sept. 1959 (W. J. Gertsch and V. Roth), 1 male, 2 females (AMNH),
Tamarak Flat, Yosemite Natl. Park, 1 1 Sept. 1959 (W. J. Gertsch and V. Roth), 1
male (AMNH); Mariposa Co., 1.6 km S Fish Camp, 23 Sept. 1961 (W. Ivie and W.
J. Gertsch), 1 female (AMNH). CANADA. BRITISH COLUMBIA: Queen Charlotte
Islands, Masset, 1891-7-2-16-26 (J. H. Keen), 4 females (BMNH), W of Dawson
Inlet, 29 Aug. 1961 (J3=), 1 immature (CNC), Alleford Bay (=Alliford Bay), 26 Aug.
1961 (collector unknown), 1 female (CNC), Vancouver Island, Coombs, spring 1979
(D. Salter), 1 male (JCC), Golden Ears Prov. Park, 2 Sept. 1973 (C. Holmberg), 1
female (RGH), Alouette Lake, 24 Aug. 1973 (R. G. Holmberg), 1 immature (RGH),
Burnaby Mountain, Simon Fraser University, 16 Oct. 1971 (R. G. Holmberg), 1
male (JCC), 1 Oct. 1973 (R. G. Holmberg), 1 female (RGH), Point Gray, Vancouver,
18-28 May 1973 (J. R. Vockeroth), 1 immature (CNC), Vancouver, 25 Sept. 1965
(P. D. Bragg), 1 female (PDB), 14 Sept. 1968 (P. D. Bragg), 1 male (PDB), 28 Sept.
1968 (P. D. Bragg), 1 male (PDB), 28 Oct. 1968 (P. D. Bragg), 3 males (PDB), 23
Nov. 1968 (P. D. Bragg), 1 male (PDB).

Leptobunus aureus, new species
Figs. 4, 15-17, 34, 35, 49, 52, 63

Types. Male holotype from Inverness, 60 m elev., Marin Co., California, 11-13
Jan. 1964 (P. H. Arnaud, Jr.), CAS; seven paratypes (listed under specimens ex-
amined).

Etymology. The specific epithet from Latin, referring to the golden color of the
body.

Diagnosis and comparisons. Leptobunus aureus is one of two species in the parvulus
group. In addition to differences in the genitalia, L. aureus can be distinguished from
other Leptobunus spp. by its small size and golden color.

Distribution. Known only from Santa Cruz, Marin, and Mendocino Counties in
California (Fig. 63).

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393

51

52

Figs. 51-60. Tibiae and tarsi of male Leptobunus spp. palpi (semi-diagrammatic). 51, 53,
55, 57, 59. Lateral views. 52, 54, 56, 58, 60. Dorsal views. 51, 52. L. aureus. 53, 54. L. parvulus.
55, 56. L. pallidus, subadult. 57, 58. L. borealis. 59, 60. L. californicus.

Description. Males: Total length 2.90-3.15, greatest width 2.00-2.30, maximum
height 1.56-1.77; dorsum light golden brown with dark brown velvety reticulations
on cephalothorax, faint cross pattern on abdomen which is invaded by many golden
brown and white spots, and last 4 to 5 distal tergites and anal operculum with creamy
white stripe (Fig. 4). Ocular tubercle length 0.34-0.35, width 0.37-0.39, height 0. 1 5-
0. 1 7, distance from anterior edge of cephalothorax 0.23-0.25; white with dark brown
to black rings around eyes, rings thicker anteriorly and appearing elliptical. Coxae,
genital operculum, and abdominal stemites creamy white to light yellow brown with
numerous white blotches. Chelicerae silvery white to yellow brown, lateral surface
of basal segment and proximal half of distal segment mottled with brown; basal
portions of fingers yellow brown, tips of fingers and teeth dark brown to black;
moveable finger with conspicuous spur (Figs. 49, 50), concolorous with finger. Palpi
creamy white to light yellow brown; dark brown mottlings on femora dorsally; dark
brown stripes on dorsum and basolateral half of patellae, proximal half of dorsum

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and basolateral % of tibiae; basal portion of tarsi only slightly enlarged (Figs. 5 1 ,
52). Palpal segment lengths: femora 0.73-0.79, patellae 0.37-0.40, tibiae 0.50-0.60,
tarsi 0.94-0.98. Legs creamy white to light golden brown; femora with many ran-
domly spaced brown splotches and irregular margined dark brown band on distal
ends; patellae golden dorsally, margined on both sides with dark brown stripes in-
terrupted by brown splotches; tibiae with dark brown bands distally, less distinct
subbasal bands on I, III, IV; tarsal and metatarsal elements shaded brown basally;
metatarsi I with one pseudosegment; tibiae II with 4 or 5 pseudosegments; tibiae IV
with one pseudosegment (rarely lacking). Femora I-IV lengths (respectively): 3.00-
3.25, 6.25-6.71, 3.30-3.50, 4.89-5.00; tibiae I-IV lengths (respectively): 2.58-2.70,
5.63-5.70, 2.56-2.67, 3.81-3.86. Penis (Figs. 15-17) with truncus slightly enlarged
on subbasal surface ventrally; glans primary setae bilobed (often asymmetrically),
spaced much further apart than stylus length, subequal in length to stylus; distal end
of glans not noticeably constricted.

Females: Total length 5.00-6.81, greatest width 3.26-3.61, maximum height 3.11—
3.21; body similar to male, darker with less golden color; dorsal cross pattern obscured
by extensive brown splotching, white stripe on posterior end of abdomen and anal
operculum much thinner but extends to base of cross pattern (stripe absent on Santa
Cruz female). Ocular tubercle length 0.40-0.42, width 0.42-0.45, height 0.18-0.22,
distance from anterior edge of cephalothorax 0.28-0.30. Palpal segment lengths:
femora 0.81-0.83, patellae 0.42-0.48, tibiae 0.70-0.73, tarsi 1.17-1.20. Femora of
legs with fewer brown splotches than male; distal brown bands distinct, margins less
irregular. Leg pseudosegments as in males, except single female with 6/4 tibiae pseu-
dosegments. Femora I-IV lengths (respectively): 3.15-3.75, 6.64-7.34, ?-3.74, 5.12-
5.74; tibiae I-IV lengths (respectively): 2.80-2.94, 5.80-6.20, ?-2.87, 3.98-4.29.
Ovipositor darkly pigmented, difficult to clear in clove oil, with 24-29 rings; furca
3 segmented, second joint with 1-3 (rarely 3) slit sensilla per side (Fig. 34). Seminal
receptacle (Fig. 35) with very short primary loop; basal loops very irregular, loop
constricting in size basally; basimesal tube undetected (lacking ?).

