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PROCEEDINGS
of the

ENTOMOLOGICAL SOCIETY

of

W ASHINGTON

Volume 80

OFFICERS FOR THE YEAR 1978

President Douglas W. S. Sutherland
President-elect Donald R. Davis
Recording Secretary Wayne N. Mathis
Corresponding Secretary Donald R. Whitehead
Treasurer F. Christian Thompson
Editor Manya B. Stoetzel
Custodian Sueo Nakahara
Program Committee Chairman Victor E. Adler
Membership Committee Chairwoman Joyce A. Utmar
Delegate to the Washington Academy of Sciences Douglas W. S. Sutherland
Hospitality Chairwoman Helen Sollers-Riedel

Published by the Society
WASHINGTON, D.C.
1978

Table of Contents, Volume 80

ANDERSON, D. M.—See CLARK, W. E.

AUSTIN, D. F.—Morning glory bees and the Ipomoea pandurata complex (Hy-
menoptera: Anthophoridae)

BARROWS, E. M.—See BOYLE, H.
BARTH, S. E.—See HENDRICKSON, R. M., JR.

BEAL, R. S., JR.—Review of the dermestid beetle genus Caccoleptus with de-
scription of a new species from Colombia (Coleoptera)

BERRIOS, A.—See MALDONADO CAPRILES, J.

BOYLE, H. and E. M. BARROWS.—Oviposition and host feeding behavior of
Aphelinus asychis (Hymenoptera: Chalcidoidea: Aphelinidae) on Schizaphis
graminum (Homoptera: Aphididae) and some reactions of aphids to this
parasite

BURKE, H. R.—See CLARK, W. E.
CHARANZA, D. R.—See THOMPSON, P. H.

CLARK, W. E., H. R. BURKE and D. M. ANDERSON.—The genus Tychius
Germar (Coleoptera: Curculionidae); Larvae and pupae of some species, with
evaluation of their characters in phylogenetic analysis

COPELAND, T. P.—A new genus and two new species of Eosentomoidea (Protura;:
Eosentomidae)

COPELAND, T. P. and C. WHITE.—A new species of Eosentomon (Protura:
Eosentomidae) from North Carolina and Tennessee

DEAHL, K. L.—See HILL, A.
DEAHL, kK. L.—See HILL, A.
DEAHL, K. L.—See HILL, A.

DEEMING, J. C.—Two new species of Phlebosotera Duda (Diptera: Asteiidae)
from northern Nigeria

EADS, R. B.—Sandfly distribution in the United States, with a first record for
Colorado (Diptera: Psychodidae)

ELLIOTT, N. B. and F. E. KURCZEWSKI.—Geographic variation in Tachysphex
terminatus (Hymenoptera: Sphecidae, Larrinae)

ENARI, L.—See WALKER, H. G.

FOOTE, B. A.—See ROBINSON, W. H.

GAGNE, R. J.—A new species of Asphondylia (Diptera: Cecidomyiidae) from
Costa Rica with taxonomic notes on related species

GAGNE, R. J.—New synonymy and a review of Haplusia (Diptera: Cecidomyi-
idae)

GONZALEZ, R. H.—A new species of xenocaligonellid mite from the Galapagos
Islands (Acari)

GORDH, G.—Taxonomic notes on Zagrammosoma, a key to the Nearctic species
and descriptions of new species from California (Hymenoptera: Eulophidae)

GREGG, E. L.—See THOMPSON, P. H.

GROGAN, W. L., JR—See WIRTH, W. W.

GROGAN, W. L., JR. and W. W. WIRTH.—Description of the male of Neurobez-
zia granulosa (Wirth) (Diptera: Ceratopogonidae)

li

626
397
441
210

171
380

44]

626

296

626

473

485
317
330
335

51

538

103
575
388
514
517

191

344
296
94

548

HAMBLETON, E. J.—Three new Neotropical Rhizoecus (Homoptera: Pseudo-
coccidae)

HARPER, P. P. and R. F. KIRCHNER.—A new stonefly from West Virginia
(Plecoptera: Chloroperlidae)

HENDRICKSON, R. M., JR. and S. E. BARTH.—Notes on the biology of
Diglyphus intermedius (Hymenoptera: Eulophidae), a parasite of the alfalfa
blotch leafminer, Agromyza frontella (Diptera: Agromyzidae)

HENRY, C. S.—An evolutionary and geographical overview of repagula (abortive
eggs) in the Ascalaphidae (Neuroptera)

HENRY, T. J.—TIwo new Ceratocapsus Reuter 1876, from the eastern United
States (Hemiptera: Miridae)

HENRY, T. J.—Description of a new Polymerus, with notes on two other little
known mirids from the New Jersey Pine-Barrens (Hemiptera: Miridae)

HENRY, T. J. and J. L. HERRING.—A new genus and species of Cardiastethini
from Peru (Hemiptera: Anthocoridae)

HERRING, J. L._—See HENRY, T. J.

HILL, A. and K. L. DEAHL.—Description and life cycle of a new species of
Histiostoma (Acari: Histiostomidae) associated with commercial mushroom
production

HILL, A. and K. L. DEAHL.—Two new species of Tarsonemus (Acari: Tarsonemi-
dae) associated with commercial mushroom production

HILL, A. and K. L. DEAHL.—A new species of Pseudopygmephorus (Acari:
Pygmephoridae) associated with commercial mushroom production

HOLMES, J. W., JR.—See THOMPSON, P. H.

KIRCHNER, R. F.—See HARPER, P. P.

KNUTSON, L. and K. VALLEY.—Biology of a Neotropical snail-killing fly,
Sepedonea isthmi (Diptera: Sciomyzidae)

KORMILEV, N. A.—Two new species of American Aradidae (Hemiptera)

KRAFSUR, E. S.—Aggregations of male screwworm flies, Cochliomyia hominivorax

(Coquerel) in south Texas (Diptera: Calliphoridae)

KRAUTER, P. C—See THOMPSON, P. H.

KURCZEWSKI, F. E.—See ELLIOTT, N. B.

LAMBDIN, P. L.—See MILLER, D. R.

LIN, N.—Defended hunting territories and hunting behavior of females of
Philanthus gibbosus (Hymenoptera: Sphecidae)

MAIER, C. T.—The immature stages and biology of Mallota posticata (Fabricius)
(Diptera: Syrphidae)

MALDONADO CAPRILES, J. and A. BERRIOS.—The allotype of Arenasella
maldonadoi and change of depository for two of Fennah’s holotypes (Homop-
tera: Tropiduchidae and Issidae)

MARSH, P. M.—The braconid parasites (Hymenoptera) of Heliothis species
(Lepidoptera: Noctuidae)

MASON, W. R. M.—A new genus, species and family of Hymenoptera (Ichneu-
monoidea) from Chile

MATHIS, W. N.—A synopsis of Neotropical Eleleides Cresson (Diptera: Ephydri-
dae)

ili

156

403

MATHIS, W. N. and W. W. WIRTH.—A new genus near Canaceoides Cresson,
three new species and notes on their classification (Diptera: Canacidae)

MILLER, D. R. and P. L. LAMBDIN.—A new genus and two new species of
asterolecaniid scale insects on palm from Colombia and Trinidad (Homoptera:
Coccoidea: Asterolecaniidae)

MOCKFORD, E. L.—New species, records and key to Texas Liposcelidae
(Psocoptera)

MUESEBECK, C. F. W.—A new egg parasite (Hymenoptera: Scelionidae) of the
elm spanworm, Ennomos subsignarius (Lepidoptera: Geometridae)

NORTON, R. A.—Notes on Nathan Banks’ species of the mite genus Carabodes
(Acari: Oribatei)

PITKIN, B. R.—Lectotype designations of certain species of thrips described
by J. D. Hood and notes on his collection (Thysanoptera)

POLHEMUS, J. T.—See SMITH, C. L.

RATANAWORABHAN, N. C.—See WIRTH, W. W.

ROBINSON, W. H—Terminalia of some North American species of Megaselia
(Aphiochaeta) and descriptions of two new species (Diptera: Phoridae)

ROBINSON, W. H and B. A. FOOTE.—Biology and immature stages of
Antichaeta borealis (Diptera: Sciomyzidae), a predator of snail eggs

ROGERS, C. E., T. E. THOMPSON and M. B. STOETZEL.—Aphids of sun-
flower: Distribution and hosts in North America (Homoptera: Aphididae)

SABROSKY, C. W.—Tachinid parasites of Heliothis in the Western Hemisphere
(Diptera: Lepidoptera)

SAUSEDA, R. A.—See THOMPSON, P. H.

SCARBROUGH, A. G.—Ethology of Cerotainia albipilosa Curran (Diptera:
Asilidae) in Maryland: Predatory behavior

SCARBROUGH, A. G.—Ethology of Cerotainia albipilosa Curran (Diptera:
Asilidae) in Maryland: Courtship, mating and oviposition

SIRIVANAKARN, S. and G. B. WHITE.—Neotype designation of Culex quin-
quefasciatus Say (Diptera: Culicidae)

SKILES, D. D.—Taxonomy and description of two prionine Cerambycidae from
southern Arizona: A new species of Stenodontes and new status for Neomallodon
arizonicus (Coleoptera)

SMITH, C. L. and J. T. POLHEMUS.—The Veliidae (Heteroptera) of America
north of Mexico—Keys and check list

STEYSKAL, G. C.—What is systematic entomology?

STEYSKAL, G. C.—Synopsis of the North American Pyrgotidae (Diptera)

STEYSKAL, G. C.—A new species of Hylemya from Wyoming (Diptera: An-
thomyiidae)

STOETZEL, M. B.—See ROGERS, C. E.

STOETZEL, M. B.—See WALKER, H. G.

STOLTZFUS, W. B.—Life history and descriptions of the immature stages of
Jamesomyia geminata (Diptera: Tephritidae)

THOMPSON, P. H.—Parasitism of adult Tabanus subsimilis Bellardi (Diptera:
Tabanidae) by a miltogrammine sarcophagid (Diptera: Sarcophagidae)

iv

524

THOMPSON, P. H.—An infestation of miltogrammine Sarcophagidae (Diptera:

Sarcophagidae) in a population of Hybomitra lasiophthalma (Macquart) (Diptera:
Tabanidae)

THOMPSON, P. H., E. J. GREGG, D. R. CHARANZA, R. A. SAUSEDA and
J. W. HOLMES, JR.—Habitats of larval Tabanidae (Diptera) in south Texas

THOMPSON, P. H. and P. C. KRAUTER.—Rearing of Texas Tabanidae ( Diptera).
I. Collection, feeding, and maintenance of coastal marsh species

THOMPSON, T. E.—See ROGERS, C. E.

TODD, E. L.—A checklist of species of Heliothis Ochsenheimer (Lepidoptera:
Noctuidae)

VALLEY, K.—See KNUTSON, L.

WALKER, H. G., M. B. STOETZEL and L. ENARI.—Additional aphid-host re-
lationships at the Los Angeles State and County Arboretum (Homoptera:
Aphididae)

WHITE, C.—See COPELAND, T. P.

WHITE, G. B.—See SIRIVANAKARN, S.

WIRTH, W. W.—See MATHIS, W. N.

WIRTH, W. W.—See GROGAN, W. L., JR.

WIRTH, W. W. and W. L. GROGAN, JR.—Notes on the systematics and biology
of the biting midge, Forcipomyia elegantula Malloch (Diptera: Ceratopogonidae)

WIRTH, W. W. and N. C. RATANAWORABHAN.—Studies on the genus
Forcipomyia. V. Key to subgenera and description of a new subgenus related
to Euprojoannisia Bréethes (Diptera: Ceratopogonidae)

Notes

BOUSEMAN, J. K.—Oxaea austera Gerstaecker in Bolivia, with a new host record
(Hymenoptera: Apoidea)

DENNIS, D. S. and J. A. GOWEN.—A “nocturnal” foraging record for Diogmites
neoternatus (Diptera: Asilidae)

FROESCHNER, R. C.—Cylindrocnema plana Mayr, 1865, a senior synonym
of Nopalis crassicornis (Reed), 1898 (Hemiptera: Acanthosomatidae)

GOWEN, J. A.—See DENNIS, D. S.

GURNEY, A. B.—William Dwight Pierce, biographical notes and a review of
his book The Deadly Triangle

HALL, R. D.—See TOWNSEND, L. H., JR.

HARDY, D. E.—Replacement name for Drosophila nigricolor Hardy (Diptera:
Drosophilidae)

IRWIN, M. E.—Holotype deposition of Henicomyia bicolor

LAVIGNE, R. J.—A case of homonymy in the genus Machimus (Diptera: Asilidae)

LEWIS, P. A.—On the use of pectinate maxillary spines to separate Stenonema
and Stenacron (Ephemeroptera: Heptageniidae)

NAKAHARA, S.—Ceroplastes denudatus, junior synonym of C. rusci (Homop-
tera: Coccoidea: Coccidae)

PERRY, D. R.—Paratropes bilunata (Orthoptera: Blattidae): An outcrossing
pollinator in a Neotropical wet forest canopy?

PRICE, R. D.—A new name for Colpocephalum abbotti Price (Mallophaga:
Menoponidae)

TODD, E. L.—New synonymy in Nerthra Say (Hemiptera: Gelastocoridae)

TODD, E. L.—New distributional records for two species of Nerthra Say from
Mexico (Hemiptera: Gelastocoridae)

TOWNSEND, L. H., JR., R. D. HALL and E. C. TURNER, JR.—Human oral
myiasis in Virginia caused by Gasterophilus intestinalis (Diptera: Gasterophilidae)

TURNER, E. C., JR—See TOWNSEND, L. H., JR.

Vi

128

313

Book Reviews

FOOTE, R. H.—Controlling Fruit Flies by the Sterile-Insect Technique

GORDH, G.—Mechanical Design in Organisms

GORDH, G.—A Revision of Western Nearctic Species of Torymus Dalmon
(Hymenoptera: Torymidae)

GORDH, G.—A Bibliography of Quantitative Ecology

GORDH, G.—Insect Flight

GURNEY, A. B.—The Windows to His World. The Story of Trevor Kincaid

GURNEY, A. B.—Theodore D. A. Cockerell. Letters from West Cliff, Colorado

GURNEY, A. B.—Entomofauna Cubana, Tomo III. Subclase Polyneoptera

KNUTSON, L.—The Tachydromiinae (Diptera: Empididae) of Fennoscandia and
Denmark. Vol. 3, Fauna Entomologica Scandinavica

KRAMER, J. P.—Taxonomic Study of the Cicadellinae (Homoptera: Cicadellidae).
Part 2. New World Cicadellini and the Genus Cicadella

Obituary

The Rev. Thomas Borgmeier—W. W. WIRTH, W. H ROBINSON and W. W.
KEMPF

Vii

134
133

138
315
659
135
136
658

139

456

141

Index to New Taxa, Volume 80

adetocorymbus Miller and Lambdin (Gram-
mococcus), 243

aicen Mathis and Wirth (Paracanace), 533

alleni Mockford (Belaphotroctes), 558

Apozyx Mason, 606

aracoma Harper and Kirchner (Alloperla),
403

aurantiacus Henry (Ceratocapsus), 385

Boreothrinax Steyskal, 151

corymbus Miller and Lambdin (Grammo-
coccus), 257

decora Robinson (Megaselia (Aphiochaeta)),
219

dichaetus Steyskal (Boreothrinax), 152

divaricatus Hambleton (Rhizoecus), 157

Dolostethus Henry and Herring, 520

droozi Muesebeck (Telenomus), 91

enterolobii Gagné (Asphondylia), 515

erwini Copeland (Eosentomon), 479

evansi Steyskal (Hylemya (Delia)), 553

galapagus Gonzalez (Xenocaligonellidus),
193

Grammococcus Miller and Lambdin, 240

heinemanni Hill and Deahl (Histiostoma),
318

hirsutoides Mockford (Liposcelis), 562

hoguei Mathis and Wirth (Paracanace), 527

indica Marsh (Microplitis), 29

intermedium Gordh (Zagrammosoma), 348

inuwa Deeming (Phlebosotera), 54

laffooni Robinson (Aphio-
chaeta)), 220

lebam Mathis and Wirth (Paracanace), 530

(Megaselia

Vili

leei Wirth and Ratanaworabhan (Forcip-
omyia (Saliohzlea)), 498

lukoschusi Hill and Deahl (Tarsonemus),
332

madericus Skiles (Stenodontes), 414

melinum Gordh (Zagrammosoma), 350

mercedesae Hill and Deahl (Tarsonemus),
330

olmuensis Hambleton (Rhizoecus), 159

oregonicus Kormilev (Aradus), 229

ornatus Mockford (Liposcelis), 565

pallidus Mockford (Liposcelis), 567

Paracanace Mathis and Wirth, 524

penai Mathis (Eleleides), 466

penyai Mason (Apozyx), 606

pseudoyosemitense White (Eosentomon),
485
pubescens Henry and Herring (Dolo-

stethus), 522
rostratus Henry (Polymerus), 543
sabroskyi Deeming (Phlebosotera), 51
Saliohelea Wirth and Ratanaworabhan
(Forcipomyia), 494
shewelli Steyskal (Boreothrinax), 153
smileyi Hill and Deahl (Pseudopygme-
phorus), 335
spinosus Henry (Ceratocapsus), 383
stami Wirth and Ratanaworabhan (For-
cipomyia (Saliohelea)), 502
Styletoentomon Copeland, 473
styletum Copeland (Styletoentomon), 473
usingeri Kormiley (Aradus), 230
variabilis Hambleton (Rhizoecus), 161
wicki Beal (Caccoleptus), 174

eZ
a7
VOL. 80 JANUARY 1978 NO. 1

=

PROCEEDINGS.:s 7

of the

ENTOMOLOGICAL SOCIETY
of WASHINGTON

DEPARTMENT OF ENTOMOLOGY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY

CONTENTS

Two new species of Phlebosotera Duda (Diptera: Asteiidae) from northern
Nigeria ; J. C. DEEMING 51

Geographic variation in Tachysphex terminatus (Hymenoptera: Sphecidae,
Larrinae ) N. B. ELLIOTT and FRANK E. KURCZEWSKI 103

An evolutionary and geographical overview of repagula (abortive eggs) in the
Ascalaphidae (Neuroptera ) Gh S:, HENRY” 735

The braconid parasites (Hymenoptera) of Heliothis species (Lepidoptera: Noc-
tuidae ) P. M. MARSH 15

A new egg parasite (Hymenoptera: Scelionidae) of the elm spanworm, Ennomos
subsignarius (Lepidoptera: Geometridae ) C. F. W. MUESEBECK 91

A checklist of species of Heliothis Ochsenheimer (Lepidoptera: Noctuidae )
Be iG LOD 1

Tachinid parasites of Heliothis in the Western Hemisphere (Diptera; Lepidoptera )
C. W. SABROSKY 37

Ethology of Cerotainia albipilosa Curran (Diptera: Asilidae) in Maryland:
Predatory behavior A. G. SCARBROUGH 113

The Veliidae (Heteroptera) of America north of Mexico—Keys and check list
C. L. SMITH and J. T: POLHEMUS 56

What is systematic entomology? CrlCISTEYSKAL, 743

(Continued on back cover)

ENTOMOLOGICAL SOCIETY

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OFFICERS FOR 1978

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Publications Committee
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PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 1-14
A CHECKLIST OF SPECIES OF HELIOTHIS OCHSENHEIMER
(LEPIDOPTERA: NOCTUIDAE)

Ei. Todd

Abstract.—This cross-indexed list of 154 species-group names in Heliothis
Ochsenheimer (Lepidoptera: Noctuidae) includes the original combination
and reference of each name, distribution and indications of references con-
taining illustrations of valid species and subspecies. Heliothis neckerensis
n. n. is proposed for Helicoverpa pallida Hardwick, secondary homonym of
Heliothis pallida (Butler). Twelve new combinations are noted.

This cross-indexed alphabetical checklist of the species-group names of
the genus Heliothis Ochsenheimer has been developed from the literature
and does not reflect a personal opinion as to the placement, validity or status
of the entities included except as noted. The list has been prepared primarily
for use by workers concerned with studies of the economically important
species of the genus, especially with biological control of such species.

Hardwick (1965a:10; 1970:30) has been followed in the inclusion in
Heliothis of certain species formerly in Timora Walker, 1856 and Canthylidia
Butler, 1886. Other species that have been included in those genera and
other related genera remain to be studied to determine whether they are
congeneric. Undoubtedly a supplementary list will need to be prepared at
some future date.

Hardwick (1970:18) feels that if the corn earworm group (Helicoverpa
Hardwick, 1965) is included in Heliothis, the species of Schinia Hubner,
1818 should also be transferred to Heliothis. That premise does not hold
if the characters on which his classification was based are given a different
weight or significance. Accordingly, Helicoverpa Hardwick is treated as a
synonym of Heliothis, and the species of Schinia have not been included in
this list.

A few species have been either deliberately excluded or tentatively in-
cluded. Noctua scutosa Denis & Schiffermiller, 1775, a species commonly
placed in Heliothis, but now standing in Protoschinia Hardwick, 1970 has
not been included. Hardwick believes that it is an intermediate species be-
tween Heliothis and Schinia. Conversely, one included species, Chloridea
chilensis Hampson, 1903 may not belong in the genus. It has been placed
in the genus Schinia by Hardwick (1966:870), but I am not convinced
his arguments are valid. Poaphila? turbata Walker, 1858 and its junior
synonym, Perigea albidentina Walker, 1865 have usually been placed in
Heliothis, but the former has been treated in recent years as an unrecognized
or “lost” species. Mr. Alan Hayes of the British Museum (Natural History)

2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

has examined and has sent colored photographic slides of the types of those
two names for my examination. He is of the opinion that these are older
names for the species described by Grote, 1875 as Heliothis lupatus. I agree
with that opinion and the names are so treated in this list. Chloridea dis-
tincta Schaus, Aspila tergemina Felder and Rogenhofer and Chloridea
molochitina Berg with its synonym, Thyreion olivofusa Dognin, have usually
been placed as junior synonyms of Heliothis virescens (F.) by most workers.
I am convinced they are good species and have treated them as such in the
checklist.

The generic name, Heliothis, is masculine in gender, but has usually been
treated as feminine. In addition, names proposed in feminine genera have
been transferred to Heliothis unchanged. All necessary changes to masculine
endings have been made in this paper, and the incorrect feminine spellings
are shown in parentheses for convenience of workers wishing to conduct
bibliographic searches.

In the past some species of the corn earworm complex have been misiden-
tified as Heliothis armigera Hiibner or Heliothis obsoleta (F.). These nu-
merous misidentifications have not been included in this list as they can be
found in the extensive bibliography prepared by Hardwick (1965a:133-
208).

The numbers in parentheses and brackets following each species listed
refer to the numbered list of references. The numbers in parentheses in-
dicate references where illustrations of adults may be found; those in
brackets refer to illustrations of genitalia, male and/or female.

Genus Heliothis Ochsenheimer
Heliothis Ochsenheimer, 1816. Schmett. Eur. 4: 91.

Type-species: Phalaena dipsacea Linnaeus, 1767. Syst. Nat. (Edn. 12),
1:856 (= Phalaena viriplaca Hufnagel, 1766. Berlin. Mag. 3(4):406)
by subsequent designation by Samouelle, 1819. Entomologist’s use-
ful Compendium. P. 252.

Chloridea Duncan [& Westwood], 1841. In Jardine, Naturalist’s Library
(Edn. 1). 33(Ent. 7):198.

Type-species: Phalaena rhexiae J. E. Smith, 1797. In Smith & Abbott,
Nat. Hist. Rarer Lepid. Insects Georgia. P. 199 (= Noctua virescens
Fabricius, [1777]. Genera Insect. P. 282) by monotypy.

Aspila Guenée, 1852. In Boisduval & Guenée, Hist. Nat. Insectes. Sp.
Gén. Lépid. 6(Noct. 2):174.

Type-species: Noctua virescens Fabricius, [1777]. Genera Insect. P.
282 by subsequent designation by Hampson, 1903. Cat. Lepid. Phala-
enae Br. Mus. 4:34.

VOLUME 80, NUMBER 1 3

Timora Walker, 1856. List of specimens of lepidopterous insects in the col-
lection of the British Museum. Pt. 9, pp. 69, 132.

Type-species: | Nonagria senegalensis Guenée, 1852. In Boisduval &
Guenée, Hist. Nat. Insectes. Sp. Gén. Lépid. 5(Noct. 1):110 by mono-
typy.

Heliocheilus Grote, 1865. Proc. Entomol. Soc. Phila. 4:328.

Type-species: Heliocheilus paradoxus Grote, 1865. Ibidem. 4:329 by
monotypy.

Rhodosea Grote, 1883. Can. Entomol. 15:4.
Type-species: Rhodosea julia Grote, 1883. Ibidem. 15:5 by monotypy.
Dysocnemis Grote, 1883. Trans. Amer. Entomol. Soc. 10:263.

Type-species: Melicleptria belladonna Hy. Edwards, 1881. Papilio. 1:20
by original designation.

Canthylidia Butler, 1886. Trans. R. Entomol. Soc. Lond. 1886, p. 406.

Type-species: Canthylidia pallida Butler, 1886. Ibidem. 1886, p. 406
(= Leucania invaria Walker, 1856. List of specimens of lepidoterous
insects in the collection of the British Museum. Pt. 9, p. 111) by
monotypy.

Helicoverpa Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40, p. 9.

Type-species: Noctua armigera Hubner, [1803-1808]. Samml. Eur.
Schmett. 4: pl. 79, fig. 370 by original designation.

acesias Felder & Rogenhofer—Jr. syn. of phloxiphagus.
adauctus (adaucta) Butler—Jr. syn. of viriplaea.
afer (afra) (Hardwick )—Subsp. of assulta. Africa.
albidus (albida) Cockayne—Ab. of maritimus warnecki.
albidus (albida) Fuchs—Form of viriplaea.
albidentina (Walker )—Jr. syn. of turbatus.
albivenatus (albivenata) (Hampson). Somaliland (8, 11).
Chloridea albivenata Hampson, 1916. Proc. Zool. Soc. Lond. 1916, p. 103,
pls ls figs 3:
alphea (Cramer )—Jr. syn. of peltiger.
anartoides (Strand)—Jr. syn. of belladona.
angarensis (Draudt )—Subsp. of viriplaea. China.
aresca (Turner)—Jr. syn. of rubrescens.
armiger (armigera) armiger (Hiibner). Eastern Hemisphere (3, 9, 12, 21,
22, 25, 26) [3, 12, 15, 25].
Noctua armigera Hubner [1803-1808]. Samml. Eur. Schmett. 4: pl. 79,
fig. 370.

4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Noctua barbara Fabricius, 1794. Entomol. Syst. 3, 2:111. Hardwick
(1965b, ref. 13) considered this a senior synonym, but applied to In-
terational Commission on Zoological Nomenclature for rejection as a
nomen oblitum.

Heliothis pulverosa Walker, 1857. List of specimens of lepidopterous in-
sects in the collection of the British Museum. Pt. 11, p. 688.

Heliothis uniformis Wallengren, 1860. Weiner Entomol. Monatschr.
aswell

Heliothis obsoleta Auctorum nec (F.) (in part). Misidentifications.

Chloridea obsoleta ab. rufa Warren, 1911. In Seitz, Die Gross-Schmetter-
linge der Erde. 3:246.

Heliothis armigera form fusca Cockerell, 1889. Entomologist. 22:4.

armiger commoni (Hardwick) n. comb. Canton Island.

Helicoverpa armigera commoni Hardwick, 1965. Mem. Entomol. Soc.
Can. No. 40, p. 101, figs. 116-117.

armiger confertus (conferta) Walker. Indonesia, Australia & Oceania.

Heliothis conferta Walker, 1857. List of specimens of lepidopterous in-

sects in the collection of the British Museum. Pt. 11, p. 690.
assulta assulta Guenée. Asia, Africa, Australia & Oceania (3, 8, 9, 12,

22 97) [3, 12].

Heliothis assulta Guenée, 1852. In Boisduval & Guenée, Hist. Nat. Insectes.
Sp. Gen. Lépid. 6(Noct. 2):178.

Heliothis separata Walker, 1857. List of specimens of lepidopterous insects
in the collection of the British Museum. Pt. 11, p. 691.

Heliothis succinea Moore, 1881. Proc. Zool. Soc. Lond. 1881, p. 362.

Heliothis temperata Walker, 1857. List of the specimens of lepidopterous
insects in the collection of the British Museum. Pt. 11, p. 689.

assulta afer (afra) (Hardwick) n. comb. Africa.

Helicoverpa assulta afra Hardwick, 1965. Mem. Entomol. Soc. Can. No.

A0M p13. he. 138
atacamae (Hardwick) n. comb. Chile & Peri (12) [12].
Helicoverpa atacamae Hardwick, 1965. Mem. Entomol. Soc. Can. No.
40, p. 81, figs. 106-107.
aurantiacus (aurantiaca) (Strand )—Ab. of peltiger.
barbarus (barbara) (F.)—Jr. syn. of peltiger vide Hampson, syn. of
armiger vide Hardwick.
belladona (Hy. Edwards). Western USA (9, 16) [15].
Melicleptria belladona Hy. Edwards, 1881. Papilio. 1:20.
Dysocnemis anartoides Strand, 1914. Arch. Naturgesch. 80A, 2:161.
borealis (Hampson). Eastern Canada (9).

Dysocnemis borealis Hampson, 1903. Cat. Lepid. Phalaenae Br. Mus.

4:24, pl. 55, fig. 6.
bracteae (Hardwick) n. comb. Paraguay, Argentina & Pert (12) [12].

VOLUME 80, NUMBER 1 5

Helicoverpa bracteae Hardwick, 1965. Mem. Entomol. Soc. Can. No.
40, p. 74, figs. 27-29, 102.
bulgaricus (bulgarica) (Draudt)—Subsp. of maritimus. SE Europe.
caesius (caesia) (Warren). Queensland (27).
Chloridea caesia Warren, 1926. In Seitz, Die Gross-Schmetterlinge der
Erde. 11:308, pl. 27, row i.
canariensis (Warren )—Ab. of viriplaea.
centralasiae (Draudt)—Subsp. of maritimus. Asia.
charmione (Stoll)—Jr. syn. of peltiger.
chilensis (Hampson). Chile (4, 9, 14) [14]. In Schinia vide Hardwick,
1966.
Chloridea chilensis Hampson, 1903. Cat. Lepid. Phalaenae Br. Mus. 4:48,
pl. 55, fig. 24.
clarissimus (clarissima) (Turati)—Form of peltiger.
commoni (Hardwick )—Subsp. of armiger. Canton Island.
condolens Schawerda—Ab. of peltiger.
confertus (conferta) Walker—Subsp. of armiger. Indonesia, Australia &
Oceania.
confusus (confusa) (Hardwick) n. comb. Hawaii (12) [12].
Helicoverpa confusa Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,
p. 116, figs. 123-124.
conifer (conifera) (Hampson). Transvaal.
Chloridea conifera Hampson, 1913. Ann. Mag. Nat. Hist. (8). 12:580.
cystiphorus (cystiphora) (Wallengren). Panama & Galapagos Islands (4).
Anthoecia cystiphora Wallengren, 1860. Wiener Entomol. Monatschr.
4:172 ¢.
Anthoecia inflata Wallengren, 1860. Ibidem. 4:172 ¢.
Anthoecia onca Wallengren, 1860. Ibidem. 4:172 °.
dejeani Oberthiir. China (18, 26).
Heliothis dejeani Oberthiir, 1893. Et. Entomol. 18:44, pl. 3, fig 40.
desertus (deserta) (Sohn-Rethel)—Var. of nubiger.
designatus (designata) (Brandt). Iran (2).
Heliocheilus designata Brandt, 1947. Mitt. Miinchn. Entomol. Ges. 31:
853.
dipsaceus (dipsacea) (L.)—Jr. syn. of viriplaea.
distinectus (distincta) (Schaus). Peru (4).
Chloridea distincta Schaus, 1898. J. N.Y. Entomol. Soc. 6:117.
errans Walker—Jr. syn. of seutuliger.
ferrugineus (ferruginea) Spuler—Ab. of maritimus.
fervens Butler. Japan & China (9, 22).
Heliothis fervens Butler, 1881. Trans. R. Entomol. Soc. Lond. 1881,
p. 186.
Heliothis foveolatus Staudinger, 1888. Stett. Entomol. Zeit. p. 263.

6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Heliocheilus fervens ab. macularis Warren, 1911. In Seitz, Die Gross-
Schmetterlinge der Erde. 3:245.
fimbria (Williams )—Jr. syn. of proruptus.
flavescens (Janse). Natal.
Chloridea flavescens Janse, 1917. Ann. Durban Mus. 1:472.
flaviger (flavigera) flaviger (Hampson). Africa (25) [25].
Chloridea flavigera Hampson, 1907. Ann. Mag. Nat. Hist. (7). 19:242.
flaviger insularis Viette. Madagascar.
Heliothis flavigera insularis Viette, 1967. Faune de Madagascar. 20(2):772,
figs. 603, 605, 611.
fletcheri (Hardwick) n. comb. Western Africa (12) [12].
Helicoverpa fletcheri Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,
p. 129, figs. 132-133.
florentinus (florentina) (Esper)—Jr. syn. of peltiger.
foveolatus Staudinger—Jr. syn. of fervens.
fumatus (fumata) (Lucas)—Jr. syn. of invarius.
fuscimacula (Janse). Natal.
Chloridea fuscimacula Janse, 1917. Ann. Durban Mus. 1:469.
fuscus (fusca) Cockerell—Form of armiger.
gelotopoeus (gelotopoeon) (Dyar). Southern South America (4, 12)
e223):
Thyreion gelotopoeon Dyar, 1921. Insec. Inscit. Menst. 9:60.
PThyreion giacomelli Koehler, 1939. Physis. 17:451.
PThyreion lizeri Koehler, 1939. Physis. 17:450.
giacomelli (Koehler )—Jr. syn. of gelotopoeus.
grandis (Druce)—Jr. syn. of julia.
guidellii Costantini. Italy.
Heliothis guidellii Costantini, 1922. Neue Breitrage Syst. Insektenk. 2:99.
hawaiiensis Quaintance & Brues. Hawaii (12, 28, 29) [12, 28, 29].
Heliothis obsoleta var. hawaiiensis Quaintance & Brues, 1905. USDA,
Bur. Entomol. Bull. 50:12.
Chloridea obsoleta signata Warren, 1926. In Seitz, Die Gross-Schmetter-
linge der Erde. 11:308.
Chloridea armigera ab. hawaiiensis (Strand), 1916. Arch. Naturgesch.
81(A), 12:143. Preoccupied.
hawaiiensis (Strand)—Jr. syn. of hawaiiensis Quaintance & Brues. Pre-
occupied.
helenae (Hardwick) n. comb. St. Helena Island (12) [12].
Helicoverpa helenae Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40, p.
106, figs. 118-119.
hungaricus (hungarica) (Kovacs )—Jr. syn. of maritimus bulgaricus.
hyalostictus (hyalosticta) Hampson. India & Burma (9, 27).
Heliothis hyalosticta Hampson, 1896. Moths of India. 4:507.

VOLUME 80, NUMBER 1 i

hyperchroa Turner—Jr. syn. of rubrescens.
hyperfuscus (hyperfusca) (Strand)—Ab. of paradoxus.
inflatus (inflata) (Wallengren)—Jr. syn. of eystiphorus.
insularis Viette—Subsp. of flaviger. Madagascar.
insulatus (insulata) (Navas )—Var. of peltiger.
intensivus (intensiva) (Warren)—Ab. of ononis.
interjacens Grote—Jr. syn. of phloxiphagus.
invarius (invaria) invarius (Walker). Queensland (9) [15].
Leucania invaria Walker, 1856. List of the specimens of lepidopterous in-
sects in the collection of the British Museum. Pt. 9, p. 111.
Canthylidia pallida Butler, 1886. Trans. R. Entomol. Soc. Lond. 1886,
p. 406, pl. 9, fig. 9.
Leucania fumata Lucas, 1889. Proc. Linn. Soc. N. S. Wales. (2). 4:1098.
invarius pallescens (Warren). West Australia.
Canthylidia invaria pallescens Warren, 1926. In Seitz, Die Gross-Schmet-
terlinge der Erde. 11:312, pl. 28, row a.
jefferyi (Janse). Transvaal.
Chloridea jefferyi Janse, 1917. Ann. Durban Mus. 1:470.
julia (Grote). Western USA & México (9, 16) [15].
Rhodosea julia Grote, 1883. Can. Entomol. 15:5.
Adisura grandis Druce, 1890. Proc. Zool. Soc. Lond. 1890, p. 516.
lizeri (Koehler)—Jr. syn. of gelotopoeus.
lugubris Klemensiewicz—Ab. of ononis.
lupatus (lupata) Grote—Jr. syn. of turbatus.
luteitinctus (luteitincta) Grote—Form of phloxiphagus.
macularis (Warren)—Ab. of fervens.
maritimus (maritima) maritimus Graslin. Palearctic (7) [7].
Heliothis maritima Graslin, 1855. Ann. Soc. Entomol. Fr. (3). 3:68, pl. 7.
Heliothis spergulariae Lederer, 1857. Noct. Eur. p. 230.
Heliothis maritima ab. ferruginea Spuler, 1908. Schmett. Eur. 1:281.
Heliothis maritima form obscura Lempke, 1941. Tijdschr. Entomol. 84:
BOT.
maritimus bulgaricus (Draudt). Southeast Europe.
Chloridea maritima bulgarica Draudt, 1938. Entomol. Rdsch. 55:307 and
1938. Rev. Frang. Lepidopt. 9:90.
Chloridea maritima hungarica Kovacs, 1950. Folia Entomol. Hung. (N. S.).
3:68.
maritimus centralasiae (Draudt). Asia.
Chloridea maritima centralasiae Draudt, 1938. Entomol. Rdsch. 55:308
and 1938. Rev. Frang. Lepidopt. 9:90.
maritimus warneckei (Boursin). Europe.
Chloridea maritima warneckei Boursin, 1963. Linn. Belg. 2:125 and
1964. Bull. Mens. Soc. Linn. Lyon. 33:240.

8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Heliothis maritima septentrionalis Hoffmeyer, 1938. Flora og Fauna.
44:6. Preoccupied.
Heliothis maritima septentrionalis ab. albida Cockayne, 1951. Entomol.
Rec. 63:160.
marmada (Swinhoe)—Jr. syn. of punctiger.
mekranus (mekrana) (Brandt). Iran (2).
Heliocheilus mekrana Brandt, 1947. Mitt. Miinchn. Entomol. Ges. 31:852.
metachriseus (metachrisea) (Hampson). Madagascar (25) [25].
Chloridea metachrisea Hampson, 1903. Cat. Lepid. Phalaenae Br. Mus. 4:
42, pl. 55, fig. 18.
minutus (minuta) (Hardwick) n. comb. Lisianski Island (12) [12].
Helicoverpa minuta Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,
p. 118, figs. 125-126.
molochitinus (molochitina) (Berg). Southern South America.
Chloridea molochitina Berg, 1882. Ann. Soc. Argentina. 14:282.
Thyreion olivofusa Dognin, 1907. Ann. Soc. Entomol. Belg. 51:230.
nana (Hampson). Tibet (10).
Chloridea nana Hampson, 1906. Proc. Zool. Soc. Lond. 1906, p. 487.
neckerensis n. n. for Helicoverpa pallida Hardwick. Necker Island
(12) EAE
Helicoverpa pallida Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,
p. 89, fig. 111. Preoccupied by Heliothis pallida (Butler), 1886.
neckerensis nihoaensis (Hardwick) n. eomb. Nihoa Island (14).
Helicoverpa pallida nihoaensis Hardwick, 1966. Can. Entomol. 98(8):
S67, figs 2:
nihoaensis (Hardwick )—Subsp. of neckerensis. Nihoa Island.
nubiger (nubigera) Herrich-Schaffer. Palearctic (7, 26).
Heliothis nubigera Herrich-Schiffer, 1851. Syst. Bearbeit. Schmett. Eur.
2:366.
Heliothis perigeoides Moore, 1881. Proc. Zool. Soc. Lond. 1881, p. 361.
Chloridea nubigera var. deserta Sohn-Rethel, 1929. Iris. 43:14.
obscurus (obscura) Lempke—Form of maritimus.
obscurus (obscura) (Lempke )—Form of peltiger.
obsoletus (obsoleta) (F.)—Syn. of zea.
ochraceus (ochracea) Cockerell—Form of zea.
oenotryx (Boursin)—Subsp. of viriplaea. Spain.
olivarius (olivaria) Graeser. Amurland.
Heliothis olivaria Graeser, 1888. Berlinger Entomol. Ztschr. 32:365.
olivofusus (olivofusa) (Dognin)—Jr. syn. of molochitinus.
omicronatus (omicronata) Richardson—Ab. of peltiger.
onca (Wallengren)—Jr. syn. of eystiphorus.
ononidis Guenée—Jr. syn. of ononis.
ononis (Denis & Schiffermiiller). Holarctic (4, 7, 18, 21, 26).

VOLUME 80, NUMBER 1 9

Noctua ononis Denis & Schiffermiiller, 1775. Ankiindung eines syste-
matischen Werkes von den Schmetterlingen der Weinergregend . . .
Wien. P. 89.

Chloridea ononis ab. intensiva Warren, 1911. In Seitz, Die Gross-Schmet-
terlinge der Erde. 3:245, pl. 50, row k.

Melicleptria septentrionalis Hy. Edwards, 1884. Papilio. 4:45.

Heliothis ononis ab. lugubris Klemensiewicz, 1912. Spraw. Kom. Fizyogr.
Krakow. 46:14.

Heliothis ononidis Guenée, 1852. In Boisduval & Guenée, Hist. Nat.
Insectes. Sp. Gén. Lépid. 6(Noct. 2):182. An invalid emendation of
ononis.

oregonicus (oregonica) (Hy. Edwards). Western USA (1, 9, 16).

Melicleptria oregonica Hy. Edwards, 1875. Proc. Calif. Acad. Sci. 6:

135.
pacifieus (pacifica) (Hardwick) n. eomb. Jarvis Island (12) [12].

Helicoverpa pacifica Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,

psHllOy fie 128:
pallescens (Warren)—Subsp. of invarius. West Australia.
pallidus (pallida) (Butler )—Jr. syn. of invarius.
pallidus (pallida) (Hardwick)—Syn. of neckerensis. Preoccupied.
pallidus (pallida) (Lempke )—Form of viriplaca.
pallidus (pallida) Cockerell—Form of peltiger.
paradoxus (paradoxa) (Grote). USA (4, 9, 16, 17) [15].

Heliocheilus paradoxus Grote, 1865. Proc. Entomol. Soc. Phila. 4:329, pl.
2, figs. 3-5.

Chloridea paradoxa ab. hyperfusca Strand, 1916. Arch. Naturgesch.
81(A), 12:143.

pauliana Hardwick—Jr. syn. of pauliani.
pauliani Viette. Amsterdam Island & Indian Ocean (24).

Heliothis pauliani Viette, 1959. Bull. Soc. Entomol. Fr. 64(2):27.

Heliothis pauliana [sic], Hardwick, 1966. Can. Entomol. 98(8):869.
Lapsus pro pauliani.

peltiger (peltigera) (Denis & Schiffermiiller). Palearctic (7, 21, 26).

Noctua peltigera Denis & Schiffermiller, 1775. Ankindung eines syste-
matischen Werkes von den Schmetterlingen der Wienergend . . .
Wien. P. 89.

Phal. [aena] Bomb. [yx] alphea Cramer, 1780. Papillons Exotiques . . .
3:99, 173 [Index], pl. 250, fig. F.

Phal. |aena] Noct. [ua] charmione Stoll, 1790. Supplement a L’Ouvrage,
intitulé les Papillons Exotiques . . . Pp. 162, 182 [Index], pl. 36, fig.
10:

Noctua florentina Esper, 1788. Die Schmetterlinge in Abbildungen nach
der Natur. Erlangen. 4: pl. 135, fig. 2.

10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Phalaena straminea Donovon, 1793. Brit. Insects. 2: pl. 61.
Noctua barbara Fabricius, 1794. Entomol. Syst. (3). 2:334.
Heliothis peltigera [f.] pallida Cockerell, 1889. Entomologist. 22:4.
Heliothis peltigera ab. condolens Schawerda, 1914. Verh. Zool.-Bot. Ges.
Wien. 64:365.
Chloridea peltigera ab. aurantiaca Strand, 1915. Arch. Naturgesch. 81A,
12:142.
Chloridea peltigera var. insulata Navas, 1924. Publ. Junta Cien. Nat.
4(10):35.
Chloridea peltigera form clarissima Turati, 1924. Atti Soc. Ital. Sci. Nat.
63:101.
Heliothis peltigera ab. omicronata Richardson, 1958. Entomologist. 91:
212:
Chloridea peltigera form obscura Lempke, 1966. Tijdschr. Entomol. 109
(9):236.
perigeoides Moore—Jr. syn. of nubiger.
phloxiphagus (phloxiphaga) Grote & Robinson. North America (4, 9).
Heliothis phloxiphagus Grote & Robinson, 1867. Trans. Amer. Entomol.
Soc. 1167.
Heliothis acesias Felder & Rogenhofer, 1872. Reise Novara... Atlas,
Inhalts-Verzeichniss Heterocera. P. 11, pl. 108, fig. 42.
Heliothis luteitinctus Grote, 1875. Proc. Acad. Nat. Sci. Phila. P. 426.
Heliothis interjacens Grote, 1880. Bull. Brooklyn Entomol. Soc. 3:30.
Heliothis phlogophagus Grote, 1882. List N. Amer. Moths. P. 36.
phlogophagus Grote—Jr. syn. of phloxiphagus.
posttriphaenus (posttriphaena) (W. Rothschild). Madagascar (25) [25].
Chloridea posttriphaena W. Rothschild, 1924. Ann. Mag. Nat. Hist. (9).
14:310.
prasinus (prasina) (Walker )—Jr. syn. of virescens.
proruptus (prorupta) Grote. Western USA (9).
Heliothis proruptus Grote, 1873. Trans. Amer. Entomol. Soc. 4:294.
Melicleptria fimbria Williams, 1905. Entomol. News. 16:237.
Melicleptria venusta Hy. Edwards, 1875. Proc. Calif. Acad. Sci. 6:133.
pulverosus (pulverosa) Walker—Jr. syn. of armiger.
punctiger (punctigera) Wallengren. Australia (3, 12) [3, 12].
Heliothis punctigera Wallengren, 1860. Wiener Entomol. Montaschr. 4:
Ale
Chloridea marmada Swinhoe, 1918. Ann. Mag. Nat. Hist. (9). 2:65.
pyraloides (Walker )—Jr. syn. of virescens.
rhexiae (J. E. Smith)—Jr. syn. of virescens.
richinii (Berio). Eritrea.
Chloridea richinii Berio, 1939. Boll. Soc. Entomol. Ital. 71:164.
rosario (Barnes )—Jr. syn. of toralis.

VOLUME 80, NUMBER 1 ll

rubrescens (Walker). Australia (3, 27) [3].
Thalpophila rubrescens Walker, 1858. List of the specimens of lepidop-
terous insects in the collection of the British Museum. Pt. 15, p. 1681.
Chloridea aresca Turner, 1911. Ann. Queensland Mus. 10:60.
Heliothis hyperchroa Turner, 1920. Trans. R. Soc. S. Aust. 44:131.
rufus (rufa) Warren. New Guinea (27).
Chloridea rufa Warren, 1926. In Seitz, Die Gross-Schmetterlinge der
Erde. 11:308, pl. 27, row i.
rufus (rufa) (Warren)—Ab. of armiger.
salmantinus (salmantina) (Fernandez )—Form of viriplaca.
scutuliger (scutuligera) Guenée. Africa (8, 9).
Heliothis scutuligera Guenée, 1852. In Boisduval & Guenée, Hist. Nat. In-
sectes, Sp. Gen. Lépid. 6(Noct. 2):180.
Heliothis errans Walker, 1865. List of the specimens of lepidopterous in-
sects in the collection of the British Museum. Pt. 33, p. 769.
senegalensis (Guenée). Senegal.
Nonagria senegalensis Guenée, 1852. In Boisduval & Guenée, Hist. Nat.
Insectes, Sp. Gén. Lépid. 5(Noct. 1):110.
separatus (separata) Walker—Jr. syn. of assulta.
septentrionalis (Hy. Edwards )—Jr. syn. of ononis.
septentrionalis Hoffmeyer—Syn. of maritimus warneckei. Preoccupied.
signatus (signata) (Warren)—Jr. syn. of hawaiiensis.
spectandus (spectanda) Strecker—Jr. syn. of virescens.
spergulariae Lederer—Jr. syn. of maritimus.
stramineus (straminea) (Donovon)—Jr. syn. of peltiger.
stombleri Okumura & Bauer. Western USA (19) [19].
Heliothis stombleri Okumura & Bauer, 1969. Bur. Entomol., Calif. Dept.
Agric. Occ. Pap. No. 18, p. 2, figs. 1-12.
sturmhoefeli Draudt. Brazil (4).
Heliothis sturmhoefeli Draudt, 1927. In Seitz, Die Gross-Schmetterlinge
der Erde. 7:335, pl. 47, row m.
subflexus (subflexa) (Guenée). Western Hemisphere (17) [6].
Aspila subflexa Guenée, 1852. In Boisduval & Guenée, Hist. Nat. In-
sectes, Sp. Gén. Lepid. 6(Noct. 2):175.
succineus (succinea) Moore—Jr. syn. of assulta.
temperatus (temperata) Walker—Jr. syn. of assulta.
tergeminus (tergemina) (Felder & Rogenhofer). South America (5).
Aspila tergemina Felder & Rogenhofer, 1872. Reise Novara . . . Atlas,
Inhalts-Verzeichniss Heterocera. P. 3, fig. 55.
tertius (tertia) Roepke. Java (20).
Heliothis tertia Roepke, 1941[1945]. Zool. Meded. 23:14, pl. 2, fig. 7.
tibetensis (Hardwick) n. comb. Tibet (12) [12].

12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Helicoverpa tibetensis Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,
pelo, fie 36;
titicacae (Hardwick) n. comb. Pert (12) [12].
Helicoverpa titicacae Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,
p. 75, figs. 23-26, 101.
toddi (Hardwick). Africa & Madagascar (12) [12, 25].
Helicoverpa toddi Hardwick, 1965. Mem. Entomol. Soc. Can. No. 40,
p. 128, figs. 133-135.
toralis (Grote). Southwestern USA.
Botis toralis Grote, 1881. Papilio. 1:178.
Prothymia rosario Barnes, 1904. Can. Entomol. 36:264.
Pseudacontia unicolor Walters, 1928. Proc. Entomol. Soc. Wash. 30:
138.
translucens (Felder & Rogenhofer). Himalayas (27).
Heliocheilus translucens Felder & Rogenhofer, 1872. Reise Novara. . .
Atlas, Inhalts-Verzeichniss Heterocera. P. 11, pl. 108, fig. 49.
tristis Stauder—Form of viriplaca.
turbatus (turbata) (Walker). Florida to Maryland (4).
Poaphila? tubata Walker, 1858. List of specimens of lepidopterous in-
sects in the collection of the British Museum. Pt. 15, p. 1834.
Perigea albidentina Walker, 1865. Ibidem. Pt. 32, p. 680.
Heliothis lupatus Grote, 1875. Can. Entomol. 7:224.
umbrosus Grote—Jr. syn. of zea.
unicolor (Walters )—Jr. syn. of toralis.
uniformis Wallengren—Jr. syn. of armiger.
venustus (venusta) (Hy. Edwards )—Jr. syn. of proruptus.
virescens (F.). Western Hemisphere & Hawaii (4, 16, 17, 28) [6, 15, 28].
Noctua virescens Fabricius, 1777. Genera Insect. P. 282.
Phalaena rhexiae J. E. Smith, 1792. In Smith & Abbott, Ins. Georgia.
2:199, pl. 100.
Xanthia prasina Walker, 1856. List of specimens of lepidopterous insects
in the collection of the British Museum. Pt. 10, p. 465.
Xanthia pyraloides Walker, 1856. Ibidem. Pt. 10, p. 466.
Xanthia viridescens Walker, 1856. Ibidem. Pt. 10, p. 465.
Heliothis spectanda Strecker, 1875. Lepid. Rhop. & Het. P. 122.
viridescens (Walker )—Jr. syn. of virescens.
viriplaca viriplaca (Hufnagel). Palearctic (5, 22, 26) [5, 12, 15].
Phalaena viriplaca Hufnagel, 1766. Berlin. Mag. 3(4):406.
Noctua dipsacea Linnaeus, 1767. Syst. Nat. (Edn. 12). 1:856.
Heliothis adaucta Butler, 1878. Ill. Het. Brit. Mus. 3:19, pl. 45, fig. 4.
Chloridea salmantina Fernandez, 1928. Relig. & Cultur. 2:216.

VOLUME 80, NUMBER 1 13

Heliothis dipsacea var. albida Fuchs, 1904. Jahrb. Nass. Ver. Nat.
57:38.
Chloridea dipsacea ab. canariensis Warren, 1911. In Seitz, Die Gross-
Schmetterlinge der Erde. 3:245, pl. 50, row i.
Heliothis dipsacea form tristis Stauder, 1923. Entomol. Anz. 3:44.
Chloridea viriplaca form pallida Lempke, 1966. Tijdschr. Entomol. 109
(9):234.
viriplaca angarensis (Draudt). China.
Chloridea dipsacea angarensis Draudt, 1938. Entomol. Rdsch. 55:372.
viriplaca oenotryx (Boursin). Spain.
Chloridea viriplaca oenotryx Boursin, 1963. Bull. Mens. Soc. Linn. Lyon.
oo: 262:
warneckei (Boursin)—Subsp. of maritimus.
xanthiatus (xanthiata) Walker. Africa (8, 9).
Heliothis xanthiata Walker, 1865. List of specimens of lepidopterous in-
sects in the collection of the British Museum. Pt. 33, p. 770.
zea (Boddie). Western Hemisphere & Hawaii (4, 12, 15, 17, 28, 29) [3,
3 228,29).
Phalaena zea Boddie, 1850. South. Cultivator. 8:132.
Bombyx obsoleta Fabricius, 1793. Entomol. Syst. 3, 2:456.
Heliothis umbrosus Grote, 1862. Proc. Entomol. Soc. Phila. 1:219.
Heliothis armigera [form] ochracea Cockerell, 1889. Entomologist. 22:4.

Literature Cited

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Lepidoptera. Contrib. Nat. Hist. Lepid. N. Amer. 1(4):40, pl. 18, figs. 18-19.

2. Brandt, W. 1941[1947]. Beitrag zur Lepidopteren-Fauna von Iran (3) Neue
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3. Common, I. F. B. 1953. The Australian species of Heliothis (Lepidoptera:
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4. Draudt, M. 1927. In Seitz, Die Gross-Schmetterlinge der Erde. Stuttgart. 7:
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5. Felder, C., and A. F. Rogenhofer. 1872[1875]. In Felder, Felder and Rogenhofer,
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6. Forbes, W. T. M. 1954. Lepidoptera of New York and neighboring states, Pt.
3 Noctuidae. Cornell Univ. Agric. Exp. Sta. Mem. 329. P. 28, figs. 35-36.

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104-105, pl. 11.

9. Hampson, G. F. 1903. Catalogue of the Lepidoptera Phalaenae in the British
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14

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13.

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Todd, E. L. 1955. The distribution and nomenclature of the corn earworm.
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PROC. ENTOMOL. SOC. WASH.

80(1), 1978, pp. 15-36
THE BRACONID PARASITES (HYMENOPTERA) OF
HELIOTHIS SPECIES (LEPIDOPTERA: NOCTUIDAE)

Paul M. Marsh

Abstract—A key and a brief diagnosis of each species are given for the
braconid parasites of Heliothis species in North America. The braconid
parasites which have been introduced but which are not established are
included. A table listing Old World records of parasites is also given.
Microplitis indicus, n. sp., is described from India; Chelonus texanus is
placed in synonymy with C. insularis.

The purpose of this paper is to present information on the identity of
the braconid parasites of Heliothis species as an aid to biological control
workers in North America. An illustrated key to the North American
species is presented and includes species that have been introduced but not
established; a brief diagnosis and discussion are given for each species. Also
presented is a list of Old World species that have been recorded from
Heliothis species.

Most of the introduced parasitic braconids have come from India, but from
the list of parasites in the Old World, there are plainly many other possible
parasites in southern Europe, Asia and Africa. No parasites have been in-
troduced from South America though several North American parasites
also occur in the Neotropical Region. I therefore feel this largely unexplored
area offers the best opportunity for further searches for Heliothis parasites.

To avoid repetition, I have listed the names and authors of the lepidop-
teran hosts mentioned in this paper. The spellings for specific names in
Heliothis are based on a study by Todd (1978).

Chilo plejadellus Zinck. Heliothis subflexus (Guen. )

Chilo zonellus (Swin.)
Colias eurytheme Bdvl.
Corcyra cephalonica Stain.
Diatraea saccharalis (¥F.)
Elaphria nucicolora (Guen.)
Ephestia elutella (Hbn.)
Faronta diffusa (Wlkr.)
Feltia subterranea (F.)
Heliothis armiger (Hbn.)
Heliothis assulta Guen.
Heliothis dipsaceus (L.)
Heliothis obsoletus (F.)
Heliothis ononis (D. & S.)
Heliothis peltiger (D. & S.)

Heliothis virescens (F.)
Heliothis viriplaca ( Hut.)
Heliothis zea (Boddie)
Leucania latiuscula (H.-S.)
Lineodes integer (Zell. )
Lineodes interruptus Zell.
Loxostege sticticalis (L.)
Ostrinia nubilalis (Hbn.)
Papaipema nebris (Guen.)
Pectinophora gossypiella (Saund.)
Peridroma saucia (Hbn.)
Phthorimaea operculella (Zell.)
Pilemia periusalis (Wlkr.)
Platynota stultana (W1sm.)

16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pseudaletia unipuncta (Haw.) Spodoptera ornithogalli (Guen.)
Semiothisa punctolineata (Pack.) Spodoptera praefica (Grote)
Spodoptera eridania (Cram.) Sylepta derogata (F.)

Spodoptera exigua (Hbn.) Symmetrischema heliopum (Low)
Spodoptera frugiperda (Sm.) Trichoplusia ni (Hbn.)

I wish to thank D. E. Bryan, U.S. Department of Agriculture, Agricultural
Research Service, National Program Staff, for reading the manuscript and
offering many helpful suggestions. Scanning electron microscope time was
supported in part by the Electron Microscope Central Facility, Center of
Material Research, University of Maryland, College Park.

New World Braconidae Parasitic on Heliothis Species

Fifteen species of Braconidae have been recorded as parasites of Heliothis
in North America. Of the 11 native species, only three—Cardiochiles nigri-
ceps Viereck, Microplitis croceipes (Cresson), and Chelonus insularis Cres-
son (= texanus)—are important in the control of Heliothis, and they have
been studied extensively. The other native species are only occasional
parasites of Heliothis, and some of the records need to be confirmed. How-
ever, they should perhaps be examined further and are, therefore included
in the key. The four introduced species have not become established. They
have been included in the key in the event that further studies are made
and that they do become established. One of the introduced species is
described as new. Brief mention is made for two foreign species which
were not introduced against Heliothis but for which there is data to indicate
possible association with Heliothis.