Immatures: Similar to adult female, except abdominal cross pattern distinct. One
specimen with 1 and 3 pseudosegments in metatarsi I, all others have single pseu-
dosegment; tibiae II with 3 pseudosegments; tibiae IV occasionally with 1 pseudoseg-
ment.

Natural history. Juveniles are known from late June and August, whereas adults
are known from September to November. The majority of specimens were collected
at 60 m elevation, but others are from 150 and 460 m. This species appears to be
restricted to lowland forests of the lower Humboldtian and Santa Cruzian biotic
districts of the Pacific Maritime Province (Schick, 1965). A female from Santa Cruz
was captured by beating the branches of redwoods.

Specimens examined. Male holotype and female paratypes. U.S.A. CALIFORNIA:
Mendocino Co., Mendocino, 20 June 1955 (J. R. Heifer), penultimate female (UCB);
Marin Co., Inverness, 19 Oct. 1946 (E. S. Ross), 2 males (CAS), 11-31 Jan. 1964
(P. H. Amaud, Jr.), 1 male (CAS), Lagunitas, 15 Sept. 1946 (E. S. Ross), 1 female
(CAS), North Toll Bridge, Bolinas Ridge, 3.2 km NW Bolinas, 13 Aug. 1965 (V. Lee
and T. Briggs), 2 immatures (CAS); Santa Cruz Co., University of California, Santa
Cruz Campus, 26 Nov. 1981 (A. Moldenke), 1 female (JCC).

1984

REVISION OF LEPTOBUNUS

395

Fig. 6 1 . Distribution of Leptobunus spp. in Mainland Alaska, and Aleutian and Bering Sea
Islands. Circles = L. borealis, star = L. borealis and L. pallidus.

SYSTEMATIC RELATIONS AND HIGHER CATEGORIES

There has been confusion regarding the date of publication of the family-group
name Leptobunidae in the past. Banks described Leptobunus in 1893 and placed it
in the tribe Liobunini Banks. Roewer (1910, 1923) cited Banks (1894a) as the author
of Leptobunini, and stated that it appeared on page 165. Silhavy (1960) claimed
Banks described it in 1893, and Rambla (1967) listed “Banks 1894.” Nowhere in
Banks (1 893, 1 894a) does the name Leptobunini appear; furthermore the latter article
ends on page 164. Likewise, Banks (1894b, 1894c, 1898, 1899, 1900a, 1900b) did
not mention the Leptobunini. The first reference to the tribe Leptobunini is Banks
(1901). In that paper, Banks did not discuss the name, simply listing it in the key to
the Phalangiidae. Nonetheless, Banks (1901) is to be credited with the establishment
of the family-group name Leptobunini (Stoll, 1961, art. 16). Roewer (1910) was the
first to diagnose and describe the “Leptobunini” as a subfamily. The following clas-
sification, proposed by Roewer (1910, 1923), has been essentially followed up to the
present:

Family Phalangiidae Simon
Subfamily Leptobuninae Banks
Leptobunus Banks
Cosmobunus Simon

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Protolophus Banks
Trachyrhinus Weed
Leurony chus Banks
Homolophus Banks

Recent studies of the harvestman subfamily Leptobuninae Banks (Leptobunidae
of Rambla, 1977; Star$ga, 1978; and Leiobunidae of Silhavy, 1960, 1965, 1976)
reveal that it is polyphyletic. This problem appears to be largely due to the hesitancy
of recent workers to give up the use of some of the traditional characters used by
Banks and Roewer to define taxa (see Roewer, 1923). Also the lack of comparative
material of North American species in European museums has slowed progress
towards a more natural classification. As noted by Cokendolpher (198 lb), the “catch
all” Leptobuninae must be dismantled with the type genus ( Leptobunus Banks) being
placed in the Phalangiidae (Phalangiinae). The complex species synonymies must
await the publication of much needed generic revisions.

Traditionally, members of the “Leptobuninae” have been reported as having smooth
palpal claws. Examination of numerous specimens reveals this is not entirely true.
Apparently denticles are often overlooked (Asian species referred to Homolophus
Thorell, some Protolophus Banks, Leptobunus Banks, and New World species referred
to Cosmobunus Simon) or the claws reported as having only a few minute denticles
on the base of the claw ( Leuronychus Banks). In the genus Protolophus, palpal claw
teeth may be present or absent in individuals of a single species. In some other cases,
the immatures have well developed palpal claw teeth which presumably wear down
or break off with age (see Suzuki, 1973). Additionally, two genera ( Trachyrhinus
Weed and Cosmobunus ) have bare palpal claws but agree in all other details with
members of the Gagrellidae (Leiobuninae and Gagrellinae), which characteristically
have denticulate claws. “Atypical” palpal claws are also known in other subfamilies.
Lawrence (1962) remarked on some of the problems with the African genus Cheops
Soerenson. The North American genus Liopilio Schenkel (see Cokendolpher, 1981b)
and the South American genus Thrasychirus Simon also have atypical palpal claws.
The “Dentizacheinae” (=Phalangiinae) of Silhavy (1960) likewise suggest the use of
alternate characters as pointed out by Star^ga (1973). Therefore, the presence or
absence of denticles on the ventral surface of the palpal claw is of limited taxonomic
use.

Traditionally, the lack of a cheliceral hook was used to characterize the subfamily
Phalangiinae, but this character appears to be plesiomorphic. Martens (1978) main-
tained two species in the genus Paroligolophus Lohmander: P. agrestis (Meade) with
a cheliceral hook, and P. meadii (Pickard-Cambridge) without it. Cokendolpher
(1981b) pointed out the similarity of Leptobunus to Liopilio, but noted that Lepto-
bunus always has a cheliceral hook, whereas in Liopilio the hook varies in size due
to age and sex of the specimen, and may be entirely lacking. Thus, the presence or
absence of a hook or tooth ventrally on the first joint of the chelicera is of restricted
value.

The angle of insertion of the endites of coxa II is often difficult to ascertain. All
genera referred to the Leptobuninae are reported as having the endites straight,
parallel to the lip of the genital operculum. In reality, several of these genera have
the endites slanted anteriorly as in the Phalangiinae. As the angle of insertion of the

1984

REVISION OF LEPTOBUNUS

397

Fig. 62. Distribution of Leptobunus parvulus in western North America (Alaska to Cali-
fornia).

endites is altered, to some extent, by the method of preservation and the age and
sex of the animal, it should be used with great caution.

Comparative lengths of the legs has been used to distinguish genera in the Phalangi-
oidea. Although often useful in verifying identifications, it should not be used as the

398

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Fig. 63. Distribution of Leptobunus spp. in central California and western Nevada. Circles =
L. californicus, triangles = L. aureus.

sole character to delineate taxa. Particularly problematic is the ratio of leg lengths
to the body length. Although not as variable in males, female total length is greatly
influenced by enlargement of the abdomen during feeding and reproductive periods.
Traditionally, Leptobunus and Leuronychus have been separated by the lengths of
the legs. As Leptobunus is now known to be a member of the Phalangiidae and
Leuronychus a member of Gagrellidae, the use of leg lengths seems trivial. Unfor-
tunately, up to now Leuronychus parvulus Banks has remained in the wrong genus,
and, even though it has relatively long legs it is clearly a species of Leptobunus.

The following placements are tentative and may be altered after thorough taxo-
nomic revisions have been completed. The family classification is essentially that of
Silhavy (1960), modified slightly based on descriptions of genital morphology by
Martens (1978).

As already noted the genus Leptobunus is a member of the Phalangiinae (Pha-
langiidae). Members of the genus, as well as others incorrectly placed in the genus,
are discussed earlier in this paper in the context of a generic revision.

Cokendolpher (1981b) reported that the monotypic East Asian genus Tchapinius
Roewer is most similar to Mitopus Thorell. A revision of Mitopus currently underway
will further document this affinity. Both genera are members of the Phalangiinae
(Phalangiidae).

Leuronychus Banks is a member of the Leiobuninae (Gagrellidae). The Gagrellidae

1984

REVISION OF LEPTOBUNUS

399

is equivalent to the Leiobunidae of some authors, but as noted by Star^ga (1972),
the latter name is a junior synonym. Presently, Leuronychus pacificus (Banks) is the
only described species correctly placed in that genus. As pointed out by Cokendolpher
(1980) Leuronychus gertschi Schenkel is a junior synonym of Leiobunum exilipes
(Wood). Examination of the female type of “ Liobunum” fulviventris O. Pickard-
Cambridge revealed that Roewer (1910, 1923) erroneously referred this species to
Leuronychus. This species is known only from two females which have been pinned
and dried, so information on the genitalia is not available. Ultimate placement will
have to await revisions of the New World species referred to Leiobunum C. L. Koch,
Nelima Roewer, and Paranelima di Caporiacco. As already noted, Leuronychus
parvulus Banks is properly a member of Leptobunus. At least two undescribed western
North American Leuronychus species await formal description.

The genus Homolophus Banks is polyphyletic. Unfortunately, I have been unable
to examine any specimens of the type species, Homolophus arcticus Banks. The New
World Homolophus biceps (Thorell) is not congeneric with the Asian Homolophus
iranus Roewer and Homolophus afghanus Roewer. Levi and Levi (1955) correctly
synonymized Togwoteeus granipalpus Roewer with H. biceps. The Homolophus species
known to me are properly members of the Leiobuninae (Gagrellidae).

Goodnight and Goodnight (1942a) revised the western North American genus
Protolophus Banks; however, a new revision is needed because large series of addi-
tional specimens are now available. The genital morphology, presence of low ab-
dominal tubercles and the position of the scent gland pores place it in the Sclero-
somatinae (Gagrellidae). As this taxa differs some from typical Sclerosomatinae, the
Protolophinae of Banks (1893) may have to be resurrected. Traditionally, the mem-
bers of Protolophus have been diagnosed by the pedipalp segment lengths. Protolophus
longipes Schenkel and often Protolophus niger Goodnight and Goodnight have the
male palpal tarsus longer than the tibiae, whereas the male tarsus is shorter than the
tibiae on all other Protolophus species. All immatures and females of Protolophus
spp. examined have the palpal patellae extended on the inner margin, as well as a
slight to moderate swelling on the inner distal margin of the palpal tibiae.

The Old World Cosmobunus granarius (Lucas) is not congeneric with New World
species referred to Cosmobunus, but all are members of the Leiobuninae (Gagrellidae).
New World species appear to be congeneric with a large complex of species presently
referred to Leiobunum. Ultimate placement of Nearctic “ Cosmobunus ” must await
a thorough taxonomic revision of New World species referred to Leiobunum. The
Old World Cosmobunus were revised and redescribed by Rambla (1970).

Trachyrhinus was recently revised, and although Cokendolpher (1981a) did not
state to which family Trachyrhinus belongs, the presence of pseudoarticulary nodules
of femora II was noted. Trachyrhinus was subsequently transferred to the Gagrellinae
(Gagrellidae) by Cokendolpher (1984a).

Star^ga (1978) suggested that the North American genus Globipes Banks may be
a member of the Leptobuninae. Globipes is properly a member of an assemblage of
phalangioid genera which is as yet unnamed (Cokendolpher 1984b; Cokendolpher
and Cokendolpher 1984).