Key to the North American Braconid Parasites of Heliothis Species
(Including Those Introduced but not Established)

1. Circular opening present between clypeus and mandibles, labrum

concave (as in Fig. 1) 2
— Without a circular opening between clypeus and mandibles, labrum

not concave (as in Fig. 2) 4
2. Occipital carina present Rogas perplexus Gahan
— Occipital carina absent 3

3. Thorax and abdomen finely granular and dull; antenna more than
20-segmented, the flagellomeres longer than wide
Bracon platynotae (Cushman)
— Thorax smooth and shining; abdomen with some reticulation but
shining; antenna less than 20-segmented, the flagellomeres about
as wide as long Bracon brevicornis Wesmael
4. Abdominal terga fused into a carapace 5
— Abdominal terga not fused, with several visible segments i

VOLUME 80, NUMBER 1 i

5.

Ge

dil

14.

Antenna of female 16-segmented; carapace of male with transverse
opening at apex (Fig. 4) Chelonus (Microchelonus) heliope Gupta
Antenna of female more than 20-segmented; carapace of male
without such an opening at apex 6
Hind femur black; parastigma yellow, apical % of fore wing strongly
infuscated (Fig. 11) Chelonus (Chelonus) narayani Subba Rao
Hind femur orange at least on apical %; parastigma black, fore
wing uniformly lightly infuscated (Fig. 13)

Chelonus (Chelonus) insularis Cresson

Abdomen petiolate 8
Abdomen sessile 9
Ventral margins of petiole meeting at base and touching for nearly

% length of petiole (Fig. 19) Meteorus laphygmae Viereck

Ventral margins of petiole meeting considerably beyond base and
touching for only short distance (Fig. 20)

Meteorus autographae Muesebeck
Third segment of radius in fore wing distinct and strongly recurved;
2nd segment of radius longer than Ist; fore wing with 3 cubital cells,
2nd longer than wide (Fig. 12) Cardiochiles nigriceps Viereck
Third segment of radius absent or weakly sclerotized and straight;
2nd segment not longer than Ist if present; fore wing with 2 or 3
cubital cells, if with 3, then 2nd a small areolet and not longer than

wide 10
Second cubital cell of fore wing a small areolet (as in Fig. 16) ay
Second cubital cell absent (as in Fig. 17) 13

Vertex, temples, and mesonotum smooth and shining; hind femur
short and stout, not more than 3x as long as greatest width

Microplitis croceipes (Cresson)
Vertex, temples, and mesonotum finely punctate and dull; hind

femur slender, more than 3x as long as wide 12
Tegula and abdominal terga 1-4 yellow in female, tegula of
male yellow Microplitis indica, new species

Tegula and abdominal terga of female and male brown or black
Microplitis melianae Viereck
Ovipositor long, about as long as abdomen; areola present on pro-
podeum (Fig. 3); stigma clear medially; radius longer than inter-
cubitus (Fig. 18) Apanteles angaleti Muesebeck
Ovipositor short, shorter than Ist abdominal tergum; propodeum
without areola; stigma brown; radius not longer than inter-

cubitus 14
First abdominal tergum wider at apex than at base, coarsely rugose
(Fig. 5) Apanteles marginiventris (Cresson)

First abdominal tergum narrower at apex than at base, smooth
and polished (Fig. 6) Apanteles militaris (Walsh)

18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 1 19

Genus Apanteles Foerster

The only key for North American species is Muesebeck (1920). Wilkinson
(1928) provides a key to the Indo-Australian species; Nixon (1972, 1973,
1974, 1976), northwestern European species; Telenga (1955), Russian spe-
cies; Wilkinson (1932a), Ethiopian species.

Apanteles angaleti Muesebeck
Apanteles angaleti Muesebeck, 1954:61.

Diagnosis —Length of body, 2.5-3.0 mm, ovipositor about 1 mm. Color:
Body black, apex of fore femur, fore tibia and tarsus, basal % of mid
tibia, mid tarsus, and basal % of hind tibia yellow, wings and veins hyaline,
stigma hyaline with brown border, metacarpus brown, legs of male with
more extensive dark coloration. Body: Head shallowly punctate and shining;
mesonotum shallowly punctate, the punctures confluent along lines where
notauli would be if present; scutellum flat, smooth and shining; propodeum
rugose with a short median basal carina and areola strongly margined on
apical end (Fig. 3); Ist abdominal tergum narrower at apex than at base
and strongly punctate on apical %, rest of terga smooth; ovipositor about
as long as hind tarsus, hypopygium extending beyond apex of abdomen.
Wing: Fore wing venation as in Fig. 18, radius perpendicular to anterior
margin of wing and longer than intercubitus.

In Muesebeck’s (1920) key to North American Apanteles, angaleti will
run to epinotae but is distinguished by the first abdominal tergum being
narrowed at the apex and the more distinctly punctate mesonotum. In
Wilkinson’s (1928) key to the Indo-Australian species, angaleti is similar
to araeceri from Java, Malaya and India but is distinguished by the less
densely sculptured mesonotum, slightly shorter ovipositor, and narrower
first abdominal tergum.

Distribution —India, Sumatra. Introduced into Arizona, Florida, Missis-
sippi, New Jersey, North Carolina, South Carolina, Tennessee, and Texas
but not established.

Hosts—Pectinophora gossypiella, Corcyra cephalonica, Sylepta derogata.
Also released against Heliothis virescens and H. zea but not established.

Additional references (mostly biology on Pectinophora gossypiella).—
Narayanan, Subba Rao and Gangrade (1956), Narayanan, Subba Rao and

—

Fig. 1. Bracon platynotae, face. Fig. 2. Chelonus insularis, face. Fig. 3. Apanteles
angaleti, propodeum. Fig. 4. Chelonus heliope, apex of abdominal carapace. Fig. 5.
Apanteles marginiventris, propodeum and first abdominal tergum. Fig. 6. Apanteles
militaris, propodeum and first abdominal tergum.

20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

} Bracon
| (8) brevicornis

Rogas

(10) Bracon
platynotae

aN
\ Cardiochil
ardiochiles
Chelonus (C.) | (2) nigriceps
narayani |
parastigma 7
brslntnlay apg a
SS ee ao SEE 7 =
Sead Se

Microplitis
Chelonus (C.) croceipes

insularis

Meteorus
(5) autographae

Ist intercubitus

Apanteles
angaleti

Apanteles
@) marginiventris

Meteorus mK Meteorus ><
laph “aN autographae ~~ \
phygma me if \\ 2 Og
|
/
/

Figs. 7-18. Wings of Heliothis parasites (stippling indicates shading only, wing hairs
not indicated). Figs. 19-20. Ventral view of petiole in Meteorus spp.

VOLUME 80, NUMBER 1 a1

Kaur (1959), Subba Rao and Gopinath (1961), Narayanan, Subba Rao and
Thontadaraya (1962).

Comments.—This species was introduced into the United States as a pos-
sible biological control agent against both Heliothis spp. and Pectinophora
gossypiella, but it did not become established on either pest. Nevertheless
it appears to prefer the pink bollworm. More laboratory tests should be
made to establish its use against Heliothis.

Apanteles marginiventris (Cresson)

Microgaster marginiventris Cresson, 1865:67.

Apanteles grenadensis Ashmead, 1900:278.

Apanteles laphygmae Ashmead, 1901:36. Nomen nudum.
Apanteles (Protapanteles) harnedi Viereck, 1912b:580.

Diagnosis —Length of body, 2.0-2.5 mm. Color: Head and thorax black,
antennal flagellum dark brown, scape yellow basally, tegula yellow, abdo-
men varying from entirely black to entirely orange, legs yellow, hind
coxa often black, wings hyaline, veins brown. Body: Head punctate, dull;
mesonotum punctate, dull, punctures confluent along lines of notauli; scutel-
lum punctate; propodeum rugose with distinct median longitudinal carina
(Fig. 5); abdominal terga 1-3 strongly rugose, occasionally weakly so on
3rd tergum, Ist tergum broader at apex than at base (Fig. 5); hind coxa
punctate; inner spur of hind tibia longer than outer one; ovipositor about
as long as last segment of hind tarsus. Wing: Fore wing venation as in
Big 17.

This species is recognizable by the sculpturing on the abdomen and hind
coxa, color, and length of the hind tibial spurs.

Distribution—Eastern United States west to Wisconsin and Texas;
Arizona, California. Mexico, West Indies, northern South America, Hawaii.

Host.—Heliothis virescens and H. zea plus numerous other Noctuidae,
most often Spodoptera frugiperda.

Additional references (mostly concerning biology on Spodoptera frugi-
perda).—Luginbill (1928), Vickery (1929), Boling and Pitre (1970).

Comments.—This species is most often found as a parasite of the fall
armyworm. It is frequently reared from Heliothis spp., but its significance
in control is not well established.

Apanteles militaris (Walsh)
Microgaster militaris Walsh, 1861:369.

Diagnosis —Length of body, 1.75-2.50 mm. Color: Body black, antenna
brown, tegula yellow, abdomen light brown ventrally at base and oc-

22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

casionally laterally on terga 1 and 2, legs yellow, wings hyaline, veins brown.
Body: Head and mesonotum shiny with scattered shallow punctures;
scutellum smooth and shiny with scattered hair pits; propodeum (Fig. 6)
finely rugose without median carina; abdomen smooth and polished, Ist
tergum (Fig. 6) long and narrow, apex narrower than base; ovipositor barely
exerted. Wing: Radius of fore wing perpendicular to anterior margin of
wing and slightly shorter than intercubitus.

This species is easily distinguished by the narrow first abdominal tergum,
smooth abdominal terga, and yellow legs.

Distribution—North America, Puerto Rico, Argentina.

Hosts.—Pseudaletia unipuncta, Heliothis zea, and numerous other Noctui-
dae.

Additional references (concerning biology on Pseudaletia unipuncta ).—
Guppy (1967), Calkins and Sutter (1976).

Comments.—This species is a common parasite of the armyworm, and
there are only a few records from Heliothis zea. It is therefore not con-
sidered important in the control of Heliothis.

Genus Bracon Fabricius

Muesebeck’s (1925) key to species is the only one available for North
America.

Bracon brevicornis Wesmael

Bracon brevicornis Wesmael, 1838:23.
Habrobracon brevicornis (Wesmael). Cushman, 1922:122.
Microbracon brevicornis (Wesmael). Muesebeck, 1925:33.

Diagnosis —Length of body, 2-3 mm. Color: Mostly black with yellow
markings on head, prothorax, mesonotum, and venter of abdomen, abdomen
occasionally entirely yellow, legs brown with coxae black, wings infuscate
on basal %. Body: Vertex and frons granular, rest of head smooth; thorax
smooth and shining; abdomen finely granular and shining; antenna 16- to
19-segmented in female, 20- to 27-segmented in male; circular opening pres-
ent between clypeus and mandibles (as in Fig. 1); occipital carina absent; ovi-
positor about as long as abdominal terga 2 and 3 combined. Wing: Fore
wing venation as in Fig. 8.

Distribution—Europe, Asia, Africa. Introduced into the United States
and Canada but not established.

Hosts.—This species has a long list of hosts. It was purposely introduced
into this country in the 1930’s against Ostrinia nubilalis and subsequently
against Pectinophora gossypiella, Heliothis zea, and H. virescens. No re-
coveries have been made.

VOLUME 80, NUMBER 1 23

Comments.—This species is a common parasite of the European corn
borer and does not appear to be important in the control of Heliothis spp.
There is still some question whether Bracon brevicornis and B. hebetor are
distinct species, but that is beyond the scope of this paper.

Bracon platynotae (Cushman)

Habrobracon platynotae Cushman, 1914:104.
Bracon platynotae (Cushman). Muesebeck and Walkley, 1951:167.

Diagnosis —Length of body, 2-3 mm. Color: Head and thorax black,
face marked with yellow; abdomen and legs varying from entirely black to
entirely honey yellow, wings dusky on basal %, veins and stigma brown.
Body: Entire body granular; ocelli small, ocellocular distance about 2
diameter of lateral ocellus; circular opening present between clypeus and
mandibles (Fig. 1); occipital carina absent; antenna 22- to 27-segmented,
segments longer than wide; ovipositor about % as long as abdomen. Wing:
Fore wing venation as in Fig. 10.

Distribution —Arizona, southern California, Georgia, Louisiana, Texas;
Mexico.

Hosts.—Heliothis zea, Lineodes integer, L. interruptus, Pectinophora gos-
sypiella, Pilemia periusalis, Platynota stultana.

Comments.—This is a common parasite of the pink bollworm and only
rarely has been reared from Heliothis spp.

Bracon kirkpatricki (Wilkinson)

This species is a primary parasite of Pectinophora gossypiella, but I have
seen one reference to it as a parasite of the bollworm (Bryan et al., 1973).
If kirkpatricki does attack Heliothis, it apparently does so at such a low rate
as to be insignificant. Further studies should be made to establish the
relationship between kirkpatricki and Heliothis.

Genus Cardiochiles Nees

The North American species of Cardiochiles can be identified by Mao's
(1949) key. Mao (1945) provides a key to the Mexican species; Fischer
(1958), Egyptian species; Telenga (1955), Russian species; De Saeger (1948),
Belgian Congo species.

Cardiochiles nigriceps Viereck

Cardiochiles nigriceps Viereck, 1912b:578.

Diagnosis —Length of body, 6-8 mm. Color: Head and antenna black,
thorax with venter and propodeum always black, scutellum and mesonotum

24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

posteriorly always orange, pronotum dorsally, mesonotum anteriorly, meso-
pleuron dorsally varying from black to orange, abdomen orange, fore and
mid legs black, tibiae and tarsi sometimes brown, hind coxa and _ tro-
chanters black, hind femur orange, hind tibia orange or brown, hind tarsus
brown, wings strongly infuscated. Body: Head broad, transverse, temples
bulging; body smooth and shining except propodeum which is strongly
rugose with distinctly margined diamond-shaped areola in middle; hind
femur swollen; ovipositor short and barely exerted. Wing: Fore wing
venation as in Fig. 12, radius strongly arched, 1st intercubitus distinctly
angulate below middle.

This species is not likely to be confused with other North American spe-
cies; its body color and angled first intercubitus of the fore wing are
distinctive.

Distribution.—District of Columbia south to Florida, west to Kansas and
Louisiana; Mexico. Also introduced into the Philippines but apparently not
established.

Hosts.—Heliothis subflexa, H. virescens. Also recorded on H. assulta in
the Philippines.

Additional references—Chamberlin and Tenhet (1926), Lewis and Brazel
(1966), Vinson and Lewis (1965), Lewis et al. (1967), Lewis and Vinson
(1968a, 1968b), Vinson (1968, 1969), Lewis and Vinson (1971), Hays and
Vinson (1971), Vinson (1972), Lewis et al. (1972).

Comments.—This is the most important and most studied braconid at-
tacking Heliothis. However, it seems to be restricted to H. virescens. There
is only one record of it developing on H. subflexa (Lewis et al., 1967). It
will attack H. zea but will not complete its development.

Genus Chelonus Panzer

There are no keys to North American species of Chelonus (in the strict
sense); McComb (1968 (1967)) provides a key to the species of the sub-
genus Microchelonus for North America; De Saeger (1948) provides keys
for species of both subgenera in the Belgian Congo.

Chelonus (Microchelonus) heliope Gupta

Chelonus heliope Gupta, 1955:209.
Chelonus (Microchelonus) heliope Gupta. McComb, 1968(1967):71.

Diagnosis —Length of body, 3-4 mm. Color: Body black, scape yellow,
basal flagellomeres brown, apical ones black, basal % of abdominal cara-
pace yellow, fore and mid legs honey yellow except coxae and last tarsal
segment which are black or brown, hind coxa black, hind trochanters brown,
hind femur black, hind tibia black on apical and basal 4% with middle %

VOLUME 80, NUMBER 1 25

yellow, hind tarsus yellow except last tarsal segment which is black. Body:
Strongly sculptured; antenna of female 16-segmented, male 24- to 26-
segmented; propodeum with strong tubercles at apical corners; Ist 3 ab-
dominal terga fused into a strongly sclerotized carapace which encloses
rest of abdominal segments, apex of abdominal carapace in male with deep
transverse groove and tubercle in center of groove (Fig 4). Wing: Fore wing
venation as in Fig. 9.

This species is somewhat similar to pectinophorae Cushman and _ black-
burni Cameron, both of which have been introduced but not established,
but it differs by having all flagellomeres at least slightly broader than long
and in having the wing entirely hyaline. It is also similar to the endemic
fulgidus McComb and shenefelti McComb but differs from both in having
a shorter malar space which is equal to the basal width of the mandible.

Distribution —India. Introduced into Arizona, Florida, Mississippi, New
Jersey, North Carolina, South Carolina, Tennessee, and Texas but not estab-
lished.

Hosts—Symmetrischema heliopum, Pectinophora gossypiella, Corcyra
cephalonica (in laboratory). Also introduced against Heliothis zea, H. vires-
cens, and Diatraea saccharalis but not established.

Additional references—Charpentier (1956, 1958), McGough and Noble
(1957), Patel et al. (1958).

Comments.—This species was introduced in the United States mainly
for control of the pink bollworm, but some introductions were also made
against Heliothis. It has not been established and appears not to be an
important parasite of Heliothis.

Chelonus (Chelonus) insularis Cresson

Chelonus insularis Cresson, 1865:61.

Chelonus texanus Cresson, 1872:179. New synonymy.
Chelonus texanoides Viereck, 1905:286. New synonymy.
Chelonus exogyrus Viereck, 1905:287. New synonymy.
Chelonus bipustulatus Viereck, 1911b:476. New synonymy.

Diagnosis —Length of body, 4.5-5.0 mm. Color: Black, abdominal cara-
pace usually with 2 white lateral spots on basal %, carapace rarely entirely
black or entirely orange, fore and mid legs orange except coxae, base of
femora and apical tarsal segments which are black, hind coxa black, hind
femur usually black, occasionally orange, hind tibia orange with base and
apex black, wings lightly infuscated, stigma and parastigma black, veins
brown or light brown near base of wing. Body: Entirely, strongly sculp-
tured, sculpture rugose but not regularly areolated; antenna 26- to 32-
segmented; propodeum with strong tubercles at apical corners; Ist 3 ab-
dominal terga fused into a rigid dorsal carapace which encloses rest of

26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

abdominal segments, apex of carapace of male without groove or impres-
sion. Wing: Fore wing venation as in Fig. 13.

This species is similar to narayani but can be distinguished by the black
parastigma, mostly orange hind femur, hyaline wings, and white spots at base
of the carapace. I have seen the type of insularis Cresson and there is no
doubt that it is the same as texanus Cresson.

Distribution—North America, Central America, South America, West
Indies. Introduced into Hawaii and South Africa.

Hosts (for North America).—Ephestia elutella, Feltia subterranea, Helio-
this zea, Loxostege sticticalis, Peridroma saucia, Spodoptera eridania, S.
exigua, S. frugiperda, S. ornithogalli, S. praefica, Trichoplusia ni.

Additional references (selected for Heliothis).—Pierce and Holloway
(1912), Wolcott (1951 (1948)), Ullyett (1949), Botrell et al. (1968), Vinson
(1975).

Comments.—This species, which occurs throughout the New World, is
one of the common native species of Heliothis parasites, though it has been
more often studied with Spodoptera frugiperda.

Chelonus (Chelonus) narayani Subba Rao
Chelonus narayani Subba Rao, 1955:63.

Diagnosis —Length of body, 3.5-4.5 mm. Color: Black, all coxae, tro-
chanters and femora black except apex of fore and mid femora which are
occasionally orange, fore tibia and basitarsus orange, rest of fore tarsus
brown, mid tibia brown at apex and base, yellow in middle, mid basitarsus
yellow, rest of mid tarsus brown, hind tibia black at apex and base, yellow
in middle, hind basitarsus yellow, rest of hind tarsus brown, fore wings
hyaline on basal %, strongly infuscate on apical 4%, stigma black, para-
stigma yellow, abdominal carapace very rarely with weak orange spots
at base. Body: Strongly sculptured, face and thorax appearing areolated;
antenna 24- to 29-segmented; propodeum with strong spines at apical cor-
ners; Ist 3 abdominal terga fused into rigid dorsal carapace, apex of male
carapace without groove or impression. Wing: Fore wing venation as in
Fig. 11.

This species is similar to insularis but can be distinguished by the yellow
parastigma and infuscate apex of the fore wing, the black hind femur, the
totally black abdominal carapace, and the more areolated sculpturing of the
face and thorax. It is also similar to iridescens Cresson but is smaller, has
a relatively shorter carapace, and has the sculpturing more areolated on the
face and thorax.

Distribution.—India. Introduced into Arizona, Florida, Louisiana, Missis-
sippi, New Jersey, North Carolina, Pennsylvania, South Carolina, Tennessee,
Texas, and Utah but not established.

VOLUME 80, NUMBER 1 27

Hosts.—Heliothis armiger, H. zea, Chilo zonellus, Corcyra cephalonica
(in laboratory). Also introduced into Texas against Pectinophora gossypiella,
into Pennsylvania against Papaipema nebris, into Louisiana against Diatraea
saccharalis and Chilo plejadellus, and into Utah against Phthorimaea
operculella but also not established.

Comments.—This is apparently a good species on Heliothis and more
studies should be made to determine its effectiveness in North America.

Chelonus (Microchelonus) blackburni Cameron

This species was introduced into the United States against Pectinophora
gossypiella but did not become established. According to Bryan et al.
(1973), this species “could (italics mine) parasitize” Heliothis zea and H.
virescens. Further field studies must be made to determine whether it will
attack Heliothis.

Chelonus (Chelonus) gossypii Viereck

This species is not included in the key. It was described (Viereck, 1912a)
from Brazil as a parasite “on cotton worm.” If this refers to Heliothis, then
gossypii is a possible additional parasite to be considered.

Genus Meteorus Haliday

The only key to North American species is that of Muesebeck (1923);
Nixon (1943) provides a key to the Ethiopian species.

Meteorus autographae Muesebeck
Meteorus autographae Muesebeck, 1923:30.

Diagnosis—Length of body, 4-5 mm. Color: Female usually entirely
honey yellow, sometimes mesonotum, propodeum and Ist abdominal seg-
ment (petiole) dark brown, male usually honey yellow with vertex, meso-
notum, propodeum, and all abdominal terga except the 2nd dark brown,
stigma uniformly pale yellow, transparent. Body: Eyes large; ocelli large,
ocellocular distance about 2 diameter of lateral ocellus; mesonotal lobes
smooth; ventral margins of petiole meeting beyond base of petiole and touch-
ing for only a short distance (Fig. 20). Wing: Fore wing venation as in
Fig. 15, lst segment of radius at most % as long as 2nd segment, recurrent
vein entering base of 2nd cubital cell.

This species is very similar to laphygmae but is distinguished by the
ventral margins of the petiole and the wing venation.

Distribution—Newfoundland south to Florida, west to Ontario and
Louisiana.

28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Hosts.—Many species of Noctuidae but primarily a parasite of Spodoptera
frugiperda.

Comments.—This is not an important parasite of Heliothis. I have seen
only one specimen reared from Heliothis zea.

Meteorus laphygmae Viereck
Meteorus laphygmae Viereck, 1913:560.

Diagnosis —Length of body, 3.5-5.0 mm. Color: Entirely honey yellow,
rarely petiole with black markings at apex. Body: Eyes large; ocelli large,
ocellocular distance about equal to diameter of lateral ocellus; mesonotal
lobes weakly punctate; ventral margins of petiole meeting at base of petiole
and touching for nearly % the length of petiole (Fig. 19). Wing: Ist seg-
ment of radius of fore wing nearly as long as 2nd segment, recurrent vein
interstitial with Ist intercubitus.

Distribution—Texas and New Mexico south to northern South America;
introduced into Hawaii.

Hosts.—Autographa sp., Colias eurythene, Elaphria nucicolora, Feltia sub-
terranea, Heliothis zea, Leucania latiuscula, Peridroma saucia, Pseudaletia
unipuncta, Semiothisa punctolineata, Spodoptera eridania, S. exigua, S.
frugiperda.

Comments.—This species is occasionally reared from Heliothis species
though it has been studied most frequently as a parasite of the fall army-
worm.

Genus Microplitis Foerster

Muesebeck (1922) provides a key to the North American species; Wilkin-
son (1930), Indo-Australian species; Telenga (1955), Russian species; Nixon
(1970), northwestern European species.

Microplitis croceipes (Cresson)

Microgaster croceipes Cresson, 1872:183.
Microplitis nigripennis Ashmead. In Quaintance and Brues, 1905:122.

Diagnosis—Length of body, 3.5-4.5 mm. Color: Head, antenna, and
thorax black, abdomen usually orange, rarely black, legs orange except
coxae which are black, wings strongly and entirely infuscated. Body: Very
smooth and shining; propodeum coarsely rugose with median longitudinal
carina; 1st abdominal tergum weakly punctate along edges; temples bulg-
ing beyond eye margins; hind femur short and stout, not more than 3x
as long as greatest width; ovipositor barely exerted. Wing: Fore wing
venation as in Fig. 14.

VOLUME 80, NUMBER 1 29

This species and longicaudus are distinct from all other Microplitis in
North America by their smooth and shining head and thorax and short and
stout hind femur. From longicaudus this species is distinguished by the
shorter ovipositor and infumated wings.

Distribution.—New Jersey south to Georgia, west to Utah and Arizona;
Oregon. Probably also occurs in northern Mexico.

Hosts.—Heliothis subflexus, H. virescens, H. zea.

Additional references.—Bryan et al. (1969), Lewis (1970), Jones and
Lewis (1971).

Comments.—This is one of the more important parasites of Heliothis
species. It has not been reared from any hosts other than those listed.
D. E. Bryan (personal communication) has indicated that biological data
presented by Bryan et al. (1969) and Jones and Lewis (1971) do not agree
in all respects with the indication that croceipes of the eastern United
States may not be the same as the western forms. Further study is needed to
clarify this matter.

Microplitis indica Marsh, new species

Description —Length of body, 2.5 mm. Color: Head including antenna
and thorax black, abdominal terga 1-4 in female and 1-3 in male orange,
rest black, 1st abdominal tergum of male sometimes darker, legs of female
orange except tarsi always and hind coxa occasionally black, legs of male
orange except coxae and tarsi which are black, tegula orange, wings evenly
infumated, veins brown, stigma brown with yellow spot at basal %. Body:
Head distinctly punctate, dull; antenna 18-segmented and longer than
body in male; mesonotum distinctly punctate and dull; notauli not im-
pressed and only weakly indicated by confluent punctures along their
course; scutellum dull, punctate, scutellar furrow wide and deep with 3-5
carinae; mesopleuron strongly punctate except smooth shining area above
the crenulate sternaulus; propodeum strongly rugose with weak indication
of median carina; lst abdominal tergum as long as greatest width, parallel
sided for 7s its length, narrowing on apical 4, finely rugose laterally, smooth
apically; ovipositor sheaths barely exerted, hypopygium not extending be-
yond apex of abdomen; hind tibial spurs equal in length. Wing: Fore
wing venation as in Fig. 16.

This species is similar to pallidipes Szepligeti which occurs in southeast
Asia but differs by the darker tarsi and antennae, less distinct notauli,
and weakly indicated median carina on the propodeum. It is also similar
to rufiventris Kokujev that occurs in southern Russia (Uzbeck, Turkmen),
but it can be distinguished by the black tip of the abdomen and black tarsi.
In North America, indicus runs to laticinctus or bradleyi in Muesebeck’s
(1922) key but is distinguished from laticinctus by its less sculptured first

30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

abdominal tergum and black antennae and from bradleyi by its orange ab-
dominal terga 1-4. It might also be confused with melianae but is dis-
tinguished by its yellow tegula, orange abdominal terga 1-4, narrower
and more rugose first abdominal tergum, and infumated wings.

Holotype female.—Kulu, India; lab. reared; Ship’t., no. 5, [X-64; ex
Heliothis sp. Paratypes, 22, 24, same data as type. All types deposited
in USNM.

Distribution.—India.

Host.—Heliothis assulta.

Comments.—This species was originally determined as Apanteles sp. and
later as Microplitis sp. near pallidipes. It was reared in laboratories in New
Jersey and Georgia but never released.

Microplitis melianae Viereck
Microplitis melianae Viereck, 1911a:185.

Diagnosis —Length of body, 2.5-3.0 mm. Color: Head, thorax and abdo-
men black, tegula brown, abdominal terga 1 and 2 brown laterally and
ventrally, coxae and trochanters black or brown, rest of legs yellow, tarsi
brown in male, wings hyaline, stigma with basal % yellow, rest brown.
Body: Head and thorax coarsely punctate and dull; mesopleuron smooth and
shining; propodeum rugose with distinct median carina; abdominal terga
smooth and shining, Ist abdominal tergum usually punctate at apex; ovi-
positor barely exerted.

This species might be confused with indicus if that species were to be
established in North America, but melianae is distinguished by its darker
tegula and abdomen, hyaline wings, and smoother abdomen.

Distribution.—Illinois, Iowa, Kansas, Michigan, Minnesota, New York,
Ohio, Ontario, Tennessee.

Hosts.—Faronta diffusa, Heliothis zea, Pseudaletia unipuncta.

Comments.—This species is a frequent parasite of the armyworm, but
there is at least one record from Heliothis zea. It is not considered an im-
portant parasite of Heliothis.

Genus Rogas Nees
The only reliable keys to species are Telenga (1941) for Russia, Fahringer
(1931) for Europe, and Granger (1949) for Madagascar.
Rogas perplexus Gahan
Rogas perplexus Gahan, 1917:205.

Diagnosis —Length of body, 4-5 mm. Color: Mostly brown with the
following parts honey yellow, vertex, orbits around eyes, face occasionally,

31

VOLUME 80, NUMBER 1

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eset ‘Azeas ‘7S6T ‘ysnbog
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LSet ‘ysnbog

LS6t ‘usnbog ‘gc6Tt ‘ebueToL
GS6lT ‘LE6T ‘ebuUeTOL

90UetI Foy

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etssny uzreyANOS
etssny uzreYyANOS
edoing
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STAIZUSATJNA STAZITdoOAOTW

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ee

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"T eTqeL

32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

along lines of notauli on mesonotum, scutellum, apex of 1st abdominal
tergum, and all of terga 2 and 3, and all legs; wings hyaline, veins brown,
stigma brown with base and apex yellow. Body: Entire body granular;
eyes and ocelli large, ocellocular distance about equal to diameter of lateral
ocellus; circular opening present between clypeus and mandibles (similar
to Fig. 1); occipital carina present; abdomen with median longitudinal
carina along terga 1-3; ovipositor barely exerted. Wing: Fore wing venation
as in Fig. 7.

Distribution.—Arizona, southern California, New Mexico, Texas.

Hosts.—Heliothis zea, Peridroma saucia, Trichoplusia ni.

Additional references.—Butler (1958), rearing records.

Comments.—Pupation of species in this genus takes place within the
mummified host larva, which retains its shape. This is an uncommon species
and insignificant as a parasite of Heliothis.

Old World Records of Braconidae Parasitic on Heliothis Species

Table 1 lists Braconidae from the Old World that have been recorded
as parasites of Heliothis species. This is not an exhaustive list of all records
that I have seen but represents what I believe to be the most reliable records.
In many of the records there is no biological information other than a rear-
ing record. Some of the records are undoubtedly incorrect; others indi-
cate some species are occasional parasites of Heliothis. These records are
presented to indicate potential species of interest and/or areas where
parasites might be collected in the future.

Literature Cited

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St. Vincent and Grenada, with additions to the parasitic Hymenoptera and a list
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—. 1901. In Chittenden, The fall armyworm and the variegated cutworm.
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Bhatnagar, S. 1950(1948). Studies on Apanteles Foerster (Vipionidae: parasitic
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Bogush, P. P. 1957. Parasites of the cotton boll-worm, Chloridea obsoleta F. (Lepidop-

tera, Agrotidae) in Turkmenistan. Entomol. Obozr. 36:98—107.

1959. Materials on parasitic insects of Turkmenia. Zool. Z. 38:189-195.

Boling, J. G., and H. N. Pitre. 1970. Life history of Apanteles marginiventris with de-
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Bottrell, D. C., J. H. Young, R. G. Rice, and R. H. Adams. 1968. Parasites reared
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Bryan, D. E., C. G. Jackson, and R. Patana. 1969. Laboratory studies of Microplitis

VOLUME 80, NUMBER 1 33

croceipes, a braconid parasite of Heliothis spp. J. Econ. Entomol. 62:1141—
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Butler, G. D. 1958. Braconid wasps reared from lepidopterous larvae in Arizona,
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Calkins, C. O., and G. R. Sutter. 1976. Apanteles militaris and its host Pseudaletia
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Chamberlin, F. S., and J. N. Tenhet. 1926. Cardiochiles nigriceps Vier., an im-
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Charpentier, L. J. 1956. 1954 studies of parasites for sugarcane borer control in
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—. 1958. Recent attempts to establish sugarcane borer parasites in Louisiana.
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Fahringer, J. 1931. Opuscula Braconologica. Bd. III, Lief. 3, pp. 161-240.

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——-. 1951. Notes on the introduction and biology of Microplitis demolitor Wilk.
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34 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Lewis, W. J., A. N. Sparks, R. L. Jones, and D. J. Barras. 1972. Efficiency of
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Lewis, W. J., and J. R. Brazzel. 1966. Biological relationships between Cardiochiles
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———. 1968b. Immunological relationships between the parasite Cardiochiles nigri-
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VOLUME 80, NUMBER 1 35

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36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

———. 1912a. Contributions to our knowledge of bees and ichneumon-flies, includ-
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—. 1972. Competition and host discrimination between two species of tobacco
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381-384.

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Res. 20:103-114.

——.. 1930. A revision of the Indo-Australian species of the genus Microplitis
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——. 1932a. A revision of the Ethiopian species of genus Apanteles (Hym. Bracon.).

Trans. R. Entomol. Soc. London 80:301—344.

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749-975.

Systematic Entomology Laboratory, I[BIII, Agric. Res. Serv., USDA,
c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 37-42
TACHINID PARASITES OF HELIOTHIS IN THE WESTERN
HEMISPHERE (DIPTERA; LEPIDOPTERA)

Curtis W. Sabrosky

Abstract.—Search of the literature and personal records of identifications
yielded a total of 61 species of Tachinidae recorded from Heliothis in the
Western Hemisphere, plus 3 species recently introduced from India but not
yet known to be established. The table includes the geographic region,
the species of Heliothis involved (H. zea, H. virescens, “others, or sp.”), and
a brief summary of the known host relationships of the tachinids. Many
of the species are uncommon, or at least rarely reared. Almost all the com-
mon species are polyphagous, especially on Noctuidae.

I present here a preliminary list of tachinid flies known to parasitize
species of the noctuid genus Heliothis in the Western Hemisphere. It
is probably complete for the major parasites because much attention has
been and is being given to parasites of Heliothis, but minor, occasional,
aberrant, or rare species of parasites will undoubtedly be added in the
future. The present list is based on card catalogs in the Systematic En-
tomology Laboratory, my personal records of identifications, the manu-
script of “A Host-Parasite Catalog of North American Tachinidae” by Paul
H. Arnaud, Jr., soon to be published by the U.S. Department of Agriculture,
and the recently published “Host-Parasite and Parasite-Host Catalogue of
South American Tachinidae (Diptera)” by J. H. Guimaraes (1977, Arq. Zool,
Sao Paulo, 28(3):1-131). For commonly reared species, host relations are
summarized from the available information. Many of the tachinids are
uncommon or at least rarely reared. Almost all of the common species are
polyphagous, especially on Noctuidae.

Abbreviations: NA = North America, SA = South America, WI = West
Indies; * = personal identification; ** = both personal identification and
published records.

Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv., USDA, c/o
U.S. National Museum, Washington, D.C. 20560.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

38

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39

VOLUME 80, NUMBER 1

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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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41

VOLUME 80, NUMBER 1

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PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 43-50

WHAT IS SYSTEMATIC ENTOMOLOGY?!
George C. Steyskal

Abstract.—Systematic entomology is defined and its position as the basic
regimen of entomology is discussed. Systematics is indispensable as it serves
the general need, biological control problems, ecological and environmental
studies, veterinary entomology, etc. Examples of the difficulties and rewards
for the systematic entomologist are given.

Most systematic entomologists have at least a fairly good idea of the aims
and content of their branch of science, but among laymen and even scientists
of physical and chemical disciplines there is some misapprehension. Some
scientists believe that systematic entomology is not even a science, ap-
parently because it does not ordinarily require a laboratory filled with ex-
pensive and complex apparatus and because, again ordinarily, controlled ex-
periments are not performed. However, systematic entomology is indeed not
only a science but the basic regimen of entomology.

Let us define our terms. The “preliminary definition” of science by Caws
(1965:11) should suffice for our purpose, inasmuch as it is in the restricted
sense we require and is even more philosophically rigorous than the dic-
tionary definitions, while at the same time in agreement with them:
“science is the explanation of nature on its own terms, together with all
that follows from doing that successfully, such as the ability to predict
how things will behave and hence to control them.” Caws’ 354-page
book, entitled “The Philosophy of Science, a Systematic Account,” is a
commentary on this definition.

The word research is defined in the scientific sense in the great Oxford
Dictionary as “a search or investigation directed to the discovery of some
fact by careful consideration or study of a subject; a course of critical or
scientific inquiry.” The Third New International Dictionary of the English
Language (Webster's) is more prolix: “a studious inquiry or examination;
esp critical and exhaustive investigation or experimentation having for its
aim the discovery of new facts and their correct interpretation, the revision
of accepted conclusions, theories, or laws in the light of newly discovered
facts, or the practical application of such new or revised conclusions,
theories, or laws.” It is thus evident that controlled experimentation is not
a necessary part of research in its broad sense, but that hypothesis and
testing in some form, even when the latter is not feasible by direct experi-
mentation, definitely are.

The definition of systematic is not quite so easy. The word is an adjective
on which the noun systematics is based. These words are in turn derived
from the noun system, which is merely a way of referring to the fact that in

44. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

science the various parts of the study of a subject agree with each other
and hang together in a cohesive whole. Science, after all, is only a syste-
matic way of considering nature.

Systematics is also known as taxonomy. A distinction is often made be-
tween these two terms, but in general they are virtually synonymous. The
dictionaries define both as “the science of classification” and cite them as
synonyms of each other. The term classification, however, when narrowed
to its meaning in biology, is too narrow and entails unfortunate connotations
carried over from its use in non-biological fields.

Let us, therefore, refer to a few texts on systematics (or taxonomy) pub-
lished since 1960, viz. Simpson (1961), Cain (1963), Hennig (1966), Black-
welder (1967), Mayr (1969), Michener (1970), and Ross (1974). There is
much more in the literature referring to the basics of zoological science, but
these works can be considered representative and authoritative enough to
use in drawing conclusions.

Simpson (1961) says that “systematics is the scientific study of the kinds
and diversity of organisms and all relationships among them.” He also
quotes Hennig’s Grundziige der phylogenetischen Systematik (1950), the
precursor of his revised work published in English in 1966, “. . . one must
first make clear that there is a systematics not only in biology, but that
it constitutes an integrating component in any science whatsoever.”

Cain (1963): “In recent years it has been gradually realized that taxonomy
is not merely a necessary pigeon-holing but also one of the most important
activities in biology, requiring a synthesis of all other biological pursuits
for its proper performance ... .”

Hennig (1966) states, “. . . systematics in the most general sense is equiva-
lent to order, rationalization, and in a certain context explanation of the
world of phenomena; and . . . in this sense systematics is a very broad task
of all natural sciences, and particularly of all biological disciplines,” and at
another place, “. . . systematics fundamentally means any investigation of
relations between natural things and natural processes insofar as they have
the character of conformity to law.”

Blackwelder (1967) gives a definition on the first page of his text: “Both
taxonomy and classification, and all the other aspects of dealing with
organisms and the data accumulated about them, are included in sys-
tematics, which is the general term that covers all aspects of the study of
kinds. Therefore, systematics is the study of the kinds and diversity of
organisms, their distinction, classification, and evoluton.”. He uses as a
prefatory quotation the following sentence from Simpson (1945): “taxonomy
is at the same time the most elementary and the most inclusive part of zool-
ogy, most elementary because animals cannot be discussed or treated in
a scientific way until some taxonomy has been achieved, and the most in-
clusive because taxonomy in its various guises and branches eventually

VOLUME 80, NUMBER 1 45

gathers together, utilizes, summarizes, and implements everything that is
known about animals, whether morphological, physiological, psychological,
or ecological.”

Mayr (1969) follows Simpson (1961), already quoted, and distinguishes
taxonomy as the “theory and practice of classifying organisms.”

Ross (1974) considers that the term taxonomy is a synonym of systematics
and that “systematics has the majority vote as the inclusive name to be
applied to the total field of investigation that we are discussing.” He uses
the newer term “biosystematics . . . as the investigational field of systematics
based on any scientific information that can be brought to bear on the
problems of the evolution of species, whether they concern speciation or
phylogeny.” Inasmuch as systematics is not restricted to biology (see Hennig,
1950), the term biosystematics is nothing more than a restrictive term refer-
ring to systematics in biology, and is superfluous when used in purely
biological context.

In a remarkably clear, balanced, and suggestive article on “Diverse Ap-
proaches to Systematics,” Michener (1970) cites Simpson’s definition of
systematics. He also states, “one might think that this (Simpson’s defini-
tion of systematics) includes everything in population biology ... but the
systematic approach is less broad” (my italics). I would consider that it
is less broad only if it concerns a part of biosystematics, one dealing only
with certain kinds of organisms. The systematics of all biology certainly
must include population biology, at least as a source of data.

There is much agreement among these definitions. Sifting them out, one
may come to the conclusions that 1) the term systematics is becoming
accepted as the term including both taxonomy and classification, whether or
not some synonymy is involved, and 2) systematics is becoming continually
broader in scope, using ever more various ways of looking at organisms.

Systematics in biology is based upon two assumptions, 1) that all bio-
logical science is founded on the species concept, and 2) that all kinds
of data are grist for its mill.

It is the basic concern of systematics to place the concept of an organism
in proper relationship within the species concept—is it that of a species,
subspecies, of a higher or lower taxonomic rank, and, establishing the rank,
what relationships in time, space, and kind does it bear to those of other
organisms, especially those closest to it?

The last few decades have seen the field of systematics expand enor-
mously; some have even said that biology is changing from many more or
less well-defined disciplines into one all-embracing subject. The concept
of holomorphology was stressed by Hennig as early as 1950. Anything
evenly remotely to be considered as morphological (even cell and chromo-
some structure), physiological characteristics (because organic chemicals are
substances and have form), and proteins, all fall within the scope of mor-

46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

phology because they have form (morphé). The use of the genitalia of both
sexes and their associated structures has now become commonplace, even
considered necessary, in the study of any kind of insect. Internal structures
are becoming increasingly important. Arnold (1972a, 1972b) has recently
been pursuing the study of insect haemocytes with reference to systematics.

The proceedings of an international conference on systematic biology
have been published in a volume (Internatl. Conf. Syst. Biol., 1969) that
includes papers on systematics of populations, ecological aspects of sys-
tematics (in plants and animals), molecular data in systematics, systematic
significance of isolating mechanisms, comparative animal behavior in sys-
tematics, comparative cytology in systematics, biometric techniques in sys-
tematics, and computer techniques in systematics. Mayr, in the opening
paper in the volume, closes with the statements “Finally, let us remember
that in virtually every taxonomic finding certain generalizations that are
of value and broad interest to biology as a whole are implicit” and “It is
my sincere belief that systematics is one of the most important and in-
dispensable, one of the most active and exciting, and one of the most re-
warding branches of biological science. I know of no other subject that
teaches us more about the world we live in.”

Edward O. Wilson, in the final paper summarizing that conference, asks,
“What is a pure systematist? He might be defined as a biologist who works
on such a large number of species that he has only time enough to con-
sider classification and phylogeny. If he narrows his focus, his unique
knowledge provides him with a good chance to make discoveries in genetics,
ecology, behavior, and physiology. But then we come to know him as a
geneticist, or an ecologist, or a behaviorist, or a physiologist. It clearly will
not do to define systematics as classification plus all these other fields, be-
cause that would be robbing the discipline of its true meaning. I think
it would be appropriate just to acknowledge everything that systematists
do for the rest of biology (and that is a great deal indeed) ....” He
also notes “It has occasionally been said that the perfect experimental biol-
ogist selects a problem first and then seeks the organism ideally suited
to its solution. In contrast, the typical systematist selects the organism first
—for the love of it. Now this is a great strength, for the systematist de-
votes his career to the organism and thereby often comes upon problems
of general significance that would be discovered otherwise; .. .”

In a paper to be delivered at the BARC Symposium II, at the Beltsville
Agricultural Research Center, on 8-11 May 1977, Foote (in press) has
brought together an imposing list of references and much data on the
importance of systematics to the general need, biological control problems,
ecological and environmental studies, veterinary entomology, etc.

Mathematics and its offspring computer technology are becoming more
and more used in systematics. Numerical taxonomy, which burst upon the

VOLUME 80, NUMBER 1 47

scene with great eclat as a new solution to the problems of systematics, is
now taking its more logical position as taxonomic mathematics. Mathe-
matics, even of the most sophisticated kind, is still the handmaiden and
not the director of the sciences.

Many new ways of working with insects are coming to hand, new ways
of getting at them and new ways of looking at them. Electron microscopy
is allowing entomologists to see things they never saw before. But en-
tomologists, especially those practicing systematics, are too few and have
too much to do to take full advantage of these new ways and means. The
best solution of a problem is often too time-consuming and costly; the sys-
tematist must therefore be content in many instances with a solution that is
merely good enough at the moment. Perhaps he may be able to do some-
thing better later. At the worst, other entomologists, some perhaps better
situated, will find in more detail, discover new facts, and make new hy-
potheses sooner or later.

Heiser (1966) has remarked that the “process of character selection, re-
jection, and weighting by taxonomists is one of the great mysteries to
some non-taxonomists and to beginning taxonomists... .” I would say that
it is part of the taxonomist’s way of experimenting, his process of hypothesiz-
ing, testing, and rehypothesizing repeated until a better hypothesis is found.

A systematic entomologist, then, studies all aspects of insect life by
any feasible means in order to add to and correct or refine our knowledge
concerning insects, and thereby advances the possibilities of dealing with
them in the fields of prediction and control.

Let us now consider a few concrete examples, selected more or less
fortuitously, to illustrate how the systematist works and what he does or
may accomplish. It should not be forgotten, in considering these examples,
that by far the greatest number of insects species have close relatives that
are not easy to distinguish from them, although sometimes their habits
may be quite different. Even entomologists who are not systematists
often lose sight of this fact.

1) The imported fire ants in the United States—For over 30 years it had
been supposed that only 1 species of Solenopsis, perhaps with two varieties,
had entered the United States. Buren (1970) showed that two distinct
species, S. richteri Forel, 1909, and S. invicta Buren, 1970 are present, with
little or no hybridizing. He was able to do this by careful analysis of
larger amounts of material and comparing it with more South American
material than others had before his study. He also was able to consider
differences in the chemical constituency of the venoms of the two species.
He was thus able to set up a new hypothesis concerning the systematics of
the organisms, one that agrees better with additional observed data.

2) North American species of Rhagoletis (Diptera: Tephritidae ).—Prob-

48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lems of the taxonomic status of morphologically very similar fruitflies feed-
ing upon different hosts were largely solved in a revision of the North
American species by Bush (1966), whose objective was “to incorporate as
much . . . biological information as possible into a reevaluation of these
so-called host races. Additional observations made in three years of field
and laboratory work on such aspects as the problem of chromosome cytology,
courtship behavior, distribution, and host relations have also been in-
cluded.” Chromosome morphology aided considerably in defining the genus.
He also stated that “certain aspects in the adaption to new hosts, such as
the genetics and chemistry of host selection, conditioning, and mating be-
havior, have yet to be studied. The hosts of several species are still un-
known and the distribution of most species, including those of economic
importance, is yet to be definitely established.”

3) The genus Muscidifurax (Hymenoptera: Pteromalidae).—Muscidifurax
raptor Girault and Saunders, 1910, a parasite of the house fly, Musca
domestica L., was considered to be the sole species of its genus and to
occur in the southwestern United States, tropical America, Europe, and
Africa, until Kogan and Legner (1970) made reciprocal crossing experiments
which showed that some of the populations were reproductively isolated
and noted other biological differences. Their study revealed five species with
several morphological differences, even in the eggs. Scanning electron
microscopy proved helpful in working with these small insects, the largest
of which is 2.8 mm long. Van den Assem and Povel (1973), in studying three
of these species, found differences in courtship behavior sufficient to act as a
barrier to crossmating, at least in sympatric species,

4) Species of Procecidochares (Diptera: Tephritidae) forming galls on
weeds of the genus Ageratina——In 1947, Stone described Procecidochares
utilis, which was collected in Mexico, the original source of the weed Agera-
tina adenophora, and introduced into Hawaii for biological control of the
weed. When a second species of the weed, A. riparia, also became trouble-
some in Hawaii, it was found that Procecidochares utilis would not use it
as a host. A search was then made, again in Mexico, for gall-forming flies
on Ageratina riparia. Another species of Procecidochares was found and
brought to Hawaii, where it was found to refuse Ageratina adenophora as
as host. Adults of the flies on A. riparia were referred to me for determina-
tion. At first I could find only a small, doubtful difference in the pattern
of the wing between the two flies, but knowledge of the host specificity in-
duced me to look further. Finally good morphological differences were
found in postabdominal details of both sexes and in eggs removed from
ovaries. I described the species as new (Steyskal, 1974).

These examples could be extended ad nauseam. Nothing has been
said here about phylogeny and higher classification. The practicing sys-
tematist is usually not a pure systematist in the sense of Wilson because he

VOLUME 80, NUMBER 1 49

does not have time for anything but classification and phylogeny, but be-
cause he must still deal with the species-distinction problems, what has
been called “alpha” taxonomy. He usually hopes that someday he can do
the big job, when the routine or teaching load permits. But even for the
basic or elementary problems he must still be able to accumulte much ma-
terial, often having to collect it himself; he must visit many museums; he
should be able to rear his specimens; he should have the use of an electron
scanning microscope; he should have computer time available; and he
should have somebody to do some of the routine jobs, such as mounting
specimens, making slides, keeping literature up-to-date, etc., etc., then
perhaps he could do more and better primary systematics or even do large
revisions, phylogenies, etc.

Literature Cited

Arnold, J. W. 1972a. A comparative study of the haemocytes (blood cells) of cock-
roaches (Insecta: Dictyoptera: Blattaria), with a view of their significance in
taxonomy. Can. Entomol. 104:309—348.

——. 1972b. Haemocytology in insect systematics: the prospect. Can. Entomol.
104:655-659.

Assem, Van den, J., and G. D. E. Povel. 1973. Courtship behavior of some Muscidi-
furax species (Hym., Pteromalidae): a possible example of a recently evolved
isolating mechanism. Neth. J. Zool. 23:465—487.

Blackwelder, R. E. 1967. Taxonomy. John Wiley & Sons, Inc., New York. x, 470 pp.

Buren, W. F. 1970. Revisionary studies on the taxonomy of the imported fire ants.
J. Ga. Entomol. Soc. 7:1-25.

Bush, G. L. 1966. The taxonomy, cytology, and evolution of the genus Rhagoletis
in North America (Diptera, Tephritidae). Bull. Mus. Comp. Zool. 134(no. 11):
431-562.

Cain, A. J. 1963. Animal species and their evolution. Hutchinson & Co., London.
ix, 190 pp.

Caws, P. 1965. The philosophy of science, a systematic account. Van Nostrand,
Princeton, N.J. xii, 354 pp.

Foote, R. H. (in press). Museums, museum data, and the real world—the taxonomic
connection. Proc. BARC Symposium II, Beltsville, Md., 1977.

Heiser, C. B., Jr. 1966. Methods in systematic research. Bioscience 1966(Jan.):31-34.

Hennig, W. 1966. Phylogenetic systematics. Univ. Ill. Press, Urbana, Illinois. 263 pp.

International Conference on Systematic Biology, 1967. 1969. Systematic Biology,
Proceedings of an International Conference, etc. Natl. Acad. Sci., Washington,
D.C. xiii, 632 pp.

Kogan, M., and E. F. Legner. 1970. A biosystematic revision of the genus Muscidi-
furax (Hymenoptera: Pteromalidae) with descriptions of 4 new species. Can.
Entomol. 102:1268—1290.

Mayr, E. 1969. Principles of systematic entomology. McGraw-Hill Book Co., New
York. xi, 428 pp.

Michener, C. D. 1970. Diverse approaches to systematics. In Dobzhansky, T., M. K.
Hecht, and W. C. Steere eds., Evolutionary biology, Vol. 4. Appleton-Century-
Crofts (Educ. Div./Meredith Corp.), New York. ix, 312 pp., 2 unnumbered pls.
Reference on pp. 1-38.

50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ross, H. H. 1974. Biological systematics. Addison-Wesley Publ. Co., Inc., Reading,
Mass. (vi), 345 pp.

Simpson, G. G. 1945. The principles of classification and a classification of mammals.
Bull. Am. Mus. Nat. Hist. 85:1-350.

——. 1961. Principles of animal taxonomy. Columbia Univ. Press, New York.
xii, 247 pp.

Steyskal, G. C. 1974. A new species of Procecidochares (Diptera: Tephritidae) caus-
ing galls of stems of hamakua pamakani (Ageratina riparia: Asterceae) in Hawaii.
U.S. Dept. Agric., Coop. Econ. Ins. Rept. 24(32):639-641.

Systematic Entomology Laboratory, ITBIII, Agric. Res. Serv. USDA, c/o
U.S. National Museum, Washington, D.C. 20560.

Footnote

1 Address of retiring 1976 President, delivered at the Society meeting held on 3 Feb-
ruary 1977.

Announcement

The Second International Working Conference on Stored-Product En-
tomology will be held in Ibadan, Nigeria, 10-16 September 1978 at the
Conference Centre on the campus of the University of Ibadan.

The purpose of the Conference is to provide a forum for intellectual and
practical discussions on current research and future research needs in the
context of the World Food Economy.

For information, inquiries should be addressed to:
Organizers
Working Conference on Stored-Product Entomology
c/o Director, Institute of Agricultural Research and Training
F.M.B. 5029, Moor Plantation
Ibadan, Nigeria

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 51-55
TWO NEW SPECIES OF PHLEBOSOTERA DUDA
(DIPTERA: ASTEIIDAE) FROM NORTHERN NIGERIA

J. C. Deeming

Abstract—Two new species of the genus Phlebosotera, P. sabroskyi and
P. inuwa, are described from northern Nigeria.

The genus Phlebosotera was erected by Duda (1927:125) to accommodate
his new species mollis from Cyprus. Sabrosky (1956:234) modified the
generic diagnosis and (1956:235) gave a key to the Old World species, which
included four Palaearctic and two Ethiopian species. Frey (1958:48) de-
scribed a further species from the Canary Islands. A further three species
are known from the Nearctic (Sabrosky, 1943:511 and 1957:50-51) and
one from the West Indies (Sabrosky, 1957:51). Two new species are here
described from Nigeria. Both species, like the type-species, have the male
postabdomen asymmetrical and the aedeagus very large, heavily sclerotized
and of a complicated structure.

Phlebosotera sabroskyi Deeming, new species
(Fig. 1-6)

Male.—Head yellow with all hairs and bristles yellow; frons rather
brownish tinged, in profile (Fig. 2) hardly raised above eye, with slightly
sunken, subshining but ill-defined greyish-white stripe extending forwards
from internal vertical bristle almost to level of anterior ocellus and slightly
raised area bearing ocelli matt black; facialia and jowls slightly infuscate,
but ground color obscured by coarse whitish-grey dust; pair of minute in-
fuscate porelike spots present on face at slightly above % its height; occiput
greyish dusted; eye with short pale hairs; antenna yellow with part of
inner surface of 3rd segment brownish and arista black; palpus yellow with
hairs of moderate length; proboscis brown. Thorax yellow, coarsely grey
dusted dorsally, more weakly dusted laterally and ventrally; mesopleuron
with long shining brownish-black mark along its lower border; postnotum
brownish black; bristles and hairs of mesonotum and scutellum dark brown,
single sternopleural bristle and fine hairs on mesopleuron and_ sterno-
pleuron yellow; prosternum (Fig. 4) joined to mesosternum and linked
to widely-separated fore coxae by membranous bridge. Wing (Fig. 5) yel-
lowish hyaline with yellow veins; haltere yellow. Legs yellow, weakly yel-
lowish dusted and with yellow hairs. Abdomen yellow, basally appearing
a little darker, weakly sclerotized and weakly yellowish dusted, tergites
bearing short brown hairs; terminalia (Fig. 6). Length about 1.75 mm, of
wing 1.90 mm.

52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Yr
VOID TPT WWD TT OTT

Figs. 1-6. Phlebosotera sabroskyi, male; 1. Head from in front; 2. Head in profile;
3. Thorax from above; 4. Prosternum, mesosternum and inner margin of fore coxae; 5.
Wing; 6. Terminalia from beneath.

Female.—Unknown.

Holotype —2, N. NIGERIA: Katsina, 24.v.1975, (J. C. Deeming), on
Ficus foliage. Deposited in USNM No. 73939. This species is closely related
to P. cyclops Sabrosky 1956:236, but differs from it in the following respects:

VOLUME 80, NUMBER 1 53

Figs. 7-12. Phlebosotera inuwa, male; 7. Head in profile; 8. Thorax from above; 9.
Prosternum and inner margin of fore coxae; 10. Wing; 11. Terminalia from behind;
12. Terminalia from beneath.

Only a single dark pleural spot; vertical bristles greyish white, unlike the
more or less infuscate mesonotal bristles; distance between posterior ocelli
slightly less than that between posterior ocellus and eye margin; frons
between internal vertical bristles slightly less than the width of an eye; the

54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

rows of acrostical hairs, though irregular, becoming further apart post-
suturally, on the posterior % of mesonotum more widely separated from
each other than from the dorsocentral lines; wing with 2nd vein very greatly
dilated apically and 4th vein straighter, the 1st posterior cell at its widest
point less than 3X its width at wing tip.

Phlebosotera inuwa Deeming, new species
(Fig. 7-12)

Male.—Head ivory white, very weakly dusted; a large U-shaped brown
mark on occiput extending from neck to base of internal vertical bristle
on either side, this connected above to pair of rugose yellow bands that fill
frons on either side of ocellar triangle and extend forwards, becoming
progressively narrower, tapering to points *% of the way from anterior
ocellus to anterior margin of frons; ocellar prominence matt black; vertical
bristles, minute ocellars and double row of minute hairs behind ocellar
prominence black, minute hairs clothing frons brown and vibrissa and
jowlar bristles yellow; eye very sparsely, short pale haired; antenna yellow
with arista and hairs ringing 2nd antennal segment black; palpus yellow,
apically with long yellowish-brown bristle about % as long as palpus; pro-
boscis yellow with yellow hairs. Thorax yellowish white, subshining through
very weak yellowish dust and (Fig. 8) with brown and yellowish-brown
markings on mesonotum, postnotum brown and faint indication of brown
spot in front of base of single pale sternopleural bristle, mesonotal and
scutellar bristles black and pleural hairs yellow; prosternum (Fig. 9)
separate from mesosternum and only narrowly separated from the fore
coxae. Wing. (Fig. 10) yellowish hyaline with yellow veins; haltere yellow.
Legs yellow with yellow hairs, apical 3 segments of fore tarsus infuscate.
Abdomen yellow, subshining, weakly sclerotized and bearing numerous
brown to yellowish-brown hairs; terminalia (Figs. 11-12) large. Length
about 1.6 mm, of wing 2.0 mm.

Female.—Unknown.

Holotype —é, N. NIGERIA: Zaria, Samaru, 9.iii.1973, (J. C. Deeming).
Deposited in USNM No. 73938.