The South American genus Thrasychirus Simon was referred to the Leptobuninae
by Canals (1935). However, Silhavy (1970) noted Ringuelet’s (1959) synonymy of

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JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Enantiobunus Mello-Leitao with Thrasychirus, and transferred Thrasychirus to the
Enantiobuninae (Neopilionidae).

ACKNOWLEDGMENTS

I would like to thank the many individuals, curators, and their institutions (listed under
materials and methods) for gifts and loans of specimens. Valuable information on Leptobunus
and “Leptobunidae” in general was provided by R. L. Crawford, to whom I am grateful. I
would also like to thank T. S. Briggs for assistance in locating some localities and altitudes in
California. A special thanks is extended to R. G. Holmberg for sharing his knowledge of
Leptobunus in Canada, sorting pertinent material from collections at CNC, and for aiding in
mapping localities in Canada. Translations of several papers were generously provided by C.
Bayat-Makou and M. K. Rylander. My wife, Jean, typed and critically read several drafts of
the manuscript, and aided in the preparation of some of the illustrations. O. F. Francke, M.
K. Rylander, and J. K. Wangberg reviewed the manuscript, and Mary L. Peek kindly typed the
final draft of the manuscript. This study was supported by a grant from the Exline-Frizzell
Fund for Arachnological Research, California Academy of Sciences. Further support was pro-
vided by The Museum and the Department of Biological Sciences, Texas Tech University. This
is contribution No. T-10-152 of the College of Agricultural Sciences, Texas Tech University.

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Received July 6, 1984; accepted September 18, 1984.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 403-404

NOTES AND COMMENTS

OVIPOSITION BEHAVIOR OF CHRYSOPS ATLANTICUS
(DIPTERA: TABANIDAE)

The salt marsh deer fly Chrysops atlanticus Pechuman is an annoying blood-seeking
pest of man and animals along the Atlantic coast, where it often has a negative impact
in recreational and agricultural industries (Hansens, 1980). Egg masses and ovipo-
sition sites of C. atlanticus have previously been reported by Jamnback and Wall
(1959) and Magnarelli and Anderson (1979). Although these authors reported finding
egg masses on Spartina alterniflora Loisel., there have been no previous observations
of oviposition other than in the laboratory.

We observed oviposition by 12 female C. atlanticus (Table 1) at a salt marsh near
Cedarville, New Jersey. Oviposition was observed in early evening, from 1912 hr
(EDT) to 2040 hr (mean ± SE = 36 ± 8 min before sunset). Ten females (83%)
oviposited on S. alterniflora, but oviposition was also observed on Spartina patens
(Ait.) and Distichlis spicata (L.). In each case the eggs were laid over wet depressions
which supported little or no vegetation, as reported by Magnarelli and Anderson
(1979).

Female C. atlanticus were oriented in a head up position prior to oviposition, then
turned 1 80 degrees (head down). As the female walked down the blade of grass, she
repeatedly touched the leaf surface with the tip of her abdomen. When oviposition
was initiated, the fly slowly moved down the blade of grass, depositing the eggs for
approximately 20 min. The females were easily disturbed if approached by an ob-
server, and left the immediate area. Freshly deposited eggs were white and turned
dark brown after several hours. Larvae hatching from the egg masses possessed the
exertile stigmatic spine characteristic of C. atlanticus.

These data show that C. atlanticus oviposits exclusively during the early evening
hours. We have observed other fresh egg masses (i.e., white) during the early evening
hours but not during the morning or afternoon despite extensive searching. Graham
and Stoffolano (1983b) reported oviposition in Tabanus simulans Walker over an
eight hour period with peak activity during midday. We have frequently observed
Tabanus lineola F. and Hybomitra daeckei (Hine) ovipositing during the early after-
noon and late morning, respectively. The ecological significance of temporal parti-
tioning of oviposition among salt marsh tabanid species is unknown, as are the stimuli
for ovipositioning. The narrow time span for oviposition of C. atlanticus (only 88
min) appears to be unusual for salt marsh tabanids.

In previous reports, S. alterniflora was the only plant on which the eggs of C.
atlanticus were found. While C. atlanticus oviposited predominantly on this plant
in our study, other plants were also suitable. The dominance of S. alterniflora around
wet depressions may account for the number of ovipositions observed on this plant.
Stimuli associated specifically with wet depressions are probably more important in
the choice of an oviposition site for C. atlanticus than stimuli associated with a
specific plant. Tabanus simulans oviposited almost exclusively on S. alterniflora and

404 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Table 1. Date, time (EDT), and plant substrate of oviposition by C. atlanticus on a salt
marsh near Cedarville, New Jersey.

Date

Time

Plant

16 June 81

2010

S. patens

1 July 82

2033

S. alterniflora

7 July 82

2018

S. alterniflora

15 June 83

1931

S. alterniflora

16 June 83

1927

S. alterniflora

16 June 83

2009

S. alterniflora

23 June 83

1948

S. alterniflora

23 June 83

2001

S. alterniflora

23 June 83

2022

S. alterniflora

29 June 83

1912

D. spicata

30 June 83

2040

S. alterniflora

13 July 83

2021

S. alterniflora

may have been responding to plant specific stimuli (Graham and Stoffolano,
1983a, b).

New Jersey Agricultural Experiment Station, Publication No. D-08 1 1 5-29-83, sup-
ported by State funds and by U.S. Hatch Act.— Roy K. Sofield and Randy Gaugler,
Department of Entomology and Economic Zoology, Cook College, New Jersey Ag-
ricultural Experiment Station, Rutgers University, New Brunswick, New Jersey 08903;
RKS present address: Science Department, Tennessee Temple University, Chatta-
nooga, Tennessee 37404.

LITERATURE CITED

Graham, N. L. and J. G. Stoffolano, Jr. 1983a. Relationship between female size, type of egg
mass deposited, and description of the oviposition behavior of the sibling species Ta-
banus nigrovittatus and T. simulans (Diptera: Tabanidae). Ann. Entomol. Soc. Amer.
76:699-702.

Graham, N. L. and J. G. Stoffolano, Jr. 1983b. Oviposition behavior of the salt marsh
greenhead, Tabanus simulans (Diptera: Tabanidae). Ann. Entomol. Soc. Amer. 76:703-
706.

Hansens, E. J. 1980. Review: Tabanidae of the east coast as an economic problem. J. New
York Entomol. Soc. 87:312-318.

Jamnback, H. and W. Wall. 1959. The common salt-marsh Tabanidae of Long Island, New
York. New York State Mus. Sci. Serv. Bull. 375, 77 pp.

Magnarelli, L. A. and J. F. Anderson. 1979. Oogenesis and oviposition in Chrysops atlanticus
(Diptera: Tabanidae). Ann. Entomol. Soc. Amer. 72:350-352.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 405-406

CARRION AS AN ATTRACTANT TO THE AMERICAN
DOG TICK, DERMACENTOR VARIABILIS (SAY)

The American dog tick, Dermacentor variabilis (Say), is a three-host tick with
larvae and nymphs that feed on small mammals, and adults that feed and mate on
larger ones. Discovery of hosts is usually facilitated by dispersal to areas where the
probability of contacting vertebrates is maximized, that is, near runways, trails or
roadsides frequented by the appropriate host species. At these places ticks typically
climb vegetation where they wait for a host to brush by (Smith et al., 1946; Sonen-
shine, 1979).

The purpose of this report is to document a newly discovered behavior of D.
variabilis that could be important in describing its ability to locate large mammalian
hosts. The following observations were made while studying the consumption of
mouse carrion in a closed-canopy mixed woodland near Chapel Hill, North Carolina,
U.S.A. The area has been described earlier by Kneidel (1984).

Six mouse carcasses ( Mus musculus L.) were exposed in plastic dishes placed 20
m apart along a transect through the woodland on 1 July 1982. The objective of the
experiment was to deter colonization by blowflies (Calliphoridae) by covering the
openings of the dishes with fiberglass window-screening. Large vertebrate scavengers
were also kept from the mice by fixing an additional cover of hardware cloth over
the dishes. This extended approximately 12 cm over the dishes and 8 cm around
them, and was held in place with three aluminum stakes.

Unlike unprotected carcasses, which are almost completely consumed within 1
week and do not have a strong smell, the protected mice became bloated, decomposed
slowly and developed a very strong offensive odor. After 5 days the window-screening
over two carcasses was removed, and after 7 days for another. These carcasses still
decomposed slowly and retained their strong odor. Although small arthropods fed
on the carcasses all were largely intact when collected from the field (after 1 0 or 11
days) and none were colonized by blowflies.

Two additional carcasses were exposed on 7 July. These had been kept at room
temperature indoors for 5 days before being moved to the field and were not covered
by window-screening. These also were not colonized by calliphorids, decomposed
slowly and had a very strong odor.

On 1 1 July I noticed adult ticks crawling near some of the carcasses. Subsequently,
I carefully inspected each one, including all leaf litter and vegetation over approxi-
mately a 0.50 m2 area around each mouse. I counted 23 non-engorged adult ticks
among the eight carcasses (4 had none, 1 had 1 , 1 had 4, and 2 had 9).

To determine whether the ticks were actually being attracted by the carrion or
whether their presence was independent of it, I also sampled an equivalent area to
that surveyed around the mice at eight sites without carcasses on 1 1 July. These were
located at 20 m intervals along a parallel transect, 20 m distant from the one with
the mouse carcasses. No ticks were found at any site, so the presence of individuals
around the mouse carcasses must be attributed to their attraction to them, and not
to chance.

406

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Given the length of time the carcasses had been in the field and the fact that one
of the two that had been kept indoors for 5 days yielded nine ticks, it can be concluded
further that the attractant must have been the odors of decomposition and not
lingering odors of the living mice. Also, the mice were kept frozen before being used
in the experiment and had no unusual odor when placed in the field.

A likely explanation for the ticks’ movement toward decomposing carcasses is that
by being near carrion, discovery and colonization of large mammalian scavengers is
facilitated. Interestingly, none of the ticks were found directly on the carcasses. Instead
they were climbing about the protective screen of hardware cloth over them. Ap-
parently the ticks climb vegetation surrounding carrion (or in this case hardware
cloth) to contact scavengers when they come near.

From 1979 through 1981 I exposed 94 dead mice and cotton rats, but observed
only four ticks near one carcass. Their attraction to the eight mice discussed above
could be explained by the fact that these carcasses were unique in having such a
strong odor. Possibly, larger carcasses are visited more frequently by ticks. This
prediction, however, remains to be verified. Most studies of carrion-arthropod as-
sociations have dealt primarily with Diptera, Coleoptera, and Hymenoptera, with
limited discussion of other taxa. Reed (1958), however, in a thorough survey of the
arthropod fauna attracted to dog carrion in Tennessee, reported no ticks visiting the
carcasses.— Kenneth A. Kneidel, Curriculum in Ecology, University of North Caroli-
na, Chapel Hill, North Carolina, USA 27514.

LITERATURE CITED

Kneidel, K. A. 1 984. The influence of carcass taxon and size on species composition of carrion-
breeding Diptera. Amer. Midi. Nat. 1 1 1:57-63.

Reed, H. B., Jr. 1958. A study of dog carcass communities in Tennessee, with special reference
to the insects. Amer. Midi. Nat. 59:213-245.

Smith, C. N., M. N. Cole and H. K. Gouck. 1946. Biology and control of the American dog
tick. USDA Tech. Bull. 905, 74 pp.