Paratype—é,N. NIGERIA: Katsina Prov., Maska Fish Farm, 26.iii.1972,
(J. C. Deeming). Paratype deposited in the Inst. Agric. Res., Samaru.

Etymology.—The specific name is taken from the Hausa word for shade
or shadow, as both specimens were collected hovering in the shade of
large trees (mango and Parkia) on days of exceptional heat.

Discussion —This species is probably more closely related to P. nigriseta
Sabrosky (1956:237) than to any other described species, but it differs from
P. nigriseta in lacking large black spots on the meso-, sterno-, and hypopleura
and small spots in front of and behind the anterior (mesothoracic) spiracle,
and in having the sternopleural bristle pale and the tergites yellow.

VOLUME 80, NUMBER Il 55

Acknowledgments

My thanks are due to Mr. J. P. Dear of the British Museum (Natural
History) for allowing me to dissect the male that was tentatively identified
by Sabrosky (1956:236) as P. mollis Duda and to the Director, Institute for
Agricultural Research, Samaru for the facilities for research.

Literature Cited

Duda, O. 1927. Revision der altweltlichen Astiidae (Dipt.). Deut. Entomol. Ztschr.
1927:113-147.

Frey, R. 1958. Entomologische Ergebnisse der finnlindischen Kanaren-Expedition
1947-51, No. 15. Kanarische Diptera Brachycera p.p. von Hakan Lindberg
gesammelt. Comment Biol. 17(4):1-63.

Sabrosky, C. W. 1943. New genera and species of Asteiidae (Diptera), with a re-
view of the family in the Americas. Ann. Entomol. Soc. Am. 36:501-514.
———. 1956. Additions to the knowledge of Old World Asteiidae (Diptera). Rev.

France. Entomol. 23(4):216-243.

—. 1957. Synopsis of the New World species of the dipterous family Asteiidae.

Ann. Entomol. Soc. Am, 50:43-61.

Institute for Agricultural Research, Samaru, P.M.B. 1044 Zaria, Nigeria.

NOTE

A CASE OF HOMONYMY IN THE GENUS MACHIMUS
(DIPTERA: ASILIDAE)

Oldroyd described Machimus truncatus (1972. Pac. Insects. 14(2):289-290)
as a new species based on specimens collected on the island of Luzon in
the Philippines. Subsequently, Martin described a new species of Machimus,
which he also named truncatus (1975. Occ. Pap. Calif. Acad. Sci. No. 119:
41-42) from Cuernavaca, Mexico. Based on the published descriptions,
it is apparent that two different species are involved. Consequently, ac-
cording to Article 53 of the International Code of Zoological Nomenclature
(1961), the junior primary homonym, M. truncatus Martin, must be rejected
and replaced. Therefore, I propose that the name Machimus truncatus
Martin be replaced by the name Machimus lurettae Martin in honor of the
late Dr. Charles Martin’s second wife, Luretta Martin, who assisted him
greatly in preparing his later papers for publication.

Published with the approval of the Director, Wyoming Agricultural Ex-
periment Station, as Journal Paper no. JA-929.

Robert J. Lavigne, Entomology Section, Plant Science Division, Box
3354, Univ. Stn., University of Wyoming, Laramie, Wyoming 82071.

PROC. ENTOMOL. SOC. WASH.

80(1), 1978, pp. 56-68
THE VELIIDAE (HETEROPTERA) OF AMERICA
NORTH OF MEXICO—KEYS AND CHECK LIST

Cecil L. Smith and John T. Polhemus

Abstract.—Keys, supplemented with scanning electron micrographs and
drawings, are presented for the five genera and 35 species of Veliidae of
North America north of Mexico. Distributions and synonymies are given
in a check list. A selected bibliography pertaining to these taxa is in-

cluded.

Although the veliids are by far the most abundant of the surface-in-
habiting Heteroptera and second only to the Gerridae in species diversity,
they are usually ignored by the average collector and are poorly repre-
sented in most collections. This relative obscurity is due primarily to their
small size (1-12 mm) and their penchant for living in cryptic habitats. Only
the Rhagoveliinae are found on open stretches of water—Rhagovelia below
the ripples and eddies of streams and Trochopus along shorelines of tropical
bays and estuaries. The microveliine genus Husseyella shares the marine
habitat with Trochopus and inhabits the margins of salt marshes and sim-
ilar niches. In general, Microvelia are found on or near the margins of
practically all still waters—ponds, lakes, temporary puddles, and the quieter
portions of streams and rivers, where they secrete themselves among the
debris, venturing out onto the open water rarely. Unlike the Rhagoveliinae,
members of this genus are not totally restricted to the water surface and
are commonly found running over nearby rocks and mud flats. Even though
individuals of the veliine genus Paravelia (= Velia of authors) are the largest
of the North American veliids, they are less often encountered because they
do not usually inhabit the water surface, but are often found, instead, on
emergent vegetation, and occasionally on vegetation some distance from
water.

There are comparatively few veliid species in North America: Five
genera with 35 included species. Of these, only the genus Rhagovelia has
been adequately monographed (Bacon, 1956). The key to Rhagovelia pre-
sented here has been adapted, in part, from his paper.

The Western Hemisphere Microvelia were reviewed by Torre-Bueno
(1924a) and later revised by McKinstry (1933, unpublished). Torre-Bueno’s
paper encompassed only 22 of the currently valid 81 New World species. In
addition to Torre-Bueno’s key several regional keys to Microvelia have
since been published (Bobb, 1974; Froeschner, 1962; Herring, 1950; Usinger,
1956; Wilson, 1958). Usinger’s 1956 key to the Microvelia of California was
largely based on McKinstry’s unpublished work. A check list compiled by

VOLUME 80, NUMBER 1 57

Drake and Hussey (1955) is the most recent synoptic work on the genus.
Smith has a monographic revision of the North and Central American Micro-
velia in progress, but since it will be some time before the results are pub-
lished, the key presented here will hopefully suffice as an identification
aid in the interim. The subgenera Microvelia (Microvelia) and M. (Kirkaldya)
are separated by the vertically oriented, leaf-like, dorsal arolium found only
in the latter. This arolium is often difficult to see, so it is not used as a
character in the key. Because no consistent characters have yet been
found for successfully separating alate individuals of Microvelia sensu lato
or females of the subgenus Kirkaldya, the key is based primarily on apterous
specimens and males of Kirkaldya species. However, this should result in
only minimal inconvenience because the majority of specimens within most
populations are apterous and males are present. The elimination of winged
forms from the key has necessitated the omission of one species, M. mar-
ginata Uhler. This species is known in the U.S. only from a single alate
specimen from Key West, Florida. M. marginata is prevalent throughout
the Caribbean area, and this single U.S. record is probably based on a
hurricane transported specimen. A similar occurrence following a tropical
storm has been verified by Herring (1958) for M. cubana (= M. portoricensis
sensu Herring, not Drake). The latter species quite possibly has adapted
to and become established in southern Florida based on several collection
records subsequent to Herring’s original observation.

Other than basic descriptive work and a few extremely localized regional
keys, the genus Paravelia has, until recently, been ignored. Members of
this New World genus were considered congeneric with the Old World
genus Velia until Polhemus (1976) revised their status. Generic concepts,
distributions, and synonymies were reviewed by Polhemus but no compre-
hensive keys exist for the genus.

Key to the Genera of Veliidae of America North of Mexico

1. Middle tarsi deeply cleft, with leaflike claws and plumose hairs aris-

ing from base of cleft (Fig. 1). Rhagoveliinae 2
— Middle tarsi not deeply cleft and without plumose hairs arising
from base of cleft 3
2. Hind tarsi 2-segmented, the basal segment very short. Apterous.
Tropical America. Marine Trochopus Carpenter
[One species, T. plumbeus (Uhler) |
— Hind tarsi 3-segmented, the basal segment very short. Apterous
or macropterous. Cosmopolitan. Riffles of streams and rivers or
(rarely) lakes Rhagovelia Mayr
3. Tarsal formula 1:2:2. Microveliinae 4

— Tarsal formula 3:3:3. Veliinae Paravelia Breddin

58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

as "ke SOLE )
Figs. 1-3. Tarsi. 1. Rhagovelia obesa (61x); 2. Microvelia americana (293);
3. Husseyella turmalis (117). Figs. 4-5. Ventral view of female Microvelia (31x). 4.
M. pulchella; 5. M. americana.

a

4. Middle tarsi with 4 leaflike blades arising from cleft (Fig. 3)
Husseyella Herring
[One species, H. turmalis (Drake and Harris) |
- Middle tarsi with narrow claws arising from cleft (Fig. 2)
Microvelia Westwood

Key to Apterous Microvelia of America North of Mexico

[Except for species keyed in couplets 8 through 13, all species belong to

the subgenus Microvelia (Microvelia).]

1. Pronotum covering entire thorax to metathoracic triangles, dorsal
surface of thorax appearing 1-segmented (Fig. 6) 2

— Pronotum not covering entire thorax, at least 1 other thoracic seg-
ment exposed (Figs. 7, 8) 6

VOLUME 80, NUMBER 1 59

2.

Minute, white, vestigial wing pads present; only 6 entire abdominal

terga visible atrata Torre-Bueno
Minute, white, vestigial wing pads absent; 7 entire abdominal terga
visible 3

Blue to gray pruinose patches present on abdominal terga 2 and
3, and, usually, 6 and 7; orange-brown to dark reddish-brown
species 4
Blue to gray pruinose patches absent on all abdominal terga; color
uniform sooty black (except orange transverse band across anterior
lobe of pronotum) 5
Dorsum entirely covered with long erect pubescence, the majority
of hairs equal in length to width of hind femur; 2nd genital seg-
ment of male without laterally directed caudal spines

fontinalis Torre-Bueno
Dorsum with only short pubescence, closely appressed to body;
2nd genital segment of male with a pair of short laterally projecting
caudal spines cerifera McKinstry
Distal segment of middle tarsi 20% longer than proximal segment;
coxae dark brown; females with glabrous depressed area on lateral
margin of pronotum glabrosulcata Polhemus
Distal segment of middle tarsi subequal to proximal segment; coxae
yellowish; female lacking glabrous depressed area on lateral margin

of pronotum austrina Torre-Bueno
Pronotum short; dorsal surface of thorax apparently consisting of 3
segments (Fig. 8) i
Pronotum longer; dorsal surface apparently 2-segmented (Fig.
) 14

Hind tibiae of males curved; females with wide groove between
front coxae for reception of rostrum, interior edges sloped grad-
ually, divergent posteriorly (Fig. 4); front coxae widely separated;
length normally less than 2 mm (1.25-2.25 mm) pulchella Westwood
Hind tibiae of males straight; females with narrow groove, barely
wider than rostrum, interior edges more vertical, parallel (Fig.
5); front coxae close together; length greater than 2 mm (2.00-

3.75 mm) (subgenus Kirkaldya—key for males only) 8
Distal ventral margin of Ist genital segment glabrous or with only
very short pubescence (Figs. 11-14) 9

Distal ventral margin of Ist genital segment with either a row or
tufts of long hairs (Figs. 15-19) (note that the tuft of hairs on M.
americana may occasionally be shortened and, thereby, obscured) 10
Distal ventral margin of Ist genital segment with a triangular
glabrous area (Fig. 11) torquata Champion

60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

SS
oe

SS

mm

Figs. 6-8. Dorsal thoracic segments—Microvelia. 6. M. fontinalis; 7. M. signata; 8.

M. americana. Figs. 9-10. Terminal ventral abdominal segments—Paravelia. 9. P.
summersi; 10. P. alvaradana.

— Posterior ventral margin of Ist genital segment with only a narrow,
transverse, glabrous area (Figs. 12-14) paludicola Champion

10. Hairs on ventral surface of Ist genital segment arranged in 2
fairly distinct lateral tufts (Figs. 18, 19) 11

VOLUME 80, NUMBER 1 61

Shas, Le
Aaa

los,

Figs. 11-19. Microvelia (Kirkaldya) male genital capsules (56). 11. M. torquata;
12. M. paludicola—Miss.; M. paludicola—Ga.; 14. M. paludicola—Tex.; 15. M.
americana; 16. M. gerhardi; 17. M. californiensis; 18. M. beameri; 19. M. fasciculifera.

— Hairs on ventral surface of Ist genital segment aligned in a con-
tinuous pattern (Figs. 15-17) 12
11. Venter of last abdominal segment with a short erect tubercle
fasciculifera McKinstry
— Venter of last abdominal segment without tubercle | beameri McKinstry
12. Distal ventral margin of Ist genital segment with a raised trans-

verse ridge (Figs. 16, 17) 13
— Distal ventral margin of Ist genital segment without a raised

transverse ridge (Fig. 15) americana (Uhler)
13. Middle of front femora distinctly swollen and blackened; inner

surface of front femora flattened gerhardi Hussey
— Middle of front femora not swollen and blackened; inner surface

not flattened californiensis McKinstry

14. Antennal segment IV subequal (90%+) to width of head through
eyes; middle % of 2nd abdominal tergum with darkened area;
males with large acute tubercle on 2nd abdominal sternum

albonotata Champion
— Antennal segment IV not longer than 75% of width of head through
eyes; 2nd abdominal tergum without darkened area; males without
ventral tubercle 15

15. Last 3 abdominal terga with broad shining areas, covering 25-90%

of at least 1 segment 16

62

16.

li

to

bo

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Last 3 abdominal terga with at most a thin, shiny, medial line
present on last 2 segments (width of line, at most, 10% of maximum

width of segment) 17
Silvery pubescence present on head around eyes and base of an-
tennae, and shoulders of pronotum buenoi Drake
Silvery pubescence absent on head around eyes and base of an-
tennae, and shoulders of pronotum signata Uhler
Tibia of mesothoracic leg subequal in length (90%+) to width of
head through eyes cubana Drake
Tibia of mesothoracic leg less than 75% as long as width of head
through eyes hinei Drake

Key to Parvelia of America North of Mexico
(= Velia of American Authors)

Body slender; genital segments of both sexes acuminate; wing pads
or wings basally brown, without white markings

stagnalis (Burmeister)
Body robust; genital segments button-like (2?) or forming a blunt
angulate projection (4) but not acuminate; wing pads or basal wing

spot white, conspicuous 2
Collar projecting, angulate behind eyes; pronotum not set off from
collar by conspicuous row of pits beameri (Hungerford)
Collar not projecting, not angulate behind eyes; pronotum set off
from collar by a more or less conspicuous row of pits 3

First antennal segment approximately 1.25 as long as width of head
through eyes, and 1.5 as long as 2nd antennal segment (south-
eastern U.S.) brachialis (Stal)
First antennal segment approximately 1.5 as long as width of head
through eyes, and 2x as long as 2nd antennal segment (south-
western U.S.) 4
Ventral projection of 7th abdominal sternum blunt (Fig. 9)
summersi (Drake)

Ventral projection of 7th abdominal sternum acuminate (Fig. 10)

alvaradana (Drake and Hottes)

Key to Rhagovelia of America North of Mexico

Genital segments in both sexes mucronate; female connexiva pro-
duced caudad from 7th tergum as long mucronate processes

becki Drake and Harris
Genital segments bluntly rounded or triangular but not mucronate;
female connexiva not produced caudad as long mucronate pro-
cesses 2
Female midfemur transversely constricted at middle; males with-

VOLUME 80, NUMBER 1 63

out median shining areas on dorsum of abdominal segments

choreutes Hussey

— Female midfemur may be dorsoventrally flattened but not trans-

versely constricted at middle; males with at least 7th (last) abdominal
segment having median dorsal shining area 3

3. Female midfemur dorsoventrally flattened; connexiva reflexed for

the last 4 segments. Males with dorsal median shining areas on at

least the last 3 abdominal segments; if shining areas are small,
posterior femur is not greatly incrassate 5

— Female midfemur not flattened; abdomen tapering evenly to apex,

connexiva not reflexed. Males with dorsal median shining areas on
only abdominal segment 7; posterior femora greatly incrassate 4

4. Venter of abdomen orange brown over at least last 3 segments,

thickly dotted with minute black conical setae. Males with abdominal
venter carinate over Ist 3 segments varipes Champion

— Venter of abdomen blackish grey over all but 7th segment, without

evident minute black setae. Male abdominal venter not carinate
torreyana Drake and Hussey

5. Apterous female with apex of pronotum continued into a long, ele-

vated process. Apterous male pronotum triangular, its apex extend-

ing over metanotum, mesonotum exposed at sides —_ oriander Parshley
— Apterous female with pronotum not produced at apex. Pronotum
of apterous male not extending over metanotum 6

6. Connexiva of apterous female diverging over apex of last abdominal
segment, apex of connexiva rounded as seen from side. Venter of
last abdominal segment of male flattened medially with prominent
hairy ridges at each side distincta Champion

— Connexiva of apterous female close together or meeting at apex;
apex of connexiva forming at least a 90° angle as seen from side.
Male venter not flattened and without hairy ridges on 7th seg-
ment 7

7. Abdominal dorsum of apterous male with only traces of median
shining areas on a few segments in addition to segment 7. Meso-
notum of apterous female tumid rivale Torre-Bueno

— Abdominal dorsum of apterous male and dorsum of metanotum with
broad median shining areas; posterior trochanter of male armed
with several small teeth. Mesonotum of apterous female tumid
only at sides or not tumid 8

8. Connexival margins of apterous female with lst 2 segments curved;
exposed portion of mesonotum longer than exposed portion of
metanotum obesa Uhler

— Connexival margins of apterous female with Ist 2 segments straight;
exposed portion of mesonotum shorter than exposed portion of
metanotum knighti Drake and Harris

64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Check List—Veliidae of America North of Mexico
Genus Husseyella Herring 1955

turmalis (Drake and Harris) 1933 Southern Florida,
Mexico, Caribbean,
Middle America

Genus Microvelia Westwood 1834

albonotata Champion 1898 Canada, US. east of
Rocky Mts., Mexico
to Peru, Caribbean

americana (Uhler) 1884 Eastern U.S., west to
Nebraska and Texas

atrata Torre-Bueno 1916 Georgia and Florida,
west to Louisiana

austrina Torre-Bueno 1924a Southeastern U.S.,

parallela Blatchley 1925 Mexico

beameri McKinstry 1937 Southwestern U.S.,
Northern Mexico,
Jamaica

buenoi Drake 1920a Northern half of U.S.,
California, Canada, Alaska,
( Florida? )

californiensis McKinstry 1937 California, Oregon,

Baja California

cerifera McKinstry 1937 Iowa, Kansas, Nebraska,
Colorado, New Mexico,
Utah, Arizona, Nevada,

California

cubana Drake 195la Southern Florida, Cuba,
Dominican Republic

fasciculifera McKinstry 1937 Texas, New Mexico,
Arizona, Mexico

fontinalis Torre-Bueno 1916 U.S. east of Mississippi
River

gerhardi Hussey 1924 Western U.S.,

americana Uhler 1895 Northern Mexico
glabrosulcata Polhemus 1974 Arizona, Mexico

hinei Drake 1920b Canada to Argentina

VOLUME 80, NUMBER 1

marginata Uhler 1893
pudoris Drake and Harris 1936

paludicola Champion 1898
alachuana Hussey and Herring 1950

pulchella Westwood 1834
capitata Guérin-Meneville 1857
borealis Torre-Bueno 1916
incerta Kirby 1890
robusta Uhler 1894

signata Uhler 1894b
oreades Drake and Harris 1928
setipes Champion 1898

torquata Champion 1898

65

Southern Florida,
Caribbean, Mexico,
Panama, Venezuela, Peru
Southeastern U.S.,
Kentucky, Tennessee,
Arkansas, Oklahoma,
Kansas, Texas, New
Mexico, Mexico,

Middle America, Caribbean
Canada to Argentina,
Caribbean

Southwestern U.S.,
Mexico, Middle America

Southwestern U.S.,
Mexico, Middle America

Genus Paravelia Breddin 1898

alvaradana (Drake and Hottes) 1952

beameri (Hungerford) 1929
brachialis (Stal) 1860
australis (Torre-Bueno) 1916
stagnalis (Uhler) 1894
(nec. Burmeister )
stagnalis (Burmeister) 1835
paulineae (Wilson) 1953
watsoni (Drake) 1919

summersi (Drake) 1951b

Northwestern Mexico
(Arizona?)

Arizona

Southeastern U.S.,
Oklahoma, Texas,
Mexico, to (?) Argentina,
Caribbean

Eastern U.S., Cuba

Arizona

Genus Rhagovelia Mayr 1865

becki Drake and Harris 1936
choreutes Hussey 1925

distincta Champion 1898
excellentis Drake and Harris 1927

mexicana Signoret 1877 (nomen nudum)

Texas, Nevada, Mexico
Southeastern U.S. to New
Mexico, California
Western U.S., Mexico,
Middle America

66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

d. arizonensis Gould 1931
d. cadyi Gould 1931

d. harmonia Gould 1931
d. modesta Gould 1931
d. proxima Gould 1931

d. valentina Gould 1931

knighti Drake and Harris 1927 Arkansas, Missouri,
Oklahoma
obesa Uhler 1871 Eastern U.S.,
arctoa Torre-Bueno 1924b Southeastern Canada
flavicincta Torre-Bueno 1924b
oriander Parshley 1922 Midwestern U.S.
rivale Torre-Bueno 1924b Colorado, Kansas, Iowa,

Missouri, Oklahoma,
Nebraska, Texas

torreyana Drake and Hussey 1957 Western Florida
varipes Champion 1898 Arizona, New Mexico,
beameri Gould 1931 Mexico

Genus Trochopus Carpenter 1898

plumbeus (Uhler) 1894a Florida (Gulf and
marinus Carpenter 1898 Atlantic Coasts),
Caribbean, Atlantic
coasts of Mexico,
Middle America and
Northern South America

Acknowledgments

We would like to extend our gratitude to Ms. Diane A. Hurd, of the
University of Georgia, for her kind assistance in the preparation of the scan-
ning electron micrographs, and to Drs. H. H. Ross and W. T. Atyeo, also
of the University of Georgia, for their most helpful suggestions on improv-
ing the manuscript.

Literature Cited

Bacon, J. A. 1956. <A taxonomic study of the genus Rhagovelia (Hemiptera, Veliidae)
of the western hemisphere. Univ. Kans. Sci. Bull. 38, Pt. 1(10):695-913.

Bobb, M. L. 1974. The insects of Virginia: No. 7. The aquatic and semi-aquatic
Hemiptera of Virginia. Research Div. Bull. 87, Va. Polytech. Inst. and State
Univ.: 1-195.

Burmeister, H. 1835. Handbuch der Entomologie. Vol. 2. Hemiptera. Berlin.

VOLUME 80, NUMBER 1 67

Champion, G. C. 1898. Biologia Centrali-Americana, Insecta, Rhynchota (Hemip-
tera-Heteroptera). Vol. II:117-158, 338-383.

Drake, C. J. 1920a. An undescribed water-strider from the Adirondacks. Bull.

Brooklyn Entomol. Soc. 15(1):19-21.

1920b. Water striders new to the fauna of Ohio, including the description
of a new species. Ohio J. Sci. 20(6):205-208.

—. 195la. New Neotropical water-striders (Hemiptera-Veliidae). Great Basin

Nat. 11(1-2):37—42.

1951b. New water striders from the Americas (Hemiptera: Veliidae).
Revista Entomol. Rio de Janeiro 22(1—3):371-378.

Drake, C. J., and H. M. Harris. 1927. Notes on the genus Rhagovelia, with de-
scriptions of six new species. Proc. Biol. Soc. Wash. 40:131-138.

—. 1933. New American Veliidae (Hemiptera). Proc. Biol. Soc. Wash. 46:45-53.

1936. Notes on American water-striders. Proc. Biol. Soc. Wash. 49:105—108.

Drake, C. J., and F. C. Hottes. 1952. Concerning some Mexican Veliidae (Hemiptera).
Proc. Biol. Soc. Wash. 65:85-88.

Drake, C. J., and R. F. Hussey. 1955. Concerning the genus Microvelia Westwood,
with descriptions of new species and a checklist of the American forms (Hemip-
tera: Veliidae). Florida Entomol. 38(3):95-115.

1956. Notes on some American Rhagovelia, with descriptions of two new
species (Hemiptera: Veliidae). Occ. Pap. Mus. Zool. Univ. Mich. 580:1-6.
Froeschner, R. C. 1962. Contributions to a synopsis of the Hemiptera of Missouri,

Part V. Hydrometridae, Gerridae, Veliidae, Saldidae, Ochteridae, Gelastocoridae,
Naucoridae, Belostomatidae, Nepidae, Notonectidae, Pleidae, Corixidae. Amer.
Midl. Nat. 67(1):208-240.

Herring, J. L. 1950. The aquatic and semiaquatic Hemiptera of northern Florida.

Part 2: Veliidae and Mesoveliidae. Florida Entomol. 33(4):145—150.

1955. A new American genus of Veliidae (Hemiptera). Florida Entomol.
38(1):21-25.

1958. Evidence for hurricane transport and dispersal of aquatic Hemiptera.
Pan-Pac. Entomol. 34(3):174-175.

Hungerford, H. B. 1929. A new Velia from Arizona with notes on other species
(Hemiptera-Veliidae). Ann. Entomol. Soc. Am. 22(4):759-761.

Hussey, R. F. 1924. A new North American species of Microvelia (Hem.). Bull.
Brooklyn Entomol. Soc. 19(5):164—-165.

—. 1925. Some new or little-known Hemiptera from Florida and Georgia. J.
N.Y. Entomol. Soc. 33:61-69.

McKinstry, A. P. 1933. Preliminary studies in Microvelia of the Western Hemisphere.

M.A. Thesis, Univ. Kansas, Lawrence, 72 pp. + 20 pl. (unpublished).

1937. Some new species Microvelia (Veliidae, Hemiptera). J. Kans. Entomol.
Soc. 10(1-2):30-41.

Parshley, H. M. 1921. On the genus Microvelia Westwood (Hemiptera—Veliidae). Bull.

Brooklyn Entomol. Soc. 16:87-93.

1922. Report on a collection of Hemiptera-Heteroptera from South Dakota.
South Dakota State Coll. Tech. Bull. 2:1-22.

Polhemus, J. T. 1974. The austrina group of the genus Microvelia (Hemiptera;

Veliidae). Great Basin Nat. 34(3):207-217.

1976. <A reconsideration of the status of the genus Parvelia Breddin, with
other notes and a check-list of species (Veliidae: Heteroptera). J. Kans. Entomol.
Soc. 49(4):509-513.

68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

—. 1977. Type-designations and other notes concerning Veliidae (Insecta:
Hemiptera). Proc. Entomol. Soc. Wash. 79(4):637-648.

Stal, C. 1860. Bidrag till Rio Janeiro Traktens Hemipterfauna. Svenika Vet. -Ak.
Handl. 2(7):1-84.

Torre-Bueno, J. R. de la. 1916. The Veliinae of the Atlantic states. Bull. Brooklyn

Entomol. Soc. 11(3):52-61.

1923. Taxonomic characters in Microvelia Westw. Bull. Brooklyn Entomol.
Soc. 18(4):138-143.

———. 1924a. A preliminary survey of the species of Microvelia Westwood (Veliidae,

Heteroptera) of the Western World, with description of a new species from the

southern United States. Bull. Brooklyn Entomol. Soc. 19(5):186-194.

1924b. The Nearctic Rhagoveliae (Heteroptera; Veliidae). Trans. Am.

Entomol. Soc. 50:243-252.

Uhler, P. R. 1871. Notices of some Heteroptera in the collection of Dr. T. W.
Harris. Proc. Boston Soc. Nat. Hist. 14:93-109.

—. 1884. Order VI—Hemiptera: sub-order III—Heteroptera. In Standard Nat-

ural History, I1:249-296. S. E. Cassino & Co., Boston.

1893. A list of the Hemiptera-Heteroptera collected in the island of St.
Vincent by Mr. Herbert H. Smith; with descriptions of new genera and species.
Proc. Zool. Soc. London 1893:705-719.

1894a. On the Hemiptera-Heteroptera of the island of Granada, West In-

dies. Proc. Zool. Soc. London 1894:167—224.

—. 1894b. Observations upon the heteropterous Hemiptera of Lower Cali-
fornia, with descriptions of new species. Proc. Calif. Acad. Sci., 2nd Ser., 4:
223-295.

Usinger, R. L. (ed.) 1956. Aquatic Insects of California. Univ. Calif. Press, Berkeley.
508 pp.

Westwood, J. O. 1834. Mémoire sur les genres Xylocoris, Hylophila, Microphysa,
Leptopus, Velia, Microvelia et Hebrus, avec quelques observations sur les Am-
phibicorisae de M. Dufour et sur I’état imparfait, mais identique de certaines
espéeces. Ann. Entomol. Soc. Fr. (1)3:637-653.

Wilson, C. A. 1958. Aquatic and semiaquatic Hemiptera of Mississippi. Tulane
Studies Zool. 6(3):115-170.

(CLS) Department of Entomology, University of Georgia, Athens, Georgia
30602; and (JTP) 3115 S. York, Englewood, Colorado 80110.

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 69-74
PARASITISM OF ADULT TABANUS SUBSIMILIS SUBSIMILIS
BELLARDI (DIPTERA: TABANIDAE) BY A MILTOGRAMMINE
SARCOPHAGID (DIPTERA: SARCOPHAGIDAE)

Patrick H. Thompson

Abstract——Larvae of Macronychia sp. near aurata (Coquillett) were
taken from adults of Tabanus subsimilis subsimilis Bellardi which were
collected near College Station, Texas in 1976. Multiple infestations of
this tabanid suggest specific host selection by sarcophagid females. This
host-parasite association is apparently the first described instance of parasit-
ism between Sarcophagidae and Tabanidae; it is a rarity between a dip-
teran species and an adult tabanid. Also, this account illuminates a pre-
viously unrecognized instance of such parasitism in the Miltogrammini, a
tribe of cleptoparasites characteristically attacking the immobilized insect
prey of wasps and bees. Rearing methods, development, and larval be-
havior are described, as are several additional observations of sarcophagid
parasitism of Tabanidae.

Immature stages of Tabanidae have been infested by dipteran parasites
representing four genera of Tachinidae and one of Bombyliidae. Good-
win (1968) summarized the literature relating to the initial reports and de-
scriptions of these six species of parasites and added a seventh species to
the list, which also parasitized the larval stage. More recently, Spratt and
Wolf (1972) found tachinid larvae (Bactromyiella sp.) in adults.

The following account describes an unusual host-parasite association
between a dipteran species and an adult tabanid; and, apparently, docu-
ments the first such association between Sarcophagidae and Tabanidae.
In addition, this account elucidates a previously unrecognized host relation-
ship of the genus Macronychia in North America and describes an unusual
case of primary parasitism in the Miltogrammini, a tribe of Sarcophagidae
characteristically cleptoparasitic upon the provisions in Hymenoptera nests.
(The term primary, in this case, refers to that usage defined by Allen (1926),
and quoted in the conclusions of this paper.) The species discussed here
is Macronychia sp. near aurata (Coquillett) and is hereafter referred to as
Macronychia sp.

During anatomical dissection of a series of Tabanus sulcifrons Macquart
adults collected in 1975, a 1.5 mm first-instar larva of Sarcophagidae was
discovered in each of two specimens. Both parasites were located within
the abdominal cavity of the host near the terminal abdominal segments.
These T. sulcifrons flies had been frozen the week before (17 July), at which

70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

time they were removed from a Gressitt Trap located in the Navasota
River bottoms near College Station.

A second association of adult horse flies with sarcophagid larvae found
several weeks later was more remote. During separation of a wet 2-day-
old Gressitt Trap catch from this same locality, another first-instar larva
representing a second species of this family, was found on the wing of a
Tabanus nigripes Wiedemann female.

A third association provided adult specimens documenting this parasitism
in the Tabanidae, when unexplained cyclorrhaphous larvae were observed
in a glass beaker containing adults of Tabanus subsimilis subsimilis Bellardi
and T. mularis Stone; these insects were being held for later treatment with
insecticides. Seven larvae had emerged, then, through the ruptured venter
of one of two dead T. s. subsimilis females, and an eighth larva remained
in the abdominal cavity of that insect. On the same day, seven smaller
larvae were observed in a second female of T. s. subsimilis in another
beaker. These were together in a ball near the ruptured abdomen of the
inverted, dead host.

Over the following seven weeks, five more lots of larvae were removed
from more glass beakers and from one-pint cardboard ice cream con-
tainers holding Tabanus spp. females. Most of these containers of five adults
included one or more specimens of T. s. subsimilis. In one of these last
five lots of larvae, two specimens were removed from a third infested female
of this species, this specimen also showing an emergence hole in the abdo-
men ventrally.

Methods

In the first rearing attempts, sarcophagids were implanted within fe-
males of several Tabanus spp., the larger of these, such as T. sulcifrons and
T. atratus F., being most satisfactory for this purpose. After horse flies were
dealated and pinned, sarcophagid larvae were implanted individually
within the ruptured abdominal venter of their hosts; such infested speci-
mens were maintained in 10-dr vials containing damp sand. In later rearings,
field crickets were used as hosts because they were more easily collected
and contained more tissues and fluids. The heads and legs of crickets were
removed, and after larval implantation, the infested insects were stored in
stacked 68-mm Syracuse watch glasses. Sarcophagid pupae were trans-
ferred to 5-dr vials containing 4% in. of damp sand which was topped with
the same depth of fine wood shavings. After their emergence, teneral
adults were held overnight before pinning. Larval specimens from early
large lots of material were killed in KAAD and stored in 95% ethyl alcohol.
However, Sanjean (1957) noted that Dietrich’s Fluid, containing % of the
usual amount of glacial acetic acid, was more satisfactory for this purpose.

VOLUME 80, NUMBER 1 ql

Observations

Development.—Because Macronychia larvae were no less than 6 mm
when first observed, the exact period of development is unknown; how-
ever, the development remaining required two to eight days, with a median
of six days. More favorable observations showed that the pupal period,
for thirteen specimens, required sixteen days (ten larvae) and fifteen or
seventeen days (two and one, respectively); this sixteen-day period for
Macronychia sp. was considerably longer than the nine to twelve days re-
quired by ten sarcophagine species reared in the laboratory at the same
temperatures of 25°C + 1 by Sanjean (1957).

Larval behavior—Once the mouthparts of larvae were exposed to
haemolymph exuding from the host, the maggots began to feed at once.
In two cases, larvae attacked crickets through intact body areas and at-
tempted to penetrate the host through intersegmental membranes. In an-
other instance, one unusually large larva of the eight larvae found in the
first lot voraciously attacked the intact cervical membrane beneath the head
of a living T. s. subsimilis. Except for this one case of aggressive host attack,
larval behavior was scavenger-like, the maggots feeding on the dead
and putrefying bodies of the hosts until pupation.

Larvae usually fed with the anal plates exposed at the surface of the
integument; exceptionally, one larva completely buried itself within the
digestive tract of a decapitated cricket. Pupation occurred externally from
the host, except in one case, when a puparium was located within the
drying abdominal cavity of a female horse fly (the abdomen of the fly
had previously been cut to introduce the larva).

Discussion

The parasite (excerpted from Allen (1926), Downes (1965), Evans and
Eberhard (1970), and Krombein (1967 )).—Miltogrammini includes 75 spe-
cies of flies in North America, the larvae of which, with rare exception,
feed on the insect provisions captured by wasps and bees. Certain anatom-
ical specializations have been recognized in several genera of the tribe and
these help explain much about the biology of the adults. The females
of Amobia and Senotainia are equipped with enlarged eye facets, frontally,
to facilitate trailing Hymenoptera which bear prospective hosts for the
flies. Conversely, the females of other species, such as those of Phrosinella,
are non-trailing and seek the burrows of their fossorial hosts. When find-
ing those nests, they employ flattened foretarsi in implanting their progeny
in nearby pits. The proboscis is short in Phrosinella, and in other genera
the adults of which are not known from flowers; while in the Senotainia
species known to visit flowers, the proboscis is long and slender.

72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

In the female flies of the Macronychia species considered here, the
frontal eye facets and foretarsi are unmodified and the proboscis is very
short. Apparently, little host biology of the genera lacking such modifica-
tions is known. Except for several rearings of Macronychia spp. from nests
of two sphecid wasp genera (Krombein, 1964; Parker and Bohart, 1966),
the relationships of the two North American species of this genus are poorly
known. Both forms are widespread throughout the continental United
States. On the other hand, two European species have been associated
with wasps and bees and geometrid Lepidoptera (Thompson, 1951).

The host—Although Tabanus subsimilis subsimilis has been found
breeding in the marginal and riparian habitats typical of many Tabanidae—
irrigation ditches and stock ponds (Burger, 1974) and bottomland forests
(Thompson et al., 1977) —in southeast Texas, this species breeds extensively
in upland situations having mineral soils and no standing water (Thompson,
1975). These larval collections from upland sites helped to explain the
larger numbers, and the dominance, of the T. s. subsimils populations
observed in a later study comparing adjacent upland and lowland faunas
(Thompson, 1977). Most significant for the survival of Macronychia sp., this
horse fly is abundantly accessible as a host throughout much of Texas.
Also, in deference to foraging parasites and predators, the pupae of this
species commonly protrude in vertical position from the soil surface. Finally,
to complete the picture of host ubiquitousness, it is common—often, very
abundant—throughout much of the period from March through September.

The host-parasite association —Macronychia sp. was a common faculta-
tive parasite feeding upon Tabanus s. subsimilis adults in the vicinity of
the airport-sewage treatment plant near College Station, Texas from 28 May
through 14 July 1976. Although the regularity of this association must
be confirmed by additional observations in following years, several facts
suggest that the relationship between larvipositing females and_ host
tabanids was selective, not accidental. First, in two of the seven lots of
parasites observed, host females suffered multiple infestations (seven and
eight larvae each); in one of these cases, seven larvae of the same size, then
in the process of drying out, were found together in a ball within the
eviscerated host abdomen. Therefore, larvae of both lots were probably
sibling individuals deposited together on, or very near, the host. Secondly,
all larvae were found in lots; i.e., at least two to eight larvae were discovered
together in containers of horse flies, most of which held only five adults. The
probability of two infested flies in one container is low, especially if this
Macronychia was predominantly parasitizing Tabanus s. subsimilis.

On the other hand, the role of many Miltogrammini as cleptoparasites
of fossorial Hymenoptera—a role in which these Sarcophagidae rely upon
their hymenopterous hosts to provide the kinds of hosts upon which their
larvae subsist—increases the biotic potential of insects which already risk

VOLUME 80, NUMBER 1 73

life only as facultative parasites. This role suggests that Tabanidae are only
one of many taxa attacked by this species.

Conclusions

In his early monograph on the Miltogrammini of North America (1926),
Allen noted:

“So far as I am aware, it has not yet been conclusively demon-
strated that any of these flies are the primary parasites of phytoph-
agous insects in their native habitat. All the records investigated
by the writer indicated that while the flies do breed in the bodies of
such hosts, they display no interest in them until after they have
become the prey of some wasp.”

In the light of Allen’s comment, the findings reported here on the biology
of a species of Macronychia are unusual; and these findings provoke interest
in a relationship which could prove illuminating to the biology of the
Miltogrammini and to their potential in managing noxious populations of
Tabanus subsimilis subsimilis and of other Tabanidae.

Acknowledgments

I gratefully acknowledge the following persons for their contributions
to this study: Mr. Joseph W. Holmes, Jr., of this laboratory, for finding
the first sarcophagid parasites in Tabanidae, and for his conscientious
library efforts; Dr. Robert L. Harris, also of this laboratory, for finding
the first larvae of Macronychia sp. in Tabanus s. subsimilis and for provid-
ing me with larval and adult specimens of that parasite; the Drs. Raymond
J. Gagne and Curtis W. Sabrosky, Systematic Entomology Laboratory,
IIBUI, ARS, USDA, for their determinations and opinions on the manu-
script; the Drs. Horace R. Burke, L. E. Ehler, and L. L. Pechuman for their
comments on the manuscript; and Dr. Darrell Bay, Texas A&M University,
for providing me with the tabanid adults remaining from his research,
several T. s. subsimilis specimens of which were infested with Macro-
nychia sp.

Literature Cited

Allen, H. W. 1926. North American species of two-winged flies belonging to the
tribe Miltogrammini. Proc. U.S. Nat. Mus. 68:1-—106.

Burger, J. F. 1974. Horse flies of Arizona II. Notes on and keys to the adult Tabanidae
of Arizona, Subfamilies Pangoniinae and Chrysopinae (Diptera). Proc. Entomol.
Soc. Wash. 76:247—269.

Downes, W. L., Jr. 1965. Family Sarcophagidae, Tribe Miltogrammini, pp. 933-940.
In A. Stone et al., A Catalog of the Diptera of America North of Mexico, USDA,
Agric. Handbook No. 276. 1696 pp.

74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Evans, H. E., and M. J. W. Eberhard. 1970. The Wasps. Univ. Mich. Press, Ann
Arbor. 265 pp.

Goodwin, J. T. 1968. Notes on parasites of immature Tabanidae (Diptera) and
descriptions of the larva and puparium of Carinosillus pravus (Diptera: Tabani-
dae). J. Tenn. Acad. Sci. 43:107—108.

Krombein, K. V. 1964. Natural history of Plummers Island, Maryland: XVIII. The
hibiscus wasp, an abundant variety, and its associates (Hymenoptera: Sphecidae).
Proc. Biol. Soc. Wash. 77:73—112.

———. 1967. Trap-nesting Wasps and Bees: Life Histories, Nests, and Associates.
Smithsonian Press, Washington, D.C. 570 pp.

Parker, F. D., and R. M. Bohart. 1966. MHost-parasite associations in some twig-
nesting Hymenoptera from Western North America. Pan-Pac. Entomol. 42:
91-98.

Sanjean, J. 1957. Taxonomic studies of Sarcophaga larvae of New York, with notes
on the adults. Cornell Agric. Exp. Sta. Mem. 349. 115 pp.

Spratt, D. M., and G. Wolf. 1972. A tachinid parasite of Dasybasis oculata Ricardo
and Dasybasis hebes Walker (Diptera, Tabanidae). J. Aust. Entomol. Soc. 11:
260.

Thompson, P. H. 1975. Larval habitats of Tabanus subsimilis subsimilis Bellardi in
southeast Texas (Diptera: Tabanidae). Proc. Entomol. Soc. Wash. 77:494—
500.

——. 1977. Tabanidae (Diptera) of Texas. VII. Comparisons of upland and
lowland tabanid populations in southeast Texas. Ibid. 79:564—574.

Thompson, P. H., E. J. Gregg, D. R. Charanza, R. A. Sauseda, and J. W. Holmes, Jr.
1977. Habitats of larval Tabanidae (Diptera) in south Texas. Ibid. In press.

Thompson, W. R. 1951. A catalogue of the parasites and predators of insect pests.
Hosts of Coleoptera and Diptera. Sect. 2, Pt. 1. Commonwealth Inst. Bio.
Cntrl.

Veterinary Toxicology and Entomology Research Laboratory, Agric.
Res. Serv., USDA, College Station, Texas 77840.

NOTE
HOLOTYPE DEPOSITION OF HENICOMYIA BICOLOR

The holotype of Henicomyia bicolor Lyneborg [ 6 specimen, number 1212,
Brazil, Nova Teutonia, 27°11’S, 52°23’W, xi-1944, Fritz Plaumann] was
mistakenly deposited in the United States National Museum of Natural
History [In Lyneborg, L. 1972. A revision of the Xestomyza-group of
Therevidae (Diptera). Ann. Natal Mus. 21(2):368]. The specimen was
borrowed by me from the Canadian National Collection and subsequently
loaned to Lyneborg for inclusion in his monograph on the Xestomyza-group
of Therevidae. The holotype of Henicomyia bicolor Lyneborg has now been
redeposited in the Canadian National Collection, Ottawa, Ontario, Canada.

Michael E. Irwin, University of Illinois and Illinois Natural History
Survey, Urbana, Illinois 61801.

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 75-86
AN EVOLUTIONARY AND GEOGRAPHICAL OVERVIEW OF
REPAGULA (ABORTIVE EGGS) IN THE
ASCALAPHIDAE (NEUROPTERA)

Charles S. Henry

Abstract—Eggs and repagula (abortive eggs) were examined in seven
previously unstudied species of Ascalaphidae from both the Old and New
World. In those forms that possess repagula, the ratio of egg length to
repagulum length is shown to be an unreliable taxonomic character. The
absence of repagula in all Old World split-eyed taxa so far examined is
interpreted as a secondary loss in a monophyletic assemblage of owlflies
derived from a rather primitive New World ancestor—a view substantiated
by ovariole number and larval morphology. New World ascalaphids show
progressive specialization in repagulum form and function from primitive en-
tire-eyed genera like Byas and Ascaloptynx to derived split-eyed ones like
Ululodes and Cordulecerus. Entire-eyed Episperches and split-eyed Colo-
bopterus are judged transitional.

The Reverend L. Guilding (1827) coined the term “repagula” to describe
the small, glossy, rod-shaped bodies that accompany the egg masses of
certain split-eyed (ascalaphine) Ascalaphidae. These structures encircle
each egg-bearing twig like a fence immediately below the fertile eggs: ap-
propriately, the meaning of the Latin word repagulum is “little barrier.”
Henry (1972) verified the protective function of repagula against insect
predators in Ululodes mexicana (McLachlan), which possesses eggs with very
thin shells.

McClendon (1902) demonstrated that the repagula of Ululodes are in
fact highly modified abortive eggs that develop in their own specialized
ovarioles. More recently, New (1971) determined that abortive eggs are also
characteristic of entire-eyed (neuroptyngine) Ascalaphidae, but that in these
latter forms the structural differences between infertile and fertile eggs
are slight. Henry (1972) showed that the abortive eggs of at least one
neuroptyngine, Ascaloptynx furciger (McLachlan), serve as food for the
newly enclosed larvae and perhaps provide them with the nourishment
necessary for group defense against ants and other predators. Repagula of
Ascaloptynx, then, are not true barriers and in fact lack the repellant fluid
coating found on the repagula of Ululodes (Henry, 1972). Other studies
have shown that in the neuroptyngine genus Byas Rambur the repagula
share a common morphology with those of Ascaloptynx, while ascalaphines
such as Cordulecerus Rambur, Colobopterus Rambur, and Ascalorphne
Banks, possess repagula very similar to those of Ululodes (New, 1971;
Henry, In Press). Repagula of intermediate form are characteristic of
the specialized neuroptyngine genus Episperches Gerstaecker (New, 1971).

76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

All ascalaphid genera mentioned above are restricted to the New World,
but both the split-eyed and entire-eyed owlfly subfamilies have nearly
world wide distributions. Early European insect anatomists—e.g., Brauer
(1854) and Dufour (1860)—neither discuss nor figure anything resembling
modified ovarioles in Ascalaphus macaronius Scopoli or A. meridionalis
Charpentier. Other works that describe the egg masses of Old World
Ascalaphidae make no mention of egg attendants (Westwood, 1888; Ghosh,
1913; Fraser, 1957). The implication is that none of the Old World owl-
flies oviposits repagula, while members of both subfamilies in the other
hemisphere do so.

The present study introduces data on egg and repagula production in
several other species of Ascalaphidae, including Helicomitus dicax (Mc-
Lachlan), H. festivus (McLachlan), Suphalomitus malayanus javanensis
Weele, Hybris subjacens (Walker), Ascalaphus libelluloides Schaffer, Cor-
dulecerus alopecinus Burmeister, and Colobopterus sp. near trivialis and
consors Gerstaecker. In each case, the presence or absence of repagula
is noted, based either upon dissection of gravid females or upon study of
field-collected egg masses. Numbers of normal and modified ovarioles
per ovary are given whenever possible, and measurements of the propor-
tions of eggs and attendants are tabulated. These data are compared with
those of previous authors. The possible biogeographical and phylogenetic
significance of the results is then discussed.

Methods and Materials

Adult ascalaphids suitable for dissection were preserved in 75% EtOH
and examined under a WILD*™ M65 stereoscopic dissecting microscope
fitted with 10 eyepieces. Eggs and other structures were figured and
measured by means of a camera lucida attachment for this microscope.

Material for the study came from several sources. The adult specimens
from Shanghai, Madagascar, Java, and Paraguay were part of an extensive
alcoholic collection of poorly labeled insects deposited in the Museum of
Comparative Zoology (Harvard University) by the late G. C. Crampton.
Egg masses of Ascalaphus libelluloides were collected by my wife and
myself in the vicinity of Penne, Tarn et Garonne, France, during the last
half of July, 1974. Eggs and repagula of the Panamanian ascalaphid
Colobopterus sp. A were obtained from a captive female by R. E. Silberglied
and A. Aiello on 26 May 1976, on Barro Colorado Island, Canal Zone,
Panama. The identities of all adults were determined from Weele’s (1908)
monograph, while larvae of Ascalaphus libelluloides were identified from
the keys of Rousset (1973). Comparative data on Ululodes mexicana and
Ascaloptynx furciger were drawn from material assembled for an earlier
study (Henry, 1972).

VOLUME 80, NUMBER 1 Ctl

Results

Results are summarized chiefly in Table 1. The Ascalaphinae species
studied belong to four of the seven accepted tribes of the subfamily:
Suhpalacsini (Helicomitus and Suphalomitus), Hybrisini (Hybris), Asca-
laphini (Ascalaphus), and Ululodini (Cordulecerus and Colobopterus). The
Old World tribes Proctarrelabrini, Acmonotini and Encyoposini are not
represented. Females of Old World taxa all have ten egg producing ovarioles
per ovary; no specimens contained abortive eggs. On the other hand, the
two New World species possess repagula within specialized ovarioles and
display marked differences between the proportions of the fertile eggs
and those of the egg attendants. Eggs of Cordulecerus alopecinus were
by far the largest in the ascalaphines studied, while the most spherical
(broadest) were found in Ascalaphus libelluloides. Repagula of Colobop-
terus sp. A were proportionately much longer and thinner than those of
Cordulecerus, being only fractionally shorter than the eggs themselves
(ratio of egg length/repagulum length = 1.10). Egg size varied markedly
among individuals in both species of Helicomitus and in Hybris subjacens
but did not appear to be closely correlated with maternal dimensions
except in H. dicax (Table 2).

It may be argued that the sizes of eggs and repagula obtained from dis-
sected females are not comparable to those of the same structures when ovi-
posited. To test this, eggs and attendants obtained by dissection were
compared with those attached to twigs in Ululodes mexicana, an ululodine
ascalaphine species from southern North America. The results (Table 3) indi-
cate that sizes are insignificantly different in each of the two classes of data
and that the pooling of data for both unlaid and oviposited eggs or repagula
of Ascalaphidae is probably valid.

Table 4 compares the ratio of egg length to repagulum length in masses
obtained from several different individuals of both Cordulecerus alopecinus
and Ululodes mexicana, in order to test New’s (1971) hypothesis that this
ratio is a reliable taxonomic character separating the egg masses of closely
related species of New World Ascalaphinae. The ratio was found to vary
widely in U. mexicana, ranging from 1.65-2.19 in the five masses reliably
measured. The ratios obtained from four Cordulecerus alopecinus dis-
sections ranged from 1.73-1.99. The ovariolar number is seven in Ululodes
mexicana and eight in Cordulecerus alopecinus; both species possess four
additional repagula-bearing ovarioles in each ovary.

Discussion

The results strengthen the hypothesis that repagula production is limited
to Ascalaphidae of the New World. Previously, it was suspected only
that members of the Old World genus Ascalaphus lacked abortive eggs

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

78

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VOLUME 80, NUMBER 1 79

Table 2. Major morphological measurements of individual ascalaphid adults com-
pared with the dimensions of eggs obtained from the same individuals by dissection.
Adult measurements include the lengths of body, forewing, and antenna and the
width of the head across the eyes.

Adult measurements (mm) Egg measurements (mm)

Body Fore- Anten- Head Meanlength Mean breadth
Le aewine Lpeonalil, WwW. + SD + SD N

Helicomitus dicax
A 22.0 27.0 18.5 5.5 1.48 = .058 0.80 = .036
AA 22.0 26.5 18.0 5.3 1.28 + .027 0.73 = .045
A-3 23.0 29.0 19.0 6.0 1.90 = .043 1.14 = .023

ao ws

Helicomitus festivus
B 27.0 28.0 23.0 57 125 = 1-30 0.80 2
BB 25.0 27.0 22.0 Bu) 6373075 0.85 + .044 8
B-3 27.0 29.0 20.0 Dal 1.09 + .075 0.65 + .024 7

Hybris subjacens
D 28.0 35.0 30.5 5.6 [29% s4in OF> == 0457 410
DD 29.0 33.0 30.0 5.6 159 = 203 0.87 = .033 a

and associated modified ovarioles. The present work extends this suspicion
to representatives of the ascalaphine tribes Suhpalacsini and Hybrisini and
confirms the absence of repagula in the major constituent and nominate
genus of Ascalaphini. Unfortunately, nothing is known of repagula forma-
tion in Old World Neuroptynginae; however, Fraser (1922) does not men-
tion attendants accompanying the fertile eggs of Balanopteryx umbraticus
Fraser in Madagascar.

Repagula production in New World ascalaphine owlflies is best in-

Table 3. Dimensions of eggs and repagula obtained from five dissected females of
Ululodes mexicana compared with those obtained from three oviposited egg masses
of the same species.

No. of Mean
diff. indivs. breadth
or egg Sample Mean length ae oD)
Structure masses size + SD (mm) (mm)
From Eggs 5 females 40 1.89 + .159 0.94 + .083
Dissected Females Repagula 5 females 50 0.99 + .176 0.22, =5,.028
From Oviposited Eggs 3 masses 14 1.90 + .099 1.06 + .049
Egg Massess Repagula* 2 masses Is 0.93 + .084 0.23 + .024

“From masses preserved in alcohol; dried egg masses possess eggs of normal size
but withered repagula.

80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

OW NEW WORLD OLD WORLD

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CRETACEOUS

abortive eggs with trophic functions;
12+l2 ovarioles/ovary

JURASSIC

Fig. 1. Phylogeny of Ascalaphidae based upon structure of adult eyes, ovarioles,
and repagula.

terpreted as a specialization within a monophyletic assemblage of insects
(Fig. 1). All New World ascalaphids of the split-eyed subfamily are
closely similar to one another in wing venation, abdominal terminalia, and
antennal morphology and have been placed—in my view correctly—in a
single tribe, the Ululodini (Weele, 1908). Similar types of fluid-covered
repagula are found in representatives from every genus of the tribe. The
effectiveness of the repagula against potential predators remains to be
tested in taxa other than Ululodes.

New (1971) speculated that the repagulum of Ululodini is derived from
one like that in the Neuroptynginae; the unavoidable implication from
his paper is that the transition from a “simple” abortive egg like that of
Byas to the specialized “rods” or “dumb-bells” of Ululodini occurred through
some more specialized neuroptyngine genus (like Episperches) possessing

VOLUME 80, NUMBER 1 81

repagula of intermediate morphology. Such views receive additional sup-
port from data given in Table 1 and in two previous papers (Henry, 1972 and
In Press). However, the absence of any sort of abortive eggs in Old World
ascalaphids is difficult to reconcile with a scheme that derives split-eyed
owlflies from entire-eyed ancestors. Presumably, abortive egg production
is a shared specialization of all Ascalaphidae, with further modification of
abortive eggs into true repagula (predator barriers) occurring within certain
Neuroptynginae that then gave rise to Ascalaphinae; this is the hypothesis
borne out by data on New World forms. If this is true, one would expect to
find egg attendants in some Old World taxa regardless of the evolutionary
relationships among family members. For example, generalized Old World
members of both subfamilies should possess unspecialized abortive eggs if
one assumes that New World forms evolved from Old World ancestors
that had already differentiated into two types with different eye morphology.
Alternatively, the Old World ascalaphines, neuroptyngines, or both, might
be derived from New World ancestors; in these cases, it is likely that split-
eyed Old World taxa would retain and perfect some sort of specialized
repagulum similar to that of Ululodini. Loss of repagula in derived taxa
is possible, but the greater adaptive value to the eggs of attendants once
the latter assume a protective function renders this hypothesis unattractive,
as well. Unfortunately, too little is known of other aspects of ascalaphid
morphology and biology to clarify the phylogenetic puzzle. However, larval
morphology (Henry, 1976) and the orientation of eggs in naturally-oc-
curring egg masses (Henry, 1972) suggest that New World Ascalaphinae
are highly specialized in comparison with their Old World relatives; among
other implications, this makes the secondary loss of egg attendants in the
latter group seem even less likely.

If one assumes that repagula have not been lost secondarily, one must
conclude that monophyly either of split eyes or of repagula production is an
incorrect interpretation of ascalaphid evolution. Either aspect of this con-
clusion is difficult to accept. On the one hand, comparative morphology
of the sulcate eyes of Old and New World Ascalaphinae indicates that
the bisection is almost certainly homologous in representatives from the
two hemispheres (Fig. 2). On the other hand, as far as is known, production
of abortive or trophic eggs from modified ovarioles is unique to Ascalaphi-
dae within the Neuroptera and very rare in other insect orders; the con-
vergent evolution of such an apomorphy separately in Ascalaphinae and
Neuroptynginae of the Western Hemisphere seems unlikely. What appears
to be progressive increase in specialization of repagula from entire-eyed
to split-eyed taxa in the New World is further evidence against con-
vergent evolution of egg attendants in the two subfamilies; one may even
interpret the Colobopterus repagulum as intermediate between that of
Episperches and those of higher Ululodini (Fig. 3). Therefore, I feel

82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

A B

Mins”

l 3 mm J CS-Henry, uv
Fig. 2. Fronto-lateral view of the head of New World (A) and Old World (B)
ascalaphine owlflies, showing bisected eyes. A = Ululodes mexicana and B = Heli-

comitus dicax.

that it is better at the present time reluctantly to postulate loss of repagula
in Old World Ascalaphinae and to assume that all owlfly taxa in all geo-
graphical areas originated from New World neuroptyngine ancestors pos-
sessing primitive abortive eggs; bisection of the eye must also have developed
first in the Western Hemisphere, with early colonization of the Old World by
a primitive ascalaphine stock (Fig. 1). The position of African and
Asian Neuroptynginae is uncertain in such a scheme since their eggs are
so poorly known.

The presently uncertain systematic position of Colobopterus is rendered
even more problematical by repagular morphology. Although it is widely
accepted that Ululodes quadrimaculata (Say) is the female of Colobopterus
excisus Hagen in North America, the Colobopterus sp. A near trivialis and
consors discussed here is markedly divergent from any known species of
Ululodes in the form and size of its egg attendants. Additionally, the larval
morphology of another Panamanian species of the genus, Colobopterus
sp. B near mulleri Weele, is totally unlike that of any known Ululodes (un-
published data). Consequently, it is probably valid to exclude some spe-
cies of Colobopterus from Ululodes; in fact, the magnitude of repagular

VOLUME 80, NUMBER 1 83

CORDULECERUS ALOPECINUS

ULULODES MEXICANA

COLOBOPTERUS SP.

mm weed i ." 7 eo wl i a q
EGG REP. EGG REP. EGG REP.
Fig. 3. The eggs and repagula of three New World owlflies of the tribe Ululodini,
drawn to the same scale. REP. = repagulum.

and larval differences just mentioned suggests that some Colobopterus
species are less closely related to Ululodes than is Cordulecerus, which
possesses a Ululodes-like repagulum (Fig. 3).