Sonenshine, D. E. 1979. Ticks of Virginia. Res. Div. Bull. No. 139, V.P. I., Blacksburg, Virginia.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 407-410

A MORPHOLOGICAL DISTINCTION BETWEEN THE
LARVAE OF CHEUMATOPSYCHE CAMPYLA AND C. PETTITI
(TRICHOPTERA: HYDROPSYCHIDAE)

Thirty-nine Nearctic species of the caddisfly genus Cheumatopsyche Wallengran
have been described, illustrated, and keyed (Gordon, 1974). Most Cheumatopsyche
larvae are distinctive in having a median notch on the anterior margin of the fron-
toclypeal apotome (a median cephalic sclerite) (Wiggins, 1977). Although adult-larval
associations have been made for several species, characters distinguishing larvae
have been difficult to find (Ross, 1944; Wiggins, 1977).

Of the two species used in this study, C. campyla larvae prefer larger rivers, seldom
inhabiting small streams (Ross, 1944). This species is most abundant through the
Com Belt states but is found over most other parts of the continent as well. Cheu-
matopsyche pettiti ( =ana/is Banks, sensu Ross, 1944) show a preference for small
streams but also occur in larger rivers (Ross, 1944). This species exhibits a wide
geographical distribution, from the Atlantic to the Pacific through the northern states,
and south to Oklahoma and Georgia.

Described below is a qualitative morphological distinction between the larvae of
C. campyla and C. pettiti. This distinction is based on the comparative examination
of 40 associated specimens of C. campyla and 2 1 associated specimens of C. pettiti.
As supporting evidence, comparisons were also made with associated specimens
from the scientific collection of the Illinois Natural History Survey.

MATERIALS AND METHODS

From May through August 1978, and April through June 1979, weekly field col-
lections of caddisfly larvae and pupae were made by handpicking rocks in riffle areas
at two locations; Horse Creek, 0.8 miles from its confluence with the Kankakee River
near Custer Park, Illinois, and the Kankakee River between Custer Park and Rest-
haven, 0. 1 miles downstream from the mouth of Horse Creek. The pupae were placed
in 3 -dram vials which were approximately half-filled with stream water and then
placed on ice for transport to the laboratory for rearing (Smith, 1984). The larvae
were preserved in 70% ethyl alcohol (EtOH) for later study.

Laboratory-reared adults were identified and preserved in 70% EtOH with their
pupal cases, which contained the hard parts of their larval exuvia. The “metamor-
photype method” of identification (Milne, 1938) was employed for those pupae which
did not emerge. With this procedure, associations between larva, pupa, and adult
can be made by using the imaginal genitalia for adult species identification, and
removing the larval sclerites from the case for identification of the larva. However,
most of the non-emerging pupae in this study were not mature enough for species
determination. Of those pupae which were identified, the larval sclerites and man-
dibles were slide mounted in polyvinyl lactophenol and examined.

The following larval characters were compared:

1 . The number of teeth on the left and right mandibles. A “mandibular formula”

408

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

Figure 1

Figure 2

Figure 3

Figs. 1-3. 1. Frontoclypeal apotome of Cheumatopsyche pettiti showing the shape of the

notch on the anterior margin. 2. Frontoclypeal apotome of Cheumatopsyche campyla showing
the shape of the notch on the anterior margin. 3. Frontoclypeal apotome of specimens from
the scientific collection of the Illinois Natural History Survey showing the difference in the

1984

NOTES AND COMMENTS

409

was used in making this comparison: (number of teeth on outer margin of man-
dible) + (number of teeth on inner margin of mandible) (e.g., 2 + 6).

2. Presence, or absence, and relative length and position of mandibular brushes.

3. Setal arrangement on setal areas 1, 2, and 3 (SA1, SA2, and SA3) (Wiggins, 1977),
on the meso- and metathoracic sclerites.

4. Presence or absence of setae and setal patterns on the head capsule.

5. Length/width ratio of the frontoclypeal apotome. The length was taken to be the
distance from the base of the median notch on the anterior margin of the fronto-
clypeus to the posterior tip of the apotome. The distance between the parallel
sides of the apotome was taken as the width.

6. Bristle patterns on the frontoclypeal apotome (Krafka, 1923).

7. Structural comparison of the shape of the anterior notch on the frontoclypeal
apotome.

The length/ width measurements were taken with an ocular micrometer at 63 x ,
all other characters were examined at 63 x and 160x .

The only morphological distinction observed between the larvae of C. campyla
and C. pettiti was a difference in the shape of the notch on the anterior margin of
the frontoclypeal apotome. The other six characters examined gave no information
useful in distinguishing between the larvae of these two species. Based on results
from 21 associated specimens, the anterior notch of C. pettiti was found to be a
shallow, crescent-shaped depression (Fig. 1). In contrast, the notch on all of the 40
associated specimens of C. campyla was larger and deeper, the lateral margins were
more angled, and the base of the depression was straight (Fig. 2). This character
difference was found to be consistent for all of the reared specimens during this study,
and also for the associated museum specimens which were examined (Fig. 3). The
fact that the museum specimens were collected from a wide range of geographical
locations lends further support to this character.

The Cheumatopsyche spp. larvae which were collected and preserved were also
compared. By using the above described character, as a general observation, it was
possible to separate the larvae as to the site where they had been collected, Horse
Creek or the Kankakee River. These results coincided with the distinction observed
between the associated specimens and with the locations where the associated spec-
imens were collected. Thirty-nine of the associated specimens of C. campyla were
collected from the Kankakee River location, and one was collected from Horse Creek.
Of the 2 1 associations made for C. pettiti, 1 9 were collected from Horse Creek and
two were taken from the Kankakee River.

The median notch on the anterior margin of the frontoclypeal apotome could be
an important character in further studies involving the larval taxonomy of the genus
Cheumatopsyche, although there is no doubt that variations within a population do

shape of the anterior notch, a. C. pettiti, Burton Creek, Quincy, Illinois, 1940. b. C. pettiti,
Piney Grove, Maryland, 1938. c. C. pettiti, South River, Palmyra, Missouri, 1938. d. C. cam-
pyla, Devils River, Devils River, Texas, 1939. e. C. campyla, Indian Creek, Serena, Illinois,
1938.

410

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

exist. Due to the qualitative nature of this distinguishing feature, it is suggested that
it be used only as a supporting character, providing one or more additional, possibly
quantitative, characters can be found.

ACKNOWLEDGMENTS

The author wishes to express thanks to Dr. R. W. Larimore for helping make this study
possible and to Dr. John D. Unzicker for confirming the species identifications of Cheuma-
topsyche adults. A special thanks goes to the numerous people who helped with both field and
laboratory work.— Martin H. Smith, University of Maryland, Munich Campus, Zoology De-
partment, APO New York 09407.

LITERATURE CITED

Gordon, A. E. 1974. A synopsis and phylogenetic outline of the Nearctic members of Cheu-
matopsyche. Proc. Acad. Nat. Sci. Philad. 1 26(9): 1 17-160.

Krafka, J., Jr., 1 923. Morphology of the head of Trichopterous larvae as a basis for the revision
of family relationships. J. New York Ent. Soc. 31:31-52.

Milne, M. J. 1938. The “metamorphotype method” in Trichoptera. J. New York Ent. Soc.
46:434-437.

Ross, H. H. 1944. The caddis flies, or Trichoptera, of Illinois. Bull. Illinois Nat. Hist. Surv.
23(1).

Smith, M. H. 1984. Laboratory-rearing of stream-dwelling insects. Antenna 8(2):67-69.
Wiggins, G. B. 1977. Larvae of the North American Caddisfly Genera (Trichoptera). Uni-
versity of Toronto Press, Toronto and Buffalo, 401 pp.

NEW YORK ENTOMOLOGICAL SOCIETY
92(4), 1984, pp. 411-413

BOOK REVIEW

A Field Guide to the Beetles of North America.— Richard E. White. Houghton-

Mifflin, Boston. 368 pp. $10.95 (paper).

White has produced a masterful field guide to the Coleoptera, brimming with more
than six hundred illustrations of the quality we have come to expect from the co-
author (with Donald J. Borror) of A Field Guide to the Insects (also of the Peterson
series by Houghton-Mifflin). White’s book could not have been more timely. It is a
first, surpassing other attempts at North American field guides for the beetles, and
is a badly needed source of information for amateurs who now cannot purchase
copies of Arnett’s Beetles of the United States or Crowson’s A Natural Classification
of the Families of Coleoptera. It cannot be said to replace either, but it serves an
immediate identification need by non-specialists and students in college courses, as
well as offering a useful pocket guide for the professional entomologist. Before ex-
tolling any more virtues of the tome, let me dispense with the problems I have found
in it.

Some of my complaints are genuine errors, pulled from some groups of special
interest to me, and I can only assume that comparable ones exist in treatments of
other taxa. Others, however, including the families chosen and the taxa included in
them, are more a matter of preference. In the book. White adopts a conservative
classification that overlooks many relations that now seem well established. For
example, the genus Dasycerus is included in the Lathridiidae (as was traditionally
the case, e.g., Leng, 1920), ignoring its placement in the Staphylinoidea by Crowson
(1955) and masking its fascinating relationships that almost certainly lie within Staph-
ylinidae (Wheeler, 1984).

The discussion of the numbers of North American species and biological habits
of the major genera of Leiodidae are essentially correct in the text (the “powdery
fungus’’ being Myxomycetes: Blackwell, 1984), but the figures are mislabeled. A
drawing of what appears to be Anisotoma discolor (Melsheimer) is labeled as “ Leiodes ”
and a Leiodes sp. as “ Anisotoma .” The nomenclatural problems of these two genera
are sufficiently monumental to justify White’s mistake, and I suspect that the national
collection may still be organized in the outmoded classification (see Hatch, 1929).

The family name Lymexylonidae is used instead of the grammatically correct
Lymexylidae (Barber, 1952). Just as in the case of Cerylonidae (cf. Cerylidae; Kuschel,
1979) the root does not include the -on ending on the genus name ( Lymexylon ,
Cerylon ). This, like the previous problem, is with published precedent and is not a
serious problem. I found that I could key out the lymexylid genus Hylecoetus in the
picture key on the front and rear endpapers, but its antennal structure is at odds with
the text description (p. 214). Melittomma, on the other hand, agrees with the text,
but does not easily key out. Also, the lymexylids are described simply as wood-
boring, overlooking their fascinating symbiotic relationship with ambrosia fungi that
predates comparable habits in scolytine weevils and may well represent the earliest
evolution of truly fungus-growing habits in the Coleoptera (Wheeler, ms; Wilson,
1971).

The literature cited at the end of the book excludes Roy A. Crowson’s classic
Biology of the Coleoptera (1981, Academic Press). I realize that Crowson’s book

412

JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 92(4)

probably appeared after the “guide” had gone into production, but a note in proof
of so important an addition to the literature on beetles would have been fully jus-
tifiable. And finally, I was a little amused and disappointed by the drawing of a
compound microscope boldly crossed out in the book. While it is not advisable for
an aspiring amateur beetle collector to run down to her local department store and
purchase a compound microscope, no one interested in beetles should be turned
from compound microscopy. Much of the detailed work necessary to adequately
describe small external structures, many genitalia and mouthparts, and many larvae
can only realistically be done with compound microscopy, and the general quality
of beetle taxonomy would only benefit from more widespread application of com-
pound microscopy by professional taxonomists and their user community.

I guess that many years have gone by since I purchased my last copy of Borror
and White’s insect field guide, and I was shocked to learn that the paperback edition
has more than doubled to nearly eleven dollars.

Most of the complaints that I have can be mitigated by an honest look at the size
of the project that White has completed. In North America alone there are more
than 30,000 species of beetles, and there is no way that the brief write-ups possible
in a field guide can do justice to all of them. Also, the field of coleopterology is
rapidly changing and no truly “up to date” publication of this scope is possible. As
an illustration, the associations of Eucinetidae with fungi were largely assumed at
the time of White’s writing, and in the interval North American species have been
associated with boletes, wood-rotting Basidiomycetes, and slime molds, confirming
and adding to our knowledge of mycophagy in that primitive polyphagan family
(Wheeler and Hoebeke, 1 984). The illustrations are superb. The blurbs on each family
are concise and largely accurate. The introductory chapters are helpful, and the
discussion of collecting methods includes more than the standard, general techniques.
An expanded treatment of the morphology of beetles would have been helpful, but
given the scope and goals of the volume, the existing balance seems reasonable.
Finally, the physical production of the book is good, surpassing that of some com-
peting field guide series.