Comparative measurements of eggs and repagula (Table 4) do not sup-
port New’s (1971) contention that “the ratio of repagula [sic] length:egg
length affords an index for separation of the egg masses of closely related
species.” In neither U. mexicana nor Cordulecerus alopecinus is such a ratio
constant. At one extreme, the ratios for Ululodes mexicana are found to
overlap extensively with those calculated from New’s data for both U.
macleayana near limbata Burmeister and U. aurifera McLachlan. At the
other extreme, egg/repagulum ratios for New’s Cordulecerus alopecinus are
significantly different from my own measurements on the same species
(1.37 vs. 188 + .11, N = 4). Unfortunately, New gives neither means
nor standard deviations of his data and does not make clear how many
different egg masses his sample sizes represent.

The phylogenetic significance of ovariole number in Ascalaphidae has
not been assessed. Within the Neuroptera as a whole, primitive taxa seem to
exhibit larger numbers of ovarioles. For example, alderflies (Megaloptera:
Sialidae) have up to 100 egg tubes per ovary (Matsuzaki and Ando,
1977), while green lacewings (Planipennia: Chrysopidae) possess only
12-15 (Hwang and Bickley, 1961; Rousset, 1976). New (1971) reported

84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 4. Comparisons of egg and repagulum lengths and of ovariole numbers for
nine dissected individuals of Ululodes mexicana and four of Cordulecerus alopecinus.
The ratio of egg length to repagulum length is calculated for each insect.

No. of
ovarioles
amen / Ratio of
eggs
bear- length:
bear- ing Mean length + SD (mm) repay
ing repag- ulum
eggs ula Eggs (N) Repagula (N) length
Ululodes
mexicana
Skel. = = 1.95 + .038 (5) 0.89 + .053 (10) 2.19
DS-1 >? 4 1.83 + .087 (10) 1O4==2 0390) 1.76
DS-2 > 4 AO == lSin(9) 1.03 + .063 (10) 1.65
DS-3 ? ? 1.86 + .095 (3) 0.72 + .034 (13) 2.58
DS-4 7 AB Rae ThaOUAM EN GONE) ee
DS-5 7 4 2.04 + .073 (10) 7 == 086a i) 1.74
DS-6 7 4 2.02 + .087 (8) 1.08 + .036 (9) 1.87
DS-8 7 Ad NF Oe te ee ThagWOO EWE OU RS)
DS-9 0 AEE We ema > ee ee rHTOWOO EY US) a
Cordulecerus
alopecinus
E = = 2.29 + .078 (8) 1LSX0) a= (083, (GUL) 1.99
EE = = 2.34 + .046 (6) 1.24 + .045 (8) 1.89
E-3 8 4 ORO == OAT) 1.16 + .042 (9) 1.91
E-4 8 4 2.26 + .083 (7) LS le== 070) lens

fairly high ovariole numbers of 22-24 in most of the owlflies he dis-
sected, although repagula occupied twelve of the tubes. The neuroptyngine
Episperches arenosus (Walker) showed a reduction from twelve to ten in
the number of egg-producing ovarioles per side, while Ululodes aurifera of
the Ascalaphinae exhibited only eight “fertile” and four “modified” tubes
in each ovary. The present study confirms that these numbers need not
be constant in a genus, since the ovary of U. mexicana has one less fertile
ovariole than that of aurifera (Table 4)'. The discrepancy noted within
Cordulecerus alopecinus between my own counts of ovarioles (8 + 4 per
side) and those of New (12 + 12) indicates almost certainly that the two
studies were not based on conspecific individuals. This might explain the
differences obtained in egg/repagulum ratios, as well.

Old World ascalaphine forms examined here possess only ten ovarioles
per ovary, as do members of the Eurasian genus Ascalaphus (Brauer,
1854; Dufour, 1860). Such a reduced ovariolar number is best interpreted
as a specialization; the fact that it is shared by representatives of the tribes

VOLUME 80, NUMBER 1 85

Suhpalacsini, Hybrisini, and Ascalaphini from Europe, Asia and Africa
strongly suggests close relationship among these taxa. The extant Old World
Ascalaphinae, then, may indeed represent an adaptive radiation from the
single primitive New World stock postulated earlier in the discussion. That
this and all other evolutionary events within the Ascalaphidae took place
after the separation of South America from Australia is strongly indicated
by the total absence of Neuroptynginae from the latter continent; probably
Africa and South America were in close proximity when owlflies with
bisected eyes originated. Subsequent isolation of African Ascalaphinae
by drifting of the continents in the Cretaceous produced the morphologically
and behaviorally distinct Old World forms that we see today (Dietz and
Holden, 1970). In the New World, repagula capable of repelling predators
certainly arose subsequent to the establishment of a dominant ant fauna
or “mosaic”: at the earliest, within the Upper Cretaceous (Leston, 1973).
This gives an independent approximate dating of certain later events of
ascalaphid evolution synchronous with that derived from drift theory.

Acknowledgments

This work was made possible by a grant from the Research Foundation
of The University of Connecticut (Storrs) and an N.S.F. institutional grant
to The George Washington University (Washington, D.C.). Collection of
important specimens of Colobopterus spp. is due to the efforts of R. E.
Silberglied and A. Aiello (Harvard University). I thank N. Henry, Pro-
fesseur A. Haget (Université de Bordeaux), and V. Roth (Southwestern
Research Station) for their valuable assistance in the field work, and Drs.
D. Leston, C. W. Rettenmeyer, C. W. Schaefer, and J. A. Slater for con-
structive comments on the manuscript.

Literature Cited

Brauer, F. 1854. Beitrage zur Kenntniss des inneren Baues und der Verwandlung
der Neuropteren. Verh. Zool.-Bot. Ver. Wien 1854(4):463—472.

Dietz, R. S., and J. C. Holden. 1970. The breakup of Pangaea. Sci. Am. 223:30-31.

Dufour, M. L. 1860. Recherches anatomiques sur l’Ascalaphus meridionalis. Ann. Sci.
Nat. Zool. Biol. Anim. 13(4):193-—207.

Fraser, F. C. 1922. Some oriental Ascalaphidae in the Indian Museum. Rec. Indian
Mus. 34:511-520.

—. 1957. Two new species of Ascalaphidae from Madagascar (Neuroptera).
Nat. Malgache. 9(2):247-250.

Ghosh, C. C. 1913. Life-history of Helicomitus dicax Walker. J. Bombay Nat. Hist.
Soc. 22:643-648.

Guilding, L. 1827. Communication on Ascalaphus macleayanus. Trans. Linn. Soc.
London 15:509-512.

Henry, C.S. 1972. Eggs and repagula of Ululodes and Ascaloptynx (Neuroptera: Asca-
laphidae): a comparative study. Psyche J. Entomol. (Camb. MA.) 79:1-22.

86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

—. 1976. Some aspects of the external morphology of larval owlflies (Neurop-
tera: Ascalaphidae), with particular reference to Ululodes and Ascaloptynx.
Ibid. 83:1-31.

—. In Press. The egg, repagulum, and larva of Byas albistigma (Walker)
(Neuroptera: Ascalaphidae): morphology, behavior, and phylogenetic signif-
icance. Syst. Entomol.

Hwang, J. C., and W. E. Bickley. 1961. The reproductive system of Chrysopa
oculata (Neuroptera: Chrysopidae). Ann. Entomol. Soc. Am. 54:422-429.

Leston, D. 1973. The ant mosaic, tropical tree crops and the limiting of pets and
diseases. PANS. 19(3):311-341.

Matsuzaki, M., and H. Ando. 1977. Ovarian structures of the adult alderfly,
Sialis mitsuhashii Okamoto (Megaloptera: Sialidae). Int. J. Ins. Morph. Em-
bryol. 6(1): 17-29.

McClendon, J. F. 1902. The life history of Ulula hyalina Latreille. Am. Nat. 36:
42-49.

New, T. R. 1971. Ovariolar dimorphism and repagula formation in some South
American Ascalaphidae (Neuroptera). J. Entomol. Ser. A Gen. Entomol. 46:
73-17.

Rousset, A. 1973. Morphologie externe et charactéres distinctifs des larves de trois
especes d’Ascalaphes (Névropteres, Planipennes). Bull. Soc. Entomol. Fr.
78:164-178.

——. 1976. Les stades du developpement ovarien de Chrysopa perla (Neuroptera).
Etude histologique sur les femelles soumises 4 des conditions trophiques optimales.
Ann. Soc. Entomol. Fr. 12(3):405-417.

Weele, H. W. van der. 1908. Ascalaphiden: Monographisch Bearbeitet. Coll. Zool.
Selys-Longchamps fasc. 8:1—326.

Westwood, J. O. 1888. Notes on the life-history of various species of the Neuropterous
genus Ascalaphus. Trans. R. Entomol. Soc. London 1888: 1-12.

The Biological Sciences Group, The University of Connecticut, Storrs,
Connecticut 06268.
Footnote

1An earlier study (Henry, 1972) reporting eight fertile egg tubes in U. mexicana
is in error.

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 87-90
LIFE HISTORY AND DESCRIPTIONS OF THE IMMATURE
STAGES OF JAMESOMYIA GEMINATA
(DIPTERA: TEPHRITIDAE)

W. Bryan Stoltzfus

Abstract—The biology of Jamesomyia geminata (Loew) is discussed
and the first-, second-, and third-instar larva and puparium are described
and illustrated. Its seasonal distribution and relationship to its host plant
Lactuca canadensis are included.

Jamesomyia was erected by Quisenberry (1949) for one species,
Jamesomyia geminata (Loew). Quisenberry gave a genus and _ species
description of J. geminata along with a record of its distribution.

The general life cycle is presented here, along with descriptions of the
immature stages.

Life History

Females were found during July on the host plant (Lactuca canadensis)
searching for flower buds of the proper size in which to oviposit. The fe-
male walked up and down the flower stalks, stopping intermittently to
flex its wings. Elongate buds of about 5.5 mm were chosen for oviposition.
The female inserted the ovipositor between the bracts at, or near the apex
of the flower bud and laid a single egg among the florets.

Males are territorial and were found “patrolling” the upper leaves and
flower buds of the host plant. As females came into the males’ territory,
the male exhibited courtship behavior, using stylized wing and body
movements to attract the female. The male extended one wing slowly per-
pendicular to the body, then the other wing. Sometimes both wings were
extended simultaneously. Quick side-ways body movements were also em-
ployed in attracting the female.

Larvae require about a month to be fully grown. They feed on the
achenes, consuming all but an outer shell of the small number of achenes
available (Fig. 7). The uneaten achenes and larval excrement are “glued”
together to form a protective “puparium case” in which the mature third-
instar larva overwinters. During August the host plant’s seeds are dis-
persed and the puparium case falls to the ground. The mature larva
usually pupates the following spring and adults begin emerging the last of
June in northeastern Ohio.

Occasionally larvae pupated in August and adults emerged the same
season. Whether adults of the second generation used the same species
of plant as a host, or did not reproduce, was not determined.

88 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The host plant, Lactuca canadensis L., occurred sparsely along old
fence rows or the borders of a woods. Jamesomyia geminata was also in-
frequently found.

Of 100 heads of Lactuca canadensis examined, 38 contained larvae. Of
these, 25% were parasitized by a wasp, Habrocytus sp. (Pteromalidae). An
anthomyiid fly infested the same flower buds and seemed to be as common
as Jamesomyia geminata. Occasionally the two species of larvae were
found infesting the same head. If there is insufficient food for both larvae,
the anthomyiid usually wins the competition. Frequently one of the two
species was parasitized by a wasp but never both species.

Descriptions of Immature Stages

Third-instar larva —Length 2.6-3.1 mm, width 1.4-2.2 mm (Fig. 1). White
to light yellow as early third-instar larva, to nearly black, especially 1st
and last segments of overwintering larva. Mature larva barrel-shaped, head
and 8th abdominal segments dark brown, middle segments yellowish brown.
Integument shagreened; no spinule patches on venter. Two irregular rows
of strong spinules on dorsum of 6th and 7th abdominal segments, smaller
spinules on other segments. Eighteen sensilla distributed around each
segment. Pseudocephalic segment invaginated into the thorax. Overwin-
tering larva with scalloped posterior border around 8th abdominal segment,
posterior part being greatly darkened.

Anterior spiracles (Fig. 2) fan-shaped, with tubules, dorsal and ventral
lobes branched, yellow to light orange. Stigmatic chamber distinctly di-
vided into several cells, 3 across; about 5 lengthwise.

Cephalopharyngeal skeleton (Fig. 3) heavily sclerotized, bearing 1 ac-
cessory tooth midway on ventral margin of mouth hooks. Mouth hooks, 0.15
mm long, slightly longer than wide. Intermediate sclerite slightly longer
than mouth hooks. Labial sclerite long and narrow. Cibarial phragma dark
except distal portion which bears window.

Maxillary palp (Fig. 4) slightly longer than wide, bearing perhaps 5 or
6 short peglike segments distally. Antenna slightly longer and more conical.

Posterior stigmatic disc (Figs. 5, 6) smooth, bearing no lobes or ridges.
Posterior stigmatic area dark, slits and stigmatic scar set off by much lighter
stigmatic plate. Slits oval with bars extending inward % of way from each
side, peritremes dark with very small opening at distal end. Slits about 0.056
mm long. Stigmatic chamber distinctly divided into 6 or 7 cells across and

ss

Figs. 1-7. Jamesomyia geminata mature third-instar larva and plant infestation.
1. Larva; ASp, anterior spiracle; 2. Anterior spiracle; StC, stigmatic chamber; T,
tubule; 3. Cephalopharyngeal skeleton; CF, clypeofrontal phragma; CP, cibarial phragma;

VOLUME 80, NUMBER 1 89

IS, intermediate sclerite; LS, labial sclerite; MH, mouth hook; W, window; 4. Antenna,
An; maxillary palp, MP; 5. Posterior spiracular disc; PSP, posterior spiracular plate; StA,
stigmatic area; StS, stigmatic scar; 6. Posterior spiracular plate; IP, interspiracular
process; SS, stigmatic slit; StC, stigmatic chamber; Tr, trabeculae; 7. Pupation site
among achenes of Lactuca canadensis.

90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

3 or 4 deep. Interspiracular processes composed of 4 very small, short
spikes, 1 on each side of a slit. Anal plate pale yellow, not shagreened and
bearing no tubercles. Anal plate set off ventrally by shagreened area and
dorsolaterally on each side by 2 papillae.

Second-instar larva—Length 1.32-2.38 mm; width 0.61-1.32 mm. White
to pale yellow, anterior end pointed, posterior end rounded. Integument
not shagreened but spinules present as in third instar except no strong
spinules on 6th and 7th abdominal segments. Anterior spiracles lighter in
color and bear the same number of tubules as in third instar.

Posterior spiracular disc light yellow. Slits about 0.024 mm long.

Cephalopharyngeal skeleton dark except for lighter area at point of
mouth hooks and distal ends of clypeofrontal and cibarial phragmata. Mouth
hooks (about 0.1 mm) slightly longer than intermediate sclerite. No win-
dow in cibarial phragma.

First-instar larva.—Length 0.8-1.5 mm; width 0.4-0.61 mm. Mouth hooks
and intermediate sclerite light brown, clypeofrontal phragma large and
dark, cibarial phragma lighter and smaller. No labial sclerites; labial
sclerite fused to intermediate sclerite. Mouth hooks about 0.018 mm long.
No anterior spiracles. Two barely visible oval slits, which are only slightly
pigmented, represent the posterior spiracles.

Egg.—Length 1.18-1.27 mm; width 0.24-0.27 mm. Dull white, micropylar
end lanceolate, opposite end more rounded. No noticeable reticulation or
other markings.

Puparium.—Length 3.45 mm; width 1.86 mm. Barrel-shaped, ends trun-
cate as in third-instar larva. Puparium very thin and dark; black at ends;
center dark brown. Anterior spiracles slightly raised; tubules as in third-
instar larva. Posterior spiracular plates only very slightly raised. Spinules,
when visible, as in third-instar larva.

Literature Cited

Quisenberry, B. F. 1949. A new genus of Tephritidae near Xanthomyia (Diptera).
Bull. Brooklyn Entomol. Soc, 44:49-52.

Biological Science Department, William Penn College, Oskaloosa, Iowa
SABI

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 91-93
A NEW EGG PARASITE (HYMENOPTERA: SCELIONIDAE)
OF THE ELM SPANWORM, ENNOMOS SUBSIGNARIUS
(LEPIDOPTERA: GEOMETRIDAE)

Carl F. W. Muesebeck

Abstract—Telenomus droozi, new species, a parasite in the eggs of the
elm spanworm, Ennomos subsignarius (Huebner), is described from Penn-
sylvania.

The species described below, which develops as a parasite in the embryo-
nated eggs of the elm spanworm, Ennomos subsignarius (Huebner), has
been confused with another parasite of geometrid eggs, Telenomus alsophilae
Viereck. Recent studies (Drooz et al., 1976) have shown that it is distinct
biologically and is not a parasite of eggs of the fall cankerworm, Alsophila
pometaria (Harris), the normal host of Telenomus alsophilae. As a result
of that disclosure, critical studies were made of both forms, and some mor-
phological differences were discovered. Although these differences are
subtle and sometimes not readily recognized, they appear to distinguish the
two species.

Telenomus droozi Muesebeck, new species

This species may usually be distinguished without much difficulty from
specimens of T. alsophilae that have developed in eggs of the fall canker-
worm. It is consistently a little larger; the head is slightly narrower rela-
tive to the width of the thorax, and as seen from above it is a little flatter
in front (Fig. 1); normally there is a continuous, narrow strip of delicate
microsculpture along the inner eye margin (in alsophilae the frons medially
is completely smooth and polished); there are slight but recognizable dif-
ferences in the basal segments of the antennal flagellum (Figs. 4 and
5); and when the insect is viewed from the side, the level of the dorsum
of the abdomen normally appears to be nearly on a line with that of the
mesonotum whereas in alsophilae it is usually much lower. A. T. Drooz and
certain of his associates at the Southeastern Forest Experiment Station,
U.S. Department of Agriculture Forest Service, have found that T. alsophilae
will develop in the eggs of a number of Geometridae other than Alsophila
pometaria. When the eggs of the host species are larger (as in the case
of Abbotana clementaria (J. E. Smith) and Oxydia trychiata (Guenée)), the
resulting parasites are noticeably larger than those emerging from eggs
of Alsophila; they are as large as specimens of Telenomus droozi and the
antennae very closely resemble those of that species. However, the other
differences mentioned above are apparent in these larger specimens also.

92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ee

Figs. 1, 4, 5. Telenomus droozi. 1. Dorsal view of head; 4. Female antenna; 5.
Male antenna. Figs. 2, 3, 6. Telonomus alsophilae. 2. Dorsal view of head; 3. Fe-
male antenna; 6. Male antenna.

VOLUME 80, NUMBER 1 93

No significant differences in the male genitalia of the two species have
been noted.

Female.—Length approximately 1 mm. Head just about as wide as max-
imal width of thorax, in frontal view broadly subtriangular, in dorsal view
as shown in Fig. 1; eyes weakly setose; surface of vertex with delicate
reticulate sculpture; weaker microsculpture narrowly along inner eye mar-
gins; lower part of frons with delicate reticulate sculpture like that of vertex;
elsewhere the frons smooth and polished; sharp margin bordering the eyes
behind not continued as a ridge across vertex, which is rather smoothly
convex from front to back; lateral ocelli situated on vertex at eye margins
and barely above level of median ocellus; antenna as illustrated (Fig. 4).

Thorax with mesoscutum sculptured like vertex; disc of scutellum strongly
transverse, smooth and polished; metanotum short, usually smooth but some-
times with a little microsculpture across base; propodeum short, very shiny,
and with a little indefinite sculpture, especially laterally; forewings over-
reaching abdomen by hardly % length of the latter; hindwings about 5x
as long as their maximal width; stigmal vein nearly as long as width of
hindwing and % as long as postmarginal vein, the terminal end of which
is rather vague but is determined by the row of 10 or 11 evenly spaced
bristles arising from it.

Abdomen a little longer than thorax, apical segments usually not re-
tracted; in lateral view level of dorsum of abdomen nearly on a line with
level of posterior part of mesonotum; Ist segment of gaster dorsally with
a row of foveae or short striae across base; large 2nd segment smooth and
polished but sometimes with a row of minute punctures across extreme
base.

Coloration Black; antennae, including scapes, black; legs largely black
or blackish, the tarsi usually paler; wings hyaline.

Male.—Essentially like the female except for the antennae (Fig. 5), a
smaller abdomen and paler legs.

Holotype —USNM No. 74012.

Described from 40 females (one, the holotype) and 41 males reared 2
June 1976, by A. T. Drooz from eggs of Ennomos subsignarius (Huebner)
which had been collected at Pottersdale, Pennsylvania.

Literature Cited
Drooz, A. T., G. F. Fedde, and J. A. Copony. 1976. Egg parasite of the elm span-

worm is not Telenomus alsophilae. Environ. Entomol. 5:492—494.

Cooperating Scientist, Systematic Entomology Laboratory, IIBIII, Agric.
Res. Serv., USDA, c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 94-102
NOTES ON THE SYSTEMATICS AND BIOLOGY OF THE
BITING MIDGE, FORCIPOMYIA ELEGANTULA MALLOCH
(DIPTERA: CERATOPOGONIDAE)

Willis W. Wirth and William L. Grogan, Jr.

Abstract—The immature stages of Forcipomyia (Forcipomyia) elegan-
tula Malloch are described and illustrated for the first time. Adults are rede-
scribed and illustrated, and all stages are compared with the closely re-
lated European species P. pulchrithorax Edwards. Biological notes are
given on this species which was found breeding under the bark of dead
cottonwood trees on Plummers Island, Maryland. The structure and
function of the peculiar hygroscopic hairs of the larva and their defensive
value against predators are discussed.

We are indebted to Theodore J. Spilman for the discovery, during June
1976 on Plummers Island, Maryland, near Washington, D.C., of a large
population of the biting midge Forcipomyia (Forcipomyia) elegantula
Malloch. Large numbers of all stages of this strikingly marked midge were
found under the thick bark on the trunks of large cottonwood trees
(Populus deltoides Bartram) that had been felled by resident beavers and
had lain on the river bank for a year or more. The colonies of the midge
lived in the layers of tissue-thin inner bark next to the cabium where there
was enough moisture to support a copious growth of microorganisms. A
complex and rich biota of coleopterous and dipterous larvae and Collembola
fed on these microorganisms or preyed on the herbivores. Fungal spores
appeared to make up the bulk of the gut contents of the Forcipomyia
larvae.

Malloch (1915) described Forcipomyia elegantula from adults collected
at his laboratory window in Urbana, Illinois. Nothing has since been re-
ported on this species. It is very similar to the Palearctic F. pulchrithorax
Edwards (1924), described from specimens bred from wood debris and from
sap flowing from wounds of elm and chestnut trees in England. Saunders
(1924) described the immature stages of F. pulchrithorax and the species
has since been reported from Hungary, Belgium, Germany, and Estonia. We
take this opportunity to present for the first time descriptions and illustra-
tions of the immature stages of F. elegantula, figures and redescriptions of
the adults and comparisons of all stages with those of F. pulchrithorax.

We are especially indebted to Ethel L. Grogan for preparing the illus-
trations. We also extend thanks to M. E. Taylor for aid in preparing the
scanning electron micrographs. The junior author carried on the field
work on Plummers Island during the summer of 1976 with financial support
from a research grant from the Washington Biologists’ Field Club.

VOLUME 80, NUMBER 1 95

Forcipomyia (Forcipomyia) elegantula Malloch
(Figs. 1-4, 6, 12-20)

Forcipomyia elegantula Malloch, 1915:311 (male, female; Llinois); Johann-
sen, 1952:157 (in key to spp. NE USA).

Forcipomyia (Forcipomyia) elegantula Malloch; Johannsen, 1943:777 (in
list N. Amer. spp.); Wirth, 1965:125 (USA distribution).

Diagnosis.—A large yellowish species most closely resembling the Palae-
arctic F. pulchrithorax; distinguished from all other Nearctic Forcipomyia
(Forcipomyia) by the following combination of characters: Adults with
3 black stripes on scutum and 2-3 black spots on pleuron; hind femur with
black band on distal 4%; tibiae of female with hastate setae. Females differ
from F.. pulchrithorax in having a black band on hind femur and oval rather
than pyriform spermathecae; males differ from that species in having a
shield-shaped aedeagus with a concave basal arch and pointed tip, the
aedeagus of F. pulchrithorax being triangular with a convex basal arch.
Pupa with respiratory horn with reticulate surface and 25-30 spiracular
papillae, and much shorter thoracic and abdominal processes than in PF.
pulchrithorax. Larva with large hastate a hairs; a hairs smaller and only
slightly hastate in F. pulchrithorax.

Female.—Wing length 1.45 mm; breadth 0.60 mm. Head: Frontoclypeus,
scape and pedicel of antenna yellow; vertex, flagellum of antenna, and
palpus yellowish brown. Proximal 8 flagellomeres globose, possessing
multiple sensoria; distal 5 elongate, lacking sensoria; apical flagellomere
with distinct apical papilla; flagellomeres with lengths in proportion of
18-14-15-15-15-15-15-15-21-21-21-22-29; antennal ratio 1.07. Palpus with seg-
ments in proportion of 11-13-32-17-11; 3rd segment expanded basally with
distinct deep pit bearing numerous minute sensilla; palpal ratio 2.46.
Mandible reduced. Thorax: Yellowish. Scutum (Fig. 1) with 3 black stripes,
the median 1 bifid posteriorly; scutellum and postscutellum black; pleuron
(Fig. 2) with 2-3 round black spots, black stripe on sternum. Legs yellow;
hind femur with black band on distal “4; tibiae with large hastate setae;
femora and tibiae with long setae. Wing about 2.5x longer than broad,
covered with dense macrotrichia; pattern (Fig. 3) with lightly infuscated
area on and distad of costa; costal ratio 0.45. Halter stem pale, knob white.
Abdomen: Yellowish; terga and last 3 sterna brown, giving abdomen a
banded appearance. Spermathecae (Fig. 4) oval, subequal with very short
necks.

Male.—Similar to female, differing as follows: Flagellar plume brown;
tibiae lacking hastate setae; wing more slender. Genitalia: As in Fig. 6.
Ninth sternum about 2% as broad as long; caudomedian margin convex; 9th
tergum tapered distally, posterior margin rounded, cerci very reduced.
Basistyle nearly straight, 2.5x longer than broad; dististyle curved slightly

96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

15

Figs. 1-15. Forcipomyia elegantula (Figs. 1-4, 6, 12-15) and F. pulchrithorax
(Figs. 5, 7-11). 1. Color pattern of scutum; 2. Color pattern of pleuron; 3. Color
pattern of female wing; 4-5. Spermathecae; 6. Male genitalia; 7. Aedeagus; 8, 12. Pupal
respiratory horn; 9, 13. Thoracic processes of pupa; 10, 14. Abdominal processes of
pupa; 11, 15. Abdominal hairs a, b, c, d of larva.

VOLUME 80, NUMBER 1 97

distally, 0.67 length of basistyle, tip darkened and slightly pointed. Aedea-
gus lightly sclerotized, shield-shaped; basal arch concave; basal arm short
with rounded, slightly recurved tip; distal portion with pointed tip.
Claspettes divided, lightly sclerotized; basal arm thickening slightly dis-
tally; distal portion slender, tapering distally with slightly recurved tip.

Pupa.—Retains larval exuviae. Respiratory horn (Fig. 12) with broadened
base, narrowed proximally, then much broader distally; scanning electron
micrograph reveals that surface is reticulate (Fig. 18), reticulations not
visible in slide-mounted specimens; apex with 25-30 spiracular papillae;
scanning electron micrograph reveals that apertures of papillae are much re-
duced (Fig. 19). Thorax with 4 pairs of dorsal setose thoracic processes;
anterior pair (Fig. 13a) with terminal seta; 2nd and 3rd pair (Fig. 13b)
similar but lacking terminal seta; 4th pair (Fig. 13c) similar to 2nd and 3rd
but shorter; also a pair of short setose posterior processes present (not fig-
ured). Abdomen with a pair of long setose dorsal processes (Fig. 14a); a
pair of short setose dorsal processes (Fig. 14b); and a slightly longer pair of
setose lateroventral processes (Fig. 14c). Female terminal segment 1.6
longer than broad; surface covered with scattered fine-pointed tubercles;
apicolateral processes appressed, surface wrinkled. Male terminal segment
1.6x longer than broad; surface smooth; apicolateral processes greatly
appressed, surface wrinkled; ventral genital processes appressed, greatly
wrinkled.

Larva (4th instar) —Body golden yellow, lightly sclerotized; internal
structures of head sclerotized. Scanning electron micrograph of head (Fig.
16) indicates chaetotaxy as follows: p hair with broadly hastate tip; q hair
similar to p hair but hastate tip slightly narrower; t hair long, slender;
antenna 3-segmented. Prothoracic pseudopod entire; apex with 4 pairs
of long sclerotized, slightly curved hooks. Midabdominal segments with
hyaline, broadly hastate a hairs (Fig. 15a) connected dorsally by a narrow
sclerotized bar to the a hair of the opposite side; b hair (Fig. 15b) heavily
sclerotized, anterior sides branched; c hair (Fig. 15c) very short, branched
on anterior side; d hair (Fig. 15d) twice as long as c hair, slightly branched
on anterior side; surface of segments covered with small pointed tubercles.
Terminal segment small; anal blood gills short, hyaline; anal pseudopod
with double row of sclerotized hooks, 8 on each side.

Type.—Holotype, 2°, allotype, ¢, Urbana, Champaign Co., Illinois, 28.vi.
and 5,12.viii.1915, on window in basement of Natural History Building,
J. R. Malloch. Through the courtesy of Donald W. Webb the types were
borrowed from the Illinois Natural History Survey and the allotype male
was mounted on a slide for examination of the genitalia. The Plummers
Island adults agreed in all respects with the types of F. elegantula.

Distribution.—Alberta and Saskatchewan south to Mississippi and east
to Maryland; locality records plotted in Fig. 20.

New records—ALBERTA: Medicine Hat, 30.viii.1949, L. G. Saunders,

98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 16-19. Scanning electron micrographs of Forcipomyia elegantula. 16. Larval
head, 150%; 17. Abdominal a hair of larva, 1,150; 18. Pupal respiratory horn, 550;
19. Enlarged view of spiracles of pupal respiratory horn showing extremely small
apertures of spiracular papillae, 2,100.

all stages, reared from sap of black poplar tree. DISTRICT OF COLUM-
BIA: 3.vi.1935, A. Stone, 12 (pinned). KANSAS: Riley Co., ix,x.1964, N.
Marston, malaise trap, 1¢, 12. MARYLAND: Montgomery Co., Plummers
Island, 3.vi.1976, T. J. Spilman; 10.vi.1976, Wirth and Grogan; 17.vi.1976,
Grogan, all stages. MISSISSIPPI: Washington Co., iv.1962, R. H. Roberts,
light trap, 1¢. SASKATCHEWAN: Saskatoon, 3.ix.1950, L. G. Saunders,
ex tree sap, 22, 1 larva, 1 pupa.

Biology—We reared adults from pupae collected 3,10,17.vi.1976 on
Plummers Island. A second generation may occur later in summer as Mal-
loch collected specimens in Ilinois in late June and again in August. The
second generation would then have approximately two months to develop

VOLUME 80, NUMBER 1 99

Fig. 20. North American locality records of Forcipomyia elegantula.

and offspring would probably be able to develop to third or fourth instars
before overwintering.

We found pupae in circular aggregations of a dozen or more individuals.
This behavior was observed by Saunders (1924) in many other species of
Forcipomyia (Forcipomyia). Apparently during pupation they assemble
in this circular manner with heads directed towards the center and tails
outward. This behavior would seem to be evolutionarily disadvantageous
for the species; those aggregating individuals being more or less an easily
available concentration of food for any predators present. But when the
larval structure and habits are studied more closely, the reason for this
behavior becomes apparent.

Living larvae of F. elegantula, as in other species of the subgenus that
have been observed, nearly always have conspicuous droplets of liquid on
the tips of the hastate a, p, and q hairs of the body and head. According
to Hinton (1955), Keilin (1918) was the first to comment on the biological
significance of these modified hairs. Keilin “supposed that a hygroscopic
substance on these hairs collected water from the atmosphere, which then
wetted the cuticle and in some way facilitated respiration. Since the larvae
are apneustic, the cuticle must be relatively permeable to air, and a
hygroscopic film over the surface might be expected to tide the larvae over
when exposed for short periods to lethal humidities. Keilin’s explanation

100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of the function of the setae was repeated by Saunders (1924), and Keilin
(1944) quotes Saunders as an authority for the view first published by
himself” (Hinton, 1955).

Frew (1923) in a very detailed anatomical and histological study of
the larva of F. (F.) picea (Winnertz), was apparently the first to con-
firm the secretory nature of the hastate body hairs of Forcipomyia. He
reported (p. 436): “The dorsolateral and dorsal setae of the body are
lodged over special hypoderm cells which are enormously hypertrophied.
Their nuclei differ from those of ordinary hypoderm cells in having the
chromatin scattered throughout the nucleus instead of concentrated round
its periphery; they contain a large nucleolus. The dorsolateral setae have
a central cavity extending almost to their tips, and this cavity is occupied by
an axial protoplasmic filament given off from the basal cell. In the dorsal
setae the central cavity extends right to the tip of the seta in the spear-
shaped head of which it becomes somewhat enlarged, the whole cavity
being occupied by a protoplasmic continuation of the basal cell. It is ob-
viously this cell which secretes the drop of liquid found at the apex of
each dorsal seta in the living larva. The large cephalic setae resemble
the dorsolateral body setae. The remaining setae of the body are solid and
are seated upon small thickenings of the hypoderm composed of several
slightly hypertrophied cells instead of one greatly hypertrophied cell. The
dorsal, dorsolateral and lateral setae of the body are attached to the body
wall by distinct articulations, but this does not appear to be the case
for the ventrolateral setae of the minute ventral setae.”

Hinton (1955) gave details of a very interesting series of experiments
performed by himself and his students to investigate the nature of the sub-
stance secreted through the specialized setae. They concluded that the
substance is defensive in function and appears to have no significance in
respiration. Using a laboratory colony of the ant Lasius niger L., they found
that the ants would attempt to attack larvae of Forcipomyia (F.) nigra
(Winnertz) when the latter were placed within their enclosure. Usually,
whenever the ants came close enough to bite the midge larvae, they touched
one or more droplets of liquid on the hastate setae, at which time they im-
mediately dropped the larvae and usually spent several minutes cleaning
themselves. Experiments at different relative humidities showed that at
low humidities, when droplets were not formed, larvae were dorpped less
quickly by the ants than at high humidities when the setae had large drops
on their apices. If the last larval skins were removed from midge pupae, the
ants would carry the pupae to their nest; they also succeeded in carrying
away pupae if they could attack them from the front and dislodge them
from the larval cuticle. But if the ants approached the pupae from the
side or rear, they always became smeared with the hygroscopic substance
remaining on the larval cuticle and would retire and clean themselves.

VOLUME 80, NUMBER 1 101

When the midge pupae form their usually complete circular aggregations,
they form nearly a perfect defensive barrier against such attacks by pred-
ators. The chemical nature and mode of action of the hygroscopic sub-
stance remain a mystery.

Forcipomyia (Forcipomyia) pulchrithorax Edwards
(Figs. 5, 7-11)

Forcipomyia pulchrithorax Edwards, in Saunders, 1924:209 (male, female;
England; bred from larvae in tree sap, tree wounds, wood debris, etc.);
Saunders, 1924:202 (larvae, pupae; from granular, solidifying sap from
open wounds of elm, chestnut, and ash trees; figs.); Remm, 1961:173
(descriptive notes on male, female; figs.; Estonia); Goetghebuer, 1950:2
(Belgium; fig. male genitalia); Neindorff, 1959:31 (descriptive notes on
all stages; habitat and life history notes; fig. wing; German); Zilahi-
Sebess, 1940:21, 29 (Hungary; male, female redescribed; figs.); Havelka,
1976:232 (Germany; male, female diagnosis, figs.; seasonal distr.).

Diagnosis —For detailed descriptions of all stages see Saunders (1924).
Forcipomyia pulchrithorax is a darker species and differs from F. elegan-
tula also in lacking the black band on distal “4 of hind femur; females
with pyriform spermathecae (Fig. 5); males with triangular aedeagus (Fig.
7) with a convex basal arch; pupa with a smaller tuberculate respiratory
horn (Fig. 8) bearing 15-20 spiracular papillae, shorter thoracic processes
(Figs. 9a—c) and shorter abdominal processes (Figs. 10a—c); larva with much
smaller, slightly hastate a hair (Fig. lla), shorter, stouter, doubly-branched
b hair (Fig. 11b), stouter c hair (Fig. llc), and shorter, stouter, fringed d
hair (Fig. 11d).

Specimens examined—AUSTRIA: Tirol, Igls, 900 m, 15.ix.1953, J. R.
Vockeroth, 14. ENGLAND: Cambridge, 31.viii.1922, L. G. Saunders,
from sap in elm wound, 1°, 4 larvae, 5 pupae. ESTONIA: Luua, 28.viii.
1960, H. Remm, 1¢, 29.

Literature Cited

Edwards, F. W. 1924. Appendix. Description of adults of two new species of
Forcipomyia. Parasitology 16:208—211.

Frew, J. G. H. 1923. On the larval and pupal stages of Forcipomyia piceus Winn.
Ann. Appl. Biol. 10:409-441.

Goetghebuer, M. 1950. Ceratopogonidae et Chironomidae nouveaux ou peu connus
d'Europe (Quatorzieme Note). Bull. Inst. R. Sci. Nat. Belgique 26:1-15.

Havelka, P. 1976. Limnologische und systematische Studien an Ceratopogoniden
(Diptera: Nematocera). Beitr. Entomol. 26:211—305.

Hinton, H. E. 1955. Protective devices of endopterygote pupae. Trans. Soc. British
Entomol. 12:49-92.

Johannsen, O. A. 1943. <A generic synopsis of the Ceratopogonidae (Heleidae) of the

102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Americas, a bibliography, and a list of the North American species. Ann. Entomol.

Soc. Am. 36:763-791.

1952. Guide to the Insects of Connecticut. Part 6. The Diptera or true
flies. Fasc. 5. Midges and gnats. Heleidae (Ceratopogonidae). Bull. Conn. St.
Geol. Nat. Hist. Surv. 80:149-175.

Keilin, D. 1918. Sur quelques modes particuliers de résistance des larves de
Diptéres contre la dessiccation. Bull. Soc. Entomol. Fr. 1918:102—104.

—. 1944. Respiratory systems and respiratory adaptations in larvae and pupae
of Diptera. Parasitology 36:1-66.

Malloch, J. R. 1915. Some additional records of Chironomidae for Illinois and notes
on other Illinois Diptera. Bull. Illinois St. Lab. Nat. Hist. 11:305-363.

Neindorff, U. von. 1959. Beitriige zur Kenntnis der Heleiden-Fauna (Dipt., Nemat.)
Berlins. Mitt. Deutsch. Entomol. Gesel. 18:31-35.

Remm, H. 1961. A survey of species of the genus Forcipomyia Meigen (Diptera,
Heleidae) from Estonia (In Russian, English summary). Eesti Nsv. Tead. Akad.
Juures Asuva Loodusuurijate Seltsi. 54:165-196.

Saunders, L. G. On the life history and the anatomy of the early stages of Forcipomyia
(Diptera, Nemat., Ceratopogoninae). Parasitology 16:164—-213.

Wirth, W. W. 1965. Family Ceratopogonidae (Heleidae). Pp. 121-142. In Stone,
A., et al. A Catalog of the Diptera of America North of Mexico. U.S. Dept.
Agr. Handbook 276. 1696 pp.

Zilahi-Sebess, G. 1940. Magyarorszag Heleidai. Folia Entomol. Hungarici 5:10—-133.

(WWW) Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,
USDA, c/o U.S. National Museum, Washington, D.C. 20560; and (WLG)
Department of Entomology, University of Maryland, College Park, Mary-
land 20742.

NOTE

REPLACEMENT NAME FOR DROSOPHILA NIGRICOLOR HARDY
(DIPTERA: DROSOPHILIDAE)

Drosophila nigricolor Hardy (1977. Proc. Entomol. Soc. Wash. 79(1):92)
is preoccupied by D. nigricolor de Meijere (1911. Tijdschr. Entomol. 54:
399) and D. nigricolor Strobl (1898. Mitt. Naturwiss. Ver. Steiermark
(1897). 34:266). I propose Drosophila picea Hardy as the new name for D.
nigricolor Hardy.

D. E. Hardy, Department of Entomology, University of Hawaii at
Manoa, Honolulu, Hawaii 96822.

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 103-112
GEOGRAPHIC VARIATION IN TACHYSPHEX TERMINATUS
(HYMENOPTERA: SPHECIDAE, LARRINAE)

Nancy B. Elliott and Frank E. Kurczewski

Abstract —Males of Tachysphex terminatus (Smith) were collected from
a number of populations during the summers of 1968, 1969, and 1970. Color,
size and punctation characteristics were compared statistically. All in-
dividuals collected east of the Rockies had red-tipped abdomens, but many
individuals from the West were all black. Increased melanization of the
genitalia was evident in individuals from coastal areas, both in the East
and West. There was no significant size variation, except for a sample of
very large individuals from coastal California, and one of small individuals
from Idaho. Samples from western Kansas consistently showed lower values
for vertex punctation than many of the other samples. Additional work with
museum specimens from throughout the species range has tended to sub-
stantiate many of these findings.

Geographic variation in animals has been documented for external color
patterns, and morphometric, chromosomal, physiological, and ecological
characters. There is much literature on geographic variation in insect groups
including Collembola (Christiansen and Culver, 1968 and 1969), Orthoptera
(Craddock, 1970; Masaki, 1967), Homoptera (Sokal, 1952 and 1962; Sokal
and Rinkel, 1963; Sokal and Thomas, 1965), Hemiptera (Slater and Knop,
1959), Coleoptera (Mason, 1964; Willis, 1967; Young, 1960), Lepidoptera
(Creed et al., 1962; Lucas, 1969), Diptera (Sokoloff, 1965), and Hymenoptera
(Alpatov, 1929; Michener, 1947). Within the Hymenoptera, intraspecific
variation in the Sphecidae has also been treated anecdotally as by Fernald
(1926) or in taxonomic revisions, as by Bohart and Menke (1963). Our paper
presents results of a study of geographic variation in the digger wasp
Tachysphex terminatus (Smith). This wasp is found throughout North
America from Alaska to Mexico, except Florida, nests in bare sand, and
may be locally abundant. Areas of suitable habitat are often separated by
vegetation, affording the local population some degree of isolation.

Tachysphex terminatus belongs to a group of closely related North
American species of Tachysphex including the following: Tachysphex
apicalis Fox, T. fusus Fox, T. similis Rohwer, T. plesia Rohwer and T.
linsleyi Bohart. Group members share certain diagnostic characters, in-
cluding a wide least interocular distance, relatively long erect setae on the
vertex, and prominent convexities behind each posterior ocellus. Tachysphex
terminatus may be distinguished from other group members by the fol-
lowing set of characteristics: a ratio between least interocular distance
and head width of .29-.33, a smooth punctate frons, and an unnotched

104. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Collection localities for males for 1968, 1969 and 1970. (01) Scullville, New
Jersey 1968, 12; (02) New Gretna, New Jersey 1969, 16; 1970, 13; (03) Colonie, New
York 1968, 13; (04) Clifton Park, New York 1969, 20; (05) Albany, New York 1970, 14;
(06) Chittenango, New York 1968, 24; 1969, 23; 1970, 20; (07) Presque Isle, Pennsyl-
vania 1968, 7; (08) Wamego, Kansas 1969, 14; 1970, 15; (09) Lakin, Kansas 1968, 12;
(10) Kendall, Kansas 1969, 25; 1970, 11; (11) Hoback Junction, Wyoming 1970, 8; (12)
St. Anthony, Idaho 1970, 18; (13) Mendocino Co., California 1970, 19.

clypeus. The behaviors of several species in the T. terminatus group
have been studied by Kurczewski (1964, 1966), allowing correlation with
findings on other aspects of their biologies. There are indications that
patterns of geographic variation could complicate taxonomic studies of the
group. Western individuals of T. terminatus, for example, appear super-
fically different from their eastern conspecifics. The range of morphological
variation in each species is unknown, and it is desirable to study the extent
of variation in each, before undertaking further ethological or taxonomic
studies.

Methods and Materials

Adults were collected during the summers of 1968, 1969, and 1970.
Collection localities are shown in Fig. 1. Preliminary comparisons between
successive generations in a single locality (Chittenango, New York) in
1968 revealed significant mean differences in several characters for fe-
males, but not for males (Elliott and Kurczewski, 1974a). Since seasonal
variation might complicate the study of geographic variation, only males

VOLUME 80, NUMBER 1 105

were collected for the remainder of the study. Specimens from museum
collections were used to supplement data based on field-collected specimens
for localities from which samples of five or more males were available. The
collections localities for museum specimens and the number of specimens
for each locality are as follows: Ithaca, New York (23); southern New
Jersey (24); Riley and Pottawatomie Co., Kansas (13); Wisconsin (19);
Lewisville, Arkansas (12); Kill Devil Hills, North Carolina (13); Virginia
(10); Galveston, Texas (12); Mendocino Co., California (5); San Francisco,
California (18); Lassen Co., California (9); St. Anthony, Idaho (6); southern
Quebec (12).

Measurements and counts on specimens were made using a binocular
dissecting microscope with a grid and ocular micrometer. A factor analysis
on 18 morphometric characters indicated head width was a good indicator
of general size. Of the characters which loaded heavily in size, it was the
easiest to measure accurately (coefficient of reliability = .999 for remeasure-
ments). For this reason it was compared as a general size measure through-
out the remaining analyses. Puncture counts per grid unit on the vertex
were transformed for statistical analysis by computing the mean of the
square root of the two counts on each individual. Genitalia were removed
from males collected in 1970, mounted on slides, and the color of the volsella,
coded on a scale from 1-3 (light-dark), was recorded. The percent of each
sample with red-tipped abdomens was also recorded.

Variation between annual samples was tested with one-way analysis of
variance for size-related characters and puncture counts. Homogeneity of
variances was first tested with Bartlett's Test (Sokal and Rohlf, 1969:370).
For homogeneous variances, means were tested with Model 1 ANOVA
to show actual mean differences, and variation was also partitioned into that
due to variation within and between groups (Model 2 ANOVA). An F
approximation was applied for unequal variances (Sokal and Rohlf, 1969:
372). Multiple comparisons were made with Student-Newman-Keuls Test
(Sokal and Rohlf, 1969:242).

Results

Table 1 shows partition of variance (Model 2 ANOVA) for samples and
years. During 1968 and 1969, when all samples were collected east of
the Rockies, there was no significant size-related variation. In 1970, when
populations from the West were also included, the Model 1 ANOVA in-
dicated significant differences for head width (F = 9.55; P < .05). The
distribution of probit means for head width in 1970 is shown in Fig. 2A.
This distribution was characteristic of those for a number of size-related char-
acters (Elliott, 1971). The lowest mean value, that for a sample from
St. Anthony, Idaho, was significantly different from those values for

106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Partition of variance in Tachysphex terminatus.*

% Variance % Variance
Characters (localities) (individuals)
1968
Head width 1.05 98.90
Punctation/vertex 34.35 65.65
1969
Head width .64 99.35
Punctation/vertex 26.00 74.00
1970
Head width 37.36 62.64
Punctation/vertex 12.15 87.85

*Summaries of ANOVAs may be obtained from senior author.

all other samples except the one from southern New Jersey. The highest
value, from Mendocino Co., California, was significantly different from all
others, except that from western Kansas. Museum samples showed sim-
ilar trends in geographic variation (F = 3.34; P < .05; see Fig. 2B). But
differences in sample size made it difficult to demonstrate statistical sig-
nificance in the multiple comparisons tests. Means for head widths for
specimens from Mendocino County still ranked among the largest, along
with those for specimens from Wisconsin and Quebec. Many of the smaller
specimens were from St. Anthony, Idaho. Some of the larger males among
the museum specimens were from Mendocino Co., California, while some of
the smallest came from nearby San Francisco.

Vertex punctation varied significantly during all years of the study (in
1968, F = 8:77; P <.05; in 1969, F = 7.70; P*< 705-41) 1970 }ee eee
P < .05). (See Fig. 3.) In 1968, samples from Colonie, New York and
Presque Isle St. Park, Pennsylvania, had the lowest mean values for this
character. These values were significantly different from those for cen-
tral New York samples. A low mean value for this character in the sample
from western Kansas was significantly different from the means from central
New York samples. In 1969, the western Kansas sample had a signif-
icantly lower value for this character than all other samples. In 1970, the
sample from this locality again had the lowest mean value for the charac-
ter, and a sample of individuals from St. Anthony, Idaho had a significantly
higher mean value for the character than all other samples.

All specimens collected in the East during 1968 and 1969 had red-tipped
abdomens. In 1970, a number of all black males were collected from the
West. None of the males from Mendocino Co., California were red-tipped;

VOLUME 80, NUMBER 1 107

Fig. 2. Size variation in Tachysphex terminatus males.

A. Distribution of probit means for head width in 1970 samples. SNK Test:
3.36 4.21 4.74 4.87 5.56 5.77 6.63

B. Probit means for head width in museum specimens. SNK Test:
3.52 3.72 3.90 4.10 4.43 5.05 5.10 5.23 5.50 5.68 5.82 6.36 6.60

108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Probit means for vertex punctation.

A. 1968 samples. SNK Test:
3.89 3.94 5.50 5.82 5.84

B. 1969 samples. SNK Test:
3.34 4.75 4.85 5.28 5.40 6.38

C. 1970 samples. “SNK Test:
3.52 4.09 4.20 5.16 5.31 5.50 5.66 6.59

VOLUME 80, NUMBER 1

Fig. 5. Distribution of probit means for volsella color (1970).

110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

56% of the Idaho males and 12% of the males from Wyoming were red-
tipped. Figure 4 shows the distribution of red-tipped males in the 1970
field samples. Similar patterns were evident in museum specimens. All
males from the East were red-tipped, whereas males from western localities
were frequently all black. Among museum specimens, there were no red-
tipped males from St. Anthony, Idaho or Mendocino Co., California, while
63% of the males from Lassen Co., California and 11% of those from San
Francisco were red-tipped.

All black specimens showed the greatest degree of melanization on the
volsella, with those from coastal California having the darkest volsellae
(Fig. 5). The greatest melanization in red-tipped males occurred in speci-
mens from coastal New Jersey. In fact, they showed more melanization
than all black males from Wyoming.

Discussion

The most distinctive geographic patterns in size variation showed in
samples collected in 1970. Excluding the samples from Idaho and Wyoming,
there seemed to be a cline of increasing size from the smallest individuals
collected near coastal New Jersey to the largest individuals from the Cali-
fornia coast. Specimens from Idaho and Wyoming, which were smaller
than those from other localities, seemed to show a second effect acting upon
size, possibly related to altitude. These localities had the highest altitudes
of any sampled (4,900’ and 6,000’, respectively). Thus an inverse relation-
ship between size and altitude appeared to be superimposed on the general
clinal pattern of increasing size from east to west. While no distinct longi-
tudinal cline of size showed up in museum samples, the inverse relationship
between size and altitude was indicated. The smallest specimens came from
Lassen Co., California, at an altitude of approximately 5,000’. The fact
that specimens from western Kansas consistently exhibited relatively sparser
vertex punctation than those from other areas seemed to indicate selection
acting on this or a related character. High density in vertex punctation in
the Idaho specimens, which were the smallest individuals, suggested a
predictable inverse relationship with size. The most obvious variation in
T. terminatus was in abdomen color. Western males were often all black,
although females occurring with them were frequently red-tipped. Males
from coastal California conformed to the stereotype of specimens from
coastal climates (see Fernald, 1926). They were large and dark with much
melanization on the volsella. Coastal New Jersey wasps, although smaller
and red-tipped, also showed increased melanization on the volsella, presum-
ably an effect of moist coastal climates. Elliott and Kurczewski (1974b) have
already reported that the presence or absence of potential competitors in
the habitat may affect the morphology of related Tachysphex species. The

VOLUME 80, NUMBER 1 lll

present study suggests that a number of environmental factors act in-
dependently to produce morphological variation throughout the range of T.
terminatus. These factors may include effects of altitude in relation to
size, humidity to increase melanization in coastal environments, and various
effects acting on punctation characteristics throughout the range.

Literature Cited

Alpatov, W. W. 1929. Biometrical studies on variation and races of the honeybee
(Apis mellifera L.) Quart. Rev. Biol. 4:1-58.

Bohart, R. M., and A. S. Menke. 1963. A reclassification of the Sphecinae with a
revision of the Nearctic species of the tribes Sceliphronini and Sphecini (Hyme-
noptera: Sphecidae). Univ. Calif. Publ. Entomol. 30:91-182.

Christiansen, K. A., and D. Culver. 1968. Geographic variation and evolution in
Pseudosinella hirsuta. Evolution 22:237-255.

——. 1969. Geographical variation and evolution in Pseudosinella violenta (Folsom).
Evolution 23:602-621.

Craddock, E. 1970. Chromosome variation in a stick insect Didymuria violescens
(Leach). Science 167:1380-1382.

Creed, E. R., W. H. Dowdeswell, E. B. Ford, and K. G. McWhiter. 1962. Evolu-
tionary studies on Maniola jurtina: the English mainland, 1958-1960. Heredity
17:237—265.

Elliott, N. B. 1971. Morphological variation in Tachysphex terminatus (Smith) and
Tachysphex similis Rohwer. (Hymenoptera: Sphecidae, Larrinae). SUNY Col-
lege of Environmental Science and Forestry, Syracuse, N.Y. Ph.D. Thesis.

Elliott, N. B., and F. E. Kurezewski. 1974a. Seasonal variation in Tachysphex termi-
natus (Smith) (Hymenoptera: Sphecidae, Larrinae). J. N.Y. Entomol. Soc.
82:268-270.

—. 1974b. Character displacement in Tachysphex terminatus and T.. similis
(Hymenoptera: Sphecidae, Larrinae). Ann. Entomol. Soc. Am. 67:725-727.

Fernald, H. T. 1926. Climate and coloration in some wasps. Ann. Entomol. Soc. Am.
19:88—92.

Kurczewski, F. E. 1964. A comparative ethological study of some Nearctic digger

wasps of the genus Tachysphex Kohl. (Hymenoptera: Sphecidae, Larrinae).

Ithaca, New York. Cornell Univ. Ph.D. Thesis.

1966. Comparative behavior of male digger wasps of the genus Tachysphex
(Hymenoptera: Sphecidae, Larrinae). J. Kans. Entomol. Soc. 39:436—453.
Lucas, A. M. 1969. Clinal variation in pattern and color in coastal populations of

the butterfly Tisiphone abeona (Donovan) (Lepidoptera: Satyridae). Aust. J.
Zool. 17:37-48.

Masaki, S. 1967. Geographic variation and climatic adaptation in a field cricket
(Orthoptera: Gryllidae). Evolution 21:725-741.

Mason, L. G. 1964. Geographical temporal variation in natural populations of
Tetraopes tetraophthalmus. Syst. Zool. 13:161-181.

Michener, C. D. 1947. A character analysis of a solitary bee, Hoplitis albifrons
(Hymenoptera: Megachilidae). Evolution 1:172-185.

Slater, J. A., and N. L. Knop. 1959. Geographic variation in the North American
milkweed bugs of the Lygaeus kalmii complex. Ann. Entomol. Soc. Am. 52:1221-
1232.

Sokal, R.R. 1952. Variation in a local population of Pemphigus. Evolution 6:296-315.

112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

—. 1962. Variation and co-variation of characters of alate Pemphigus
populi-transversus in eastern North America. Evolution 16:227-245.

Sokal, R. R., and R. C. Rinkel. 1963. Geographic variation of alate Pemphigus
populi-transversus in eastern North America. Univ. Kans. Sci. Bull. 44:467-
507.

Sokal, R. R., and F. J. Rohlf. 1969. Biometry. The Principles and Practice of Sta-
tistics in Biological Research. W. H. Freeman and Co., San Francisco. Pp. 776.

Sokal, R. R., and P. A. Thomas. 1965. Geographic variation of Pemphigus populi-
transversus in eastern North America: stem mothers and new data on alates.
Univ. Kans. Sci. Bull. 46:201-252.

Sokoloff, A. 1965. Geographic variation of quantitative characters in populations
of Drosophila pseudoobscura. Evolution 19:300-310.

Willis, H. L. 1967. Bionomics and zoogeography of tiger beetles of saline habitats
in the central United States (Coloptera: Cicindelidae). Univ. Kans. Sci. Bull.
A47:145-313.

Young, F. N. 1960. Regional melanism in aquatic beetles. Evolution 14:277-283.

Department of Entomology, SUNY College of Environmental Science
and Forestry, Syracuse, New York 13210.

Present address—(NBE), Department of Biology, Hartwick College,
Oneonta, New York 13820.

NOTE

NEW SYNONYMY IN NERTHRA SAY
(HEMIPTERA: GELASTOCORIDAE)

Two specimens of predaceous bugs of the family Gelastocoridae inter-
cepted in Miami, Florida in an air shipment of ferns from Colombia on
22 August 1975 by F. Matthews confirm a suspected synonymy (Todd.
1955. Univ. Kans. Sci. Bull. 37(1), no. 11, p. 389). The specimens, a male
and a female, represent respectively Nerthra ater (Melin, 1929) and Nerthra
rudis (Melin, 1929) described in Zoologiska Bidrag Fran Uppsala, Band
12, p. 185 and p. 182. In 1955 I suggested that if the two names did apply
to the two sexes of one species, Nerthra ater (Melin, 1929) would fall as the
junior synonym. This comment was based on the concept of page priority
current at that time. Since the shape of the male paramere is a better
recognition character than the overlapped basal part of the right ovipositor
lobe of the female, I place Nerthra rudis (Melin, 1929) as the junior
synonym of Nerthra ater (Melin, 1929) [New synonymy].

E. L. Todd, Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,
USDA, c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(1), 1978, pp. 113-127
ETHOLOGY OF CEROTAINIA ALBIPILOSA CURRAN
(DIPTERA: ASILIDAE) IN MARYLAND:
PREDATORY BEHAVIOR!

A. G. Scarbrough

Abstract.—A field study of the predatory behavior of Cerotainia albipilosa
Curran is reported. The asilid foraged from leaves and stems of sunlit plants
found along the margins of clearings and paths. The forage flight patterns
are discussed. Only 14% of the flights were successful. Most prey (93%)
were captured within 2 m of foraging sites. Cerotainia albipilosa exhibited
specialized methods for capturing prey. The hypopharynx was inserted
into the dorsum or one end of the body. The asilid immobilized prey in
the air before returning to foraging sites. Individual feeding times averaged
5 minutes but varied considerably depending on prey characteristics and
weather conditions. Interfeeding times ranged from 0-35 minutes. While
feeding, the asilid frequently hovered in front of a perch and manipulated
the prey. Over 94% of the prey belonged to five insect orders (Diptera,
Coleoptera, Hymenoptera, Psocoptera and Hemiptera-Homoptera) and
67% to two (Diptera and Hemiptera-Homoptera). Prey were usually soft-
bodied, weak flyers and averaged 1.6 mm in length and 0.05 mg in weight.
Females captured larger prey than males. Mean predator to prey size and
weight ratios were 3.7 and 7.3, respectively. Several factors which influenced
prey selectivity are discussed. This asilid was sometimes preyed on by
spiders, vespid wasps and other asilids.

In the first part of this study, Scarbrough and Norden (1977) reported
on the diurnal activity rhythm and seasonal distribution of Cerotainia
albipilosa Curran in Maryland. They reported that most of the species
flight activities were centered around predation and reproductive behaviors.
The purpose of this paper is to report on the predatory behavior of the
species. A third paper will deal with reproduction.

Methods and Procedures

General methods and procedures for observations made in the field and
the study site were described in a previous paper (Scarbrough and Norden,
1977). Observations were recorded in a notebook at the study site or on
a Sanyo solid state tape recorder model 2212 and were later transcribed
into a notebook. When possible, photographic records of different behaviors
were taken. A Honeywell Pentax 35 mm, single lens reflex camera in con-
junction with telephoto lenses and extension tubes was used to obtain
close-up pictures.

114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Collections of prey were made during the summers of 1973 and 1974. Care
was taken to collect prey from each segment of the species diurnal activity
period through the season to obtain a maximum variety of prey types. Most
prey were obtained by capturing asilids seen with prey with a 15 dram snap
top vial as I walked through an observation site. The asilid usually ceased
feeding immediately and dropped the prey in the vial, although some
continued to feed, only to drop them at a later time. This procedure mini-
mized damage to prey specimens. Once the prey was dropped, the asilid
was identified and usually released. Some prey were recovered from
leaves below feeding sites where asilids had dropped them. The prey were
then transferred to vials containing 70% ETOH with the following informa-
tion: Sex of predator, date, location, and collector.

Prey were analyzed by making two types of body measurements: Size and
weight. To determine prey size, each specimen was measured for total
body length to the nearest 0.5 mm using an ocular micrometer. From a
reference collection made at the study site, prey of selected lengths were
oven dried to 38°C for 30 minutes and weighed to the nearest 0.01 mg.
Similar measurements of the asilid were obtained by measuring and weigh-
ing ten specimens in the same way as described for prey.

Some prey were identified by the author with the aid of the
collection housed at the Towson State University Museum. However, most
prey were shipped to the Systematic Entomology Laboratory (SEL), IIBII,
Agric. Res. Serv., USDA, Beltsville, Maryland and to the Smithsonian In-
stitution (SI), Washington, D.C. A few specimens were also identified by
recognized specialists in other parts of the country.

Results and Discussion

Foraging and feeding.—Cerotainia albipilosa foraged under bright skies
from leaves and stems of vegetation along margins of clearings and paths,
and at air temperatures above 19°C (Scarbrough and Norden, 1977). These
flies were usually active on foraging sites, rapidly turning their bodies to
face one direction and then another. Each movement was preceded by
quick movements of the asilid’s head as potential prey flew in front of or to
the side-of the foraging site. Presumably this behavior enables the asilid
to better perceive potential prey and, at the same time, it is in a more
suitable position to make a direct forage flight (Dennis and Lavigne,
1975). Similar foraging behavior is exhibited by Stichopogon trifasciatus
Say (Lavigne and Holland, 1969) and by several species of Holopogon
(Lehr, 1972; Dennis and Lavigne, 1975).

Most foraging flights were directed toward potential prey in the air
after they flew near the asilid. Upon perceiving prey, the asilid “leaped”
into the air, flew at an oblique angle to the prey’s path, converged and

VOLUME 80, NUMBER 1 115

intercepted it a short distance from the foraging site. Prey were usually
captured at the same height or slightly above the foraging site. Adjust-
ments in flight patterns occurred once the asilid took flight. In most cases,
after leaving a perch, the asilid’s flight path was initially below the prey,
although a noticeable elevation in its flight was detected near the intercep-
tion point. Such behavior facilitated capture, and subsequent immobiliza-
tion of the prey, in flight. The asilid also compensated for overestima-
tions of flight speed of prey by looping about 5-7 cm above the prey’s path,
hesitated in this position, and then attacked as the prey passed below at
the interception point. They frequently flew toward potential prey only
to turn away a few cm from foraging sites or near the interception points.
Dennis and Lavigne (1975) termed these flights “investigatory” since they
permit discrimination between unsuitable and suitable prey before actually
attacking. In a few cases the asilid fluttered its wings and tilted the body
forward without leaving a perch in response to passing insects. Unless
disturbed, the asilid returned to the same foraging site following each
flight.

Females of C. albipilosa often flew after insects and falling objects that
were several times (>3%) larger than themselves. Several females on separate
occasions foraged after butterflies (Pipilio glaucus L.) and ichneumonid
wasps (Ophion sp.), but invariably they turned away from them near the
interception points. Two other females attempted to capture a_halictid
bee (Augochloropsis sp.) and a Japanese beetle (Popillia japonica Newman).
They attacked the insects and held onto their wings for about 1-2 seconds
before releasing their grasps and returning to foraging sites. Falling leaves
were also frequently pursued by females. Similar observations of asilids
flying toward falling objects have been reported by other investigators
(Melin, 1923; Parmenter, 1952).

Foraging flights were directed at potential prey flying within a range
from 8 cm-4 m in front of or to one side of a perched asilid. Most flights
occurred when prey were within 2 m of foraging sites. Flights that re-
sulted in prey captures were made when prey were at the following distances
from foraging sites: 68% at 8 cm-l m, 25% at 1-2 m and 7% at 2-4 m.
Ninety percent of the capture flights within 1-20 cm resulted when prey
flew from the vegetation and to one side of a perched asilid. Prey cap-
tured at greater distances were invariably flying in front of foraging sites.
Furthermore, prey captured beyond 2 m by females were characteristically
large (>3 mm) whereas those taken at closer ranges represented all size
classes. Males captured smaller prey (<2 mm) and were rarely successful
(<1%) beyond 2 m. Thus, it is assumed that the visual capability of C.
albipilosa is more acute at short range.

Few foraging flights resulted in prey capture. Only 14% of the flights
were successful regardless of distances from foraging sites. However males

116. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Cerotainia albipilosa feeding on Aleurochiton sp. (Hemiptera-Homoptera:
Aleyrodidae).

were more successful than females with 17% and 11%, respectively. This
forage success differential is directly related to disruptions of the female’s
foraging behavior caused by courting and mating males (Scarbrough and
Norden, 1977) and to females foraging longer distances in pursuit of larger
prey than males. It is also interesting to note that if these flights were
separated into no-contact flights (investigatory) and contact-flights (forag-
ing), as defined by Dennis and Lavigne (1975), then foraging success is
much greater (91%). Nevertheless, if one considers the amount of energy
which is expended in foraging (investigatory and contact flights), the
tendency to fly after most moving objects appears to be quite wasteful. At
the same time, since asilids can probably detect only a rough outline of a
moving dark body (Melin, 1923), this behavior would maximize the
probability that they pursue and capture suitable prey.

Like many asilid species (Dennis and Lavigne, 1975; Shtakel’berg, 1950;
Fackler, 1918; Horning and Barr, 1970; Scarbrough and Sipes, 1973;
Schmid, 1969; Wallis, 1913; Zinovera, 1959), C. albipilosa has specialized
methods of capturing prey and common sites for inserting the hypopharynx.
The asilid usually captured prey along their dorsolateral surfaces, and
impaled them on the dorsum or at one end of the body (Fig. 1). Large
bodied insects with hard wings, i.e. the first pair of wings of beetles and
cicadellids, were captured with their wings extending in a flight position,

VOLUME 80, NUMBER 1 117

exposing the soft underlying integument. The hypopharynx was inserted
into this area, usually near the junction of the thorax and abdomen.
Staphylinid beetles and formacine ants were similarily impaled through thin
or soft areas of the integument; i.e. eyes and conjunctiva between sclerites
at the end of the abdomen. The remaining prey had “soft” integuments,
and the hypopharynx was inserted at various places on the dorsum.

Once prey were captured, C. albipilosa usually immobilized them in the
air before returning to foraging sites. Most immobilizations occurred in
flight as soon as they were captured. When prey were large (>2.0 mm),
the asilid hovered in front of the foraging site, manipulated them using
all six tarsi, and inserted its hypopharynx. Immobilizations were rare
events at foraging sites since only one successful and four attempts were
observed in over 1,200 observation hours. In these instances, the asilids
landed on a leaf, fell on one side, and used all six tarsi to manipulate the
prey. The prey were large, and these behaviors were preceded by unsuccess-
ful immobilizations in the air before landing.

Like other asilid species (Dennis and Lavigne, 1975), C. albipilosa some-
times foraged with prey impaled upon its hypopharynx. On several occasions
the asilids returned with two prey impaled upon their mouth parts, one was
the original and the second a new prey. The hypopharynx had been forced
completely through the former, and the latter was projecting from its
apex. Usually when two prey were captured, the last captured was lost
or dropped during manipulation.

Unless disturbed, C. albipilosa remained at its initial foraging site until
feeding was completed. Males were more “nervous” than females and were
easily disturbed. When feeding was completed, the asilid moved its fore
tarsi in alternating sequences and disengaged the prey from the hypo-
pharynx. Prey were also discarded by the asilid in flight for another prey,
but without using its tarsi. Other investigators (Lavigne and Holland,
1969; Dennis and Lavigne, 1975) have also observed this behavior and have
suggested that the hypopharynx is merely retracted into the labrum, allowing
gravity to “pull” the prey off the proboscis. While feeding, the asilid con-
tinued to observe other moving “prey” which flew near its perch with rapid
movements of the head and body.

When feeding, C. albipilosa frequently hovered in front of a perch and
manipulated the prey. The asilid used all six tarsi to disengage, to rotate
and to re-insert the hypopharynx at a new location in the prey. It then
returned to the feeding site, re-oriented its body into a foraging position
and continued to feed. Prey were sometimes dropped during manipula-
tion in which case the asilid usually returned to the foraging site. However
foraging flights were occasionally initiated from the hover position when
prey were dropped. The mean time for manipulating prey was 6.1 seconds.

Cerotainia albipilosa fed on prey for an average of 5 minutes. The

118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

length of individual feedings varied considerably, ranging from 1-81 minutes
depending upon the size and shape of prey and weather conditions. Aphids
such as Aphis sp., Macrosiphum sp., and Myzus sp., which had soft bulbous
bodies and average lengths of 2 mm, were fed on for 1-2 minutes. Dipterans
and psocopterans, which also had similar integuments and average body
lengths, had tubular shapes and were fed on for an average of 3 minutes.
Larger prey, such as reproductive ants (Ponera pennsylvanica Buckley
and Lasius sp.) which have hard exoskeletons and constricted “waists,”
were fed on for an average of 20 minutes. The asilid frequently manip-
ulated them and alternated the points at which the hypopharynx was
inserted from the eyes to the ends of their abdomens. In other large hard
bodied prey, such as Gymnetron pascuorum (Gyll.) and Pseudopentarthrum
sp. (Colopetera: Curculionidae) and Macrosteles sp. (Homoptera: Cicadel-
lidae), the hypopharynx was alternated from the dorsum below the elytron
to the end of the abdomen. Manipulations of aphids, dipterans, psocopterans
and other prey with similar body characteristics were rarely observed.
Weather conditions frequently influenced the duration of feeding. One
asilid fed on Ponera pennsylvanica (3.0 mm) for 81 minutes under over-
cast skies. Feeding on various prey of a similar size under sunlit conditions
ranged from 10-20 minutes. Similar observations were reported by Dennis
and Lavigne (1975).

The average interfeeding time for C. albipilosa was 6 minutes with a
range between 0 and 35 minutes. The 0 minute interfeeding time was for
individuals who dropped prey and immediately captured new ones. Based
upon this data, the average time for a complete feeding and the average
9 hr foraging period, C. albipilosa could theoretically feed on a maximum
of 49 prey/day. These calculations, like those reported for other asilid
species (Lehr, 1964; Musso, 1971; Dennis and Lavigne, 1975, 1976a,
1976b), overestimate the number of prey fed upon by this asilid, since it did
not forage and feed continually but often engaged in other activities such
as courtship and mating (Scarbrough and Norden, 1977). A more reasonable
estimate would be 25-30 prey/day.

Prey.—Most prey of C. albipilosa, like other aslid species (Hobby, 1931;
Brues, 1946; Poulton, 1906; Cole and Lovett, 1921: Melin, 1923; Dennis and
Lavigne, 1975), belong to a few orders of insects. Of the ten orders of prey re-
corded for this species, over 94% belong to five insect orders and formed
98% of the prey dry weight (Table 1). Prey belonging to Diptera and
Hemiptera-Homoptera formed the major prey orders, comprising over
67% of the prey captured and 63% of the prey dry weight. The orders
Coleoptera, Hymenoptera and Psocoptera were less significant, forming
27.4% of the total number of prey and 34.1% of the prey dry weight. These
results are consistent with prey availability. When sweep samples (N = 25)
of vegetation were taken at various times at the study sites, Diptera and

VOLUME 80, NUMBER 1 119

Table 1. Dietary composition of Cerotainia albipilosa at the Loch Raven Watershed
in Baltimore County, Maryland.

Percentage
No. Percentage Dry total
Order prey prey weight (mg) dry wt.
Diptera 274 37.1 8.2 24.1
Hemiptera—

Homoptera 225 30.4 S'S 39.7
Hymenoptera 88 ILLS) 3:5 10.3
Coleoptera 60 8.1 4.2 12.3
Psocoptera 55 7.4 3. HES
Others 37 5.0 0.7 2.0

Totals 739 34.0

Hemiptera-Homoptera were more abundant per sample than other prey
orders (x*/sample; 31.9% Diptera; 39.3% Hemiptera-Homoptera; 19.87%
Coleoptera, Hymenoptera, Psocoptera; 9.0% others).

The major prey orders consisted of a wide diversity of species and
families (see list of prey). At least 56 and 63 species belonged to 15 and
10 families of Diptera and Hemiptera-Homoptera, respectively. In Coleop-
tera, Hymenoptera and Psocoptera, the prey consisted of a relatively small
number of species (20, 25, 8, respectively) but belonged to several families
(12, 13, 7, respectively). It is also interesting to note that within the two
major orders, most prey belong to three categories. Over 61% of Diptera
belong to Nematocera, and at least 62% of Hemiptera~-Homoptera belong
to Cicadellidae (22%) and Aphididae (40%). These results were not sur-
prising since the majority of the prey are some of the most abundant plant
feeders inhabiting woodland ecosystems.

The average prey captured by C. albipilosa was 1.6 mm in length and
0.05 mg in weight (Table 2). The mean predator to prey size and weight
ratios were 3.7 and 7.3, respectively. However prey varied considerably
ranging from 1-4 mm in length and from 0.01-0.12 mg in weight. Males
were rarely found with prey larger than 2 mm in length and 0.09 mg in
weight whereas females were found with prey of all class sizes. Most
of the prey of males belonging to Diptera, Hemiptera~-Homoptera and
Hymenoptera consisted of nematocerans, aphids and Apocrita parasitoids,
respectively. In addition to the latter prey, females captured a dispropor-
tionately greater sample of larger prey, i.e. cicadellids and reproductive
formicids, than males.

The tendency of several asilids to capture and feed upon soft-bodied
insects is assumed to be directly related to the inability of weak mouth
parts to penetrate hard cuticles (Melin, 1923; Martin, 1968; Dennis and
Lavigne, 1975). The predaceous behavior and prey records of C. albipilosa

120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Mean body lengths and weights of prey arranged according to the sex of
the predator.

Predator sex“

x prey lengths (mm) x prey dry weights (mg)

Orders of Both Both

prey Male Female sexes Male Female sexes
Diptera 1.4 1s 1.4 0.02 0.03 0.03
Hemiptera—

Homoptera 1.6 2.0 1.9 0.05 0.07 0.06
Hymenoptera IES 2.1 1.8 0.01 0.06 0.04
Coleoptera 1.8 Led Ls 0.06 0.07 0.07
Psocoptera 1.4 15) Ly) 0.06 0.07 0.07
Miscellaneous 1.4 Osa 1.0 0.02 0.02 0.02

Means 1k5 tay 1.6 0.04 0.05 0.05

“ Predator lengths (mm) 5.6 6 6, 6.492 2; weights (mg) 0.234 ¢, 0.432 9.

support this assumption. Over 73% of the prey were soft-bodied. Among
the remaining prey, which included Coleoptera, Hymenoptera, Strepsiptera
and Cicadellidae, the predator utilized specific techniques to capture and to
immobilize them. The hypopharynx was invariably inserted in areas where
the cuticle was thin or soft.

Other investigators (Lehr, 1958; Hobby, 1931; Dennis and Lavigne,
1975) have suggested that differences in predatory habits of the two sexes
were the result of temporal segregation of activity patterns, and differential
densities and nutrition requirements of females. This study supports these
suggestions. Males of C. albipilosa spend less time foraging and feeding in
afternoons, and more time searching for and mating with females (Scar-
brough and Norden, 1977). Females foraged throughout the day, and
thus were found more frequently with prey than males. Furthermore sev-
eral prey types captured by females were active and abundant at times
when males were involved in other activities. Females captured most of
the coleopterans and psocopterans between 1:00 and 4:00 PM. Flying
reproductive ants swarmed and were captured during hot humid afternoons
which were preceded by rain. Females, which had well-developed ovaries,
evidenced by the presence of numerous eggs, weighed much more than an
equal number of males (N = 20, x = 0.43 mg 2 2, 0.23 mg ¢ 2). However,
females whose ovaries and eggs were removed by dissection, weighed
about the same (N = 20, x = 0.28 mg) as males. Thus the difference in body
weights of the sexes is indirectly related to differences in predatory habits.
Females must spend more time foraging and feeding to obtain additional
nutrition for the continuous production of eggs.

VOLUME 80, NUMBER 1 121

Movement of prey evokes feeding behavior in numerous predatory ani-
mals, provided that the object falls within certain size limits (Marler and
Hamilton, 1956). Melin (1923) suggested that the vision of asilids is not
well developed and that they perceive prey as dark moving objects.
Cerotainia albipilosa appears to be “programmed” to forage after almost
any small moving object that passes near its perch. Motion of insects in
flight and falling leaves were sufficient to stimulate the asilid to leave a
perch in pursuit. All prey, except spiders, were winged and captured
in flight. The spiders were immature, and in effect “flying” since they were
either “ballooning” or moving at the ends of suspended silk threads. Wings
of most prey were large and extended to or beyond their bodies. Further-
more, prey moving in front of a perched asilid interrupt the rays of
light, casting a shadow upon itself. The fluttery motion produced by
large wings of soft-bodied prey, together with a dark shadowed body,
greatly increase the total size of the prey’s body, and undoubtedly form the
major cues which stimulate the asild to leave its perch in pursuit.

A few asilid species are apparently capable of utilizing color to select
prey (Linsley, 1960; Bohart, 1958). Dennis et al. (1975) showed that
when Efferia frewingi Wilcox was presented with black, orange and
white models of various sizes and shapes, the asilid responded preferentially
to black, oblong ones. However use of colors to select prey by C. albipilosa
was not detected. Color of prey integuments varied considerably, ranging
from light yellow to dark brown. Color detection would be difficult at best
and probably could not occur until the predator was about to, or was in
contact with the prey because 1) the flight path of the asilid is slightly
below that of the prey until the two arrive at the interception point, and
2) the asilid approaches the prey on its shadowed side.

The following is a list of prey taken by C. albipilosa at the Lock Raven
Watershed, in Baltimore County, Maryland. All prey were collected at
the study site between 30 June and 30 August of 1973 and 1974. In some
instances prey is presented only to the order or family level since specific
identifications are not yet available. Each notation of prey refers to a
single record in the list of prey unless followed by a number in paren-
theses. All prey are adults except for Araneida. W. B. Peck (Araneida);
D. R. Smith (Isoptera); E. L. Mockford (Psocoptera); K. O'Neill (Thysanop-
tera); J. L. Herring, L. M. Russell, J. P. Kramer, R. C. Froeschner (Hemip-
tera-Homoptera); C. W. Sabrosky, W. W. Wirth, R. J. Gagné, G. C. Steyskal,
L. Knutson (Diptera); A. S. Menke, B. D. Burks, P. M. Marsh, D. R. Smith
(Hymenoptera); D. M. Anderson, R. E. Warner, R. E. White. J. M. Kingsol-
ver, P. J. Spangler (Coleoptera); D. R. Davis (Lepidoptera); are thanked
for the identification of their respective groups. Froeschner, Spangler and
Peck are with the Smithsonian Institution.

Prey captured by C. albipilosa. ARANEIDA, Unidentified (1) 31.VII.74;

122 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Theridiidae, Oecobius sp. 7.VII.73; Linyphiidae (2) 31.VIII.74; Araneidae
(2) 31.VII.74; Agelenidae (3) 30.VII.74, 1.VIII.74, 3.VIII.74; Clubionidae,
Clubioninae (3) 31.VII.74; Thomisidae, Philodrominae (2) 31.VII.74, 12.
VIII.74; Salticidae (3) 10.VII.74, 16.VII.174, 17.VII.74; ISOPTERA, Rhino-
termitidae, Reticulitermes flavipes (Kollar) (6 winged reproductives) 1.VII.
73; PSOCOPTERA, Amphipsocidae, Polypsocus corruptus (Hagen) 16.VII.
74, 30.VII.74; Caeciliidae, Caecilius aurantiacus (Hagen) 19.VII.74,
31.VII.74, 6.VIII.74; Ectopsocidae, Ectopsocopsis cryptomeriae (EndlIn.)
(2) 20.VIL.73,. (7) 12.VIL.74, (9) 17.VIL74, (2) 22.Vi.74,.(3) 1. Vii
3.VIII.74; Lachesillidae, Lachesilla pallida (Chapman) (1) 20.VII.74; Peri-
psocidae, Peripsocus quadrifasciatus (Harris) (1) 1.VII.73; P. alboguttatus
group (1 sp.) 30.VI.74; Philotarsidae, Aaronella sp. (2) 15.VII.74, (2) 19.
VIL.74, (7) 31.VII.74; Psocidae, Trichadenotecnum alexanderae Somm.
19.VI1.73, 2.VIII.73, 19.VII.74; Unidentified (2) 16.VII.74, 31.VIL74, 2.
VIIL.74, 15.VU1I.74: THYSANOPTERA, Aecolothripidae, Acolothrips vitti-
pennis Hood (2) 9.VII.74, Thripidae Frankliniella tritili (Fitch) 10.VII.74,
F. runneri (Morgan) 12.VII.74, Anaphothrips obscurus (Mueller) 15.VII.74,
Limothrips cerealium (Haliday) 16.VII.74, Chaetanaphothrips sp. near or =
orchidii (Moulton) 17.VII.74, Unidentified 19.VII.74, Phlaeothripidae, Lepto-
thrips sp. near or = mali (Fitch) (3) 31.VII.74, Hopolothrips fieldsi Craw-
ford (2) 1.VII.74, Liothrips sp. 1.VUI.74: HOMOPTERA-HEMIPTERA,
Aleyrodidae, Aleyrodinae 22.VII.74, 31.VII.74, 1.VIII.74, Aleurochiton sp.
(?) 3.VIL.73; Aphididae, Amphoraphora sp. (2) 8.VII.73, A. sensoriata Mason
7.VII.73, Anoecia cornia (F.) (3) 30.VII.74, A. querci (Fitch) 3.VIII.74, (2)
15.VIII.74, Aphis sp. 27.VII.73, 1.VIII.74, A. gossypii Glov. 27.VII.74;
A. rubifolii (Thos.) 29.VII.74, A. sambucifoliae Fitch 12.V11.74, Capitophorus
elaeagni (Del Guer.) 19.VII.74, 22.VII.74, 1.VIII.74, Chaitophorus sp.
1.VIIL.74; C. hippophaes (Wik.) 17.VI1.74, 3.VIII.74, C. pusillus Hottes and
Frison 27.VII.74, Drepanosiphinae 30.VII.73, Drepanaphis sp. 25.VI1.73,
D. acerifoliae (Thos.) 29.VII.74, D. saccharini Smith and Dillery 6.VIII.74,
Dysaphis sp. 16.V1I.74, D. radicola (Mord.) 31.VII.74, Essigella pini Wilson
22.VIII.74, Hyadaphis foeniculi (Pass.) 1.VIII.74, Hyalopterus pruni (Geof.)
15.VIII.74, Macrosiphon sp. 30.V1.74, M. liriodendri (Monell) (3) 30.V1I.74,
12.VII.74, 16.VII.74, 19.VII.74, (2) 30.VII.74, M. avenae (F.) 30.VI.74, 16.
VIIL.74, 31-VIL.74, 1.VIII.74, M. euphorbiae (Thomas) 1.VIII.74, Mastopoda
pteridis Oestlund 27.VIII.74 Monellia sp. 30.V1.74, M. costalis (Fitch) 1.
VII.73, Monelliopsis sp. (2) 19.VII.74, 31.VII.74, 20.VIII.74, Myzocallis sp.
(2) 1.VIII.74, 15.VIII.74, (2) 23.VIIII.74, M. asclepiadis (Monell) (2) 1.
VIII.74, 15.VIII.74, (2) 22.VIII.74, 23.VIII.74, Myzus persicae (Sulz.) 1.
VIII.74, Masonovia sp. 7.VIII.74, Ovatus phyloxae (Sampson) 12.VII.74,
Prociphilus fraxinifolii (Riley) 9.VIL74, 15.VIIL74, 18.VII.74, 29.VII.74,
Rhopalosiphum sp. 1.VIUI.74, R. maidis (Fitch) 16.VII.74, 19.VII.74, 22.
VIL.74, 30.VII.74, 31.VII.74, 1.VIII.74, Schizolachnus sp. 16.VI1.73, Thecabius

VOLUME 80, NUMBER 1 123

sp. 12.VII.74, Therioaphis trifolii (Thos.) 15.V1I.74, Tinocallis kahawaluoka-
lani (Kirk.) 17.VII.74, 19.VII.74, T. ulmifolii (Monell) 7.VII.74, 9.VII.74,
12.VII.74, 16.VII.74, 17.VII.74, 19.VII.74, 22.V1I.74; Unidentified 30.V1.74,
(2) 12.VI1I.74, (4) 17.VII.74, (2) 18.VII.74, (4) 19.VII.74, 27.VII.74, (2) 29.
VII.74, (3) 31.VIL74, (5) 1.VIII.74, 15.VIII.74, 22.VII1.74; Cicadellidae,
Agallia constricta Van Duzee 16.VII.74, Alebra albostriella (Fallen) (2)
22.V11.74, Aphrodes sp. 1.V1I.73, 7.VI1I.73, (4) 8.VII.73, 16.VII.73, 10.VII.74,
19.VII.74, Balclutha sp. (2) 7.VII.73, Coelidia olitoria (Say) 20.VII.74,
Deltocephalinae 23.VII.73, Dikraneura sp. 7.VII.73, 12.VII.74, 15.VII.74,
18.VII.74, D. mali (Provancher) 16.VII.74, 1.VIII.74, Empoasca sp. 16.
VIL.74, 17.VI1.74, 20.VII.74, E. bifurcata DeLong (2) 25.VII.73, Erythro-
neura sp. 7.VII.73, 8.VII.73, (2) 12.VII.74, 15.VII.74, (3) 16.VII.74, (2)
20.VII.74, 29.VII.74, (2) 31.VII.74, E. tricincta Fitch 14.V11.74, E. vulnerata
Fitch 12.VI11.73, Forcipata loca DeLong and Cardwell 20.VI1L.73, 1.VIII.74,
Graminella nigrifrons (Forbes) (2) 29.VII.73, 14.VII.74, 30.VII.74, 31.VII.74,
1.VIII.74, 3.VIII.74, Macrosteles fascifrons (Stal.) 8.VII.73, 10.VII.73, 20.
VII.74, 30.VII.74, M. slossoni (Van Duzee) 12.VII.73, Onecpsis verticis (Say)
30.VII.74, 2.VIII.74, Scaphytopius sp. 30.VI.74, S. acutus (Say) 31.VII.74,
S. ampius DeLong and Mohr 3.VIII.74, Typhlocybinae 30.VII.74, 31.VII.74,
Xestocephalus pulicarius Van Duzee 30.VII.74; Cixidae 20.VII.74; Del-
phacidae, Delphacodes sp. 16.V1I.73, D. puella Van Duzee (2) 27.VII.74,
Pissonotus sp. 22.VII.74; Derbidae, Cedusa gedusa McAtee 29.VII.74;
Miridae, Lygus lineolaris (P. daB.) 7.VI.73, Unidentified (2) 15.VII.74,
(3) 16.VII.74, (3) 29.VII.74, (4) 31.VII.74, (3) 1.VIII.74, (5) 15.VII.74, (4)
23.VIII.74; Phylloxeridae, Phylloxera (3) 15.VIII.74, Psyllidae, Craspedolepta
sp. 16.VII.74, C. fumida Caldwell 16.VII.74, Psylla annulatus Fitch 1.VIII.
73, Unidentified 7.VII.73, 8.VII.73; Tingidae, Corythucha arcuata (Say)
29.VII.74, (2) 3.VIIL74, LEPIDOPTERA, Tineidae 31.VII.74, Unidentified
(2) 1.VIII.74: DIPTERA, Unidentified 16.VII.74, (2) 30.VII.74, (18) 31.
VII.74, (13) 1.VIII.74, (2) 12.VII1.74, 15.VIII.74, (3) 23.VIII.74; Agromyzi-
dae, Cerodontha dorsalis (Loew) 29.VII.74, C. (Poemyza) muscina (Meigen)
17.VI1.74; Cecidomyiidae, Cecidomyiidi 9.VII.74, 10.VII.74, (2) 12.VII.74,
(3) 20.VII.74, 1.VII.74, Anarete sp. 29.VII.73, 27.VII.74, 2.VIII.74, 3.VIII.74,
(2) 20.VIII.74, A. pritchardi Kim 20.VII.73, Asteromyia sp. 12.VII.74,
Atrichopogon levis (Coq.) 20.VII.74, Contarinia sp. 19.VII.74, Culicoides
paraensis (Goeldi) 20.VII.74, 22.VII.74, Dasineura sp. 17.VII.74, 22.VII.74,
27.V1I.74, (2) 31.VII.74, Dasyhelea sp. 22.V1II.74, 30.VII.74, 31.VII.74, 3.
VIUI.74, Forcipomyia sp. 30.V1.74, F. brevipennis (Macq.) 30.V1.74, Hyper-
diplosis sp. 15.VII.74, Lasioptera sp. (2) 16.VII.74, Lestremia sp. (3) 10.
VII.73, 28.VII.73, Lestodiplosis sp. 12.VII.74, Micromyia sp. 9.VII.73, 15.
VII.73, 18.VII.73, 25.VI1.73, (2) 17.VII.74, Neolosioptera 7.V1II.73, 12.V1I.74,
19.VII.74, Procystiphora sp. 30.V1.74, 7.VII.74, 19.VII.74, Porricondyla sp.
19.VII.74, 31.VII.74, Resseliella sp. 30.VII.73, (2) 16.VII.74, (2) 17.VII.74,

124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

22.VI11.74, 3.VIII.74; Chaoboridae, Unidentified (3) 7.VII.73, Chaoborus sp.
(2) 30.V1.74; Chironomidae, Orthocladiinae (3) 1.VII.73, 30.VII.74, (2)
7.VII.74, (3) 12.VII.74, (4) 15.VII.74, 18.VII.74, (2) 19.V1I.74, 20.VII.74, (3)
22.V1I.74, (5) 31.VII.74, 2.VIII.74, 6.VIII.74 (3) 15.VIIL.74, (2) 23.VII1.74,
Anatopynia sp. 22.VI1I.74, A. dyari (Coq.) 10.VII.73, Chironomus sp. 12.VII.
74, 17.VII.74, 20.VII.74, Cricotopus sp. (2) 20.VII.73, 25.VII.73, (2) 26.VII.
73, (2) 16.VII.74, 22.VII.74, Procladius bellus (Lw.) 16.VI11.74, P. culiciformis
(L.) 18.VII.73, Tanytarsus sp. 30.VII.74; Chloropidae, Conioscinella sp. 16.
VII.74, Elachiptera umbrosa (Lw.) 7.V11.73, Goniopsita catalpae (Mall.)
1.VIII.74, Oscinella carbonaria (Lw.) 25.VII.73, 29.VII.73, (2) 31.VIL.74,
O. painteri Sabr. 28.VII.73, Siphonella nigripalpis (Mall.) 30.VIL.74, Thau-
matomyia bistriata (Wlk.) 2.VIII.74, T. glabra (Mg.) 31.VII.74; Dolichopodi-
dae, Chrysotus sp. (5) 1.VII.73, 7.VII.74, 12.VII1.74, 18.V1I.74, Gymnopternus
debilis Loew 17.VI1.73, Thrypticus sp. 7.V1L.73, Drosophilidae, Scaptomyza
adusta (Lw.) 12.VIL74, S. pallida (Zett.) (5) 7.VIL.73, 15.VIL74, 18.VIL.74,
20.VI1I.74, 30.VII.74, 6.VIII.74, S. wheeleri Hackmann 30.VII.74; Lonchopter-
idae, Lonchoptera furcata (Fallen) 1.VHI.74; Phoridae, Unidentified (2) 16.
VIL.73, (2) 15.VII.73, Megaselia sp. (3) 7.VII.73, 7.VII.74, 12.VI1.74, 15.VII.
74, (3) 16.VII.74, (4) 17.VII.74, (3) 31.VII.74, Puliciphora sp. (6) 16.VII.74,
(2) 17.VII.74, (4) 22.VII.74, (2) 12.VIII.74; Psychodidae, Unidentified 28.VII.
73; Psychoda sp. (2) 1.VIII.74; Scatopsidae, Unidentified 8.VII.73, Scatopse
fuscipes Mq. 31.VII.74, Rhegmoclema sp. 31.VII.74, 6.VIII.74, 18.VII.74;
Sciaridae, Bradysia sp. (3) 1.VII.73, (4) 12.VII.74, (2) 15.VII.74, (4) 16.VII.
74, (2) 20.VII.74; Sepsidae, Sepsis punctum (Fab.) 12.V11.74; Sphaeroceridae,
Leptoeera sp. (5) 16:VI1.73,' (2) 22.V1L-73; 20.V1IL-74, (2) S0:iViieARee
VII.74, L. (Pterogramma) palliceps Johnson 16.V11.74, Sphaerocera pusilla
(Fallen) 7.VII.73, S. vaporarium Holaday 31.VII.74, 1.VII.74; Stratiomyidae,
Microchrysa polita (L.) (2) 31.VII.74, 1.VIII.74, Oxycera sp. 15.VIII.74,
Tipulidae, 16.VII.74: COLEOPTERA, Unidentified 30.VI.74, (2) 12.VII.
74, 22.VII.74, (3) 31.VI.74, Alleculidae, Mycetochara haldemani: LeC.
2.VIII.74, Chrysomelidae, Baliosus sp., 12.V11.74, 16.VII.74; 20.VII.74, 25.
VII.74, Chaetocnema sp. 25.VI1I.73, 30.VII.74, (2) 31.VII.74, (2) 1.VIII.74,
11.VI11.74, 14.VII1.74, 25.VU1.74; Curculionidae, Gymnetron pascuorum
(Gyll.) 16.VII.73, 17.VII.74, 19.VII.74, Microtrogus picirostris (F.) (3) 17.
VII.74, Pseudopentarthrum sp. 25.VI1.73, I.VIII.73; Hydrophilidae, Cercyon
sp. 25.VIII.74, 27.VIII.74; Lathrididae, Corticaria sp. 27.VIII.74; Leptodiri-
dae, Nemadus sp. prob. parasitus LeC. 22.VIII.74; Leiodidae, Colenis im-
punctata LeC. 28.VIII.74; Mycetophagidae, Litargus quadrispilotus LeC.
31.VIL74, L. nebulosus LeC. 31.V1I.74; Phalacridae, Stilbus sp. (2) 31.VII.74
(2) 1.VIII.74, 2.VIII.74, Olibrus sp. 28.V1I.74; Ptilodactylidae, Ptilodactyla
angustata (3) 16.VII.73; Rhipiphoridae, Rhipiphorus sp. 18.VII.74, Hy-
pothenemus sp. (3) 9.VII.74, Pityophthorus sp. 7.V 11.73, Pityogenes hopkinsis
Swaine 20.VII.74; Staphylinidae, Unidentified, (2) 15.VII.74, (5) 16.VII.

VOLUME 80, NUMBER 1 125

74, 22.VIII1.74, 23.VIII.74: STREPSIPTERA, Stylopidae, Pseudoxenos lugu-
bris (Pierce) 18.VII.74, Unidentified 31.VII.74: HYMENOPTERA, Uniden-
tified (2) 16.VII.74, 19.VII.74, (3) 31.VII.74, Aphelinidae, Unidentified 27.
VII.74, 1.VII1.74; Aphidiidae, Aphidius sp. 18.V11.74, Lysiphlebus sp. 30.VII.
74: Brachonidae, Asobara sp. 16.VII.74, 19.VII.74, 30.VII.74, Aspilota sp.
20.V1I.74, 31.VII.74, Chorebus sp. 2.VIII.73, Oenonogaster sp. 12.VIII.73,
Synaldis sp. 20.VII.74, 31.VII.74; Ceraphronidae, Ceraphron sp. (4) 17.
VII.74, Conostigmus sp. (2) 31.VII.74, Lygocerus sp. 12.VII.74, (2) 16.VII.74,
30.VII.74; Chalcedectidae, Euchrysia sp. 25.V1I.73; Cynipidae, Encoilinae
16.VIII.74, Alloxysta sp. 3.VIII.74; Dryinidae, Unidentified 31.VII.74, 1.
VIIL.74, Encyrtidae, new Encydnus sp. 10.VII.74, 12.VII.74, 16.V1I.74, 19.
VII.74, 31.VII.74, 1.VIII.74; Eulophidae, Unidentified 31.VII.74; Eurytomi-
dae, Unidentified 16.VII.74; Formicidae, Dolichoderus sp. 16.V11.74, Lasius
sp. (10) 26.VII.73, (2) 22.VII.74, (3) 27.VII.74, 15.VIIL74, Leptothorax sp.
(3) 28.VII.74, Monomorium minimum (Buckley) 30.V1.74, Paratrechina sp.
26.V1I.73, 19.VII.74, Ponera pennsylvanica Buckley 31.VUI.74, (4) 1.VIII.
74, (5) 23.VIII.74, (3) 27.VIII.74, Solenopsis molesta (Say) 1.VIII.74; Mymari-
dae, Unidentified 12.VII.74, 16.VII.74; Pteromalidae, Pteromalini 16.VII.74,
19.VII.74, (2) 31.VII.74.

Enemies.—Cerotainia albipilosa are sometimes preyed on by arthropods
larger than themselves. Thirteen instances of predation were observed:
Theridon sp. (Tetragnathidae: Araneida), Vespula sp. (2) and V. arenaria
(F.) (8) (Vespidae: Hymenoptera), Efferia aestuans (L.) and Dioctria
tibialis McAtee (Asilidae: Diptera). Several unsuccessful attacks were
directed by the following: Pisaurina mira (Walkenuer) (Pisauridae: Arane-
ida), Misunera sp. (Thomopsidae: Araneida), Laphria sicula McAtee, D.
tibialis (Asilidae: Diptera), and Vespula spp. Each predator used a specific
attack strategy. Vespula spp. flew along the margin of the study site and
hovered frequently in front of occupied perches. This behavior “flushed”
the asilid which was attacked as it attempted to escape. Vespula spp. some-
times attacked before the asilid attempted to fly away. Theridon sp. cap-
tured its prey with a web while the other araneids attacked from a con-
cealed position below a foraging site. The asilid predators, which were
usually perched near the prey, attacked C. albipilosa when it flew from a
perch. While cannibalism by this species was not observed, males and fe-
males displayed agonistic behavior toward other males (Scarbrough and
Norden, 1977).

Literature Cited

Bohart, G. E. 1958. Marked bees taken selectively by robber flies, p. 580. In
Linsley, E. Gorton. The ecology of the solitary bees. Hilgardia 27:543-599.

Brues, C. T. 1946. Insect dietary. An account of the food habits of insects. Harvard
Univ. Press. Cambridge, Mass. Pp. 266-267.

126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Cole, F. R., and A. L. Lovett. 1921. An annotated list of the Diptera (flies) of
Oregon. Proc. Calif. Acad. Sci. 11:197-344.

Dennis, S. D., and R. J. Lavigne. 1975. Comparative behavior of Wyoming robber

flies II (Diptera: Asilidae). Univ. Wyoming Agric. Exp. Sta. Sci. Monogr., No.

30, 68 pp.

1976a. Ethology of Efferia varipes with comments on species coexistence

(Diptera: Asilidae). J. Kans. Entomol. Soc. 49:49-62.

——. 1976b. Ethology of Leptogaster parvoclava in Wyoming (Diptera: Asilidae).
Proc. Entomol. Soc. Wash. 78:208—222.

Dennis, S. D., G. P. Roehrkasse, and R. J. Lavigne. 1975. Prey recognition of
Efferia frewingi (Diptera: Asilidae). Ann. Entomol. Soc. Am. 68:404—408.
Fackler, H. L. 1918. The tiger beetles of Kansas (Family Carabidae; Subfamily

Cicindellidae) Order Coleoptera. M.S. Thesis, Univ. Kansas. 51 pp.

Hobby, B. M. 1931. The British species of Asilidae (Diptera) and their prey. Trans.
Entomol. Soc. South England, No. 6, 1931 (1930), pp. 1-42.

Horning, D. S., Jr., and W. F. Barr. 1970. Insects of Craters of the National Monu-
ment, Idaho. Univ. Idaho Coll. Agric. Misc. Ser. 8:1-118.

Lavigne, R. J., and F. R. Holland. 1969. Comparative behavior of eleven species
of Wyoming robber flies (Diptera: Asilidae). Univ. Wyoming Agric. Exp. Sci.
Monogr., No. 18, 61 pp.

Lehr, P. A. 1958. The biology of robber fly feeding on locusts. Dokl. KasASKhN.
4:189-209.

—. 1964. On the feeding and significance of robber flies. Trudy Nauchno-

issled. Inst. Zasch. Rast., Alma-Ata 8:213-244.

1972. The robber flies of the genera Holopogon Loew and _ Jothopogon
Becker (Diptera: Asilidae) in the fauna of the USSR. Entomol. Rev. 51:99-109.

Linsley, E. G. 1960. Ethology of some bee- and wasp-killing robber flies of south-
eastern Arizona and western New Mexico. Univ. Calif. Publ. Entomol. 16:357—
381.

Marler, P., and W. J. Hamilton. 1956. Mechanisms of Animal Behavior. John
Wiley and Sons, Inc. New York, p. 131.

Martin, C. H. 1968. The new family Leptogasteridae (the grass flies) compared
with Asilidae (robber flies) (Diptera). J. Kans. Entomol. Soc. 41:70-100.
Melin, D. 1923. Contributions to the knowledge of the biology, metamorphosis, and
distribution of the Swedish asilids in relation to the whole family of asilids. Zool.

Bidr. Uppsala 8:1-317.

Musso, J. J. 1971. Etude preliminaire sur les activités journaliére d'une popula-
tion d’Andrenosoma bayardi Seguy. Bull. Entomol. Soc. Fr. 76:175-182.
Parmenter, L. 1952. Notes on the Asilidae (robber flies). Entomol. Record. 64:229-

234, 263-266, 295-299.

Poulton, E. B. 1906. Predaceous insects and their prey. Trans. R. Entomol. Soc.
London 1906:323—409.

Scarbrough, A. G., and A. Norden. 1977. Ethology of Cerotainia albipilosa Wilcox
(Diptera: Asilidae) in’ Maryland. Diurnal activity rhythm and seasonal dis-
tribution. Proc. Entomol. Soc. Wash. 79(4):538—-554.

Scarbrough, A. G., and G. Sipes. 1973. The biology of Leptogaster flavipes Loew
in Maryland (Diptera: Asilidae). Proc. Entomol. Soc. Wash. 75:441—-448.
Schmid, J. M. 1969. Laphria gilva (Diptera: Asilidae), a predator of Dendroctonus
ponderosae in the Black Hills of South Dakota. Ann. Entomol. Soc. Am. 62:

1237-1241.

VOLUME 80, NUMBER 1 127

Shtakel’berg, A. A. 1950. Diptera, animal life of the USSR. III. Izd. ANSSR.
Pp. 162-213.

Wallis, J. B. 1913. Robber-fly and tiger beetle. Can. Entomol. 45:134.

Zinovera, L. A. 1959. Biology and ecology of robber flies (Diptera: Asilidae) in
the Naryn Sands. Entomol. Oboz. 38:499-511.

Associate Professor of Biology, Department of Biology, Towson State
University, Baltimore, Maryland 21204.
Footnote

‘Portions of this work were supported by funds obtained from the Towson State
University Faculty Research Committee.

128 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

NOTE

OXAEA AUSTERA GERSTAECKER IN BOLIVIA, WITH A
NEW HOST RECORD (HYMENOPTERA: APOIDEA)

Oxaea austera Gerstaecker is a rather little known South American bee
that has been reported from a few collections made in northeastern and
southern Brazil (Ceara, Maranhao, Sao Paulo, and Rio Grande do Sul),
Paraguay and Mesopotamian Argentina (Ducke, 1910, Revue d’Entomol.
(Caen), 28:82). Hurd and Linsley (1976, Smiths. Contrib. Zool., 220) have
recently summarized the few available records of floral hosts for the spe-
cies. The purposes of this paper are to report the genus Oxaea and O.
austera from Bolivia for the first time, to add a floral host to those already
known for O. austera, and to report oxaeids for the first time as visitors
of Passifloraceae.

On 6 December 1973, I collected four males of Oxaea austera from
blossoms of a trailing Passiflora sp. (Passifloraceae) growing along the
roadside seven kilometers south of Warnes (ca. 17°30’S, 63°10’W, 375 m)
in the Department of Santa Cruz, Bolivia. There, these robust bees (ca.
2 cm) approached the flowers very rapidly and then braked to hover
stationarily in front of the blossoms before landing and entering them. This
same behavior in males of O. austera was noted by Schrottky (1904,
Allgemeine Zeitschr. Entomol. 9:346) at flowers of an exotic in Brazil, the
motherwort, Leonurus sibiricus L. (Lamiaceae). This plant is the only pre-
viously known nectar source for males of O. austera.

The bees that I took from the Passiflora near Warnes were peppered
with pollen and particularly so on their thoracic dorsa. It seems probable
that these large bees are effective pollinators of at least this species of
Passiflora. No Oxaea species has previously been recorded from Passiflora
(see Hurd and Linsley, supra cit., for a comprehensive summarization of the
available floral records), and the genus is generally thought to be polli-
nated by carpenter bees (Xylocopa sp.) (Faegri and van der Pijl. 1971.
The Principles of Pollination Ecology, ed. 2, Pergamon).

Thanks are due to Prof. J. S. Moure of the Universidade Federal do
Parana, Curitiba, Brazil, for his determinations of the bees and to Prof.
Richard B. Selander of the University of Illinois, Urbana, for his support of
the author's fieldwork in Bolivia. The specimens of Oxaea austera re-
ported herein will be deposited in the collection of the Section of Faunistic
Surveys and Insect Identification of the Illinois Natural History Survey
at Urbana.

John K. Bouseman, Agricultural Entomology, Illinois Agricultural Ex-
periment Station and Section of Economic Entomology, Illinois Natural
History Survey, Urbana, Illinois 61801.

VOLUME 80, NUMBER 1 129

NOTE

HUMAN ORAL MYIASIS IN VIRGINIA CAUSED BY
GASTEROPHILUS INTESTINALIS
(DIPTERA: GASTEROPHILIDAE)

The horse bot fly, Gasterophilus intestinalis (L.) is a widespread spe-
cies causing enteric myiasis in equines. Subcutaneous myiasis due to
gasterophilids, often broadly termed “creeping eruption” or “larval migrans,”
has been reported involving people associated with horses. In these instances
Ist-instar larvae burrow into the skin and produce tunnels which are ac-
companied frequently by an intense itching sensation. Infestations may end
spontaneously or by suppuration (James. 1947. USDA Misc. Publ. 631:92-
99). Zumpt (1965. Myiasis in man and animals in the Old World. Butter-
worths, London. 267 pp.) states that these misplaced larvae never reach
the second stage.

Creeping eruption in man tends to occur on the extremities (Austmann.
1926. J. Amer. Med. Assoc. 87:1196-1200). Chereshnev (1953. Dokl. Akad.
Nauk (N.S.). 91:173-176) found that rubbing or moistening of G. in-
testinalis eggs is necessary to stimulate hatching. He reported that Ist-
instar larvae could penetrate the mucous membrane, but not human skin.
However, Danilov (1973. Med. Parazitol. Parazit. Bolezn. 42:361) described
multilinear human myiasis with puritis and skin vesicles on the extremities
and stomach caused by 65 G. equi (= intestinalis) larvae.

James J. Keeble, Entomologist, Maintenance Division, Atlantic Division,
Naval Facilities Engineering Command, Norfolk, provided the following
information from a case of human myiasis in Virginia. On 22 December
1976 a 10-year-old Virginia Beach girl complained of an irritation in her
mouth. Subsequently, a physician removed a small G. intestinalis larva from
the inner lining of her mouth near the jaw. Further examination of the
oral area and tongue revealed no other larvae.

The girl received a horse as a present two months before the infesta-
tion was noticed. She was with the horse regularly and kissed it frequently.
It is not known if the infestation was a direct result of oral contact or if
the larva was transferred to the mouth indirectly. The former route seems
likely.

Sukhapesna et al. (1975. J. Med. Entomol. 12:391-392) reported that G.
intestinalis eggs deposited in late fall in Kentucky may remain viable for
several months. Hatch of field-collected eggs terminated in late January.
They stated that egg age and environmental temperatures probably in-
fluenced egg viability.

Causey (1937. J. Econ. Entomol. 30:39-40) noted that the presence of
animal myiasis in livestock poses potential public health problems for

130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

humans. While such cases may produce relatively mild discomfort, they
demonstrate that care should be exercised by persons in close contact with
domestic livestock or pets. A variety of parasites may be encountered
under similar circumstances, several of which are not so benign.

Dr. R. J. Gagné, Systematic Entomology Laboratory, IIBIII, Agric. Res.
Serv., USDA, confirmed identification of the specimen.

L. H. Townsend, Jr., R. D. Hall (Graduate Research Assistants) and E. C.
Turner, Jr. (Professor), Department of Entomology, Virginia Polytechnic
Institute and State University, Blacksburg, Virginia 24061.

NOTE

CYLINDROCNEMA PLANA MAYR, 1865, A SENIOR SYNONYM
OF NOPALIS CRASSICORNIS (REED), 1898
(HEMIPTERA: ACANTHOSOMATIDAE)

The Heteroptera that formed the basis of Edwyn C. Reed’s “Sinopsis de los
Hemipteros de Chile, Primera Parte: Heteropteros” was purchased by Carl
J. Drake who bequeathed his collection, including Reed’s specimens, to
the United States National Museum where it is now housed. All speci-
mens from the Reed collection bear a printed label “Sin. Hem. Chile
Coll., E. C. Reed” but, unfortunately, very few bear a locality label; some
of the specimen-pins bear Reed’s handwritten identification labels. Among
the latter specimens is one labeled “L. crassicornis N. S. 2°.” Even though
this specimen has no locality label it is hereby designated the lectotype of
Lanopis crassicornis Reed. The other specimen of the two that comprised
the type-series is missing; both specimens were reported in Reed’s original
description (1898. Rev. Chilefia de Hist. Nat. 2:156-157) as being from
the island of Chiloe.

Reed’s specimen agrees very well with the original description of
Cylindrocnema plana Mayr (1865. Verh. Zool.-Bot. Ges. Wien. 14:912-
913), and especially in such critical characters as two-segmented tarsi; large
size [15 mm here, 12 mm in Mayr’ description]; thickened antennal
segments, especially the second and third; angularly produced humeral
angles concave along the posterior margin; hemelytral membrane nar-
rowly produced along costal margin [to a point about opposite mid-
length of corio-membranal suture]; prosternum with a deep mediolongi-
tudinal groove [as wide as labial segment I]; and venter of abdomen
mediobasally convex but without anteriorly projecting tubercle or spine.
Comparison of Reed’s specimen with specimens labeled as C. plana re-
vealed no taxonomic differences and showed that both species share the
lack of a foliaceous carina on the mesosternum.

VOLUME 80, NUMBER I 131

Although Reed listed C. plana for Chile, apparently on Mayr’s original
data, he did not include that genus in his key to the genera of “Ditomo-
tarsina” and perhaps thus misled himself into assigning this specimen
to Nopalis where it did not agree with any of the other species.

I conclude Nopalis crassicornis (Reed), 1898, is a junior synonym of
Cylindrocnema plana Mayr, 1865, and so assign it here as NEW SYNON-
YMY.

R. C. Froeschner, Department of Entomology, U.S. National Museum,
Washington, D.C. 20560.

NOTE

THE STINK BUG PADAEUS TRIVITTATUS STAL AND NOT
PADAEUS VIDUUS (VOLLENHOVEN) IN THE UNITED STATES
(HETEROPTERA: PENTATOMIDAE)

This tropical American species was first reported for the United States
from “Florida” under the preoccupied name Padaeus irroratus (Herrich-
Schaeffer) by Van Duzee (1904. Trans. Amer. Ent. Soc., 30:78); he credited
the record to a specimen in the H. G. Barber collection. Barber (1910.
Jour. N.Y. Ent. Soc., 18:35), using the same name combination, corrected
the locality in Van Duzee’s report and gave the specimen data as “Hua-
chuca Mts., Ariz., 1899, collector R. E. Kunze.” Later, Van Duzee (1917.
Univ. Calif. Pubs. Ent., 2:44) catalogued it from “Ariz. (not Fla.).” All
subsequent listings of this species for the United States appear to be based
on these references.

Barber’s now headless specimen, still bearing the Van Duzee determina-
tion label as Padaeus irroratus, is in the National Museum of Natural
History. Examination found it to represent another tropical American
species, P. trivittatus Stal, and not P. viduus. Thus the latter name must be
deleted from the United States list and, assuming the locality is correct,
the former name must be added.

These two species can be separated readily by any of several char-
acters; three easily used color characters are compared in the following
couplet:

Abdomen dark brown to black with numerous widely scattered yellow
flecks. Costal margin (except at very base) concolorous with the
brown to black corium. Black color of connexivum broadly reach-
ing extreme lateral margins of segments and more or less enclosing
a small, median, suboval pale spot P. viduus (Vollenhoven)

132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Abdomen yellow with three longitudinal black stripes, one median
and one each side sublaterally. Costal margin along basal third
narrowly calloused yellow. Black color of connexivum separated
(often narrowly) from lateral margins of segments for full length

P. trivittatus Stal

Richard C. Froeschner, Department of Entomology, U.S. National Mu-
seum, Washington, D.C. 20560.

VOLUME 80, NUMBER 1 133
BOOK REVIEW

Mechanical Design in Organisms, by S. A. Wainwright, W. D. Biggs, J. D.
Currey, and J. M. Gosline. John Wiley and Sons, New York. 423 pp.
Cost $19.50.

This book will be of interest to functional morphologists, some physi-
ologists, and some ecologists.

Mechanical Design in Organisms is somewhat revolutionary in that it
departs from the traditional concept of rigid discipline orientation and
attempts to integrate principles of mechanical engineering with biological
structure and function. An implicit assumption throughout the book is that
if one understands the physical basis of a phenomenon, general principles
can be used to predict responses to different conditions. In this sense the
text serves as a valuable companion to Alexander's (1968) Animal Mechanics
and to his (1971) Size and Shape.

The text is divided into three parts (“materials,” “structural elements and
systems, and “ecomechanics”’) covering eight chapters. Authorship ap-
pears evenly divided with each writer contributing at least one chapter,
and some chapters are co-authored.

“Materials” constitutes half the text and provides essential physical
parameters and their mathematical derivation. This section has a moderate
mathematical basis, which is to be expected considering the nature of the
subject matter. However, the approach taken by the authors in develop-
ing the mathematical relationships is methodical and should not prove
an insurmountable obstacle to understanding. Types of biological materials
including tensile materials (chitin, cellulose, and collagen), pliant materials
(resilin, elastin, and cartilage), and rigid materials (keratin, bone, etc.) are
discussed.

“Structural elements and systems” is concerned primarily with the be-
havior of materials under conditions of stress and formulates principles
of structural optimization. Chapter 7 considers mechanical support in
organisms and develops principles for biological structural systems.

“Ecomechanics” is the shortest part of the book (one chapter) and is
largely conjectural. Ecology and mechanical design are interrelated, but
the extent of interrelationship is unknown. Among the environmental
factors that influence mechanical design are gravity, surface tension, fluid
flow, and pressure. Wainwright attempts to develop ecological predictions
based on these biomechanical data. However, much more data must be
accumulated before the interaction of environmental parameters and _bio-
logical structure is appreciated.

The only limitation that I can find with the book is that it lacks any
treatment of fluid mechanics. However, the omission is acknowledged

134 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

by the authors and is not significant in view of the enormous literature de-
veloped by engineers. The authors do not emphasize insects in their de-
velopment of theories about mechanical design. Were they to do so, the
general principles that they seek to develop would be circumscribed.

I found the book lively and well written. The organization is good, ideas
are presented concisely, and the innumerable line drawings, graphs, and
figures clarify the points made in the text. Mechanical Design in Organisms
should stand for some time as a fundamental reference in the field of
biomechanics.

Gordan Gordh, Systematic Entomology Laboratory, IITBIII, Agric. Res.
Serv., c/o U.S. National Museum, Washington, D.C. 20560. (Present ad-
dress: Department of Entomology, University of California, Riverside, Cali-
fornia 92521.)

BOOK REVIEW

Controlling Fruit Flies by the Sterile-Insect Technique. 1975. IAEA,
Vienna, Austria, STI/PUB/392. 175 pp. Available from Unipub. Inc.,
P.O. Box 433, Murray Hill Station, New York, N.Y. 10016. Price $10.00.

The following words, appearing as part of the foreword to this very
interesting book, comprise a statement which can well stand alone to serve
as the raison detre for the volume: “. . . the containment of fruit flies by
methods less reliant on insecticides has taken on a new urgency .... The
Mediterranean fruit fly, which first appeared in the 1950’s in Central
America, has surged northward throughout this area and is now poised to
invade Mexico. The economic significance of an unchecked spread of this
pest is reflected in an estimated annual loss of $6.8 million for the Central
American area excluding British Honduras and Mexico. Should the medfly
invade those two countries, their citrus industries would suffer a combined
loss of $6 million. Should the medfly go unchecked and invade the United
States, the U.S. fruit industry would lose annually about $85 million for
citrus and $200 million for deciduous fruits . ‘i

A result of a panel and research coordination meeting organized by the
Joint FAO/IAEA Division of Atomic Energy in Food and Agriculture, this
collection of authoritative papers addresses a wide range of activities within
the sterile-male technique picture. Not only is mass-rearing (the heart of
the matter) discussed thoroughly, but subjects such as genetic variation,
comparative behavior, and field observations are also included for the most
important of the world’s tephritids. Summaries of work performed at
various institutions throughout the world are included, as are concluding

VOLUME 80, NUMBER 1 135

statements and general recommendations for work to be undertaken in the
future.

The authors of the eleven major papers in the volume are widely known
in and outside their respective specialties, a feature adding materially to
the excitement inherent in the messages they deliver. It is gratifying to
realize that these scientists are able to contribute so much substantive ma-
terial to the general subject and at the same time deal effectively with the
specifics so valuable to the individual researcher in the laboratory or in the
field. It is hoped that the International Atomic Energy Agency will be able
to keep this valuable series alive in light of the severe economic implications
presented by the singular habits of these important insects.

Richard H. Foote, Systematic Entomology Laboratory, IIBIII, Agric.
Res. Serv., c/o U.S. National Museum, Washington, D.C. 20560.