I consider this the best available book for the beginning student of beetles. It is
extremely valuable in its own right, and will always be a useful companion to other
books (such as Arnett and Crowson, when these again become available). My copy
is already showing a little wear, and I suspect that a new generation of coleopterists
will benefit from this professionally prepared volume. I recommend the book highly,
and offer a thanks to Dr. White for his timely contribution. — Quentin D. Wheeler,
Department of Entomology, Cornell University, Ithaca, New York 14853.

LITERATURE CITED

Barber, H. S. 1952. Notes on Telegeusis and some relatives (Coleoptera: Lymexylidae). Pan-
Pac. Ent. 28:163-170.

Blackwell, M. 1984. Myxomycetes and their arthropod associates. Pp. 67-90 in: Q. Wheeler
and M. Blackwell (eds.), Fungus-Insect Relationships: Perspectives in Ecology and Evo-
lution. Columbia University Press, New York, 514 pp.

Crowson, R. A. 1955. The natural classification of the families of Coleoptera. Lloyd, London.
Hatch, M. H. 1929. Leiodidae in Coleopterorum catalogus, vol. pars 105.

1984

BOOK REVIEW

413

Kuschel, G. 1 979. The genera Monotoma Herbst (Rhizophagidae) and Anommatus Wesmael
(Cerylidae) in New Zealand. N.Z. Entomol. 7:44-48.

Leng, C. W. 1920. A catalog of the Coleoptera of America, north of Mexico. Sherman, Mount
Vernon, New York, 470 pp.

Wheeler, Q. D. 1984. Notes on fungus hosts of Dasyceridae of the Appalachian Mountains.
Coleopt. Bull, (in press).

Wheeler, Q. D. ms. Revision on the genera of Lymexylidae (Coleoptera, Cucujiformia). In
prep.

Wheeler, Q. D. and E. R. Hoebeke. 1984. A review of mycophagy in the Eucinetoidea,
including an association of the eucinetid beetle, Eucinetus oviformis, with a Coniophora-
ceae fungus (Coleoptera). Proc. Ent. Soc. Wash. 86:274-277.

Wilson, E. O. 1971. The Insect Societies. Harvard University Press, Cambridge, 548 pp.

1984

HONORAR Y MEMBERS

LIFE MEMBERS

Dr. F. S. Chew

Mrs. K. Firmage (SiZan Noguchi Swain)

Dr. James Forbes

Dr. John G. Franclemont

Dr. Alexander Klots

Dr. R. Vishniac

Dr. Lucy Clausen
Dr. Howard E. Evans
Dr. S. W. Frost
Mr. Irving Granek
Mr. Mark Indenbaum
Mr. Kikumaro Okano
Dr. Nellie M. Payne
Dr. L. L. Pechuman
Mr. Hubert J. Thelan
Mr. George R. Townes

SUSTAINING MEMBER

Yvonne Santiago

REVIEWERS

The Editorial Staff thanks the following individuals who reviewed manuscripts
considered for publication in 1984: J. R. Aldrich, J. Carpenter, R. C. Dalgleish, G.
B. Edwards, G. C. Eickwort, O. S. Flint, Jr., H. G. Fowler, R. C. Foreschner, B. J.
Harrington, L. H. Herman, D. D. Kopp, J. P. Kramer, J. E. McPherson, G. D.
Maddox, B. Peckarsky, L. L. Pechuman, S. Peterson, J. T. Polhemus, L. H. Rolston,
J. G. Rozen, Jr., M. D. Schwartz, W. A. Shear, J. A. Slater, R. R. Snelling, L. Sorkin,
D. W. Tallamy, D. B. Thomas, Jr., H. Topoff, D. M. Wood, R. K. Vander Meer, A.
G. Wheeler, Jr.

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Journal of the

New York Entomological Society

VOLUME 92 OCTOBER 1984

NO. 4

CONTENTS

Ormenaria rufifascia (Homoptera: Fulgoroidea: Flatidae): descriptions of nymphal
instars and notes on field biology Stephen W. Wilson and James H. Tsai

A revision of the genus Horridipamera Malipatil (Hemiptera: Lygaeidae)

James A. Slater and Le-Yi Zheng

New synonymy and a new species in the genus Mormidea (Hemiptera: Pentatomidae)

L. H. Rolston

Pellaea santarosensis (Hemiptera: Pentatomidae), a new species from Costa Rica

L. H. Rolston

Karyotype of Conomyra flava (McCook) (Hymenoptera: Formicidae)

James C. Cokendolpher and Oscar F. Francke

Key to the males of the nominate subgenus of Euschistus in South America, with
descriptions of three new species (Hemiptera: Pentatomidae) L. H. Rolston

Chemical analysis of the male aedeagal bladder in the fire ant, Solenopsis invicta
Buren David E. Ball, H. J. Williams and S. B. Vinson

Revision of the harvestman genus Leptobunus and dismantlement of the Leptobun-
idae (Arachnida: Opiliones: Palpatores) James C. Cokendolpher

Notes and Comments

Oviposition behavior of Chrysops atlanticus (Diptera: Tabanidae)

Roy K. Sofield and Randy Gaugler

Carrion as an attractant to the American dog tick, Dermacentor variabilis (Say)

Kenneth A. Kneidel

A morphological distinction between the larvae of Cheumatopsyche campyla and C.
pettiti (Trichoptera: Hydropsychidae) Martin H. Smith

Book Review

A Field Guide to the Beetles of North America Quentin D. Wheeler

Honorary, Life, and Sustaining Members
Reviewers for 1984

307-315

316-341

342-343

344-348

349-351

352-364

365-370

371-402

403-404

405-406

407-410

411-413

414

414

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