BOOK REVIEW

The Windows to His World. The Story of Trevor Kincaid. By Muriel L.
Guberlet. 1975. 287 pp., 16 photographs. Pacific Books, Publisher, P.O.
Box 558, Palo Alto, California 94302. $9.95.

The name of Trevor Kincaid (1872-1970) is little known by the current
generation of American entomologists unless it be in the Northwest where
he worked at the University of Washington as a student, professor, and
emeritus professor of zoology for about 75 years. This book is a fascinating
biography of a most unusual man whose inspired teaching and whose widely
recognized versatility in most branches of natural history made him almost
a legend to several generations of students. From early middle age on, he
was chiefly concerned with the marine life of the Puget Sound area, espe-
cially the oyster and fisheries industries. Thus he established the Puget Sound
Biological Station, which became the University of Washington Ocean-
ographic Laboratories. Nevertheless, throughout his life, but especially as
a boy in Ontario and in Washington (after he moved there in 1889) and
as a student and youthful professor at the University, he collected and
studied insects with boundless enthusiasm. This biography reports (page
45) that during 1890-94 he made a personal collection of 60,000 insects
and also sent 100,000 specimens to entomologists in all parts of the U.S. Two
expeditions to Alaska were of much importance in his early career. In 1897
he participated in a 3-month study of the fur seals of the Pribilof Islands,
in the company of David Starr Jordan and a group of Stanford University
students including R. E. Snodgrass. Then in 1899 he went on the Harriman
Alaska Expedition during which he collected 1,000 species of insects. As a
result of these collections, he spent the summer of 1900 in Washington,

136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

D.C. and worked on Alaskan insect specimens. Numerous animals (including
51 insects) have been named in his honor. For short periods of 1908 and
1909 (in the book wrongly stated as 1906 and 1907), he traveled and col-
lected widely in Japan and Russia, respectively, as a special agent of the
U.S. Department of Agriculture in search of parasites of the gypsy moth.
At the National Archives I have examined a thick folder of interesting
correspondence relating to the Japanese trip. Letters are chiefly between
Kincaid and L. O. Howard, but also included are some letters with State
officials in Massachusetts and others. Thus Trevor Kincaid’s contributions
to entomology and to biology generally are much greater than suggested
by his modest though significant bibliography of papers. In fact, his in-
terests were so broad that he thought of himself as an “omnologist.”

This book is well indexed and illustrated and is fine reading for people
interested in biography, the history of science, or the growth of higher
education in the Northwest. Mrs. Guberlet is the widow of a former uni-
versity zoologist, Prof. John Guberlet who taught in Prof. Kincaid’s depart-
ment for 17 years, so she had a long acquaintance with Kincaid. She
also had available an extensive but unpublished autobiography. Her book
is a beautifully writen and warmly personalized testimony to Kincaid’s
splendid human qualities and constant dedication to his science, his family,
and his friends. Those of us who never met him are given a glimpse of a
man not easily forgotten.

Ashley B. Gurney, Resident Cooperating Scientist, Systematic Entomol-
ogy Laboratory, IIBIII, Agric. Res. Serv., USDA, c/o U.S. National Mu-
seum, Washington, D.C. 20560.

BOOK REVIEW

Theodore D. A. Cockerell. Letters from West Cliff, Colorado. 1887-
1889. 1976. William A. Weber (Ed.) 222 pp., illus. Colorado Associated
University Press, 1424 15th St., Boulder, Colo. 80309. $8.95.

Theodore Dru Alison Cockerell (1866-1948) was a very unusual and pro-
ductive man. The letters in this book, selected from a large accumulation
sent back to England during the nearly 3-year period of his first visit in
America, are of great interest both for what they reveal of Cockerell’s
youth and as descriptive accounts of life in a part of Colorado about 90
years ago. A scion of a substantial English family, since then distinguished
by several outstanding members, “Theo” went to West Cliff (now West-
cliffe), in Wet Mountain Valley about 50 miles west of Pueblo, just prior to
his 21st birthday and lived in a rural community in which English families
were well represented. Many had gone there for health reasons, as he did to

VOLUME 80, NUMBER 1 137

counter frailness and suspected tuberculosis. The letters are addressed
chiefly to Annie Fenn, whom he married in 1891, or to her brother. In
England he had already begun a lifestyle different from that of most boys:
he was not involved in athletics, regarded many of their activities as
frivolous, and, partly due to his poor health, omitted much formal school-
ing. Instead, he was a somewhat delicate, odd, naturalist type, and by
the time he was 20 he had published 158 notes and papers on snails, insects,
plants and other subjects; his pattern of short, quickly written contribu-
tions had been established. While quite young he had become familiar,
at the British Museum, with identification practices and, when unable to
recognize the species of local plants and animals from available handbooks,
he entertained himself by writing short descriptions and inventing scientific
names for them.

At West Cliff he lived mainly with a congenial neighborhood family and
supported himself by doing light farm chores, informal teaching, sketching
and many other tasks for which his background qualified him. He
regularly received several natural history periodicals from England and
the U.S., and he corresponded actively with established workers (Alfred
R. Wallace, for whom he later worked in England; Binney and Pilsbry,
among malacologists; Riley, Scudder, Strecker, Wickham and other ento-
mologists). He referred (page 163) to a paper by Scudder on fossils from
Florissant, which is only 60 miles from Westcliffe, but evidently did not
go there to collect seriously until 1906. In 1907 one of his students, S. A.
Rohwer, found a remarkably well preserved specimen of a strange neurop-
teron with long narrow hind wings, one of the family Nemopteridae, which
does not now occur in North America. A picture of the fossil specimen was
shown on covers of Entomological News in 1908, and the species was men-
tioned in a News editorial as “the most remarkable insect made known
during the preceding year.”

The numerous Cockerell letters, which have been chosen and grouped by
subjects likely to interest a variety of readers, are illustrated by many of
the simple sketches that originally accompanied them. The editor, Dr. W. A.
Weber, a Colorado botanist, prepared the book with great care. In 1965
he published a short biography and a bibliography of 3,904 Cockerell pub-
lications (Univ. Colo. Studies, Series in Bibliography No. 1:1-124), and he
also listed about 30 biographical articles and notices about Cockerell (cus-
tomarily pronounced with emphasis on the first syllable). The bibliography
is an invaluable reference to the numerous papers on bees, coccids, fossils,
and other subjects reported on by Cockerell.

Ashley B. Gumey, Resident Cooperating Scientist, Systematic Entomol-
ogy Laboratory, Agric. Res. Serv., USDA, c/o U.S. National Museum,
Washington, D.C. 20560.

138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

BOOK REVIEW

A Revision of Western Nearctic Species of Torymus Dalman (Hymenoptera:
Torymidae). E. E. Grissell. 1976. Univ. Calif. Publ. Entomol. 79:1-120,
109 figs., 6 pls. Cost $6.00.

The Chalcidoidea presently consist of about 1,200 genera and 10,000
described species distributed over 18 families. Aside from their small to
minute size, the primary limiting factor to studying chalcidoids is a lack of
keys to the genera and species. Thus it is with some enthusiasm that I note
the publication of a taxonomic study of western species of Torymus.

About 25 genera and 180 species of Torymidae have been described in
North America; about 100 species have been described in Torymus. The
genus is poorly studied biologically, but most species for which the biology
is known are primary, external parasites of gall-forming insects. Host
specificity does not appear to be common in Torymus because several well-
known species have extensive host lists. The genus appears to be habitat
specific.

L. L. Huber (1927. Proc. U.S. Nat. Mus. 70(14):1-114) prepared the last
taxonomic study of North American Torymus. For many reasons that work is
unsuitable. During the fifty years since Huber’s study there have been no
published taxonomic studies of the North American Torymus, and most
biological accounts have been anecdotal.

Grissell’s study considers 61 species of Torymus. He places 17 species
in synonymy and described seven new species. I suspect that more new
species will be found in the area under consideration. One of the features
about the study that I find encouraging is the use of species groups in-
stead of subgenera. Grissell recognizes five species groups in western
Torymus. When the rest of the New World species are studied, the con-
cepts of these groups will undoubtedly be altered somewhat, but this
will not affect the nomenclature of Torymus. In general I find this work
to be excellent. Characters have been carefully thought out, the keys are
workable, descriptions are concise, and extensive distribution and host
data are provided. If Grissell extends his studies to Torymus found in
eastern North America, we will better understand this large and interesting
genus.

G. Gordh, Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,
USDA, c/o U.S. National Museum, Washington, D.C. 20560. (Present ad-
dress: Department of Entomology, University of California, Riverside,
California 92521.)

VOLUME 80, NUMBER 1 139
BOOK REVIEW

The Tachydromiinae (Diptera: Empididae) of Fennoscandia and Denmark.
Vol. 3, Fauna Entomologica Scandinavica. M. Chvala. 1975. Scandinavian
Science Press Ltd., Klampenborg, Denmark. 336 pp. Paperback, 98
Dkr. (= $16.00 at this writing). Subscription price: 68.60 Dkr. (= $11.00).

As the needs for reliable identifications increase and as it becomes more
and more obvious that the limited number of taxonomists cannot supply
all these needs, it is increasingly important that high-quality regional
handbooks are produced. Dr. Chvala’s study of Tachydromiinae is one
of the finest examples of this kind of publication.

“The Tachydromiinae” is the first of a projected group of six volumes
on Empididae by Dr. Chvala for this new faunistic series. Altogether, 128
species and 9 genera of Danish and Fennoscandian tachydromiines are
treated, and other species that may occur in this area are included in the
keys and are briefly diagnosed. The life history and zoogeography of the
Tachydromiinae are briefly treated. A succinct but useful description of the
adult morphology prepares the non-specialist for using the keys, and a
valuable discussion of the classification and phylogeny is included. The
author adds a sufficiently detailed section on the material (18,000 speci-
mens) used in the study.

Although only three Holarctic species of Tachydromiinae are known
(Platypalpus unguiculatus Zett., Tachypeza winthemi Zett., and Drapetis
assimilis Fall.), additional species probably will be found to be Holarctic in
the future, and the volume will be especially useful to North American
workers in that regard. It should be noted that Stilpon (p. 292) is masculine.
Also, Chersodromia difficilis Lundbeck, 1910 is a junior synonym of Tachy-
peza alata Walker, 1835 (July: not 1836), a validly described species, al-
though originally described as a variety.

This volume is nicely produced, a crisp photo-offset, and the unadjusted
right-hand margins are not particularly unpleasing but presumably saved
production costs. A special feature is the number and quality of the
illustrations: 790 very clear and useful figures.

The following comments, modified from an advertisement by the pub-
lisher, gives some detail on the Fauna Entomologica Scandinavica series.
The series was founded in 1973 and is edited by a committee appointed by
five entomological societies in Denmark, Finland, Norway, and Sweden.
Four volumes have been published, and over 60 volumes are in preparation.
The series presents comprehensive monographic treatments of the fauna
of insects and other terrestrial arthropods occurring in northern Europe
(Denmark, Finland, Norway, Sweden and adjacent areas).

Each volume deals with a systematic unit, usually a superfamily, family

140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

or subfamily. The text includes general chapters on morphology, biology,
economic importance, zoogeography and phylogeny. The treatment at the
generic and specific levels includes keys and short descriptions, and infor-
mation on local faunistics, total distribution, habitats, and biology is given
under each species.

All volumes are richly illustrated, usually with several figures per species.
The illustrations include habitus drawings, details of external morphology,
complete sets of genitalia drawings, and photos of wings and biological
features. The section on Lepidoptera will include color-plates illustrating
all species.

Each volume is concluded by a catalogue giving detailed information on
the distribution of all species in the area from Great Britain and northern
Germany to the North Cape and eastern Karelia.

All volumes will be published in English. It is planned to publish 2-5
volumes with a maximum of 600 pages each year to keep prices at an
acceptable level. The volumes will appear at irregular intervals. Volumes
appearing in the near future will include the Agromyzidae (Diptera) and
Elachistidae (Lepidoptera).

L. Knutson, Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,
USDA, c/o U.S. National Museum, Washington, D.C. 20560.

VOLUME 80, NUMBER 1 141

OBITUARY

The Rev. Thomas Borgmeier, O.F.M.
1892-1975

Thomas Borgmeier was born in Bielefeld, Westphalia, Germany on 31
October 1892 and died in Rio de Janeiro, Brazil on 11 May 1975. In the
more than 82 years between these dates Father Borgmeier lived a full
life devoted to religious activities, editorial and publishing work, and en-
tomology. Members of the Entomological Society of Washington have many
reasons to pause and pay tribute to this great man and distinguished en-
tomologist.

After graduation with distinction in the program of classics and hu-
manities in the “gymnasium” in Bielefeld, Thomas went to Brazil in 1910
to join the Franciscan Order of Friars Minor in January 1911. He studied
philosophy in Curitiba from 1912-1914 and theology in Petropolis from
1915-1918. While a student in Petropolis, Brother Thomas became in-
terested in entomology while observing the habits of ants. Spending his

142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

vacations in Blumenau, Santa Catarina, in 1917 he met and _ befriended
Professor Hermann von Jhering who had founded the Museu Paulista in
Sao Paulo. Through the generosity of a wealthy industrialist in Rio,
Thomas was able to acquire von Jhering’s large reprint library on ants
as well as a fine binocular microscope. While studying ants in Petropolis,
Thomas discovered phorid flies attacking and parasitizing them. Puzzled,
he contacted another young entomologist, the Jesuit Father Hermann
Schmitz, who as a disciple of the famous E. Wassmann, was well qualified
to introduce him to the intricacies of the taxonomy and biology of the
Phoridae. Borgmeier’s first scientific publication, on the biology of the
ant Odontomachus affinis Guérin, appeared in 1920, and in 1922 he de-
scribed his first new species, the phorid Dohrniphora brasiliensis.

Father Thomas was ordained a priest in May 1918, and from 1920-1924
was Professor of Biblical Sciences at the Major Seminary in Petropolis. At
the same time he contributed freely to the monthly cultural magazine
“Vozes de Petropolis” which was published by the friars. He quickly
mastered the Portuguese language and translated a number of religious
classics into that language. In 1922 Borgmeier began a friendship with
Dr. Arthur Neiva who obtained from the Franciscans his complete freedom
for entomology and his entrance into public service. In 1923 Thomas became
an adjunct research scientist in the National Museum in Rio de Janeiro
and in 1924 he moved definitively to the Museum. In 1927 he obtained his
Brazilian citizenship and in 1928 went to Sao Paulo as assistant in entomol-
ogy under Neiva in the newly founded Instituto Biologico. In 1933 he
returned to Rio to become head of the entomological section of the Instituto
de Biologia Vegetal in the Botanical Garden, where he remained for eight
years.

In 1931 Father Borgmeier founded the international journal, Revista
de Entomologia, which he edited and published until 1951 (when adverse
financial circumstances ended its publication). He then started the pub-
lication of entomological monographs in the series “Studia Entomologica,”
which with the support of the Conselho Nacional de Pesquisas became a
journal in 1958 and is now (1976) in its 19th volume. In 1940 the Franciscans
elected Father Thomas to the provincial government as one of the four
counsellors, and at the same time he was made director of their publish-
ing house “Vozes.” His administrative duties as head of this large publishing
company took much time and his taxonomic studies suffered until he re-
tired from “Vozes” and moved to Jacarepagua in 1952 at the age of 60
years.

Living practically in seclusion, with only nominal duties as chaplain for
the Catholic institution for blind women in Jacarepagua that he had founded
in 1947, Father Borgmeier devoted the ensuing 20 years of his life to full-

VOLUME 80, NUMBER 1 143

time systematic entomology. He closed his work on the army ants in 1955
with the publication of his monograph of the Ecitonini of the Neotropical
Region (Studia Ent. 1:1-717) and turned his ant collection over to W. W.
Kempf. He then plunged into a comprehensive taxonomic study of phorid
flies, resulting in revisions of the Neotropical species (1958-1969, about
800 pp.), North American species (1963-1965, 575 pp.), Indo-Australian
species (1966-1967, about 400 pp.), and a catalog of the Phoridae of the
world (1968, 367 pp.). During his lifetime, Father Thomas published 243
papers in entomology and related subjects, totaling over 5,000 pages, and
described more than 1,000 new species of phorid flies, about 100 species
of ants, and a number of myrmecophilous beetles. His fine private collec-
tions of phorids and mrymecophilous beetles are now in the Museum of
Zoology, Universidade de Sao Paulo, and his collection of Neotropical
Formicidae is now in the hands of W. W. Kempf.

From 1959-1965, Father Borgmeier studied nearly 10,000 specimens of
Phoridae from the United States National Museum of Natural History in
Washington from which he determined about 500 species in 40 genera.
Holotypes of 125 new species were deposited in the USNM collections. In
1965 the Smithsonian was able to purchase for a nominal sum from Father
Borgmeier several hundred additional Neotropical phorids, mostly paratypes
including more than 200 species not previously represented in the museum.
During two extensive visits to Washington in 1961 and 1964 Father Thomas
made many close personal friendships which included a number of mem-
bers of the Entomological Society of Washington.

Thomas Borgmeier received many honors during his lifetime, all of
which he accepted modestly and thankfully, as was his nature. He was
the Brazilian delegate to the 5th International Congress of Entomology ©
in Paris in 1932. He was awarded an honorary degree of Doctor of
Science from St. Bonaventure University in New York in 1945. In 1955 he
was elected as titular member of the Brazilian Academy of Science, and
in 1962 he was the first recipient of the Costa Lima prize, instituted by
the Seabra family and administered by the Academy. He was elected as a
fellow of the Brazilian National Research Council in 1958 with the office
of Pesquisador-Chefe. The Franciscan Order in 1965 conferred on him
the honorary degree of Lector Generalis Jubilatus of the Order.

Thomas Borgmeier was a great man in the true sense of the word. We
have listed here some of his innumerable tangible accomplishments. But
his greatness probably lies even more in his intangible influence on his
colleagues, friends, and students in the fields of entomology, education,
religion, and publications, in each of which he was a recognized leader.
Through personal dedication and with great sacrifice of time, effort, and
money he almost singlehandedly kept two important entomological journals

144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

going for more than 40 years. Numerous distinguished Brazilian entomol-
ogists began as his students. Those of us who had the privilege of his
guidance and his friendship join in tribute in memoriam.

W. W. Wirth, Systematic Entomology Laboratory, IIBIII, Agric. Res.
Serv., USDA, c/o U.S. National Museum, Washington, D.C. 20560, W. H.
Robinson, Department of Entomology, Virginia Polytechnic Institute and
State University, Blacksburg, Virginia 24061 and W. W. Kempf (now
deceased).

VOLUME 80, NUMBER 1 145

SOCIETY MEETINGS

840th Regular Meeting—3 March 1977

The 840th Regular Meeting of the Entomological Society of Washington was called
to order by President Ramsay at 8:00 p.m. on 3 March 1977, in the Ecology Theater
of the National Museum of Natural History. Thirty-one members and nine guests were
present. The minutes of the previous meeting were read and approved.

Membership Committee Chairman Utmar reported that 50 members were admitted
to the Society in 1976. Three life members were announced:

Hans Ulrich of Bonn, Germany, the Society’s first foreign Life Member, Lloyd
Knutson, and Joyce Utmar.

The following new applicants for membership were read for the first time by
Membership Chairman Utmar:

Albert E. Cole, Director, W. Va. Dept. of Agri., Plant-Pest Control Div., Charleston,
W. Va. 25305.

Eric M. Risher, 902 Temple Av., Long Beach, California 90804.

Wemer L. Jakob, CDC, P.O. Box 2087, Ft. Collins, Colo. 80522.

Tim L. McCabe, New York State Museum & Science Service, Albany, N.Y. 12224.

Henry E. Sprance, Dept. of Ent., Univ. of Maryland, College Park, Md. 20742.

William Wills, Pa. Dept. of Environ. Res., Med. Ent. Lab. 1, P.O. Box 1467, Harris-
burg, Pa. 17120.

President-Elect Sutherland reported that the Society’s annual banquet will be
tentatively held in June at the Chief Petty Officers’ Club, Washington Navy Yard. Those
attending will be guests of Victor Adler. There was no discussion or disagreement with
these arrangements.

President Ramsay announced that the Washington Academy of Science will meet
on April 21st. The subject of the program is “Signalling by Fireflies.”

President Ramsay requested volunteers to serve as judges for the Fairfax Co.
Regional Science and Engineering Fair.

The program for the evening, presented by Dr. John M. Kingsolver, Systematic
Entomology Lab., was entitled “Bruchidae: Specialists in Seeds.” Dr. Kingsolver
reviewed the distribution and life history of Bruchidae, or seed weevils. All bruchids
spend larval stages in or on seeds. Most of the economically important species attack
commercial legumes. They vary in host specificity; some attack one species or genus
of plant; others have members of 15 families as hosts. Morphology of adults and
affinities to Chrysomelidae were noted. Finally, Dr. Kingsolver showed slides illus-
trating plant associations, mostly from Costa Rica. Examples of toxic plant hosts and
secondary invasion of seeds by other bruchid species were shown.

Several guests were introduced.

Notes and Exhibitions

Robert Nelson said that he received a letter from Lou Davis, now living in California.
Lou recently called Mr. & Mrs. Mort Armitage.

Curtis Sabrosky reported that the Proceedings of the XVth International Congress of
Entomology will be printed and bound this month and should be in the mail by the
end of March. One copy will be mailed free to the registrants of the Congress.
Additional copies may cost more than the original estimate of $9.00.

An exhibit of insect artifacts—insect motifs in man-made objects—was arrayed on
three tables. Over 13 exhibitors showed insects on fabric, shirts, neckties, needle-
work, plates, glasses, toys, pillboxes, small sculptures, pictures and plaques, jewelry, and
other ornaments. Eleven orders of insects were represented.

146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The meeting was adjourned, after which cookies and punch were served.

Joyce Utmar, Recording Secretary, pro tem

841st Regular Meeting—7 April 1977

The 841st Regular Meeting of the Entomological Society of Washington was called
to order by President-Elect Sutherland at 8:00 PM on 7 April 1977, in the Ecology
Theater of the National Museum of Natural History. Twenty-one members and 10
guests were present. The minutes of the previous meeting were read and approved.

President Ramsay was excused due to illness.

No new applications for membership were received during the past month but 2
new life members were announced:

Robert W. Carlson, Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,

USDA, c/o U.S. National Museum, Washington, D.C.
Paul J. Spangler, Department of Entomology, Smithsonian Institution, Washington,
IDK Gy

Treasurer Thompson announced that the Society has a new contract with the printer,
Allen Press, which should allow publication of more pages at the same cost. President-
Elect Sutherland reported that the annual banquet of the combined memberships of
our Society and the Washington Insecticide Society will be held on Thursday, June 2nd
at the Chief Petty Officers’ Club, Washington Navy Yard. Tickets for the banquet
will be available soon. President-Elect Sutherland also reported that the final plans
were being made for the picnic-collecting outing to be held on the weekend of May
14th. A map to the area will be provided at the next meeting.

The main speaker for the evening was Dr. William E. Bickley, Agricultural Experi-
ment Station, University of Maryland. Dr. Bickley spoke on transmission of dog
heartworm by mosquitoes in Maryland. After presenting an overview of the life cycle, Dr.
Bickley discussed his research and that of his students on evaluating the vector potential
of three species of mosquitoes. He reported that Culex salinarius Coquillett is only mar-
ginally successful as a vector but that Aedes canadensis (Theobald) and Aedes vexans
(Meigen) are likely candidates to act as vectors of the microfilariae. Dr. Bickley’s talk
was illustrated with many Kodachrome slides and following the presentation, several
questions were asked concerning the disease.

Notes and Exhibitions: Ashley B. Gurney showed 25 slides of insects he has en-
countered on trips to South America and from around his home. Curtis Sabrosky
announced that recent earthquakes in Romania had caused considerable damage to the
museum in Bucharest as reported by Dr. Weinberg, an asilid expert at the museum.

Several guests were introduced including Dr. Roger W. Crosskey and Mr. Brian
R. Pitkin from the British Museum (Natural History). Dr. Crosskey spoke briefly on
the progress of the Afro-tropical catalogue of Diptera, which the dipterists at the
British Museum are preparing.

The meeting was adjourned at 9:34 PM, after which punch and cookies were
served.

Wayne N. Mathis, Recording Secretary

842nd Regular Meeting—5 May 1977

The 842nd Regular Meeting of the Entomological Society of Washington was
called to order by President Ramsay at 8:05 PM, on 5 May 1977, in the Ecology
Theater of the National Museum of Natural History. Twenty-two members and seven
guests were present. The minutes of the previous meeting were read and approved.

Membership Chairman Utmar read for the first time the names of the following
new applicants for membership:

VOLUME 80, NUMBER 1 147

Zehra Imran Ali, c/o K. M. Hussain, 4 Captain Drive, Watergate Apt. 311. Emery-

ville, California.

Ron Cave, 801 Jamaica Way, Bakersfield, California.

William P. Foerster, 1911 W. St. John’s Ave., Austin, Texas.

Pierre Paul Harper, Department des Sciences Biologiques, Universite de Montreal,

Quebec.
William P. Magdych, Department of Zoology, University of Oklahoma, Norman,
Oklahoma.

Arnold Mallis, 3406 Chiswick Court, Apt. 2C, Silver Spring, Maryland.

Harry Myers, Box 1034, Wiley Hall, West Lafayette, Indiana.

Sandra S. Vincent, 12 C Plateau Place, Greenbelt, Maryland.

President Ramsay announced that Rosella Warner-Spilman has been granted emeritus
status by the unanimous vote of the Executive Committee.

Program Chairman Hellman reviewed plans for the upcoming collecting/picnic out-
ing being sponsored by the Society. The outing will be held during the weekend of
May 13th and 14th at Merrit 4-H camp in Charles County, Maryland. There will be
a pot-luck dinner Friday evening and 6 or 7 cabins will be available for sleeping.
Activities other than collecting will be canoeing, fishing, and fun. Maps were available.

President-Elect Sutherland noted that the annual banquet is scheduled for June 2nd
and that the guest speaker will be Dr. Dale F. Bray, University of Delaware. He
encouraged the membership to purchase tickets early and noted that there will be
a cash bar.

Editor Stoetzel reported that there will be a new format for the Proceedings which
should allow publication of more pages for the same price. The format will have
more print per page, larger type, and different paper. She asked for suggestions or
comments from the membership concerning the format. Editor Stoetzel also an-
nounced that funds are available for another Memoir and that the Editorial Committee
was soliciting manuscripts.

The main speaker for the evening was Dr. Jeffery P. LaFage form the National
Pest Control Association. Dr. LaFage spoke on “Some New Ideas in Termite Ecology.”
Dr. LaFage described his research on two dry wood termites in Arizona and illustrated
his talk with several excellent Kodachrome slides. An interesting discussion period
followed the talk.

Notes and Exhibitions: Ted Bissel noted the existence of the Maryland Entomological
Society, showed a recent publication of the society, and encouraged attendance at
their monthly meetings at Catonville, Maryland.

Manya Stoetzel introduced John Beardsley from the University of Hawaii and also
announced that the Burkses (Barney and Kellie) will be moving to Arizona.

The meeting was adjourned at 9:34 PM, after which punch and cookies were served.

Wayne N. Mathis, Recording Secretary

843rd Regular Meeting—2 June 1977

The combined Annual Banquet of the Entomological Society of Washington and
the Insecticide Society of Washington was held 2 June 1977, at the Chief Petty Officers’
Club, Washington Navy Yard. The evening was presided over by Dr. Maynard Ramsay,
President of the ESW, and Mr. John Kennedy, Chairman of ISW.

Mr. William Helms was Master of Ceremonies. Following the social hour and
buffet dinner, 127 members and guests heard Dr. Dale F. Bray, University of Dela-
ware, discuss “Lesser Known Biological and Cultural Pest Control Techniques.” Mr. and
Mrs. F. F. Smith provided custom designed floral arrangements. Dr. John Neal con-
ducted drawings for floor prizes which were provided by members and industry.

D. W. S. Sutherland, Recording Secretary, pro tem

148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

MISCELLANEOUS PUBLICATIONS

Cynipid Galls of the Eastern United States, by Lewis H. Weld — $ 5.00
Cynipid Galls of the Southwest, by Lewis H. Weld 3.00

Both papers on cynipid galls — cette kee io eee yO Sates ee 6.00
Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman _~ 25

Unusual Scalp Dermatitis in Humans Caused by the Mite Dermato-
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A Short History of the Entomological Society of Washington, ic Aehisy
LBL, (CUPUARYES I do sett ote Noel Reg NE eckeeaens: Soli Sec eis mani P LL DON ORE eee a 50

Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae ),
by George C. Steyskal rE Tie oe Ea Lip rnd eee OF Be 2 I Bes

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

No. 1. The North American Bees of the Genus Osmia, by Grace
Sandhouse. 1939 __ : ey eee AN ee ee ee OD

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9
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. A Classification of Larvae and Adults of the Genus ee
by Adam G. Boving. 1942 . 15.00

No. 3. The Nearctic Leafhoppers, a Generic Classification and Check
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No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace
P. Murdoch and Hirosi Takahasi. 1969 oEAS EEE Bah 6)

No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and
Jeanette Wheeler. 1976 - Bore es SUM A eI ee aN el

Back issues of the Proceedings of the Entomological Society of Washington are
available at $15.00 per volume to non-members and $9 per volume to members of
the Society.

Prices quoted are U. S. currency. Dealers are allowed a discount of 10 per cent
on all items, including annual subscriptions, that are paid in advance. All
orders should be placed with the Custodian, Entomological Society of Wash-
ington, c/o Department of Entomology, Smithsonian Institution, Washington,
D.C. 20560

CONTENTS

(Continued from front cover)

Life history and descriptions of the immature stages of Jamesomyia geminata
(Diptera: Tephritidae ) W. B. STOLTZFUS 87

Parasitism of adult Tabanus subsimilis subsimilis Bellardi (Diptera: Tabanidae)
by a miltogrammine sarcophagid (Diptera: Sarcophagidae) P. H. THOMPSON 69

Notes on the systematics and biology of the biting midge, Forcipomyia elegantula
Malloch (Diptera: Ceratopogonidae) W. W. WIRTH and W. L. GROGAN, JR. 94

NOTES 55, 74, 102, 112, 128, 129, 130, iam
BOOK REVIEWS 133, 134, 135, 136, 138, 139
OBITUARY: The Rev. Thomas Borgmeier 141
SOCIETY MEETINGS 145

ANNOUNCEMENT

VOL. 80 APRIL 1978 NO. 2

545, 70673

PROCEEDINGS

of the

ENTOMOLOGICAL SOCIETY

WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY

CONTENTS

Review of the dermestid beetle genus Caccoleptus with description of a new
species from Colombia (Coleoptera) R. S) BEAT. JR:

A new species of xenocaligonellid mite from the Galapagos Islands (Acari)
R. H. GONZALEZ

Three new Neotropical Rhizoecus (Homoptera: Pseudococcidae)
E. J. HAMBLETON

Notes on the biology of Diglyphus intermedius (Hymenoptera: Eulophidae), a
parasite of the alfalfa blotch leafminer, Agromyza frontella (Diptera: Agromyzi-
dae) R. M. HENDRICKSON, JR. and S. E. BARTH

Biology of a Neotropical snail-killing fly, Sepedonea isthmi (Diptera: Sciomyzidae)
L. KNUTSON and K. VALLEY

Two new species of American Aradidae (Hemiptera) N. A. KORMILEV

Aggregations of male screwworm flies, Cochliomyia hominivorax (Coquerel) in
south Texas (Diptera: Calliphoridae) E. S. KRAFSUR

Defended hunting territories and hunting behavior of females of Philanthus
gibbosus (Hymenoptera: Sphecidae) N. LIN

A new genus and two new species of asterolecaniid scale insects on palm from
Colombia and Trinidad (Homoptera: Coccoidea: Asterolecaniidae)
D. R. MILLER and P. L. LAMBDIN

(Continued on back cover)

op WASHINGTON

DEPARTMENT OF ENTOMOLOGY
SMITHSONIAN INSTITUTION

iW |

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ENTOMOLOGICAL SOCIETY —

OF WASHINGTON
ORGANIZED Marcu 12, 1884

OFFICERS FOR 1978

Dovuctias W. S. SUTHERLAND, President HELEN SOLLERS-RIEDEL, Hospitality Chairwo
Donatp R. Davis, President-Elect Victor E. ApLER, Program Chai
Wayne N. Maruis, Recording Secretary Joyce A. Urmar, Membership Chairwoman
DonaLp R. WHITEHEAD, Corresponding Secretary SuEo NAKAHARA, Custodian
F. Cristian THOMPSON, Treasurer Maynarp J. Ramsay, Delegate, Wash. Acad. Sci.

Manya B. StTorerzEL, Editor
Publications Committee
EARLENE ARMSTRONG ASHLEY B. GURNEY
WAYNE E. CLark GEORGE C. STEYSKAL

a

Honorary President
C. F. W. MursEBECK |

Honorary Members
FREDERICK W. Poos Ernest N. Cory RayMonp A. St. GEORGE

ee ae

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STATEMENT OF OWNERSHIP
Title of Publication: Proceedings of the Entomological Society of Washington.
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Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of
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Editor: Dr. Manya B. Stoetzel, Systematic Entomology Laboratory, BARC-W, Beltsville, Maryland 20705.
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Second Class Postage Paid at Washington, D.C. and additional mailing office.

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use

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 149-155
SYNOPSIS OF THE NORTH AMERICAN
PYRGOTIDAE (DIPTERA)

George C. Steyskal

Abstract—A key to all species of Pyrgotidae known to occur north of
Mexico is given. The new genus Boreothrinax (type of genus, Oxycephala
maculipennis Macquart) is described and two new species, Boreothrinax
dichaetus (Arizona and Mexico) and B. shewelli (British Columbia to
Colorado) are referred to it.

The Pyrgotidae may sometimes cause a marked decrease in the number
of soil grubs (larvae of scarabaeid beetles) by parasitizing the adults. The
female flies have a remarkedly specialized ovipositor for inserting eggs
into the abdomen of the beetles in flight. Because most species of their
hosts fly at night, so do the pyrgotids.

Although about 330 species of Pyrgotidae are known from all major
regions of the world, only eight species in five genera are known from
America north of Mexico. References to previously described species,
synonymy, etc., may be found in Steyskal, 1965.

Key to Genera and Species of North American Pyrgotidae

1(2). Posterior apical corner of anal cell 90°, vein closing cell
straight; alula rudimentary; face with single cuplike depres-
sion, without median keel; body without strong bristles, even
on scutellum (Minnesota to North Carolina and northeast-
ward) Pyrgotella chagnoni (Johnson)

2(1). Posterior apical corner of anal cell much less than 90°; alula
well developed or absent; face with or without median keel
separating antennal grooves; body usually with strong bristles.

3(4). Face without median keel; alula lacking; vein closing anal cell
straight, but forming acute anal with posterior side of anal
cell; body without strong bristles (northward into Arizona)

Stenopyrgota mexicana Malloch

4(3). Face with median keel separating antennal grooves; alula pres-
est (Figs. 1, 2); vein closing anal cell sinuate; body with strong
bristles.

5(8). Wing (Figs. 1, 2) with large areas of plain color, not mottled;
2nd vein usually with stump vein extending backwards near
tip; hindtibia tapering to base (Fig. 4); female with strong
hook (often more or less withdrawn) below at tip of abdomen
(Fig. 3) Genus Pyrgota

150. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

9

Figs. 1-6. Details of Pyrgotidae. 1. Pyrgota undata, wing; 2. Pyrgota fenestrata,
wing; 3. Pyrgota undata, lateral view of female postabdomen; 4. Pyrgota undata, lateral
outline of hindtibia; 5. Boreothrinax shewelli, lateral outline of hindtibia; 6. Spheco-
myiella valida, lateral view of female postabdomen.

VOLUME 80, NUMBER 2 151

6(7).

10(9).

14(13).

Wing (Fig. 1) with hyaline to pale-brown posterior areas open
to margin of wing (Manitoba to Texas and eastward)
Pyrgota undata Wiedemann

Wing (Fig. 2) with hyaline to whitish areas enclosed (“Caro-
lina” to Florida) Pyrgota fenestrata (Macquart)
Wing with rather finely mottled pattern, without large areas
of plain color; 2nd vein without stump vein; hindtibia various;
female abdomen with or without apicoventral hook.
Hindtibia tapering to base (Fig. 4); ocellar and postocellar
bristles usually well developed (ocelli absent); prosternum with
lateral setae; female ovipositor sheath (Fig. 6) with strong hook
at tip below, which may be more or less withdrawn (Minnesota
to Texas and eastward) Sphecomyiella valida (Harris)
Hindtibia strongly constricted near basal % (Fig. 5); ocellar
bristles usually lacking, postocellars lacking (ocelli absent); pro-
sternum setose or bare; female ovipositor sheath (Figs. 9, 10,
12) without apicoventral hook Boreothrinax, new genus
Length of wing 9.3-12.0 mm; prosternum bare; female post-
abdomen as in Fig. 9; male postabdomen as in Fig. 8 (British
Columbia to Colorado) Boreothrinax shewelli, new species
Length of wing less than 9 mm (south and east of Colorado).
Length of wing 7.0-8.4 mm; prosternum bare; female post-
abdomen with spiracle close to tip, without apicolateral lap-
pets (Fig. 11); male postabdomen as in Fig. 7 (Maryland,
Iowa, Arizona, and southward)

Boreothrinax maculipennis (Macquart)
Length of wing 5.7-7.7 mm; prosternum with fine lateral setae;
female postabdomen with spiracle closer to midlength than to
tip, with pair of circular apicolateral lappets (Fig. 12); male
postabdomen as in Fig. 13 (Huachuca Mountains, Arizona and
Mexico) Boreothrinax dichaetus, new species

Boreothrinax Steyskal, new genus

Type of genus, Oxycephala maculipennis Macquart.

The sinuate vein at tip of apical cell, well-developed alula and median
facial keel, and irrorate wing pattern bring this member of the tribe
Pyrgotini into comparison only with Sphecomyiella Hendel, from which it
differs as in the above key. The only known species besides the type of the
genus are the new species described below. The name is of masculine
gender and similar to a few names derived from Greek thrinax by Enderlein.

152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 7-11. Details of Boreothrinax species. 7. Boreothrinax maculipennis, lateral
view of male postabdomen, less aedeagus; 8. Boreothrinax shewelli, lateral view of
male postabdomen; 9. Boreothrinax shewelli, lateral and ventral views of female post-
abdomen; 10. Boreothrinax shewelli, profile of head; 11. Boreothrinax maculipennis,
lateral and ventral views of female postabdomen.

Boreothrinax dichaetus Steyskal, new species
Figs. 12, 13

Differing from B. maculipennis and B. shewelli as in the preceding
key; general color somewhat darker than in either of these species, with
apical % of 3rd antennal segment and most of femora blackish; size in-

VOLUME 80, NUMBER 2 153

termediate, with length of wing 5.7-7.7 mm. Prosternum with about 5 fine
lateral setae.

Female postabdomen as in Fig. 12; ovipositor sheath longer and more
slender than in other species, with pair of circular lappets at latero-
ventral apex, conspicuous pair of long ventral bristles, and spiracles little
apicad of midlength of sheath.

Male postabdomen as in Fig. 13; epandrium in profile bluntly and
obliquely truncate, roughly elliptical.

Types.—Holotype, allotype, and 1é and 32 paratypes, Huach. (= Hua-
chuca) Mts., Ariz., Catal. No. 1311, Brooklyn Museum Coll. 1929 (No. 73650
in U.S. National Museum); paratypes; 22, Miller Canyon, Huachuca Moun-
tains, Cochise County, Arizona, I.V. and 19.VII.1969, 5,000 ft (R. F.
Sternitzky); 12, 5 mi W Durango, Durango, Mexico, 29.VI.1964, 6,500 ft
(J. F. McAlpine); 12, 11 mi W Durango, Durango, Mexico, 15.V1I.1964,
7,000 ft (J. F. McAlpine), all in Canadian National Collection, Ottawa.

The species-name, an adjective meaning “two bristle,” is derived from

Greek.

Boreothrinax maculipennis (Macquart, 1846), new combination
Figs. 7, lu

This species has been placed in Sphecomyiella in the North American
Catalog (Steyskal, 1965), but the constricted hindtibia and the differences
in the female postabdomen show that a new genus is required for it and the
two new species here described. It is likely that other species will be
found, especially south of the United States.

Boreothrinax shewelli Steyskal, new species
Figs. 5, 8-10

Very similar in coloration and external structure to B. maculipennis;
paler than B. dichaetus, antenna and femora usually wholly tawny; larger
(length of wing 9.3-12.0 mm) than B. maculipennis and B. dichaetus; and
differing in postabdominal structure.

Head as in Fig. 10; prosternum bare; outline of hindtibia in lateral view
as in Fig. 5; female postabdomen as in Fig. 9, base of ovipositubus much
swollen and bearing 2 patches of dense, colorless hairs just apicad of
black-setose patches at apex of ovipositor sheath; male postabdomen as in
Fig. 8, epandrium acutely pointed and details of hypandrium, aedeagal
apodeme, and cerci differing markedly from those of the other species.

Types.—Holotype and 1 paratype, $’s, Oliver, B.C., 13.V.1953 (D. F.
Hardwick); allotype, Victoria, B.C., V.1916 (R. C. Treherne); ¢ paratype,
Keremeos, B.C., 8.V.1923 (E. R. Buckell); ° paratype, Boulder, Colo., 18.
VI.1961 (W. R. M. Mason); all in Canadian National Collection, Ottawa;

154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 12-13. Boreothrinax dichaetus. 12. Lateral and ventral views of female post-
abdomen, with (A) spermatheca; 13. Male postabdomen, with (A) sperm pump, (B)
lateral view of postabdomen less sperm pump and apical part of aedeagus, and (C)
tip of aedeagus.

paratypes in U.S. National Museum: 42, 14, Stratton Experimental Water-
shed, near Saratoga, Carbon County, Wyoming, 31.V-12.VI.1972 (J. M.
Schmid), Hopkins No. 36775-J, at light; 12, Logan, Utah, 1.VI.1939 (G. F.
Knowlton, G. S. Stains), Madras, Ore. (Ore. Dept. Agr.).

VOLUME 80, NUMBER 2 155

The species is dedicated to Guy E. Shewell, who first recognized the
distinctness of the species, and in gratitude for many favors.

Stenopyrgota mexicana Malloch
1929 Ann. Mag. Nat. Hist. (10)3:259.

Malloch described this species from a female specimen (now in U.S.
National Museum) taken at an elevation of 7,300 ft at Rio Piedras Verdes,
Sierra Madre, Chihuahua, Mexico. I have examined that specimen and 1
each from Fortin de las Flores, Veracruz, Mexico (in Florida State Collec-
tion of Arthropods) and Nicaragua (U.S. National Museum). A female
specimen taken 0.7 mi SW Turkey Creek, Chiricahua Mountains, Cochise
County, Arizona, 13.VII.1966 (R. G. Beard), examined and returned to
Southwestern Research Station of the American Museum of Natural History,
is the first record of the species north of Mexico.

Literature Cited

Steyskal, G. C. 1965. Family Pyrgotidae. In Stone, A., et al., A catalog of the
Diptera of America north of Mexico. USDA, Agric. Res. Serv., Agr. Hand-
book no. 276:657-658.

Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. Educ. Admin.,
USDA c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 156-163

THREE NEW NEOTROPICAL RHIZOECUS
(HOMOPTERA: PSEUDOCOCCIDAE)

Edson J. Hambleton

Abstract—Three new species in the genus Rhizoecus Kiinckel d'Herculais,
R. divaricatus from Nicaragua, R. olmuensis from Chile and R. variabilis
from Colombia, are described and illustrated. Rhizoecus divaricatus is a pest
of coffee and is the first known species of the genus from Nicaragua.

Among several collections of mealybugs received for identification are
three undescribed species of Rhizoecus from Central and South America.
With the addition of R. divaricatus from Nicaragua, a total of 14 species of
Rhizoecus are known to occur in the American tropics on the roots of
coffee. Three other species of closely related genera of the tribe Rhizoecini
also occur on coffee. The new species, R. olmuensis and R. variabilis, are
from Chile and Colombia respectively. Rhizoecus variabilis was found on
Agave sp., but the host of R. olmuensis is unknown.

The genus Rhizoecus, with the three species described as new, contains
60 species for the Western Hemisphere. My key to the species (1976) is
revised to accommodate the new species as follows:

4(3). “Antennae 6-segmented sonomae McKenzie”

— Antennae 5-segmented A
A(4). With 1 circulus; sensory setae not clavate; tubular ducts

present californicus Ferris

— With 2 circuli; sensory setae clavate; tubular ducts absent
divaricatus, new species

and
37(36). Anal lobes sclerotized; anal ring 45-60 wide; rostrum 51-57 wu
long 38
— Anal lobes unsclerotized; anal ring 68-78 ye wide; rostrum 65-
95 yw long B

38(37). “Anal ring about 60 mu wide, its setae about 87 pw long, outer
part with 25-30 large, angular, irregularly quadrate, mostly
isolated cells; orifice of circulus narrow, less than ™ its basal
width floridanus Hambleton”

— “Anal ring about 45 mw wide, its setae about 55 mw long, outer
part with 19-20 small, elongate, oval cells almost touching end
to end; orifice of circulus wide, more than ™% its basal width
tropicalis Hambleton”
B(37). Tubular ducts absent; with 250 tritubular cerores
olmuensis, new species
— Tubular ducts present; with 45-145 tritubular cerores 39

VOLUME 80, NUMBER 2 157

39(B). Tubular ducts varying in size, complex in design; with 130-
145 tritubular cerores; apical segment of antennae less than
2x as long as wide; sensory seta on segment V short, stout
relativus Hambleton
— Tubular ducts about same size, of simple design; with 45-
50 tritubular cerores; apical segment of antennae 2x as long
as wide; sensory seta on segment V narrow, elongate
ovatus Hambleton
and
41(40). “Digitules at least 4% as long as claws, hind claw about 40 wu
long; rostrum about 80 p long and 78 m wide; with 35-40
tritubular cerores; lobe setae about same size as ring setae
maritimus (Cockerell)”
— Digitules less than % as long as claws, hind claw 24-26 pw long;
rostrum 69-73 p long; lobe setae shorter and more slender
than ring setae C
C(41). Anal ring SO p in diameter; 46-48 cells of outer part without
spicules; cephalic plate absent; circulus dome shaped, 36 pu
wide at base variabilis, new species
— Anal ring 57 mw in diameter, 24-30 cells of outer part with
spicules; cephalic plate present; circulus conical, about 19 wu
wide at base arabicus Hambleton

Rhizoecus divaricatus Hambleton, new species
Figs. 1-9

Adult female—Oval elongate, stout, broadest across abdomen. Length,
1.63-2.16 mm; width, 0.95-1.11 mm. Antennae 5-segmented, rather stout,
average length of segments in microns: I, 35; I, 23; III, 30; IV, 19; V, 75;
apical segment 2 as long as wide, with 4 stout, clavate sensory setae
that taper distally, and 1 spinelike sensory seta near apex. Interantennal
space about length of segment I. Eyes absent. Rostrum of medium size,
averaging 67 wu long, 53 uw wide; rostral loop reaching 2nd coxae. Cephalic
plate irregularly triangulate, 8-10 small body setae on or near its periphery,
length 53 yw, width, 63 px. Dorsal ostioles strongly sclerotized, with body
setae and pores bordering ostiole rims.

Legs moderately short, stout, average length of segments of hind pair in
microns: Trochanter, 46; femur, 94; tibia, 88; tarsus, 72; claw, 27; digitules
short, setose, not reaching to % length of elongate, narrow claws.

Two conical circuli, 16 « wide at base, 1 each on abdominal segments
III and IV. Anal lobes each with strongly sclerotized protruding area,
longer than width of anal ring, and 12-14 subequal elongate setae, longest
80-83 pw long. Anal ring of medium size, about 46 wide, its setae stouter
than longest anal-lobe seta, about 60 y long; outer portion of anal ring

158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

en
Dy

il

} SUD. . .

Figs. 1-9. Rhizoecus divaricatus, female. 1. Terminal segment of antenna; 2. Anal
ring, right half; 3. Cephalic plate; 4. Rostrum; 5. Circulus; 6. Tritubular ceroris, small,
dorsal; 7. Tritubular ceroris, large, lateral; 8. Hindclaw; 9. Tritubular ceroris, large,

ventral.

VOLUME 80, NUMBER 2 159

with 18 small, oval, elongate cells, each with elongate spicule; inner por-
tion of ring with 8-12 larger, more elongate cells, some serpentine shaped,
adjacent to 8-10 large, darkened, globular cells. Tritubular cerores of 2
types, 14 large ones with stout, tapering, divaricating ducts ranging between
15-24 pw long, occurring dorsally on or near body margins and along mid-
dorsal line, 8 ventrally and submarginally slightly smaller; the remaining
16-21 cerores about % size of larger type with short, stout ducts, 8-9 mu
long, occurring ventrally across abdominal segments V-IX, occasionally
1 on IV. Multilocular disk pores occurring ventrally, 21-49 scattered on
abdominal segments VIII-IX, 5-8 along posterior border of VII, 1-6 on
VI. Tubular ducts absent. Trilocular pores fairly evenly distributed but
sparse in some areas. Body setae sparse, mostly short, inconspicuous,
larger setae about 60 pw long.

Holotype female——Nicaragua: Granada, 5-I-1975, José Gonzales, on
roots of Coffea arabica; paratypes, 5 mounted on 2 slides, taken with holo-
type. All in USNM.

This interesting mealybug keys to R. californicus Ferris but differs by pos-
sessing only 38-45 cerores of 2 types, by having 2 circuli, by its stout,
clavate sensory setae, and by the absence of tubular ducts. Rhizoecus
divaricatus most closely resembles R. vitis Borchenius from the Crimea.
It differs primarily by having 2 instead of 3 circuli, a large triangulate
cephalic plate, and no tubular ducts. Rhizoecus divaricatus is the first
recorded species of the genus from Nicaragua.

Rhizoecus olmuensis Hambleton, new species
Figs. 10-15

Adult female.—Undistended body elongate. Length, 0.94 mm, width,
0.37 mm. Antennae 6-segmented, of medium size, length of segments in
microns: I, 28; II, 22; III, 28; IV, 20; V, 19; VI, 43; apical segment less than
2x as long as wide, with 3 narrow, elongate falcate sensory setae and 1
slender, spinelike sensory seta; segment V with 1 shorter, elongate sensory
seta. Interantennal space equal to length of segment I. Eyes absent. Ros-
trum 65 pu long, 57 « wide; rostral loop reaching beyond 2nd coxae. Cephalic
plate 38 yw long, 30 uw wide, irregularly quadrate, with 4-5 body setae on its
periphery. Dorsal ostioles conspicuous, sclerotized, pores and setae not
crowded near ostiole rims.

Legs of medium size, length of segments of hind pair in microns: Tro-
chanter, 40; femur, 83; tibia, 73; tarsus, 51; claw, 20; digitules slender, di-
lated apically, extending to or slightly beyond apex of stout, acute, curved
claws.

One conical circulus, about 10 w in diameter as base. Anal lobes un-
developed, unsclerotized, each with 3 elongate setae, longest about 57 yu

160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

H{ l | 5 re
: tq

aN

SS

(

Figs. 10-15. Rhizoecus olmuensis, female. 10. Terminal segments of antenna; I1.
Hind claw; 12. Rostrum; 13. Anal ring, right half; 14. Cephalic plate; 15. Tritubular
ceroris, lateral.

and several body setae. Anal ring prominent, 74 s wide, its setae stouter
and longer than anal-lobe setae, 77-82 gz long; outer portion of anal ring
with about 28 irregularly oval or triangulate cells, most with short, blunt
spicules; inner portion of ring with 18-20 elongate curved to triangulate

VOLUME 80, NUMBER 2 161

cells adjacent to area of darkened, globular cells. Tritubular cerores of
medium size, 4.5-5.0 wide, their ducts about 7 mw long, about 250 well
distributed over derm, occurring with setae and pores, forming bands
around segments, leaving clear areas intersegmentally. Multilocular disk
pores and tubular ducts absent. Trilocular pores fairly abundant, more
numerous dorsally. Body setae variable in size and length, longest about
40) » long, uniformly distributed.

Holotype female —Chile: El Granizo, Olmue, Valparizo Prov., 5-IV-1961,
L. Campos. Host unknown. In University of California at Davis.

The description of R. olmuensis is based on a mature undistended fe-
male, consequently the body measurements given are tentative. In the
revised key this species places close to R. relativus Hambleton. The absence
of tubular ducts and greater number of cerores in olmuensis readily sepa-
rate the two species. In general appearance olmuensis resembles R. ad-
venoides Takagi and Kawai from Japan but differs in the absence of eyes,
multilocular disk pores and tubular ducts.

Rhizoecus variabilis Hambleton, new species
Figs. 16-20

Adult female ——Oval elongate. Length, 1.75-2.19 mm; width, 0.85-1.20
mm. Antennae 6-segmented, elongate, average length of segments in mi-
crons: I, 34; II, 25; III, 49; IV, 20; V, 20; VI, 59; apical segment 2 as long
as wide, with 3 elongate, medium-sized falcate sensory setae and 1 shorter,
strongly tapered sensory seta; segment V with 1 smaller, weakly lanceolate
sensory seta. Interantennal space equal to combined length of segments
IV-VI. Eyes small, hemispherical. Rostrum averaging 73 mw long, 57 pw
wide; rostral loop extending beyond halfway to 2nd coxae. Cephalic plate
apparently absent. Dorsal ostioles inconspicuous, weakly sclerotized, bor-
dered by few setae and pores.

Legs moderately stout, elongate, average length of segments of hind pair
in microns: Trochanter, 51; femur, 143; tibia, 110; tarsus, 67; claw, 26;
digitules short, setose, variable in length, sometimes about as long as
slender claws.

One domed-shaped, faveolate circulus, 36 w wide at base, 15 mw across
orifice. Anal lobes undeveloped, each lobe area with 1 elongate seta,
about 88 yu long and 2 shorter setae. Anal ring large, well defined, 80 mw in
diameter, ring setae averaging 91 mp long, longer and stouter than lobe
setae; outer portion of anal ring with 46-48 oval, subtriangulate cells un-
evenly arranged; cells of inner portion of ring 24-26 in number, larger, more
irregular, some elongate, bordered by a darkened, semicircular, cellular area.
Tritubular cerores small, with finely tapered ducts, 55-65 present, widely
distributed, more common dorsally. Multilocular disk pores absent. Tubular

162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 16-20. Rhizoecus variabilis, female. 16. Terminal segments of antenna; 17.
Circulus; 18. Rostrum; 19. Anal ring, right half; 20. Hind claw.

VOLUME 80, NUMBER 2 163

ducts short, stout, their diameter less than that of trilocular pore, sparsely
distributed over derm dorsally and ventrally. Trilocular pores numerous,
fairly evenly distributed. Body setae mostly short, rather inconspicuous,
variable in length, longest about 36 wu long.

Holotype female —Colombia: Garagoa, 1-V-1973, F. Mosquera, on Agave
sp., on slide with 2 paratypes, specimen in middle is holotype; paratypes,
6 taken with holotype. In USNM.

Rhizoecus variabilis keys to R. arabicus Hambleton, also from Colombia.
However, variabilis is a larger species and is readily separated from arabicus
by the size of its anal ring, whose cellular structure is without spicules, by
the dome-shaped faveolate circulus, by the narrow claws, and by the absence
of a cephalic plate.

Literature Cited

Borchsenius, N. S. 1949. Fauna of USSR. Homoptera, Pseudococcidae. Vol. VII,
382 pp. Akad. Nauk Zool. Inst. Leningrad (In Russian).

Hambleton, E. J. 1976. A revision of the New World mealybugs of the genus
Rhizoecus. U.S. Dept. Agri. Tech. Bull. 1522. 88 pp.

Takagi, S., and S. Kawai. 1971. Two new hypogeic mealybugs of Rhizoecus from
Japan (Homoptera: Coccoidea). Kontyu. 39(4):373-378.

Cooperating Scientist, Systematic Entomology Laboratory, I[BIII, Fed.
Res., Sci. Educ. Admin., USDA (mail address: 5140 Worthington Drive,
Washington, D.C. 20016).

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 164-170
AGGREGATIONS OF MALE SCREWWORM FLIES,
COCHLIOMYIA HOMINIVORAX (COQUEREL) IN
SOUTH TEXAS (DIPTERA: CALLIPHORIDAE)

E. S. Krafsur

Abstract.—Noted, for the first time, is the occurrence of aggregations of
males of the screwworm fly, Cochliomyia hominivorax (Coquerel). The
continuous presence of males in each of two sites in south Texas was in-
dependent of the presence of cattle or of female screwworm flies. Males
captured in autumn 1975 proved to be wild; those captured in spring 1976
were released, sterile insects. The behavior of males, wild or sterile, included
vigorous conspecific and interspecific interactions. There was evidence of
territoriality and competition for favored perching sites. The observations
are consistent with an hypothesis that the male screwworm aggregations
were mating assemblies similar to those known among other cyclorrhaphan
Diptera. The significance of male aggregation to sterile fly liberations is dis-
cussed.

Sterile fly release strategy, as currently practiced in the USDA, Southwest-
ern Screwworm Eradication Program, is predicated upon three factors. One
is that sterile Cochliomyia hominivorax (Coquerel) must be applied to areas
where breeding may be occurring because the probabilities of detecting
cases of myiasis, particularly when prevalence is low, are rather poor.
Another factor upon which sterile fly dispersions is based is that of scale:
Vast areas are at risk to primary screwworm and while it is possible to
learn where breeding occurs, it is much more difficult to know where it
is not. Thus great effort must be put to distributing sterile flies in areas
that indeed have no screwworms at all. For these reasons, program aircraft
are scheduled to “grid” evenly the region at risk. Generally, an area is
treated to sterile flies packaged in units of 1,800-2,200 (%2 males) and
distributed in lanes set 5 or 10 miles (8-16 km) apart,' “doses” of sterile
flies varying with their availability and with case incidence reports sub-
mitted by ranchers, state, and federal personnel. The third consideration
underlying sterile fly release strategy is the implicit assumption? that mating
takes place after random dispersion of virgin flies from their pupation sites,
a behavioral trait assumed to have evolved in order to prevent brother-
sister matings and consequent inbreeding. This supposition justifies the
practice of distributing sterile flies in parallel lanes and requires of males
that they seek out mates up to several miles from the point of their re-
lease. That they have the ability and necessary behavioral traits necessary
to accomplish this was never demonstrated. It is therefore of very practical
importance that we learn the sequence of events leading to mating; failing

VOLUME 80, NUMBER 2 165

this, that at least the operational premise of dispersion prior to mating is
tested against such evidence as exists or can reasonably be inferred from
data on related calyptrate fly species.

Observations

Two male screwworm aggregation sites, about 40 miles apart, were
found in the thorny brush of northern Hidalgo County in south Texas. Male
aggregation and mating behavior in screwworms has been heretofore un-
recorded. The sites, wooded with Acacia spp. and mesquite (Prosopis spp.)
up to about 7 m in height, were discovered in October 1975, and one of
them (Fig. 1) was visited repeatedly until July 1976. Ladders were used
when needed to gain access to perching sites. Where it proved impos-
sible to get within a few feet of a specimen, identification was made
through the use of a telescopic lens. Numerous color photographs were
obtained. Discrimination of Cochliomyia hominivorax from C. macellaria
(Fabricus) was easily accomplished visually. Captured material was used
to confirm the identifications and some 16 crosses were made to lab-
oratory-reared female C. hominivorax under controlled conditions (see
below). Cochliomyia macellaria was commonly observed in the study lo-
cations, but males of this species were never observed to demonstrate the
behavior characteristic of C. hominivorax to be described, nor to occupy
similar resting and perching sites.

Numbers of male C. hominivorax actually observed (simultaneously)
varied from two to many on any one day, but their speed and frequency of
flight were such that they proved impossible to census accurately. Even
when three observers were present, simultaneous sighting of more than eight
flies was difficult. More males than actually enumerated may well have been
present. There was available no means which allowed us to make an
unbiased estimate of male numbers, as the males moved rapidly through
three dimensions and were not always easy to see. No screwworm
males were observed in other locations similar to the identified aggrega-
tion sites nor were any found after repeated searching throughout the areas
surrounding the two sites. Why these locations were preferred is unknown,
as there seemed to be nothing particularly distinguished about them.

The aggregations of screwworm males were not dependent on the pres-
ence of livestock. When the present observations were begun, cattle were
no longer to be found in site A, and only small, transient numbers of them
were observed nearby in the year following. Site B was the home of a sub-
stantial herd of cattle, among whom were individuals infested with screw-
worms. Pens of wounded, sentinel sheep (Davis et al., 1968) were estab-
lished in both aggregation sites. Gravid female C. hominivorax oviposit on
the wounds and the resulting number of egg masses can be taken as an
index of screwworm population density. No confusion exists between wild

166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-6. Cochliomyia hominivorax, November 1975. 1. Aggregation site A. Trees
are mesquite, Prosopis; 2-6. Male screwworms photographed in life, all on mesquite
branches and twigs. Note that the flies face the sun.

VOLUME 80, NUMBER 2 167

Table 1. Weekly screwworm egg masses, mean temperatures, and reported incidence
of myiasis in Hidalgo County, Texas, 1975.

Week beginning of
SS Vari-
10/22 10/29 11/5 11/12 11/19 11/26 12/3 12/10 Avg. ance

Egg Mass Nos.

Site A — 1 13 1 0 1 0 0) ey — BIPNE
Site B 5 18 24 9 0) 0) 0 0 HO SUH
Incidence?’ 23 33 22, 23 4 9 11 7 16.5 102.9

Aveatemps CG 21.7 23.0 23:3 183 134 154 17.3 189 — —-

“Based on samples of larvae voluntarily submitted by ranchers to the Southwestern
Screwworm Eradication Program, APHIS, USDA.

and released, irradiated females because ovarian development does not occur
in radiosterilized flies. Of 72 egg masses collected (Table 1), only four
were sterile (6%), suggesting that sterile male releases of about 1,540/km?/
week were ineffective. The number of ovipositions were few in the first
week of sampling, were greatest in the second or third weeks and declined
precipitously thereafter. Screwworm incidence in Hidalgo County, as
indicated by samples of larvae submitted to APHIS for identification, fell
off less sharply than the local populations sampled in sites A and B. It
is important to note that temperatures (Table 1) were adequate to allow
ovarian development and do not explain the virtual cessation of oviposition.
The egg-mass-sampling distributions were strongly clumped, as suggested
by high variance to mean ratios. These are typical of screwworm popula-
tions in Texas and Mexico (Krafsur and Hightower, unpublished). The
absence (in site A) or presence (in site B) of cattle seemed to have little
influence on female screwworm populations.

Males continued to be seen until early December, well after the ap-
parent dispersion of female screwworms. Three captured males were each
caged with five virgin, laboratory-reared females. That the flies were wild
was confirmed because only fertile egg masses were obtained. One of
the males was caught in site B while feeding on a fresh cow dropping.

No evidence of screwworms was obtained after the advent of cold
weather in December. Despite the continuous presence of wounded sheep
in site A, no egg masses were found until 26 April, but male screwworms
were Observed from 5 March onwards. Some 13 were caught over a 3-
month period, and their fertility was tested by crossing them to labora-
tory-reared stock. All proved to be sterile, released flies. Females caught
probing in wounds of the sentinel sheep also proved to be released flies,
their ovaries failing to show evidence of yolk deposition after being held
four days at 29°C. No male screwworms were observed on the sentinels,
nor in their pens. Both sexes of C. macellaria, however, were frequent on
sheep wounds and droppings. Nectar feeding among spring populations of

168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

screwworm males was suggested by the observation of heavy pollen deposits
on some perching specimens.

Clear behavioral patterns were noted, and these were common to autumn,
spring, and summer screwworm populations, being observed whenever
the sites were visited, from 1000-1830 h. Resting males most frequently
chose as perching sites prominent, leafless branches 3-6 m above ground
level and facing open, sunny spaces. Distal perches were favored, but when
these were occupied, exposed interstitial positions were assumed. Some
male screwworms perched in leaves of the evergreen shrub, granjeno
(Acanthoceltus spp.). Perching could easily be induced by holding up a
stick or one’s hand near a flight station or in one of the open “arenas.” The
new station usually would be taken by a nearby male. Characteristic,
aggressive poses were adopted by perching males, heads up and abdomens
down (Figs. 2-6). Cochliomyia macellaria were never seen to adopt such
attitudes. When temperatures were about 20°C or higher, perching males
periodically took off on short elliptical or circular flights; they quickly
returned to the same positions unless displaced by another male. Vigorous
interactions among screwworm flies were evident. A male occupying one
site would reconnoiter other sites, and if occupied, interception and ap-
parent conflict invariably occurred with two males flying off together in
a mass of legs and wings, only to return to their respective stations. A
perching male typically flew after screwworms, other fly species and
even butterflies when these came within about 3 m of its resting position.
Small numbers of perching sarcophagid and muscid males were occasionally
noted.

On cool days, <20°C, perching behavior was less frequently observed,
and males could more easily be found resting closely appressed to the
internodes of mesquite branches and twigs, usually exposed to the sun.
This resting behavior supports the observations of Hightower (1963), who
recorded nocturnal resting sites of marked, released screwworms. When
windy conditions prevailed in addition, flies were found resting on mesquite
twigs, within 1 m of the ground and C. macellaria were observed to
behave similarly at such times.

Discussion

Observations made simultaneously on male and gravid female screw-
worm flies are consistent with an hypothesis that local aggregation of a
cohort of females is transitory, while that of males is not. In other Diptera,
males taken from a swarm or waiting station were demonstrated to re-
turn to their aggregation sites (Downes, 1969; Hunter and Webster, 1973).
The sequence of events for most female screwworms seems to be, eclosion,
mating at 2-3 days of age, oviposition at 5-6 days (Hightower et al., 1972)
(should suitable hosts locally exist) and dispersion. This interpretation

VOLUME 80, NUMBER 2 169

is supported by the clumped distribution of ovipositions observed in this
and unpublished work conducted in Texas and Mexico. The sampling
distribution of egg masses, typically aggregated spatially and temporally,
probably arises from a similar distribution of larvae in their vertebrate
hosts a generation earlier. Random dispersion before mating is the al-
ternative. The hypothesis is unlikely. Recapture of sterile females is max-
imal on the third or fourth day after release, when the flies are 4-5 days
old and already mated (Hightower, 1969). Cochliomyia hominivorax exist
in characteristically low but highly aggregated density for much of their
breeding season and over much of their range. The probability of encounter
between the sexes of randomly dispersed screwworms, in the absence of a
specific, long-range mechanism of attraction, must surely be very low. No
such mechanism has been demonstrated, to my knowledge, among the Dip-
tera. The principal cue initiating sexual activity, in many Diptera, is move-
ment. Pheromones enabling sexual recognition over very short distances or
upon contact have been demonstrated in the house fly and in a tsetse
fly (Langley et al., 1975).

Male aggregations seem an altogether common phenomenon in many
species of calyptrate flies, and are held to function as mating assemblies
(see Downes, 1969 for review). The characteristic poses, territoriality, short,
frequent flights among screwworms and their interactions with other flies
differ little from that described for Gasterophilus sp. (Walton, 1930), the
face fly, Musca autumnalis DeGeer (Teskey, 1969), several species of
Cuterebridae (Catts, 1967; Hunter and Webster, 1973), Oestromyia sp.
(Grunin, 1958), Sarcophaga sp. (Thomas, 1950), three species of Tachinidae
and an anthomyiid fly (Lederhouse et al., 1976). Species of Syrphidae,
Tabanidae, Tachinidae, Sarcophagidae, Calliphoridae, Muscidae, and
Larvaevoridae have been observed in localized aggregations by Chapman
(1954) and by Dodge and Seago (1954).

The significance of the present observations to sterile fly release opera-
tions is clear. The efficiency of the sterile male release method will be
enhanced by maximizing the chances of putting sterile screwworms into
actual or potential breeding sites. Numerous mark, release, and recapture
studies on sterilized screwworm flies have demonstrated the extraordinary
dispersal potential of females (Hightower et al., 1965). There is, however,

no evidence at all to support the contention that released males disperse in
this way.

Acknowledgments
Messrs. L. Garcia, M. Hernandez, J. Villarreal, A. Vela, F. Gonzalez and
R. Ramirez, each of Veterinary Services, USDA-APHIS, Southwestern
Screwworm Eradication Program, Mission, Texas, assisted in making the
present observations. I thank Ms. Mary Cochran for secretarial help and
Dr. B. G. Hightower, Comision Mexico-Americana, Mexico 1, D.F., for

170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

critically reading the manuscript. Dr. Bruce M. Christensen, Iowa State
University, Ames, kindly made up the plate from my color slides.

Literature Cited

Catts, E. P. 1967. Biology of a California rodent bot fly Cuterebra latifrons Coquillett.
J. Med. Entomol. 4(2):87-101.

Chapman, J. A. 1954. Studies on summit-frequenting insects in Western Montana.
Ecology 35:41-49.

Davis, R. B., B. G. Hightower, D. A. Alley, J. E. Turner, and E. Lopez. 1968. Releases
of sterile screwworm flies in northern Veracruz, Mexico, measured by recovery
of sterile egg masses. J. Econ. Entomol. 61(1):96—101.

Dodge, H. R., and J. M. Seago. 1954. Sarcophagidae and other Diptera taken by
trap and net on Georgia mountain summits in 1952. Ecology 35:50-59.

Downes, J. A. 1969. The swarming and mating flight of Diptera. Ann. Rev. Entomol.
14:271-298.

Grunin, K. I. 1958. The biology Oestromyia marmotae. Ged. (Diptera, Hypoder-
matidae), the warble fly of the long-tailed marmot. Entomol. Rev. 37:763-767.

Hightower, B. G. 1963. Nocturnal resting places of the screwworm fly. J. Econ.
Entomol. 56(4):498—500.

—. 1969. Population dynamics of the screw-worm fly. Cochliomyia hominivorax
(Coquerel), with respect to control by the sterile-male technique. In “Insect
Ecology and the Sterile Male Technique,” pp. 25-31, I.A.E.A., Vienna.

Hightower, B. G., A. L. Adams, and D. A. Alley. 1965. Dispersal of released irradiated
laboratory-reared screwworm flies. J. Econ. Entomol. 58:373-374.

Hightower, B. G., J. J. O'Grady, Jr., and J. J. Garcia. 1972. Ovipositional behavior
of wild-type laboratory-adapted strains of screwworm flies. Environ. Entomol.
1(2):227-229.

Hunter, D. M., and J. M. Webster. 1973. Aggregation behavior of adult Cuterebra
grisea and C. tenebrosa (Diptera: Cuterebridae). Can. Entomol. 105:1301-1307.

Langley, P. A., R. W. Pimley, and D. A. Carlson. 1975. Sex recognition phermone
in tsetse fly Glossina morsitans. Nature (London). 254:51-52.

Lederhouse, R. C., R. A. Morse, J. T. Ambrose, D. M. Burgett, W. E. Conner, L. Ed-
wards, R. D. Fell, R. Rutowski, and M. Turell. 1976. Crepuscular mating
aggregations in certain Ormia and Sitophaga. Ann. Entomol. Soc. Am. 69(4):
656-658.

Teskey, H. J. 1969. On the behavior and ecology of the face fly, Musca autumnalis
(Diptera: Muscidae). Can. Entomol. 101:561—576.

Thomas, H. T. 1950. Field notes on the mating habits of Sarcophaga Meigen (Dip-
tera). Proc. R. Entomol. Soc. London (A) 25:93-98.

Walton, C. L. 1930. The occurrence of males of the horse bot fly. North Western
Natur. 5:224-226.

Department of Entomology, Iowa State University, Ames, Iowa 50011
(formerly: Veterinary Services, APHIS, USDA, Mission, Texas 78572).

Footnotes

1This practice was discontinued in early 1977. Small fly containers and narrow
swaths (~2 km) are now used. (See Krafsur, E. S., and L. Garcia. 1977. J. Med. Entomol.
14(6):687-697.)

* Made explicit by Bushland, R. C. 1975. Bull. Entomol. Soc. Am. 21:23-26.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 171-178
REVIEW OF THE DERMESTID BEETLE GENUS CACCOLEPTUS
WITH DESCRIPTION OF A NEW SPECIES FROM COLOMBIA
(COLEOPTERA)

RS, Bealiyr:

Abstract —Adult and larval stages of Caccoleptus wicki, new species from
Colombia, are described. A key is provided for the three species in the
genus. The known geographic range of C. anisotomoides Sharp is ex-
tended from Honduras to Panama.

Larvae and adults of an undescribed species within the dermestid beetle
genus Caccoleptus have been found in northern Colombia preying on eggs
and first-instar larvae of Opsiphanes cassina F., a nymphalid butterfly de-
foliator of banana plants. Because of the potential economic value of the
species in biological control, it is important to name and describe it. No
less important is the opportunity to restudy this little-known genus with a
view to gaining a better understanding of its place in the family Dermesti-
dae and of increasing our knowledge of the biology of the family.

The genus was described by David Sharp in 1902. No one appears to
have worked on the genus since the appearance of Sharp’s original study.

Recognition

Adult members of the genus may be recognized as dermestids by their
compact shape, the small, deflexed head which is retracted into the pro-
thorax, the presence of a median ocellus, and the shape of the hind coxa,
which is grooved for the reception of the femur. Members of the genus,
which falls within the tribe Megatomini (adopting the classification of
Zhantiev, 1976), are easily distinguished from most other dermestids by their
rotund shape, which gives them an appearance of tiny coccinellids. Other
characters which appear to separate adults from other genera of the
Megatomini are the following: The male antenna (observed only in C. ro-
tundus Sharp) has an 8-segmented club (Fig. 1D); the female antennal
club may be 4- or 5-segmented; the pronotum is margined laterally for
its entire length, including the anterolateral angle; the antennal fossa oc-
cupies all of the hypomeron and is margined along its entire posterior side
by a knife-like carina; the prosternum is relatively short with a broad or
narrow posterior process; the short mesosternum is completely divided by
a groove for the reception of the prosternal process with the pieces on either
side of the groove much wider than long (Fig. 1E); the metasternum is
strongly convex; the first visible abdominal segment has a single oblique
stria on each side originating near the medial edge of the trochanter and

172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

extending to the posterior margin of the segment. Sharp’s statement that
the scutellum is covered by the pronotal lobe is not entirely correct. In each
of the species the scutellum is somewhat exposed, although in C. rotundus
only a very small part of the apex is visible. The metathoracic wing (Fig.
1C) (adopting the terminology of Wallace and Fox, 1975) has a very short
Aia + Ai, (designated as the postcubitus by Beal, 1967, for species of
Megatoma), an unbranched vein formed from Aj, (designated as first
vannal vein by Beal), and a single vein formed from Ay, + Aie + Az (desig-
nated as second vannal by Beal). Vein A; is absent. Vein A, (jugal vein)
is very faint, if it is present at all.

The larval stages are known only for C. wicki. Mature larvae of this
species are easily recognized as belonging to the tribe Megatomini by
the presence of hastisetae (spear-headed setae) inserted on the nota and
terga. They are readily distinguished from known larvae of all other genera
of Megatomini by each of the following characters: (1) the reduced width
of the first abdominal segment, which is about % as wide as either the meta-
thoracic notum or the tergum of abdominal segment 2 (Fig. 2F); (2) the
absence of an antecostal suture on any of the nota or terga; (3) the absence
of spicisetae on both the anterior part of the abdominal terga (the region
that probably should be considered the acrotergite) and along the posterior
margin of the terga (Fig. 2F); (4) the elongated shape of the accessory
papilla at the apex of the second segment of the antenna (Fig. 2D).

Systematic Position

Both larval and adult characters appear to associate this genus with
Phradonoma and Trogoderma. The fact that in the larvae none of the hasti-
setae are inserted on the membrane behind any of the abdominal terga but
are all inserted on sclerotized areas of the terga places the genus in a
group with Trogoderma, Phradonoma, Globicornis, Megatoma, and Reesa.
A specialization found in Anthrenus, Thaumaglossa, Cryptorhopalum, and
Ctesias, which separates them as a group, is that all have a large cluster
of hastisetae inserted on the membrane on each side behind one or
more of the abdominal terga. The position of Caccoleptus with respect to
each genus in the former group is a little less obvious, but it seems to
have originated from a stem common to Phradonoma and Trogoderma.
Each of these has the distal group of papillae on the epipharynx enclosed
in one or more rings rather than in the center of a callosity (Ford and
Kingsolver, 1966). As is true of Phradonoma tricolor (Arrow), the distal
group of papillae on the epipharynx are divided into a group of four and
a group of two papillae and are enclosed in two rather than in a single
ring. It also has the setae at the apex of the pretarsus greatly unequal in
length in common with most Trogoderma rather than equal in length as is

VOLUME 80, NUMBER 2 173

true of Globicornis, Megatoma and Reesa. In the case of adults, the shape
of the antennal fossa, enclosed behind by a knife-like carina, is characteristic
of Trogoderma. The 8-segmented club of the male antenna is also much
more like Trogoderma than like the 1- to 3-segmented club of the latter
group. Nonetheless, the specializations found in Caccoleptus, particularly in
the larval stages, clearly warrant its separation from Trogoderma.

Type of the Genus

The type of the genus is Caccoleptus rotundus Sharp by original designa-
tion.

Key to Species of Adult Caccoleptus

1. Integument of elytra dark mahogany brown to piceous with broad
ochreous or reddish submedian and subapical bands; submedian
band usually with extension along median suture to base. Dorsal
pubescence of dark brown to piceous and light golden hairs. Golden
hairs on elytra forming broad bands coincident with light macu-
late areas. Prosternal process broad, as wide as 2* width of front
tibia at level of hind margin of front coxa wicki, new species

— Dorsal integument immaculate, yellowish brown to mahogany
brown. Dorsal pubescence of white hairs and light to dark golden-
brown hairs. Elytra transversed by narrow bands of white pu-
bescence. Prosternal process narrow, no wider than front tibia at
level of hind margin of front coxa

bo
bo

White hairs of elytron forming small basal patch, narrow, more or
less continuous submedian band, and narrow subapical band; white
hairs somewhat ensiform but not more than 12x as wide as
golden-brown hairs rotundus Sharp
— White hairs of elytron forming small basal patch near scutellum,
subbasal patch near humerus, narrow, interrupted submedian band,
narrow interrupted subapical band, and apical patch; white hairs
ensiform, about 2 as wide as golden-brown hairs and tending to
form clusters with strikingly white appearance anisotomoides Sharp

Caccoleptus rotundus Sharp

Caccoleptus rotundus Sharp, 1902:650.

The type-locality for this species is Panama. An additional male speci-
men in the U.S. National Museum of Natural History is from El Cermeno,
Panama, July to August, 1941 (J. Zetek). A label on the specimen states that
it was collected “ex fruit of Labatia standleyana Pittier.”

174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Caccoleptus anisotomoides Sharp
Caccoleptus anisotomoides Sharp, 1902:650.

The type-locality is Rio Hondo, British Honduras. Two female speci-
mens are deposited in the U.S. National Museum of Natural History, one
from Panama City, Panama (no date, E. A. Schwarz), and one from Pedro
Miguel, Canal Zone, Panama, 17 April 1911 (E. A. Schwarz).

Caccoleptus wicki Beal, new species

Adult female.—Habitus as illustrated (Fig. 1A). Dorsal pubescence sub-
recumbent, bicolorous with light golden and piceous hairs; no ensiform
setae present. Head with integument reddish brown on frons, piceous on
vertex. Ratio of width across compound eyes to interocular distance 43:26.
Punctures of vertex simple, equal in diameter to diameter of facet of com-
pound eye, separated by 1-4 diameter of single puncture. Antenna
ochreous with 4-segmented club as illustrated (Fig. 1B). Pronotum with in-
tegument reddish; light colored hairs forming transverse subapical band
and basal band; punctation of disc similar to that of vertex of head. Elytra
with integument piceous with broad median and apical ochreous bands;
light colored setae of elytra distributed on light colored bands of in-
tegument and in addition forming narrow band along base just posterior
to sides of pronotal process and forming line from base along median
suture to median band. Ventral surfaces with recumbent, light golden
setae; integument of thoracic sterna piceous; integument of abdominal
sterna reddish brown with black margin on first 4 sterna. Prosternal pro-
cess broad (Fig. 1E). Mesosternum strongly transverse (Fig. 1E). Legs
ochreous. Ratio of width (measured across humeri) to length (of pronotum
and elytra combined) 1:1.39. Length (of pronotum and elytra) 1.66 mm.

Range of observed variations: Color of bands on elytra varying from
ochreous to brick red; median band frequently produced anteriad to base
and often expanded at base to form short band as wide as pronotal process.
Visible abdominal sterna 1-4 usually black with sternum 5 reddish. Ratio
of width to length varying from 1:1.35 to 1:1.49. Length ranging from
1.60 mm to 1.82 mm.

Mature larvae.—Color of nota and terga dusky; sterna and legs hyaline.
Dorsal spicisetae long, some on lateral margins of nota % longer than
width of notum; hastisetae with apex as illustrated (Fig. 2E); hairs of
caudal brush long, some % longer than body length. Antenna as illustrated
(Fig. 2D); segment 1 bearing 1 or no setae; segment 2 without setae; acces-
sory papilla long. Epipharynx (Fig. 2B) with setae of middle setal series at
margin equally narrow; lateral setae at margin spatulate, short; distal
sensory papillae clustered in 2 margined groups with 4 papillae in an-

VOLUME 80, NUMBER 2 175

Fig. 1. Adult characters of species of Caccoleptus. A, dorsal aspect of C. wicki; B, fe-
male antenna of C. wicki; C, metathoracic wing of C. wicki; D, male antenna of
C. rotundus; E, thoracic sterna of C. wicki (Cox = coxa, Fem = femur, Msst = meso-
sternum, Prost = prosternum, Tr = trochanter).

terior and 2 papillae in posterior group; cluster not enclosed by callosity;
6 sensory cups in proximal row. Maxilla as illustrated (Fig. 2A); terminal
segment of palp relatively broad. Labial palp with single seta inserted on
ventral side of segment 1, none on segments 2 or 3; 2-3 ensiform setae in-
serted at apex of each lobe of ligula. Mandible with fringed prostheca and
6-8 lateral setae. Tergum of abdominal segment 1 about % as wide as
metathoracic notum or tergum of abdominal segment 2; antecostal suture
lacking (on all nota and terga as well); no spicisetae inserted on acrotergite
or on tergite anterior to median row of large spicisetae; no spicisetae in-
serted along posterior margin of tergum; hastisetae inserted entire width
of tergum in row posterior to median row of spicisetae (Fig. 2F). No hasti-
setae inserted on membrane behind any abdominal tergum (all inserted
on sclerotized part of tergum). Tergum of abdominal segment 9 bearing

176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Larval characters of C. wicki. A, maxilla, ventral aspect; B, epipharynx and
labral margin; C, pretarsus of mesothoracic leg; D, antenna; E, head of hastiseta from
tergum of abdominal segment 2; F, left half of tergum of abdominal segment 1 show-
ing size relative to widths of, metathoracic notum and tergum of abdominal segment 2
(circles represent sockets for insertion of spicisetae, size of the socket roughly pro-
portional to size of seta; dots represent points of insertion of hastisetae).

VOLUME 80, NUMBER 2 ia

long spicisetae forming terminal brush. Pretarsus as illustrated (Fig. 2C);
anterior terminal seta 2x as long as posterior.

Pupa.—Abdominal segments lacking gin-traps.

Holotype 2 and 13 2 paratypes——El Zulia (10 kilometers northwest of
Cuicuta), Santander del Norte, Colombia, 2 July 1976 (A. Lopez).

Other paratypes—1?, Botanical Gardens, Georgetown, British Guiana,
26 September 1918 (Harold Morrison); 12, Barro Colorado Island, Canal
Zone, Panama, April-May, 1942. Holotype deposited in the collection of
the U.S. National Museum of Natural History. Paratypes deposited in
the collections of the U.S. National Museum of Natural History, the
British Museum (Natural History), the Natural History Museum, Los
Angeles County, the California Academy of Sciences, and the collection of
the author.

The color of the integument and setal characters given in the key readily
separate adults of this species from the two previously described species
in the genus. In addition, the single male of C. rotundus available for
study has unusually prominent compound eyes. Whether this is a sexual
character common to males of all species (assuming there are males of
C. anisotomoides and C. wicki) or a distinguishing character of C. rotundus
is not known.

At the present, no males are known for C. wicki or C. anisotomoides.
This seems a little unusual, since a moderate series of C. wicki has been
collected. It suggests the possibility that C. wicki may reproduce partheno-
genetically, a trait found in the somewhat closely related genus Reesa.

Etymology.—This species is named for Dr. J. R. Wick, Chairman of the
Department of Biological Sciences, Northern Arizona University, in recog-
nition of his faithful and effective leadership in the field of biological
education.

Acknowledgments

I wish to thank F. Lance Wallace for his assistance in interpreting the
wing venation of C. wicki. Iam especially grateful to Dr. John M. Kingsolver,
Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. Educ. Admin.,
USDA, for arranging the loan of the specimens used in this study and for a
critical reading of the manuscript. I thank Dr. Jose Cuatrecasas for help
with the manuscript.

Sumario

Adultos y larvas de una especie nueva de los alrededores de Cucuta, Co-
lombia, llamada Caccoleptus wicki, se han encontrado atacando huevos y
larvas de Opsiphanes cassina F. en la primera etapa de su desarrollo post-em-
brionario. Como es sabido este es un en la defoliacién del bananero. La
especie se diferencia de C. rotundus y C. anisotomoides por la presencia

178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

en el integumento del élitro de franjas submedianas y subapicales ligera-
mente coloreadas. En las otras dos especies de Caccoleptus, las franjas del
élitro estan formadas por pelos ligeramente coloreados pero el integumento
permanece inmaculado. La larva de C. wicki se puede diferenciar de la
larva de otro género de Megatomini por la anchura reducida del primer
tergo abdominal y por la alargada papila complementaria del segundo
segmento de la antena.

Literature Cited

Beal, R. S., Jr. 1967. <A revisionary study of the North American dermestid beetles
formerly included in the genus Perimegatoma (Coleoptera). Misc. Pub. Entomol.
Soc. Amer. 5(6):279-312.

Ford, E. J., and J. M. Kingsolver. 1966. Description and biological notes on the
larvae of Phradonoma tricolor (Coleoptera: Dermestidae). Coleopterists’ Bull.
20(1):27-29.

Sharp, D. 1902. Fam. Dermestidae. In Biologia Centrali-Americana, Coleoptera
2(1):642-669.

Wallace, F. L., and R. C. Fox. 1975. A comparative morphological study of the hind
wing venation of the order Coleoptera, Part I. Proc. Entomol. Soc. Wash.
77(3):329-354.

Zhantiev, R. D. 1976. The Hide Beetles of the Fauna of the U.S.S.R. [in Russian].
Moscow University, pp. 1-182.

Northern Arizona University, Flagstaff, Arizona 86011.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 179-190
ETHOLOGY OF CEROTAINIA ALBIPILOSA CURRAN
(DIPTERA: ASILIDAE) IN MARYLAND:
COURTSHIP, MATING AND OVIPOSITION?!

Aubrey G. Scarbrough

Abstract—Male Cerotainia albipilosa (Curran) exhibit a complex aerial
courtship in front of perched females. The position of the male in flight
and subsequent behaviors are discussed. Most courtship flights by males
fail to terminate in mating. Males mount females following a rear or frontal
flight approach. Non-receptive females display agonistically toward court-
ing males. The mating position consists of male over female and lasts for
an average 11.5 minutes. Unlike many asilid species, mating pairs do not
fly in copulo. Females initiate separation of mating pairs. Most courtship
flights and matings occur between 2:00 and 5:00 PM. The egg chorion is
bright amber in color and oval in shape; one averaging 0.25 and 0.29 mm
in width and length, respectively. Elevated ridges occur on the surface of
the chorion except in the immediate area surrounding the micropyle.

Several workers have contributed significant information on the basic
reproductive habits of asilid flies (Poulton, 1906; Melin, 1923; Richards,
1929; Bromley, 1933; Wilcox and Martin, 1936; Hull, 1942; Dennis and
Lavigne, 1975, 1976; Lavallee, 1970; Lavigne, 1963, 1964, 1968, 1970a,
1970b, 1971, 1972; Lavigne and Dennis, 1975; Lavigne and Holland, 1969;
Lavigne et al., 1976; Lehr, 1970; Musso, 1972; Alcock, 1977). Many of the
asilid species reported in these studies displayed specific patterns of pre-
mating behavior. For many species, premating behavior consisted of short
rapid flights in which males searched for receptive females. Matings, fol-
lowing these flights, were initiated either in the air or on substrates. Those
initiated in the air began when males overtook females in flight. Follow-
ing a brief struggle, the pair fell to the substrate where copulation ensued.
Those initiated on substrates began when a male landed on a perched fe-
male nearby. In the former, the male landed on the female’s dorsum, faced
the same direction as the female, clasped her genitalia, and assumed a
copulation position. In the latter, the male quickly mounted the perched
female from the side or rear. In both examples, a brief struggle some-
times preceded genital contact.

Courtship behavior has been observed in some species of at least eleven
asilid genera (Heteropogon, Promachus, Diogmites, Stichopogon, Holo-
pogon, Dioctria, Stelidopogon, Cyrtopogon, Proctacanthella, Proctacanthus,
Ablautus). Courtship behavior involved a hover flight in most species, al-
though leg extension was sometimes included when the male approached

180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

a potential mate. Males of five species (Cyrtopogon glarealis Melander, C.
auratus Cole, C. montanus Loew, C. marginalis Loew and Ablautus rufo-
tibialis Back) performed courtship behavior, which also included leg ex-
tension, on substrates in front of perched females. Matings were initiated
when males landed on the female’s dorsum or mounted them from sub-
strates.

Mating positions are usually ‘tail to tail’ or ‘male over female’ although
some species may show slight modifications of either position.

Oviposition habits of females are variable, depending on specializations
of their genitalia. In many species, the posterior segments of the abdomen
are modified into slender structures for inserting eggs between and below
substrates; in others a circlet of spines is present on the 9th abdominal
tergite and is used for digging and covering eggs with soil. Still in other
species, neither specialization is present; and presumably they drop their
eggs in flight or place them on surfaces or inside fissures in plants or decay-
ing wood.

Methods and Procedures

Extensive field observations of Cerotainia albipilosa Curran were made
at the Loch Raven Watershed in Baltimore County, Maryland. General
methods and procedures for observations made in the field were described
earlier (Scarbrough and Norden, 1977). Field observations were recorded on
tape and later transcribed into a notebook. Photographic records of various
behaviors were taken whenever possible. A Yashica movie camera with
a zoom lens was used to record complex behaviors such as courtship
oscillations, attempted mountings and agonistic displays. The movies were
analyzed to obtain a more accurate understanding of these behaviors.

Results and Discussion

Courtship and mating.—Courtship flights performed by Cerotainia al-
bipilosa males involved aerial displays conducted in front of females resting
on sunlit perches. Most courtship flights occurred in mid- to late afternoon
(Scarbrough and Norden, 1977). At this time females selected perches
which ranged from 5 cm to 6 m above ground. However most females
(85.1%, N = 2,151) were concentrated on perches situated below 3 m.

=

Figs. 1-4. 1. Typical positions of a male and female resting on a perch prior to
a courtship flight. The female is at the tip of the leaf; 2. Typical posture of a non-recep-
tive female following an attempted mounting by a male. Note the positions of the
wings, hind legs and arched abdomen; 3. Typical mating position of Cerotainia
albipilosa; 4. Egg of Cerotainia albipilosa (100).

181

VOLUME 80, NUMBER 2

182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Males rested on and launched courtships from various plant perches lo-
cated 1-6 cm behind and 1-2 cm above or below perched females. A male
sometimes occupied the same perch on which the female rested (Fig. 1).

While enroute to the courtship position, the male usually flew several
cm above or below the perched female to a position about 7-10 cm in
front of her. Upon arriving at the latter position, he rotated his body 180°
while hovering and faced the female. When the male’s flight path was
to one side of and at the level of the female, she invariably flew toward
him as if he were potential prey, but she turned away before making con-
tact with him. Both sexes usually returned to their perches before the male
resumed courtship flights.

In the courtship position, male C. albipilosa employed hover flights with
leg extensions when approaching a perched female. From this hover posi-
tion, males oscillated slowly forward and rearward 3-7 cm in front of
females before attempting to mount them. During the forward phase, the
male’s fore legs were elevated to the sides of its head and extended toward
the female. The midlegs were directed below the body, and the hind legs
extended ventrolaterally and formed an angle of 45-60° from the body. The
male hesitated in the forward phase about 3 cm in front of the female and
palpated his front tarsi. During the rearward phase, his fore legs dropped
below his eyes, and mid- and hind legs remained in one of the previously
described positions. The duration of an oscillation averaged 4 s but ranged
from 1-12. The average number of oscillations per courtship flight was 5
but ranged from 1-9. Males usually rested on perches near females between
courtship flights.

Frequently during an oscillation, a male quickly retreated backward to
a position about 15 cm in front of the perched female and then darted
forward touching the female’s head, thorax and/or wing bases. However,
in some cases the male stopped his forward progress just prior to reaching
the female only to retreat again. He sometimes did this 2 or 3 times before
touching the female. The male’s mid- and hind legs were drawn up below
his body when he touched the female with the extended fore tarsi. Darting
flights were usually preceded by 2 or more forward oscillations and oc-
curred prior to the male’s attempt to mount a female.

During courtship, males of Heteropogon spp. (Lavigne and Holland,
1969; Lavigne, 1970b; Lehr, 1970; Alcock, 1977), Cyrtopogon spp. (Wilcox
and Martin, 1936; Lavigne, 1970a; Lavallee, 1970), Holopogon albipilosa
Curran (Dennis and Lavigne, 1975), Ablautus rufotibialis (Lavigne, 1972)
and Stichopogon trifasciatus (Say) (Lavigne and Holland, 1969) approach
prospective mates with leg extension in a similar manner as that performed
by male Cerotainia albipilosa. With the exception of C. willistoni (Curran),
males of the above species also move forward and touch females with their
fore tarsi. The tarsi and tibiae of all legs of C. albipilosa are covered with

VOLUME 80, NUMBER 2 183

silvery-white hairs. Parts of the fore tarsi of males of the above species, with
the exceptions of Holopogon albipilosa and Stichopogon trifasciatus, are
ornamented with white hairs. Dorsal patches of white hairs are also on
the middle tibiae of Heteropogon maculinervis James; and all legs of
Stichopogon trifasciatus are pollinose, with short appressed silvery hairs.
The position and movement of the ornamented fore and midlegs and subse-
quent contact of the female with the male’s fore tarsi undoubtedly serve
as visual and tactile stimuli which convey conspecific information to fe-
males (Lavallee, 1970; Lavigne and Holland, 1969; Alcock, 1977).

Males of Heteropogon lautus Loew (Bromley, 1933) and Stichopogon
trifasciatus (Lavigne and Holland, 1969) include a lateral movement dur-
ing the courtship hover. Soon after assuming a courtship position or fol-
lowing a rearward phase of an oscillation, Cerotainia albipilosa males
hovered for about 1-5 s before resuming forward movements. At this stage
they sometimes moved 1-2 cm to the right or left of the females. These
lateral movements appeared to be primarily associated with sudden changes
in wind speeds and shifts of their bodies produced by normal wing move-
ments. Lateral movements were more apparent on windy than on calm
days.

Non-receptive females of C. albipilosa, like Heteropogon stonei Wilcox
(Alcock, 1977), responded to oscillating males by moving parts of their bodies
singly or in combination and/or flying after males. As a male moved
toward a female, she spread her wings laterally so that the wings formed
an angle to her body of 50-60°; she frequently vibrated her wings and
tilted her body forward. Her abdomen was sometimes vibrated up and
down, although it was usually elevated and held in a stationary position
above the perch at an angle of 30-40°. She sometimes extended her fore
legs and ‘waved’ her tarsi in a boxing motion toward the male. When a
male touched a female’s dorsum, the female rocked forward, vibrated her
wings and lowered her abdomen. If a male attempted to mount, a female ele-
vated her wings and hind legs above her body and curved the tip of
her abdomen down against the perch. She then vibrated her wings and
kicked her hind legs at the male (Fig. 2). Females responded to lateral
movements of males by pivoting on their midtarsi and shifting their bodies
to face them. Frequently females flew from these crouched or tilted
positions to chase males or to forage for prey. Females forage when they
are non-receptive (Scarbrough, 1978). It is assumed that these movements
and subsequent postures of Cerotainia albipilosa females provided the
primary means of communicating non-receptivity to males and, at the
same time, placed them in the normal position from which to fly.

Males usually courted a single female for 20 or more minutes, but
periods ranged from 0-204. Zero periods represent those instances when
searching males were immediately chased away upon arriving at the site

184 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

by non-receptive females or other courting males. During this period a
male often courted a female several times (f = 6/5 min interval; R = 1-10)
with brief interruptions for attempted mountings and perchings. While
courtship flights took place during most of the diurnal activity period, most
flights occurred from 2:00-5:00 PM. Air temperatures at courtship heights
ranged from 22-34°C.

Searching C. albipilosa males sometimes appeared to court other Diptera
and the ends of twigs. Other asilid species (Holcocephala abdominalis (Say),
H. clava (Loew), Holopogon phaenotus (Loew)), syrphids (Allograpta spp.)
and tephritids (Rhagoletis sp.) were abundant, occupied perches during the
day near female Cerotainia albipilosa, and were courted frequently by
males. Larger flies, such as Musca domestica L. (Muscidae) and Sarcophaga
spp. (Sarcophagidae), were less abundant and were courted less frequently.
Asilids and syrphids were alarmed by this attention, and males usually
remained with and courted them from 1-2 min. The remaining dipterans
were easily disturbed and flew away almost immediately. No mounting
attempts were observed. Additionally, males frequently courted perched
males of the same species, and on occasion, the ends of dead twigs. These
courtships were brief, usually consisting of one or two oscillations. Thus it
is assumed that searching males cannot recognize conspecifics until the
proper responses are performed.

Most courtship flights by male Cerotainia albipilosa failed to terminate
in matings. This is true of several asilid species exhibiting aerial displays
(Lavigne and Holland, 1969; Lavigne, 1970a, 1970b; Dennis and Lavigne,
1975; Lehr, 1970). During over 400 hours of observing courting males,
only 38 successful mountings and subsequent matings were observed.
Mountings of females were accomplished from a frontal or rear approach.
From a frontal approach, a male flew to the usual courting position in
front of a perched female. He then flew directly to a position about 2 cm
above her; and while hovering, he rotated his body 180° to be in line with
hers. Upon landing, the male grasped her wings and abdomen with his
legs. The end of his abdomen looped downward, between his legs and over
the end of her abdomen. The male’s claspers grasped the female’s genitalia
and union was accomplished. From a rear approach, a male flew directly
from its perch behind the female, landed on her dorsum and grasped her
in the above manner. Prior to both approaches, males courted females
several times before attempting to mount them. When males were un-
successful in mounting, they flew to perches behind females before re-
suming courtship flights.

Because of the limited number of complete matings (38) observed and
the speed at which the observed ones occurred, it was not possible to
ascertain what signal(s) was (were) emitted by the female to indicate ac-
ceptance. The only detectable response produced by females was the

VOLUME 80, NUMBER 2 185

elevation of the ends of their abdomens prior to union of the genitalia of
the sexes. Perhaps receptivity is signaled indirectly by the absence of
agonistic behavior toward males attempting to mount. Females which
were totally non-receptive flew after and chased males away as soon as
they assumed a courtship position. In other cases, females tolerated courting
males for several minutes before chasing them away. One male oscillated
37 times in front of a female during a 15 min period (1:03-1:18 PM), while
touching her only 6 times. The female appeared to ignore the male during
much of the courting period (23 min), although sometimes she spread her
wings and vibrated them and her abdomen when he approached her. She
also foraged 3 times and fed on one prey during the courtship. The
female chased him 4 times before he departed the area. In contrast, an-
other male oscillated 42 times during a 15 min period (2:12-2:27 PM) while
darting forward and touching the female 28 times. The female also foraged
5 times and fed on 2 prey during this period but displayed her wings only
once and did not chase the male. The pair mated at 2:33 PM. Thus it
seems plausible that non-receptivity in females exists at various levels and
may be influenced with appropriate internal (satiation) and external (court-
ing males) stimulation.

While mating pairs were observed from 12:06-5:10 PM, the majority
(88%) took place between 2:00 and 5:00 PM. Most pairs were found on
sunlit upper surfaces of perches at heights ranging from 30 cm up to about
7 m above ground. When temperatures exceeded 34°C pairs moved to the
shaded margins of or below leaves. Air temperatures ranged from 25-
38°C at heights where mating pairs were observed.

Males usually attempted to mount females from a frontal approach dur-
ing courtship. Most of the successful (73.6%; N = 38) and unsuccessful

(70.9%; N = 2,261) mountings were initiated following oscillations in
front of females. The remaining successful (26.4%; N = 38) and unsuccessful
(29.1%; = 2,261) rear mountings were directed from perches and ex-

cluded an oscillatory or hover flight. In each case, the male courted the
female for several minutes prior to perching and subsequent mounting or
attempted mounting. Seven of the successful and 223 of the unsuccessful
rear mountings occurred immediately after females returned to perches
following various flight activities. Males which had been courting females
followed them in flight and mounted or attempted to mount them as
soon as they landed, without assuming a courtship position. If females were
receptive and received appropriate stimulation, they allowed males to
mount from either frontal or rear approach. However, the position assumed
by courting males in front of a female probably functioned as an orienta-
tion mechanism for the sexes. Thus males were in an effective position
from which to recognize conspecifics, to convey species specific information
and to mount females.

186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Male courtship flights and female flight activities frequently attracted
other males to the female’s perch. It was not unusual to find 2 or 3
males simultaneously attempting to court a single female. This situation
occurred several times when a single male courted a female for long
periods (>25 min). These situations were brief since the resident male
soon began to attack the intruders. The attacks consisted of two males meet-
ing in midair and flying in a tight downward directed spiral. The flights
usually terminated a few cm above the ground although a few pairs crashed
into shrubs or high grass. Resident males sometimes rammed or grappled
with intruders in air while falling to the ground. Physical damage to the
flies was not detected. The males also produced a high-pitched, audible
buzz during these encounters. The resident male usually remained in
the area and courted the female.

Most mating pairs (95.9%; N = 222) of C. albipilosa took the ‘male over
female’ position (Fig. 3), although a few pairs (4.1%; N = 222) were found
clinging to margins of leaves or twigs in a chain position. In the latter posi-
tion, both sexes were holding onto the perch, and their bodies formed
variable angles, ranging from 55-95°.

Mating pairs remained at the original female perch sites until the sexes
separated. The average duration of 38 complete matings was 11.5 minutes
(R = 10-15). Partial mating durations of 184 pairs ranged from 5-18
minutes. Females usually initiated separation by slowly pushing alternately
with their hind tarsi at the mounted males. Continued pushing eventually
dislodged males from their dorsal positions. Still connected by their gen-
italia to females, males fell backward and assumed a head down position
with their dorsal surfaces facing the same direction as the female’s head.
In this position females began to vigorously kick at males with both tarsi,
simultaneously. The males responded by either immediately separating
from the females in the suspended position or crawling to a position on
a perch beside the females before separating. Times of normal separa-
tions (female leg kicking and genital disengagement) ranged from 6-20
seconds. In some cases males released females without the usual tarsal prob-
ing. Approach of a predator (Vespula sp.) was also sufficient to stimulate
males to loosen their grasps of females and fly away.

Upon separation, members of a mating pair reinitiated behaviors which
were interrupted by mating. Males usually flew from the area, although
some remained and courted the same female or others nearby. Females
sometimes flew to another perch, but they usually remained at the mating
site where they foraged and fed and/or were courted again.

Females sometimes mated several times during the flight season. Marked
females were observed at 15 min intervals/h from 2:00-5:00 PM during
a 7 day period (11-18.VIII.76). These observations showed that of 40

VOLUME 80, NUMBER 2 187

marked females, 20% mated each day, 10% mated more than once per
day and 5% were not observed mating. In one case a female mated with
the same male twice. The male began to court the female immediately
after the first meeting. He courted her for 16 min before mating the second
time.

Unlike many asilid species, Cerotainia albipilosa does not fly while in
copulo. Mating pairs rested on fully exposed sunlit perches in an almost
motionless state. Movement was usually restricted to females’ maintain-
ing their positions on perches, discouraging intruders or probing at mounted
males near the end of mating. Intruder males usually hovered in the court-
ing position in front of mating pairs. Females responded by extending their
fore legs and slowly waving their tarsi at the approaching intruders. The
intruders then either flew away or landed near mating pairs on perches
where they probed the tips of their abdomens toward the female's genitalia
and apparently attempted to mate with them. Mating females responded
by kicking at them with their hind legs and/or crawling to new positions on
perches. Kicking by females and the inability of males to clasp their gen-
italia was usually sufficient to discourage the intruders. Continued probing
by one or more intruding males sometimes induced separation of the mating
pair. Similar reactions were elicited when a pair was touched with a pencil.
Females waved or kicked their legs at the probing pencil and then crawled
away. When captured in vials, mating pairs did not separate until they were
vigorously shaken, and females moved only to maintain a balanced position.
Mating males usually maintained a thanatosis-like state until they released
their grasp on females. Mating males occasionally elevated their wings
and vibrated them when exposed to these stimuli. This response appeared
to be more a function of maintaining balance than an agonistic display.

According to some investigators (Cloudsley-Thompson, 1961; Poulton,
1906; Hobby, 1931; Musso, 1971), some asilid males may present potential
mates with food before mating or court only feeding females. In each case
females were momentarily engaged in feeding which permitted a period
for males to mate with females and, at the same time, to escape predation
from the latter. Evidence to support this suggestion is lacking (Dennis and
Lavigne, 1975). Behavior patterns of C. albipilosa do not support this sug-
gestion since 1) males court without prey (Scarbrough, 1978), 2) males
court both non-feeding and feeding females (only 40% (N = 596) of the
courted females were feeding), 3) only 7% (N = 212) of the mating females
were feeding and 4) cannibalism was not observed.

Eggs and oviposition.—Attempts to obtain eggs in the field and from
live or decapitated adults placed in vials for oviposition were unsuccessful.
Eggs were obtained by dissecting live females in Ringer’s Insect Solution.
Females to be dissected were taken at random throughout the flight season.

188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Eggs found in calyces, oviducts and connecting genital chambers were
considered to be mature since their chorions were sclerotized and the
darkest in color. The number of developed eggs obtained from 30 females
ranged from 52-148 with most flies (17) having over 100. In addition, each
ovary consisted of about 20 ovarioles which contained 5-8 developing
eggs. Six females captured in copulo early in the season (5-10.VII.73) and
dissected lacked mature eggs, but the ovarioles contained numerous oocytes.
Although the number of eggs produced by a single female was not de-
termined, these results indicate that 1) eggs are produced continuously
during the female’s life, 2) eggs are not mature and ready for fertilization
when females emerge, but require a period for maturation and 3) females
may mate before eggs are mature.

Sclerotized chorions of mature eggs were uniform in color, shape and
size. They were bright amber, slightly oval and averaged 0.25 mm in
width and 0.29 mm in length (100 eggs, 10 from 1022) (Fig. 4). The
chorion had characteristic elevated ridges forming 4-7-sided rings with
6-sided rings being the most common. Melin (1923) reported similar
structures on eggs of Laphria spp. and Dioctria spp. At one end, the ridges
faded away to form a smooth surface which surrounded the micropyle.
Other microstructures associated with insect chorions were not visible
at 550x.

Oviposition habits of Cerotainia albipilosa females are unknown. Only
one oviposition was observed during this study. A perched female (13.
VII.74; 3:41 PM) lowered its abdomen, touched the tip to the perch and
released 3 eggs consecutively. She groomed the tip of the abdomen with
her hind legs and flew away. Air temperature at the oviposition site was
31°C. Other females (12) behaved similarly by touching the tips of their
abdomens to perches and grooming their abdomens, but eggs were not
located. The female’s ovipositor is minute and lacks specialization for insert-
ing eggs into substrates. Females of Dioctria, Laphria (Melin, 1923) and
Andrenosoma (Musso, 1971), which also lack genital specializations, typically
deposit eggs into holes or grooves on substrates or while in flight. Thus
Cerotainia albipilosa females probably behave similarly by depositing eggs
at perches or at specific sites among vegetation in the area where they
forage and mate.

Acknowledgments

I wish to thank the two reviewers, Dr. Robert Shoemaker (Depart-
ment of Biology, TSU) and Dr. Lloyd Knutson (Systematic Entomology
Laboratory, IIBIH, Fed. Res., Sci. Educ. Admin., USDA) for their critical
review of the manuscript. Dr. Robert J. Lavigne (University of Wyoming)
kindly provided a translated segment of Lehr’s work on the courtship be-
havior of Heteropogon pyrinus Hermann. For their identifications of the

VOLUME 80, NUMBER 2 189

prey listed in Scarbrough, 1978, I wish to thank the research entomologists
and cooperating scientists of the Systematic Entomology Laboratory and
the entomologists in the Department of Entomology, Smithsonian Institu-
tion.

Literature Cited

Alcock, J. 1977. The courtship behavior of Heteropogon stonei (Diptera: Asilidae).
J. Kans. Entomol. Soc. 50: 238-243.
Bromley, S. W. 1933. Courting and mating performances of an asilid fly (Hetero-
pogon lautus). Psyche. 40:144.
Cloudsley-Thompson, J. L. 1961. Animal behavior. The MacMillan Co., New York,
N.Y. 89 pp.
Dennis, D. S., and R. J. Lavigne. 1975. Comparative behavior of Wyoming robber
flies II (Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr. No. 30,
68 pp.
1976. Ethology of Efferia varipes with comments on species coexistence
(Diptera: Asilidae). J. Kans. Entomol. Soc. 49:48-62.
Hobby, B. M. 1931. The relationship between courtship and predaceous habits in
Dioctria rufipes DeG. and D. atricapilla Meig. (Diptera: Asilidae). Proc. Entomol.
Soc. London (Ser. A) 6:74—75.
Hull, F. M. 1942. The mating habits of robber flies (Diptera: Asilidae). Entomol.
News 53:132.
Lavallee, A. G. 1970. Courtship and mating habits of an asilid fly, Cyrtopogon mar-
ginalis. Ann. Entomol. Soc. Am. 63:1199.
Lavigne, R. J. 1963. Notes on the behavior of Stenopogon coyote Bromley with a
description of the eggs. Pan-Pac. Entomol. 39:103-107.
1964. Notes on the description and ethology of Efferia bicaudata (Diptera:
Asilidae), with a description of the eggs. Ann. Entomol. Soc. Am. 57:341-344.
1968. Notes on two species of Asilus (Diptera: Asilidae) associated with
animal burrows, with a redescription of Asilus gilvipes Hine. J. Kans. Entomol.
Soc. 41:334-339.
1970a. Courtship and predatory behavior of Cyrtopogon auratus and C.
glarealis (Diptera: Asilidae). J. Kans. Entomol. Soc. 43:163-171.
1970b. Courtship and predatory behavior of Heteropogon maculinervis (Dip-
tera: Asilidae). J. Kans. Entomol. Soc. 43:270-273.
1971. Backomyia seminoensis sp. n. from Wyoming with ethological notes
on B. limpidipennis. J. Kans. Entomol. Soc. 44:337—342.
1972. Ethology of Ablautus rufotibialis on the Pawnee Grasslands IBP Site.
J. Kans. Entomol. Soc: 45:271-274.
Lavigne, R. J., and D. S. Dennis. 1975. Ethology of Efferia frewingi (Diptera:
Asilidae). Ann. Entomol. Soc. Am. 68:992-996.
Lavigne, R. J., and F. R. Holland. 1969. Comparative ethology of eleven species
of Wyoming robber flies (Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn.
Sci. Monogr. No. 18, 61 pp.
Lavigne, R. J., L. Rogers, and F. Lavigne. 1976. Ethology of Efferia benedicti
(Diptera: Asilidae) in Wyoming. Proc. Entomol. Soc. Wash. 78:145—153.
Lehr, P. A. 1970. Robber fly species Heteropogon Loew and Anisopogon Loew
(Diptera: Asilidae) of Eurasia. (In Russian). Biology and Geography, Kazakh
State Univ., Alma-Ata 6:69-79.
Melin, D. 1923. Contributions to the knowledge of the biology, metamorphosis and

190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

distribution of the Swedish asilids in relation to the whole family of asilids.
Zool. Bidr. Uppsala 8:1-317.

Musso, J. J. 1971. Etude préliminaire sur les activités journaliéres d’une population
d’Andrenosoma bayardi Séguy. Bull. Soc. Entomol. Fr. 76:175—-182.

——. 1972. Etude des migrations journaliéres d’Andrenosoma bayardi Séguy
(Diptera: Asilidae). Bull. Soc. Zool. Fr. 97:45-53.

Poulton, E. B. 1906. Predaceous insects and their prey. Trans. Entomol. Soc.
London 1906:323—409.

Richards, O. W. 1927. Sexual selection and allied problems in the insects. Biol. Rev.
Cambridge 2:298-364.

Scarbrough, A. G. 1978. Ethology of Cerotainia albipilosa Curran (Diptera: Asilidae)
in Maryland: Predatory behavior. Proc. Entomol. Soc. Wash. 80(1):113—127.

Scarbrough, A. G., and A. Norden. 1977. Ethology of Cerotainia albipilosa Curran
(Diptera: Asilidae) in Maryland: Diurnal activity rhythm and seasonal dis-
tribution. Proc. Entomol. Soc. Wash. 79(4):538-554.

Wilcox, J., and C. H. Martin. 1936. A review of the genus Cyrtopogon Loew in
North America (Diptera: Asilidae). Entomol. Am. 6:1—95.

Department of Biology, Towson State University, Baltimore, Maryland
21204.
Footnote

* Portions of this work were supported by funds obtained from the Towson State
University Faculty Research Committee.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 191-196
A NEW SPECIES OF XENOCALIGONELLID MITE FROM
THE GALAPAGOS ISLANDS (ACARI)

Roberto H. Gonzalez

Abstract—A new family of prostigmatid mites, the Xenocaligonellidae
is erected to include the bizarre genus Xenocaligonellus De Leon 1959,
which was formerly placed under the Caligonellidae. A new species, X.
galapagus, is described from the Galapagos Islands off the coast of Ecuador.

De Leon (1959) erected the genus Xenocaligonellidus for X. ovaerialis, a
bizarre species from southern Florida and western Mexico. A major cali-
gonellid character, namely the peritremata entering the stylophore, was
taken into consideration for placing this genus in the family Caligonellidae
Grandjean, as defined by Summers and Schlinger (1955). However, De
Leon, recognizing other major characters which deviate from the family
characteristics, created the subfamily Xenocaligonellinae to account for
these non true caligonellid characters.

As Xenocaligonellidus shares only one major character with the true
caligonellids whilst having a number of clear cut differences, a new family
status is proposed for these odd looking mites. Reasons supporting this ac-
tion are ample, viz: Xenocaligonellidus lacks an important raphignathoid
character—the palptibial claw. In addition, non caligonellid characters
include: Coxae contiguous, not separated in groups I-II and III-IV; anal
slit distinctly ventral and contiguous with the genital opening; and em-
podium with a double row of multiple capitate hairs. The general body
shape and the outstanding development of most of the dorsal setae are
further characters which make it possible to define a new family, the
Xenocaligonellidae.

From a phenotypic standpoint, xenocaligonellid mites are remarkably
different than all known species in the families of Raphignathoidea. The
arrangement and quality of dorsal setation, the absence of the paraaxial
claw on palptibiae, the lack of transverse fisures on dorsal idiosoma, and
the tetranychid kind of empodial arrangement make those mites to be an
extremely marginal raphignathoid group. The placement of the new family
under this group has reluctantly been made as there is no other superfamily
providing for such assorted combination of distinct characters. On the other
hand, the distinctly looped peritrematal arrangement would suggest that
this family is an offshoot of the Caligonellidae.

Xenocaligonellidae, new family status

Small, flattened mites, with cheliceral bases fused into a stylophore bear-
ing a chambered, loop-shaped peritremata. Gnathosoma exposed from

192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

above. Palpi 5-joined, tibia lacking distal claw, palptarsus longer than
tibia, with a set of stubby distal setae. One pair of flagellate ventral setae
on subcapitulum. Dorsal setae on idiosoma whiplike, some of them longer
than body; other setae acicular, distally spinose. Eyes, two pairs. Coxae
II-III contiguous. Anal slit and genital opening contiguous on the ventral
side. Legs short, robust with a blade shaped empodia bearing two rows of
multiple capitate hairs.

As De Leon stated for X. ovaerialis, males and nymphs resemble the fe-
male except for the reduction in dorsal setae numbers. Larvae have fewer
pairs of dorsal setae, and the caudalmost pair of anal setae are much
longer than other dorsals.

The habits of this family are known for X. ovaerialis only. De Leon re-
ported this species to occur on Quercus, Persea, Celtis, Inga and several
unidentified plants. An interesting characteristic of Xenocaligonellidus is
the egg-laying habit. They are attached to the distal end of an erect stalk, a
feature known in insects apparently to avoid egg predation. With the find-
ing of a second species from the neotropical region, the generic charac-
teristics have been better understood and amended accordingly.

Xenocaligonellidus De Leon 1959, emend.

Description—lIdiosoma circular, flattened, with smooth integument;
stylophore broadly triangular, fused along the midline except at the distal
%; peritremata 2-shaped, without a distinct distal opening. Movable digits
styletlike. Fixed digits finely pointed. No terminal claw on _palptibia.
Palptarsus longer than palptibia with a crown of stubby, straight setae.
Two pairs of eyes. One single pair of ventral subcapitular setae. Fifteen
pairs of dorsal setae distributed as follows: 5 pairs of propodosomal setae;
10 pairs of hysterosomals. Coxae contiguous, longer than respective femora,
without coxal plates. Legs short, with genua and tibia of about equal size.
Ambulacra with strong claws and a bladelike empodia bearing 2 rows of
multiple, capitate hairs. Anal slit contiguous to the ventral vaginal open-
ing, guarded posteriorly by 2 pairs of thick, barbed, long setae.

Type-species.—Xenocaligonellidus ovaerialis De Leon 1959, by original
designation.

Terminology.—For descriptive purposes, idiosomal dorsal setae are sep-
arated into propodosomals and hysterosomals. There are 5 pairs of whip-
like propodosomal setae, the posteriormost pair termed central pro-
podosomal (cp). The hysterosomals include 1 pair of humerals (h), 2 pairs
of laterals (L;, Lz), 2 pairs of sublaterals (S;, S2) and 5 pairs of dorsocentrals
(dc) (Fig. 1). The homology of L; to the acicular setae of the dorsocentral
series and that of Lz to the sublateral pairs suggest that these lateral setae,
apart from their marginal position, share no functional characters in common.

VOLUME 80, NUMBER 2 193

Fig. 1. Xenocaligonellus galapagus, dorsum of female. Setae: cp central pro-
podosomals; h humerals; s:—s2 sublaterals; —l. laterals; dc dorsocentrals.

All measurements are given in microns; length of dorsal whiplike setae
are rounded to the nearest 5 microns.
Xenocaligonellidus galapagus Gonzalez, new species

A species with circular, flattened, idiosoma and a combination of acicular
and whiplike dorsal setae; posterolateral setae L. the longest of dorsal series,

194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 2-6. Female. 2. Leg I; 3. Dorsal acicular seta; 4. Tarsus leg II; 5. Ventral aspect;
6. Stylophore with looped peritremes, and palpi.

VOLUME 80, NUMBER 2 195

nearly 2.4x longer than diameter of body. Posteriormost 2 pairs of anal
setae, acicular, stub.

Female.—lIdiosoma 260 in diameter, leg I, 170; stylophore broadly tri-
angular 132 x 95, deeply notched at the front end. Stylets needle-like, fixed
digits finely pointed. Peritremes of caligonellid type, extend to the distal
% of stylophore, loop made up of 14 cells (Fig. 6). Number of setae on palpal
segments as follows: Femur 2; genu 1; tibia 3; tarsus 4, in addition to a
proximal bulb-shaped sensory rod and 4 stubby terminal sensory setae.
Dorsal aspect of idiosoma: 15 pairs of setae borne on strong tubercles. Five
pairs of propodosomals, anteriormost pair 222, 2nd pair 360, preoculars
270, postoculars 395, central propodosomals (cp) 425. Ten pairs of hystero-
somals of which 5 pairs are whiplike, L, being the longest of dorsal series,
660; all dorsocentrals but pair 4 and the lateral L, are acicular, distally
spinose, ranging from 90-95 each (Fig. 3).

Number of setae and special sensoria (in parenthesis) on leg segments
I through IV: Coxae 2-1-1-1; trochantera 1-1-1-1; femora 3-3-1-1; genua
3(1)-3-1-1, tibiae 4(1)-4-3-3, tarsi 11(1)-7(1)-7-7. Seta on trochanter III dis-
tinctly thickened and barbed (Fig. 5). Genu I carries a spinelike solenidion
closely associated with dorsolateral seta; tibia and tarsus I provided with
a well-developed corresponding solenidia 8 and 14 respectively (Fig. 2,
leg I). Anterolateral setae on tibiae III/IV strongly barbed, 65 and 60 re-
spectively (see Fig. 1). Tarsi II to IV provided with 4 branched terminal
setae (Fig. 4, tarsus II).

Ventral side (Fig. 5)—No distinctive plates except for the anal covers. One
pair of long, flagellate, subcapitular setae in the maxillicoxal area, mesal to
a short, spinelike seta behind palpal bases. Coxal areas I-IV contiguous; 2
pairs of anteroventral setae and 1 pair of paragenitals close to genital group.
Three pairs of subequal genital setae and 3 pairs of anals; the last 2 pairs of
anals distinctly needle-like, barbed, and longer than anogenital slit mea-
suring 50 and 88 respectively.

Holotype.—Collected on unknown plant, Wenman Island, Galapagos
Arch., 31 January 1964 (D. Q. Cavagnaro). Deposited in Acarology Collec-
tion, Faculty of Agronomy, University of Chile, Santiago.

Remarks.—The two known species of Xenocaligonellus can be sepa-
rated on the basis of the length of dorsal and ventral setae relative to the
body size and in the quality of anal setae. Xenocaligonellus galapagus
has longer whiplike setae, uniformly long acicular setae and two thick
barbed posteriormost anal setae longer than the anogenital slit. Xenocali-
gonellus ovaerialis has one anal seta of this kind only. In addition, genu
II of galapagus bears three setae instead of two and acicular seta (L,) is
marginal, not sublateral, in position.

196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Resumen

Una nueva familia de acaros prostigmatos, Xenocaligonellidae, se ha
creado para el género Xenocaligonellus De Leon 1959, el cual originalmente
se ubicé en la familia Caligonellidae. Se describe una nueva especie, X. gala-
pagus Gonzalez, colectada en las islas Galapagos en las costas del Ecuador.

Literature Cited

De Leon, D. 1959. A new genus of mites occurring in Florida and Mexico (Acari:
Caligonellidae). Fla. Entomol. 42(1):17-19.

Summers, F. M., and E. I. Schlinger. 1955. Mites of the family Caligonellidae
(Acari). Hilgardia 23(1):539-561.

Plant Production and Protection Division, FAO, United Nations, Rome,
Italy.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 197-209
BIOLOGY OF A NEOTROPICAL SNAIL-KILLING FLY,
SEPEDONEA ISTHMI (DIPTERA: SCIOMYZIDAE)

Lloyd Knutson and Karl Valley

Abstract.—Sepedonea isthmi (Steyskal) is known from the Canal Zone,
Panama, Trinidad, Venezuela, Colombia, Bolivia and Brazil. The species
is found in various fresh-water situations, where the larvae are obligate
predators of pulmonate snails. Females collected in Venezuela in April,
Trinidad in May and Colombia in June laid up to 406 eggs each in the
laboratory. The incubation period was 2-6 days; the first larval stadium
lasted 2-4 days; the second, 3-6; the third, 5-8; and the pupal period was 8-
14 days, at room temperatures. Larvae killed and ate as many as 23
aquatic snails ranging in size from 1-6 mm. Prey species included Bi-
omphalaria glabrata (Say), Drepanotrema (Fossulorbis) depressissimum
Moricand, D. (F.) lucidum Pfeiffer, Helisoma trivolve (Say), Physa venustula
Gould and Lymnaea sp. in the laboratory. The morphology of the immature
stages of the Sepedon Group is discussed; and the egg, all three larval
stages and the puparium of Sepedonea isthmi are described and figured.

The introduction of Sepedomerus macropus (Walker) from Nicaragua
into Hawaii in 1958 and 1959 for control of Lymnaea ollula Gould', the
snail host of the cattle liver fluke, Fasciola gigantica Cobbold (Chock et al.,
1961; Berg, 1973), has shown that it is possible to rear Sciomyzidae in
large quantities and that an exotic species can disperse from release sites
and build up large and persistent populations. Future use of Sciomyzidae
as biological control agents must be based not only on quantitative and
experimental field and laboratory studies of critical aspects such as preda-
tion and competition, e.g. Eckblad and Berg (1972) and Berg (1973), but
also on knowledge of the behavior of many different candidate predator
species. Most snails that are important as intermediate hosts of parasitic
worms of humans and domestic animals are pulmonate, aquatic and amphib-
ious species in tropical and subtropical regions. Sepedon Latreille and re-
lated genera are the dominant Sciomyzidae in tropical regions, and the larvae
of most reared species are highly specialized predators of aquatic snails. Thus
it is of special interest to broaden our knowledge of the basic life cycles of
species belonging to these genera.

The fundamental work on the life histories and morphology of immature
stages of Sepedon and some related genera is the treatment of ten Nearctic,
four Neotropical and two Palearctic species by Neff and Berg (1966). The
larvae of all species discussed in that paper are predators of fresh-water
pulmonate snails. Knutson et al. (1967) described the biology of the first
Sepedon known to be a parasitoid of terrestrial snails in the genus Succinea

198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Draparnaud, and they added two African species to the list of aquatic pred-
ators. Abercrombie (1970) presented extensive biological information on
six species of Sepedonea Steyskal. Abercrombie and Berg (1975) reported on
the biology of Thecomyia limbata (Wiedemann), a Neotropical member of
the Sepedon Group. The present paper treats the biology and immature
stages of a Neotropical, aquatic, predaceous species, Sepedonea isthmi
(Steyskal).

Taxonomy

The genera of Sciomyzidae related to Sepedon have been treated as a
subtribe, tribe or subfamily by various authors. Steyskal (1973) regarded
these genera as forming only a “Group” that is not particularly distinctive
from typical Tetanocerini (subfamily Sciomyzinae). Steyskal included the
following genera in the “Sepedon Group”: Sepedon Latreille, Sepedonella
Verbeke, Sepedoninus Verbeke, Thecomyia Perty, and his new genera
Sepedomerus and Sepedonea (= Sepedon lindneri Group of authors). The
Neotropical genus Sepedonea includes the species discussed herein (Sepe-
donea isthmi), seven other described species (S. barbosai Knutson and
Bredt, S. canabravana Knutson and Bredt, S. guatemalana (Steyskal), S.
guianica (Steyskal), S. lagoa (Steyskal), S. lindneri (Hendel), and S. telson
(Steyskal)), and five undescribed species known to us.

Sepedonea can be distinguished from related genera by using the char-
acters given by Steyskal (1973). Sepedonea isthmi is distinguished from
other Sepedonea by the following features: (1) hind femur with apical dark
marking but without pre-apical marks, (2) middle femur posteroventrally
with 10-12 spinules extending % distance to base, (3) wing with anterior
and posterior crossveins distinctly infumated. The male genitalia also show
important specific differences and indicate that S. isthmi is most closely re-
lated to S. guianica. The species of Sepedonea were catalogued by Knutson
et al. (1976).

Distribution

Sepedonea isthmi has been recorded from the Canal Zone, Panama, Trini-
dad, Venezuela, Colombia, Bolivia and Brazil. Steyskal (1951) described this
species from three specimens from Panama, Canal Zone, Corazal, 1
March 1912, 14 (holotype), No. 60905, USNM, 12 (allotype), and Juan
Mina, 2 September 1923, 14 (paratype), USNM. Also, we have collected or
seen the following specimens: BOLIVIA: Beni: Rurrenabaque, 175 m., 10-
23 October 1956, 1¢6, L. E. Peha, CNC. BRAZIL: Amazonas: Parana da
Cigana, Parintins, November 1969, 14, 12, Exp. Perm. Amaz., USNM.
Pard: Breves, Ilha do Marajé, September 1969, 12, Exp. Perm. Amaz.,
MZUSP. Goids: Rio Preto, 70 km NE Brasilia, 20 km S Formosa, 7 Novem-

VOLUME 80, NUMBER 2 199

LONGITUDE WEST OF GREENWICH 50

DRAINAGE

600 MILES

SOUTH AMERICA
200 400
SANSON'S EQUAL AREA PROJECTION

DISPUTED BOUNDARY
——~ CANAL

COPYRIGHT BY
McKNIGHT & McKNIGHT, BLOOMINGTON, ILLINOIS

McKNIGHT & McKNIGHT, BLOOMINGTON, ILLINOIS

Fig. 1. Collecting sites for Sepedonea isthmi.

200 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ber 1974, 1¢, L. Knutson, USNM. Sdo Paulo: S. Vicente, Parque Bitaru,
29 May 1967, 1é, J. Abercrombie and C. O. Berg, CU. Espirito Santo,
Itaguacu, October 1970, P. C. Elias, 1 damaged specimen, MZUSP. CO-
LOMBIA: Valle: 1.7 km W Cali Puerto, 11 June 1969, 36, 12; 14 June
1969, 22, K. R. Valley, CU. 5 km SE Cali, near Navarro, 11 June 1969, 2¢,
K. R. Valley, CU. 6.5 km SE Cali, Navarro, 11 June 1969, 54, 49, 14 June
1969, 83, 10°, K. R. Valley, CU. Morga, 20 km SE of University, 22-25
June 1964, 12, C. O. Berg, USNM. PANAMA: Panama, La Jagua Hunt
Club, about 32 km ENE Balboa, 1 July 1969, 12, K. R. Valley, CU.
TRINIDAD: Caroni River, 12 October 1918, 12, H. Morrison, USNM.
Princess Margaret Highway, 0.15 km south of junction with Churchill-
Roosevelt Highway, 9 km west of Port of Spain, 4, 5 May 1972, 86, 29,
F. D. Bennett, M. Yaseen, L. Knutson, USNM. VENEZUELA: Carabobo:
Valle Seco, January 1940, 16, P. Anduze, USNM. Valencia, 16 March 1971,
12, C. O. Berg, CU. Aragua: Pto. de Cata, between Caracas and Puerto
Cabello, 17 April 1972, 93, 12, C. J. Rosales and L. Knutson, USNM.
Ocumare de la Costa, 28 km NW Maracay, 14 March 1971, 26, C. O.
Berg, CU. Embalse de Guataparo, 13 April 1972, 1¢, L. Knutson, USNM.
Cojedes: Lago Taguanes, near Tinaquilla, 13 April 1972, 32, L. Knutson,
USNM. La Piedrita, 16 February 1911, 12, S. Brown, ANSP. (ANSP =
Academy of Natural Sciences, Philadelphia, CNC = Canadian National
Collection, CU = Cornell University, MZUSP = Museu de Zoologia, Uni-
versidade de Sao Paulo, USNM = United States National Museum.)

The distribution of S. isthmi is shown in Fig. 1. Sepedonea guianica has
a partially contiguous range.

Biology

Neff and Berg (1966) reared Sepedonea guatemalana in Central America,
and Abercrombie (1970) reared S. lagoa, S. lindneri, S. telson, and three
undescribed species of Sepedonea, all from southeastern Brazil and Argen-
tina. The larvae of all of these species are predators of aquatic snails in
various fresh-water situations.

Habitats of S. isthmi are typical of many Sciomyzidae. Most of the
adults that were used to start our rearings were swept from herbaceous
vegetation emerging from water about 30 cm deep in an extensive
series of roadside borrow pits (Fig. 2) near Port of Spain, Trinidad, on
4 and 5 May 1972. No other Sciomyzidae were collected at that locality,
although larvae, pupae, and adults of Sepedomerus macropus were taken
on 5 May at the edge of nearby Caroni Swamp, San Juan, Trinidad. The
only other Sciomyzidae known from Trinidad are Sepedomerus bipuncticeps
trinidadensis (Steyskal) and an undescribed species of Thecomyia.

In Venezuela, adults of Sepedonea isthmi and Sepedomerus bipuncticeps
(Malloch) were swept from weed-choked, roadside ditches at Puerto

VOLUME 80, NUMBER 2 201

Fig. 2. Habitat of Sepedonea isthmi, fresh-water marsh, six miles west of Port of
Spain, Trinidad, 4 May 1972.

de Cata, between Caracas and Puerto Cabello, on 17 April 1972. Adults of
Sepedonea isthmi, with those of S. guianica and Sepedomerus bipuncticeps,
were collected at the margin of Lago Taguanes, near Tinaquilla, Venezuela,
on 13 April 1972. The single male collected in Goids, Brazil, was found
with Sepedomerus bipuncticeps, Sepedonea barbosai, and S. canabravana.
Adults of S. isthmi were swept from dense, emergent grasses growing in
a wet pasture near Cali, Colombia between 11 and 14 June 1969. Imagines
of S. guianica, Sepedomerus bipuncticeps and Sepedomerus macropus were
usually taken with Sepedonea isthmi in various localities visited in the Cali-
area.

Immature stages were collected in Colombia. Nine puparia were found
floating in shallow water among shoots of emergent vegetation on 14
June near Cali; three produced adult flies within six days and six each
yielded a parasitoid wasp (Trichopria sp., Diapriidae) between 20 June and
3 July. A third-instar larva collected on the same date near Cali pupariated,
and an adult emerged on 29 June.

Adults collected in Trinidad, Colombia and Venezuela were placed by
pairs in small jars (6 X 12 cm) containing a substrate of damp cotton, pro-
visioned with crushed snails and a mixture of powdered milk, honey and
brewer's yeast and covered with mesh cloth. A pair of imagines collected
on 14 June in Colombia were first observed copulating 10 days later; they
mated approximately 30 minutes. Adults collected on 5 May in Trinidad
were first seen mating on 16 May, and they continued to mate infrequently

202 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

until 8 September. Flies collected on 17 April in Venezuela were seen mat-
ing on 7 May. Adults that emerged in the laboratory during August did not
mate during the following 3 months in the laboratory. The copulatory
posture is similar to that of most other Sciomyzidae (male’s front tarsi on
parafrontal area of female’s head, male’s middle tarsi on sides of female’s
thorax, and male’s hind tarsi grasping end of female’s abdomen).

One female collected in Colombia on 11 June deposited 104 eggs during
the following four days; daily egg counts ranged from 26 to 31. A laboratory
reared male and female that emerged on 21 July were placed in a breeding
container the next day, but the three eggs that the female deposited on 6
August were inviable. Four females were collected in Trinidad on 5 May,
and they respectively laid 406 eggs between 16 May and 5 October, 214
between 16 May and 23 August, 30 between 10 May and 6 June and 4 on
12 May. A female collected in Venezuela on 17 April laid 38 eggs between
6 and 25 May. Laboratory reared females from Trinidad did not oviposit.

Eggs were laid side by side in rows of 2-14 on pieces of vegetation or
on the dry walls of the breeding jars, but not on shells of crushed or living
aquatic snails. The eggs gradually darkened, and the incubation period
was 2-6 days at room temperatures (20-22°C).

Larvae killed and ate the aquatic snails Biomphalaria glabrata (Say),
Drepanotrema (Fossulorbis) depressissimum Moricand, D. (F.) lucidum
Pfeiffer, Helisoma trivolve (Say), Physa venustula Gould and Lymnaea sp.
during laboratory rearings. Biomphalaria glabrata and Physa sp. were the
dominant aquatic snails at the collecting sites in Trinidad, and Drepano-
trema depressissimum, D. lucidum, and Physa venustula were the dominant
aquatic snails at the collecting sites in Colombia. None of the larvae fed
in egg masses of Helisoma trivolve, but a few second-instar larvae pene-
trated egg masses of Physa sp. and ate the embryonic snails.

First-instar larvae preyed upon newly hatched and small snails (1-2 mm
in greatest dimension). As the larvae grew in size they attacked larger snails;
mature Helisoma trivolve were frequently killed by large, third-instar larvae.
One larva killed and fed on 22 snails (1-6 mm) during its three stadia;
another consumed parts of 23 prey individuals (1-4.2 mm). During their
5-day third stadia, two larvae killed and consumed, respectively, seven
and eight snails, measuring 4-6.5 mm. The larvae remained in the shells,
consuming relatively fresh tissue, for less than 1 day. They rested under the
wet sand or gravel in the rearing containers, with their posterior spiracles
exposed above the water surface. Larvae lived a relatively long time
without food; three first-instar larvae survived 1-4 days and nine second-
instar larvae lived 6-11 days in starvation chambers. The first stadium
lasted about 2-4 days; the second, 3-6 days; and the third, about 5-8 days.
The larval period ranged from 13-17 days.

Puparia were formed in wet sand, among pebbles of gravel and on the

VOLUME 80, NUMBER 2 203

dry undersides of lids of rearing containers; they are obviously adapted for
flotation (Fig. 17). Eighteen puparia formed between 29 July and 4 August
produced adults 10-13 days later; of 17 puparia formed between 8 and
28 July, 16 produced adults in 8-9 days and one yielded an adult after
14 days. Field collected adults lived up to 78 days in the laboratory.

Morphology of Immature Stages
Sepedon Group

Immature stages of species of the Sepedon Group are among the most
distinctive in the family, but because less than 10% of the immature stages
of acalyptrate Diptera have been described, diagnostic features can be
stated only provisionally. The major source of information on the mor-
phology of the immature stages of Sepedon and related genera is Neff and
Berg’s treatment (1966) of 16 species. Knutson et al. (1967) described the
immature stages of three African species of Sepedon, Nagatomi and Tanaka
(1967) provided a thorough description of the egg of Sepedon sauteri
Hendel, and Abercrombie (1970) characterized the immature stages of six
species of Sepedonea. Abercrombie and Berg (1975) described the develop-
mental stages of Thecomyia limbata.

Although eggs of some species of the Sepedon Group are distinctive,
there is no single character that distinguishes them reliably from other genera
of Sciomyzidae. All described eggs of this group bear a subterminal micro-
pyle, characteristic of all known eggs of the Tetanocerini. Neff and Berg
(1966) stated that the coarse longitudinal striations on eggs of reared
Sepedon are rather distinctive for this genus. The patterns of chorionic
sculpturing, color and dimensions have been useful characters for separat-
ing eggs of various members of the Sepedon Group.

Mature larvae of the aquatic Sepedon Group have a tapered, strongly re-
tractile anterior end; truncate posterior end with three or five pairs of
fleshy lobes around the spiracular disc; paired posterior spiracles each
with three slits and four profusely branched, interspiracular float hairs;
frequently a dark, patterned integument with many folds and wrinkles, and
hair patches dorsally; wartlike body tubercles with those on the ventral sur-
face functioning as pseudopods; and inconspicuous anterior spiracles.

Larvae of the Sepedon Group are further characterized by the following
features diagnostic of the cephalopharyngeal skeleton of all reared Sciomy-
zidae: An unpaired, anteriorly serrate, transverse ventral arch present
between the anteroventral margins of the mandibular sclerites, posterior
arms of hypostomal sclerite not overlapping the anteroventral process of
the pharyngeal sclerites in third instar, ventral cornu without dorsobasal
lobe and no oral grooves around the mouth opening or longitudinal ridges
in the floor of the pharynx (except subfamily Salticellinae). Also, larvae

204. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of the Sepedon Group have three to seven accessory teeth on the basal
part of the mandibular sclerite, a ventral arch with a relatively deep emargi-
nation posteriorly, no dorsal bridge between the pharyngeal sclerites, no
window in the dorsal cornu of the pharyngeal sclerites and an indenta-
tion index of less than 50.

Puparia of the Sepedon Group have the posterior end curved dorsally
so that the posterior spiracles with their hydrofuge float hairs project above
the water surface. The anterior spiracles project only slightly from the
anterolateral angle of the puparium. The body surface retains to some
extent the color pattern of the mature larva, and it frequently has a dis-
tinctive texture, with lateral bumps and grooves.

Sepedonea isthmi (Steyskal)

Egg—Length 1.28-1.38 mm; greatest width 0.30-0.36 mm. White.
Elongate-ovoid. Dorsum with 2 prominent longitudinal ridges. Small dorso-
lateral ridge visible dorsally. Anterior and posterior poles with distinct
punctures. Reticulations on chorion strongest between dorsal ridges, be-
coming faint laterally and ventrally. Based on 10 specimens.

First-instar larva (Figs. 3, 4)—Length 1.8-3.4 mm; greatest width 0.3-
0.4mm. White. Integument transparent, sparsely covered with spinules as
compared to later larval instars. Cephalopharyngeal skeleton (Fig. 3)
length 0.29-0.32 mm; mandibular sclerite composed of darkly pigmented,
small, decurved portion anteriorly, a larger intermediate portion, and a
darkly pigmented, triangular portion posteriorly, the latter connected to
intermediate portion by a thin strap; ventral arch below intermediate por-
tion; epistomal sclerite (Fig. 4) with 5 windows; hypostomal sclerite fused
to pharyngeal sclerite, hypostomal plate with 4-6 windows; pharyngeal
sclerite light brown, darker along pharyngeal indentation and at anterior
%s of sclerite; indentation index 30-37. Metapneustic. Segments 5-10 with
inner and outer dorsal tubercles contiguous, hair covered. Segment 11 with
number of hairs on dorsal tubercles greatly reduced. Spiracular plates
light tan, with B-shaped spiracular openings. Anal proleg with short de-
curved spinules. Spiracular disc, except bare central area, with many
long, fine hairs. Based on 25 specimens.

Second-instar larva (Figs. 5-9).—Length 3.8-4.5 mm; greatest width 0.6-
0.8 mm. Light tan to brown. Integument translucent, densely covered with
spinules. No middorsal stripe or dorsolateral V-shaped stripes. Cephalo-
pharyngeal skeleton (Fig. 6) length 0.48-0.58 mm; mandibular sclerite
(Fig. 7) 0.11-0.12 mm long, with 3-4 accessory teeth; ventral arch (Fig. 8)
with 21-26 teeth anteriorly; epistomal sclerite (Fig. 5) with 4 windows;
hypostomal sclerite fused to pharyngeal sclerite, hypostomal plate with 4
windows; pharyngeal sclerite without dorsal wing; indentation index 31-

VOLUME 80, NUMBER 2 205

N 12 WOODS? 6)

Figs. 3-12. Larval structures of Sepedonea isthmi. 3. First-instar larva, ceph-
alopharyngeal skeleton; 4. Same, epistomal sclerite and parastomal bars; 5. Second-
instar larva, epistomal sclerite and parastomal bars; 6. Same, cephalopharyngeal skele-
ton; 7. Same, mandibular sclerite; 8. Same, ventral arch; 9. Same, anterior spiracle;
10. Third-instar larva, cephalopharyngeal skeleton; 11. Same, mandibular sclerite; 12.
Same, anterior spiracle. The scale for figures 4, 5, 7-9, 11, and 12 is below figure 12.
(AT = accessory tooth, DC = dorsal cornu, ES = epistomal sclerite, HS = hypostomal
sclerite, LS = ligulate sclerite, MS = mandibular sclerite, P = papilla, PS = pharyngeal
sclerite, VA = ventral arch, VC = ventral cornu).

206 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

35. Anterior spiracles (Fig. 9) light tan, with long tubular basal part; rounded
distal part with 8 marginal papillae. Segments 1-10 as in third-instar larva.
Segment 11 with inner and outer dorsal tubercles greatly reduced; dorso-
lateral tubercle absent. Anal proleg without hooks. Based on 7 specimens.

Third-instar larva (Figs. 10-16).—Length 5.3-10.8 mm; greatest width 0.9-
1.8 mm. Light tan to dark brown; integument subopaque. Light to dark
middorsal stripe on segments 3-11; dorsolateral V-shaped stripes also pres-
ent. Cephalopharyngeal skeleton (Fig. 10) length 0.79-0.88 mm. Indenta-
tion index 36-38. Paired mandibular sclerites (Fig. 11) connected dorsally;
mouthhook darkly pigmented, strongly decurved; basal part with 2 win-
dows. Ventral arch (Fig. 16) below and either joined to or articulating with
basal part of mandibular sclerite; anterior margin with 22-26 short, blunt
teeth; 2 longitudinal slits anteromesially, circular to transverse slits postero-
laterally; pigmented part of arch with posteromedian indentation. Epistomal
sclerite (Fig. 13) with 4 windows, anterior margin hyaline, remainder mostly
darkly pigmented. Parastomal bars fused to paired pharyngeal sclerites
and continuing to pharyngeal indentation as salient dark lines. Hy-
postomal sclerite (Fig. 15) darkly pigmented except for dorsal wing and
hypostomal plate, the latter between anterior rami of the sclerite and with
4 windows; connected to pharyngeal sclerite by thin strap. Ligulate sclerite
darkly pigmented. Pharyngeal sclerites with medium pigmentation, except
for hyaline area at posterior tip of ventral cornu, lightly pigmented to
hyaline area along dorsal surface, and hyaline strip along ventral mid-
line.

Segment 1 longitudinally bilobed anteriorly, each lobe bearing sensory
papilla; postoral spine band posteroventrally and partially extending up
sides of segment. Segment 2 with ring of 8 setae around anterior end; bear-
ing anterior spiracles posterolaterally. Anterior spiracles (Fig. 12) light
tan, with tubular basal part; rounded distal part bearing 7-8 marginal papil-
lae. Segments 3 and 4 with 1 dorsal and 1 lateral seta, and 3 setae on small
ventral tubercle. Segments 5-10 with small, rounded inner dorsal tubercle
covered with many long, fine hairs, usually 1 longer and heavier than others;
a wide, hair-covered outer dorsal tubercle; a small, rounded dorsolateral tu-
bercle with approximately 4 short hairs; a group of 3 lateral tubercles, includ-
ing upper and lower lateral tubercles, each with 1 long seta, the former also
with a group of about 4 shorter hairs, and a middle lateral tubercle, smaller
and slightly anterior to the others and with group of about 4 short hairs; a
ventral tubercle group consisting of a conspicuous middle row of 4 tubercles,
each with tuft of about 4 short hairs, an anterior row of 2 widely separated
tubercles, each with tuft of about 4 hairs, and a posterior row apparently
devoid of distinct tubercles and appearing as creeping welt. Segment 11
with inner and outer dorsal tubercles reduced, the latter with only 2 long

VOLUME 80, NUMBER 2 207

0.05
DP Eee rAoT 7

Figs. 13-18. Larval structures and puparium of Sepedonea isthmi. 13. Third-
instar larva, epistomal sclerite and parastomal bars; 14. Same, posterior spiracular disc;
15. Same, ligulate sclerite and venter of hypostomal sclerite; 16. Same, ventral arch; 17.
Puparium, lateral view; 18. Same, dorsal view. The scale for figures 13, 15, and 16
is below figure 15. (DL = dorsal lobe, DIL = dorsolateral lobe, LL. = lateral lobe,
PB = parastomal bar, VL = ventral lobe, VIL = ventrolateral lobe.)

208 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

hairs; dorsolateral tubercle reduced (may be absent); lateral and ventral
tubercle groups as in segment 5-10, except the posterior row of ventral
tubercle group reduced or absent.

Segment 12 with small lateral tubercle above base of hookless anal pro-
leg; perianal pad bordering anal proleg posteriorly. Posterior spiracular
disc (Fig. 14) with conspicuous ventral and ventrolateral lobes, the former
long and tapered, the latter, 2-segmented, with basal segment truncate and
apical segment digitiform; dorsal, dorsolateral, and lateral lobes smaller,
rounded, and inconspicuous; lobes covered with many fine hairs; central area
of disc base bare. Paired spiracular plates at apices of sclerotized, basally
scalloped, stigmatic tubes; each plate with 3 diverging, elongate-oval spirac-
ular slits, a stigmatic scar, and 4 branching interspiracular float hairs. Based
on 5 specimens.

Puparium (Figs. 17, 18)—Length 5.3-6.0 mm; greatest width 1.9-2.3 mm.
Yellow to dark brown. Translucent to opaque. Barrel-shaped with anterior
end narrow, tapered; posterior end sharply upturned. Cephalopharyngeal
skeleton as in third-instar larva. Yellowish anterior spiracles protruding
from anterolateral corners of dorsal cephalic caps. Most of dorsum of seg-
ments 5 and 6 yellow. Dorsum with 3-4 brown, V-shaped stripes extending
to middle tubercle groups. Yellow lateral line extending from segment
5, where it is infuscated with brown to posterior end of puparium, below
junction of segments 11 and 12. Area below lateral line largely brown with
ventral tubercles yellow. Posterior spiracular disc with lobes shrunken;
float hairs inconspicuous; stigmatic tubes yellow. Anal plate invaginated;
anal proleg inconspicuous. Dorsal and dorsolateral tubercles reduced, hair
patches closely appressed to the surface. Based on 4 specimens.

Acknowledgments

Fieldwork by the junior author was supported by grant GB 11672, Gen-
eral Ecology Fund, National Science Foundation, awarded to C. O. Berg,
Cornell University. Fieldwork by the senior author was conducted, in
part, while he was with the Office of Environmental Sciences, Smith-
sonian Institution, and also was supported by ARS-NSF Interagency Agree-
ment No. 12-14-1001-203. The senior author thanks Carlo Julio Rosales,
Universidad Central de Venezuela, Maracay, for aiding with fieldwork
in Venezuela and F. D. Bennett and M. Yaseen, Commonwealth Institute of
Biological Control, Curepe, for similar aid in Trinidad. The late Robert
Beard helped with rearing work at Cornell University. Joseph Rosewater,
Smithsonian Institution, provided identifications of snails and P. M. Marsh,
Systematic Entomology Laboratory, USDA, determined the diapriid wasps.
Linda Heath, Systematic Entomology Laboratory, prepared figures 14, 17,
18. We thank C. O. Berg, Nelson Papavero, Museu de Zoologia da Univer-
sidade de Sao Paulo, Brazil, and Guy Shewell, Biosystematics Research

VOLUME 80, NUMBER 2 209

Institute, Ottawa, Canada, for making specimens of S. isthmi available to us,
and C. O. Berg and J. Abercrombie, Environmental Technology Division,
Chemical Systems Laboratory, U.S. Army Armament Research and Develop-
ment Command, Aberdeen Proving Ground, Maryland, for reviewing the
manuscript.

Literature Cited

Abercrombie, J. 1970. Natural history of snail-killing flies of South America (Diptera:
Sciomyzidae: Tetanocerini). Ph.D. Thesis, Cornell Univ. (L.C. Card No. Mic
70-23, 095), Univ. Microfilms, Ann Arbor, Mich. (Dissertation Abstr. Internat. (B)
31:3456-3457). 335 pp.

Abercrombie, J., and C. O. Berg. 1975. Natural history of Thecomyia limbata (Dip-
tera: Sciomyzidae) from Brazil. Proc. Entomol. Soc. Wash. 77:355-368.

Berg, C. O. 1973. Biological Control of Snail-borne Diseases: A review. Exptl.
Parasitol. 33:318-330.

Chock, Q. C., C. J. Davis, and M. Chong. 1961. Sepedon macropus (Diptera: Sciomy-
zidae) introduced into Hawaii as a control for the liver fluke snail, Lymnaea
ollula. J. Econ. Entomol. 54:1-4.

Eckblad, J. W., and C. O. Berg. 1972. Population dynamics of Sepedon fuscipennis
(Diptera: Sciomyzidae). Can. Entomol. 104:1735-1742.

Knutson, L. V., S. E. Neff, and C. O. Berg. 1967. Biology of snail-killing flies from
Africa and southern Spain (Sciomyzidae: Sepedon). Parasitology 57:487-—505.

Knutson, L., G. C. Steyskal, J. Zuska, and J. Abercrombie. 1976. Family Sciomyzidae.
In Museu de Zoologia, Universidade de Sao Paulo. A Catalogue of the Diptera of
the Americas South of the United States 64: 1-24.

Nagatomi, A., and A. Tanaka. 1967. Egg of Sepedon sauteri Hendel (Diptera, Sciomy-
zidae). Konty 35:31-33.

Neff, S. E., and C. O. Berg. 1966. Biology and immature stages of malacophagous
Diptera of the genus Sepedon (Sciomyzidae). Bull. Va. Agric. Exp. Sta. 566:
1-113.

Steyskal, G. C. 1951. The genus Sepedon Latreille in the Americas (Diptera: Sciomy-
zidae). Wasmann J. Biol. (1950) 8:271-297.

——. 1973. A new classification of the Sepedon group of the family Sciomyzidae
(Diptera) with two new genera. Entomol. News 84:143-146.

(LK) Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. Educ.
Admin., USDA, Beltsville, Maryland 20705; and (KV) Department of En-
tomology, Cornell University, Ithaca, New York 14853.

Present address—(KV) Bureau of Plant Industry, Pennsylvania Depart-
ment of Agriculture, Harrisburg, Pennsylvania 17120.

Footnote

1 This species is considered as a possible synonym of Galba viridis (Quay and Giaimard)
by B. Hubendick, 1951, Recent Lymnaeidae, Kungl. Svensk. Vittenskapakad, Fjardel 1,
Bde. ao. | pp:.1—223,. d» pls.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 210-215
NOTES ON THE BIOLOGY OF DIGLYPHUS INTERMEDIUS
(HYMENOPTERA: EULOPHIDAE), A PARASITE OF THE
ALFALFA BLOTCH LEAFMINER, AGROMYZA FRONTELLA
(DIPTERA: AGROMYZIDAE)

R. M. Hendrickson, Jr. and S. E. Barth

Abstract.—Diglyphus intermedius (Girault) is the most common para-
site of the alfalfa blotch leafminer, Agromyza frontella (Rondani), on alfalfa
in Delaware, southcentral Pennsylvania, and southern New Jersey. It is
a larval ectoparasite of leafminer larvae and is usually solitary; however,
occasionally 2-5 parasites develop on a single host. It paralyzed and killed
all three larval instars of its host agromyzids without ovipositing though it
usually oviposited and developed on 3rd-instar larvae. The parasite has
three instars as measured by changes in mandible length. At a constant
25.5°C, it developed from egg to adult in 11 days, lived 3-4 weeks as an
adult and had an average fecundity of 40 offspring per female.

The alfalfa blotch leafminer (ABL), Agromyza frontella (Rondani), an
introduced pest of European origin, was first observed in the USA in
Hampshire County, Massachusetts, in 1968 (Miller and Jensen, 1970). It
is usually a subeconomic pest on alfalfa throughout the northeastern
USA and adjoining Canadian provinces, but occasionally it has become a
serious pest (100-200 adults collected/sweep (Kim, 1975)). We report here
notes on the biology of the most important of the native parasites of ABL,
Diglyphus intermedius (Girault).

Diglyphus intermedius is a Nearctic species that has been reported to
parasitize Liriomyza subpusilla Frost in California and Phytomyza atricornis
Meigen in Rhode Island (Peck, 1963); Liriomyza sativae Blanchard (=
Liriomyza prosperpusilla Frost) in Florida and Illinois (Burks, 1967); Phy-
tomyza chrysanthemi Kowarz in Rhode Island (Peck, 1951); and Liriomyza
brassicae (Riley) in California (Spencer, 1973). We have collected it
from Liriomyza trifoliearum Spencer on alfalfa throughout the northeastern
USA.

Laboratory Studies

All the life history studies of Diglyphus intermedius in the laboratory were
conducted on a native agromyzid, Liriomyza trifoliearum Spencer, obtained
from field-collected alfalfa and maintained in a laboratory colony on
“Bountiful” bush snap bean, Phaseolus vulgaris L., and “Thaxter” lima bean,
Phaseolus limensis Macf., in caged plants at 25.5 + 1.1°C and 60 + 5%
RH with a photoperiod of 16L:8D. Observations on larval development

VOLUME 80, NUMBER 2 211

were made by constructing artificial mines because Diglyphus intermedius
develops entirely within the host’s mine, and development there is ob-
scured by the leaf tissue. The mine consisted of a microscope slide on which
we placed a small square of blotter paper. A circle of construction paper
with a hole in it, similar in size and shape to the gummed reinforcements
used for notebook paper, was laid on the blotter paper. In the central hole
was placed a parasitized leafminer larva that had been dissected from the
leaf mine. A coverslip was placed on top of the larva to prevent any
mobile parasite larva from leaving the artificial mine. The entire slide was
placed in a closed petri dish that had moistened plaster of Paris (Fig. 1)
or sand on the bottom to maintain a high RH. The slide was removed
from the petri dish for observation by microscope. Since the majority of
parasites handled in this way survived from egg to adult, the technique
could be applied to the study of other larval ectoparasites of leafminers.

The number of instars was determined by exposing 3rd-instar hosts to
adult parasite adults for 15 h. When the parasites were removed at 0800 h,
that hour was counted as the beginning of the first day of maturation of
the parasite though, in some cases, as much as 15 h had elapsed. Then at
0800 h on succeeding days, parasite larvae were removed and mounted on
slides in Hoyer’s medium. The instar was determined by measuring the
length of the mandibles from the tip of the “tooth” to the posterior margin
of the dorsal process (the greatest distance across the mandible). The
length of a single mandible averaged 10 mw (n = 12, range = 6-13 pz) for the
Ist instar, 17 pw (n = 7, range = 15-19 yz) for the 2nd instar, and 27 pw (n = 11,
range = 23-30 mw) for the 3rd instar (Fig. 2). These observations were cor-
roborated by measuring the mandibles of larvae in eggs shortly before
eclosion, and those in cast skins. Therefore, we concluded that the para-
site had three larval instars. A Ist-instar parasite larva feeding on a
Liriomyza trifoliearum larva is shown in Fig. 3.

From the observations of the larval parasites in artificial mines, it
was found that at 25.5°C development from egg to adult required 11
days: 1 day for the egg stage, 4 for the larval stage and 6 for the pupal
stage. The egg hatched within 24 h after it was laid, and if it was not
already attached to the host, the young parasite larva sought out the host
and began feeding immediately. The Ist instar lasted about 24 h, the 2nd
less than 24 h and the 3rd about 2% days. On the 3rd day, the larva changed
from light yellow to a lime-bluish green and soon constructed 6-8 meconial
pillars that extended from the upper to the lower surface of the mine. Since
these pillars are usually arranged in pairs on each side of the pupa (Fig. 4),
they apparently protect the pupa from being crushed if the leaf dries out
and also prevent the pupa from rolling about inside the mine. When the
parasite transformed to the adult stage (Fig. 5) in the host’s mine, it chewed
an oval hole in the upper epidermis of the leaf and left the mine.

bo
bo

1 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-5. 1. Artificial mine. Parasite and host are inside black paper ring with
coverslip placed on top. Plaster of Paris in petri dish was moistened to maintain high
humidity; 2. Mandibles of Ist-, 2nd-, and 3rd (final)-instar Diglyphus intermedius
larvae. Bar is 10 yu; 3. First-instar D. intermedius larva feeding on 3rd-instar Liriomyza
trifolicarum host. Bar is 0.1 mm; 4. Meconial pillars around Diglyphus intermedius pupa.
Tops of pillars are attached to underside of coverslip; 5. Adult female D. intermedius.
Bar is 1 mm.

VOLUME 80, NUMBER 2 213

Repeated attempts to observe mating in cages of various sizes, with and
without hosts, were unsuccessful. Mating was infrequently observed in
maintenance culture cages containing thousands of parasites.

Oviposition of Diglyphus intermedius was studied by confining parasite
adults in a petri dish with a bean leaf mined by 3rd (final)-instar hosts. The
activity could then be observed through a microscope. The searching be-
havior of the female parasite consisted of palpating the leaf until a mine
was located. Then she followed the mine by swaying back and forth, almost
in a rhythmic motion, while she palpated the mine alternately with each
antenna. Once she discovered a host, she immediately circled above it and
palpated the leaf epidermis with her antennae. The host larva apparently
sensed the presence of the parasite above the mine and often began to thrash
around within the confines of the mine, even before the parasite penetrated
the mine with the ovipositor. Within seconds after locating a host, the
parasite attempted to sting it and the host responded to each attempted
sting by making a rapid twisting motion in an apparent attempt to avoid
the ovipositor. The parasite was observed to sting and kill each of the
three host instars, but oviposition was never observed on Ist-instar larvae.
Stinging without oviposition lasted from 2-14 s. After several stings, the host
ceased movement, but the hindgut was observed to continue to function
until waste matter was expelled; then the larva became completely im-
mobile. More prolonged stings, average 39 s (n = 6, range = 25-53 s), were
assumed to be oviposition attempts. The parasite commonly oviposited
1-2 eggs/host, but on one occasion three eggs were laid. Eggs were or-
dinarily placed on the host larva, but occasionally some were placed 1-2
mm from the host but within the mine. Oviposition usually occurred on 3rd-
instar hosts, but in one case, a female was observed to sting a 2nd-instar
larva nine times and to lay one egg on the host. Since attempts to rear the
parasite on 2nd-instar host larvae were unsuccessful, we suspect that
such Liriomyza trifoliearum larvae do not usually provide sufficient food
to allow complete development of the parasite.

Fecundity of the parasite was determined by confining a single female,
less than 24 h old, with two males on “Bountiful” bush snap beans heavily
infested with 3rd-instar L. trifoliearum. The parasites were supplied with
honey as food, and fresh host material was placed in the cage weekly until
the female died. The plant material was removed and held for parasite
emergence. The average fecundity was found to be 40.2 progeny per
female (n = 6). Females in the test lived 3-4 weeks.

Parasites reared on “Bountiful” bush snap bean had a sex ratio of 1? :2.24
(n = 204). However, the ratio on “Thaxter” lima bean was 12:14 (n = 117).
A similar disparity was noted when we reared the European parasite
Diglyphus isaea (Walker) on these two bean species.

214 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Field Observations

Samples of 50 alfalfa leaflets mined by either ABL or Liriomyza spp.
were collected weekly from early May through November 1975, from each
of seven fields (one at Newark, Delaware; three near Oxford, Pennsyl-
vania; and three near Rancocas, New Jersey) unless the alfalfa was cut or
bad weather prevented fieldwork. If populations of agromyzid larvae
were low, a sample consisted of as many mined alfalfa leaflets as could
be collected in each field in 20 min. In the laboratory, the mined leaflets
were placed in petri dishes (50 mm diam with tight-fitting lids) on slightly
moistened filter paper, which retained moisture for 3-4 weeks, long enough
for the parasites to develop and emerge at 22.2 + 1.1°C. The host larvae
were separated by mine character. ABL, in the Ist and early 2nd instar,
formed a fairly straight linear mine; in late 2nd and throughout the 3rd
instar it produced a large blotch mine. Liriomyza spp. formed a serpentine,
linear mine in all three instars. The instar of the host was determined by
measuring the length of the cephalopharyngeal skeleton by transmitted light
under the microscope and comparing it to predetermined length ranges
for each instar for each species.

Diglyphus intermedius parasitized a total of 9.2% of the ABL larvae col-
lected from the field (n = 4,748), but actual field mortality due to the
parasite was probably higher than that. We did not include larvae killed
by parasite stinging without oviposition because they could not be easily
separated from those killed by predators, diseases, overcrowding in the
leaflet, or environmental factors. Total parasitism of ABL by all parasitic
species was 18.8% and D. intermedius accounted for almost 50% of this
amount. Diglyphus intermedius preferred 3rd (final)-instar larvae, although
23.7% of the hosts were 2nd-instar larvae. Thus our inability to rear the
parasite on 2nd-instar Liriomyza trifoliearum larvae in the laboratory was
probably caused by the smaller size of the host.

The parasite was usually solitary but was gregarious on 13.3% of the
larvae (2-5 adults from a single host).

Acknowledgments

We thank G. Gordh and G. C. Steyskal, Systematic Entomology Lab-
oratory, IIBIII, Agric. Res. Serv., USDA, for their respective identifica-
tion of Diglyphus intermedius and of Agromyza frontella and Liriomyza
trifoliearum.

Literature Cited

Burks, B. D. 1967. Superfamily Chalcidoidea. In K. V. Krombein and B. D. Burks,
Eds. Hymenoptera of America North of Mexico. Synoptic catalog. USDA Agric.
Monogr. No. 2, second supplement, pp. 213-282.

VOLUME 80, NUMBER 2 215

Kim, K. C. 1975. Annual Summary. Alfalfa blotch leafminer. Penn. Coop. Econ.
Ins. Rep. 35:6.

Miller, D. E., and G. L. Jensen. 1970. Agromyzid alfalfa leaf miners and _ their
parasites in Massachusetts. J. Econ. Entomol. 63:1337—1338.

Peck, O. 1951. Superfamily Chalcidoidea, pp. 410-594. In C. F. W. Muesebeck et al.,
Eds. Hymenoptera of America North of Mexico. Synoptic catalog. USDA. Agric.
Monogr. No. 2.

—. 1963. A catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera).
Can. Entomol. Suppl. 30. 1092 pp.

Spencer, K. A. 1973. Agromyzidae (Diptera) of economic importance. In E. Schimit-
schek, Ed. Series Entomologica, Vol. 9. W. Junk, The Hague. xi + 418 pp.

Beneficial Insects Research Laboratory, Agric. Res. Serv., USDA, Newark,
Delaware 19713.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 216-227
TERMINALIA OF SOME NORTH AMERICAN SPECIES
OF MEGASELIA (APHIOCHAETA) AND DESCRIPTIONS
OF TWO NEW SPECIES (DIPTERA: PHORIDAE)

William H Robinson

Abstract.—Borgmeier’s (1964, 1966) revision of North American Megaselia
lacked detailed figures of the adult terminalia. In this large genus the male
epandrium and hypandrium and the female tergum and sternum 6 provide
excellent distinguishing characters. This work presents a brief diagnosis
and figures of the male and female terminalia of the following species of
the subgenus Aphiochaeta: Megaselia californiensis (Malloch), M. capta
(Borgmeier), M. georgiae Borgmeier, M. decora new species, M. gravis
Borgmeier, M. laffooni new species, M. marginalis (Malloch), M. me-
landeri Borgmeier, M. nigriceps (Loew), M. plebia (Malloch), M. proclinata
Borgmeier, M. relicta Borgmeier, M. robusta Schmitz. Two species of this
group, M. decora and M. laffooni, are described as new.

Borgmeier’s (1964, 1966) revision of the North American Megaselia placed
the taxonomy of this genus on a firm foundation. He studied material in
several U.S. collections and was able to examine nearly all the types of
North American Megaselia. In this work Borgmeier provided descriptions
and keys for about 260 Megaselia species. However, Borgmeier’s revision
lacks detailed figures of the adult terminalia. The external terminalia of the
male (epandrium and hypandrium) and of the female (tergum 6) provide
excellent characters for distinguishing species. The proper use of these
characters along with other structural features in species descriptions could
help remove some of the difficulties in accurately determining species of
this genus.

The purpose of this work is to provide a brief diagnosis and the first de-
tailed figures of the male and female terminalia of the Megaselia species in
Borgmeier's (1964) Group I. Two new species, Megaselia (Aphiochaeta)
decora, and M. (A.) laffooni are described.

Materials and Methods

This work is based on the study of 470 Nearctic specimens of the Mega-
selia species of Borgmeier’s Group I. Specimens were seen from most states
and provinces of U.S. and Canada, but not from Greenland and Mexico.

I have examined, or received information on, the holotypes of all the
Nearctic species of Group I. Specimens of the type-series of Holarctic
species were not available for examination at this time. Information on
original descriptions and synonymies can be found in Borgmeier (1964).

The methods used for removing and preparing terminalia for study were
similar to those used by other workers. Terminalia were permanently

VOLUME 80, NUMBER 2 217

stored in 4 X 11 mm plastic vials, partially filled with glycerine and capped
with rubber stoppers, and placed on the same pin as the insect.

External Terminalia of Megaselia

The external male terminalia of Megaselia species are platyperzid-like
in appearance. A brief explanation will help to understand the simple struc-
ture of the preabdomen and postabdomen.

The preabdomen (segments 1-6) is moderately elongate and more or less
cylindrical. The terga are generally unequal in size, and the sterna are
absent.

The postabdomen of the male (Fig. 1) lies in a straight, longitudinal
relationship with the preabdomen. Terga and sterna 7 and 8 are absent. The
epandrium (EP, Fig. 1) is generally symmetrical and often has scattered
setae laterally. The hypandrium (Fig. 2) is generally asymmetrical; the
right side, and occasionally the left side, bears a setulose lobe that extends
posteriorly. The proctiger (PR, Fig. 1) is well developed; the cerci are
distinct (Fig. 1) and fused along the mid-line.

The postabdomen of the female consists of an elongate ovipositor that
telescopes within segment 6. The ovipositor is entirely membranous—never
forming a horny, nonretractile stylet. Terga and sterna 7 and 8 are re-
duced and are sometimes absent. The cerci are distinct and bear numerous
setae.

Group I Megaselia (Aphiochaeta)

The practice of partitioning the hundreds of Megaselia species into eight
“artificial” groups as frequently done by other authors (Lundbeck, 1922;
Schmitz, 1956; Borgmeier, 1964) has been followed here. These groups
provide an excellent, albeit abitrary, means of reducing the large numbers
of species into small, workable taxonomic groups. The structural charac-
ters upon which these groups are based (see Borgmeier, 1964) are rela-
tively constant and discernible on all but the poorest specimens.

Species of Group I can be distinguished by the setose mesanepisternum
and the 4-6 scutellar bristles. This group is known from males of 13 Nearctic
species, including the two new species described below; females are known
for only 4 species of the Group.

Megaselia (A.) nigriceps (Loew) and M. (A.) robusta Schmitz are the
only known Holarctic species of this group.

Megaselia (Aphiochaeta) californiensis (Malloch)
igs: 1120

Diagnosis—This species can be distinguished from other Nearctic Aphio-
chaeta species with 4-6 scutellars by the following combination of char-
acters: Mesanepisternum without bristles; halter brown; fore basitarsus

218 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 2 219

enlarged apically; costa 48-52% of wing length. Terminalia: Epandrium
(Fig. 11) short, with numerous bristles ventro- and posterolaterally; hy-
pandrium (Fig. 20) bare, except for setulae on lobe. Female unknown.
Known distribution California, Washington.
Material examined.—Two 6,12.

Megaselia (Aphiochaeta) capta Borgmeier
Figs. 18, 19

Diagnosis—The male differs from other Nearctic Aphiochaeta species
with 4-6 scutellars by the character combination: Mesanepisternum with
bristles; costa 41% of wing length. Terminalia: Epandrium (Fig. 18) curved
under proctiger, with 1 strong bristle posterolaterally; hypandrium (Fig.
19) setulose laterally, setulae longer on lobe. Female unknown.

Known distribution.—Virginia.

Material examined.—Holotype.

Megaselia (Aphiochaeta) georgiae Borgmeier
Figs. 14, 21

Diagnosis —The male differs from other Nearctic Aphiochaeta species
with 4-6 scutellars by the character combination: Scutellum with 6 bristles;
mesanepisternal bristle absent. Terminalia: Epandrium (Fig. 14) higher
than wide, with numerous bristles, left side with long, unarticulated pro-
cess directed posteroventrally; hypandrium (Fig. 21) setulose, with 1 long,
slender, setulose lobe. Female unknown.

Known distribution Georgia.

Material examined.—Holotype.

Megaselia (Aphiochaeta) decora Robinson, new species
Figs. 5, 22

Diagnosis—The male differs from other Nearctic Aphiochaeta species
with 4-6 scutellars by the character combination: Mesanepisternum without
bristles; propleuron with numerous scattered fine setae and bristles, and the
presence of 2 supra-antennal bristles. Female unknown.

Description of male-—Body yellowish brown. Frons brown, dull, as wide
as high; 1 pair of supra-antennals, more approximate than preocellars; inner
frontal bristle midway between supra-antennal and outer frontal bristle;

<

Figs. 1-9. Megaselia spp. terminalia. 1. M. nigriceps, male terminalia; 2. M. nigriceps,
male hypandrium; 3. M. nigriceps, female tergum 7; 4. M. marginalis, male epandrium;
5. M. decora, male epandrium; 6. M. marginalis, female tergum 7; 7. M. gravis, male
epandrium; 8. M. relicta, female tergum 7; 9. M. robusta, female tergum 7.

220 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

posterior frontal bristle on slightly higher level than preocellar bristle.
Parafacia with 3 bristles. Third antennal segment brown. Palpus brown.

Thorax brown; propleuron with numerous scattered setae, 1 strong ven-
tral and 3 dorsal propleural bristles; mesanepisternum with about 22 fine
setae, bristles absent. Scutellum with 4 bristles.

Abdominal terga brown; sterna yellowish brown. Terminalia long, brown;
proctiger yellow; epandrium (Fig. 5) slightly longer than tergum 6, with
scattered setae; hypandrium (Fig. 22) bilobed, lobes unequal, with setae.

Legs brownish yellow; fore tibia with small anterodorsal bristles, and
18 short, weak posterodorsal bristles; hair-seam extending 0.8 length of
tibia; middle basitarsus with 2 weak bristles on basal %. Hind femur with
9 decumbent bristles on basal ™%; hind tibia with indistinct anterodorsals, 5
strong, widely spaced posterodorsals on apical %, and 3-4 short postero-
dorsals on apical %; hind basitarsus with 2 bristles near distal %.

Wing 2.54 mm long; membrane hyaline, veins pale brown; costa 47% of
wing length, ratio of first 2 costal divisions 2:3; costal bristles long; 6
axillary bristles. Halter entirely brownish yellow.

Material examined.—Holotype.

Holotype —Male, Cowichan Lake, British Columbia, II-20-1964, J. A.
Chapman. Type No. 15131, in the Canadian National Collection.

Remarks.—The type-specimen was apparently mounted from alcohol.

Megaselia (Aphiochaeta) laffooni Robinson, new species
Figs. 17, 23

Diagnosis —The male differs from other Nearctic Aphiochaeta species
with 4 scutellars by the character combination: Mesanepisternum without
bristles; costa 47% of wing length; knob of halter and palpus blackish brown.
Female unknown.

Description of male—Body blackish brown. Frons blackish brown, dull,
as wide as high; supra-antennals subequal, lower pair closer than upper
pair to median furrow, upper pair about as far as preocellar from median
furrow; inner frontal bristle close to outer frontal bristle; posterior frontal
bristle higher than preocellar. Parafacia with 5 bristles. Third antennal
segment dark brown. Palpus blackish brown, narrow and pointed apically.

Thorax blackish brown; propleuron without scattered hairs, 3 ventral
and 3 dorsal propleural bristles in a row adjacent to anterior spiracle;
mesanepisternum with 8-10 fine setae, bristles absent. Scutellum with
4 bristles.

=>

Figs. 10-18. Megaselia spp. male epandria. 10. M. plebeia; 11. M. californiensis;
12. M. proclinata; 13. M. relicta; 14. M. georgiae; 15. M. melanderi; 16. M. robusta;
17. M. laffooni; 18. M. capta.

221

VOLUME 80, NUMBER 2

222 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Abdominal terga blackish brown; sterna brown. Terminalia small, blackish
brown; proctiger small, pale brown; epandrium (Fig. 17) with strong
bristles ventrally near base, and weak bristles posterolaterally; hypandrium
(Fig. 23) with 1 lobe bearing fine setae.

Legs dark brown; fore tibia with indistinct antero- and posterodorsal
bristles; fore basitarsus enlarged, broader than apex of fore tibia. Middle
tibia with 8 anterodorsals and 4 weak, decumbent posterodorsal bristles;
hairseam extending % length of tibia; middle basitarsus with 2 weak bristles
on basal %. Hind femur with 7 short, widely spaced fine setae on slightly
more than %; basal hind tibia with indistinct anterodorsal bristles, 14 pos-
terodorsal bristles that are weak on basal ' of row; hair seam; strongly
arched. Hind basitarsus with 1 bristle on basal %.

Wing 1.62 mm long; membrane pale brown, veins dark brown; costa 47%
of wing length, ratio of first 2 costal divisions 2:0; costal bristles long; 4
axillary bristles. Halter entirely blackish brown.

Material examined.—Holotype.

Holotype——Male, Isabel Pass, 2,900’, mile 206, Richardson Highway,
III-13-1962, P. J. Skitsko. Type No. 15132, in the Canadian National Col-
lection.

Remarks.—I take pleasure in naming this phorid for the late Dr. Jean L.
Laffoon of Iowa State University. As a scholar, teacher, dipterist and
friend he was as unique as this species.

Megaselia (Aphiochaeta) gravis Borgmeier
Figs. 7, 24

Diagnosis—The male differs from other Nearctic Aphiochaeta species
with 4-6 scutellars by the following combination of characters: Mesanepi-
sternum without bristles; costa 55% of wing length; and hind femur with
8 long bristles on basal %. Terminalia: Epandrium (Fig. 7) setulose dorsally;
hypandrium (Fig. 24) setulose laterally, lobe setulose and with long, fine
setae apically. Female unknown.

Known distribution.—Alaska.

Material examined.—Holotype.

Megaselia (Aphiochaeta) marginalis (Malloch)
Figs. 4.16, 25

Diagnosis.—This species differs from other Nearctic Aphiochaeta species
by the combination of 6 scutellar and 1 mesanepisternal bristle. Terminalia:
Epandrium (Fig. 4) subshining dorsally, setose laterally, with 4-6 bristles
ventrally near base; hypandrium (Fig. 25) symmetrical, bilobed. Oviscapt:
Tergum 7 (Fig. 6) broad, with 4 fine apical setae; sternum 7 narrow, Y-
shaped and with 2 apical setae on each fork of Y.

VOLUME 80, NUMBER 2 223

Known distribution.—Florida, Georgia, lowa, Kansas, Maryland, Michi-
gan, Missouri, New York, Tennessee, Virginia, Wisconsin.

Material examined.—Sixteen 6, 7°.

Remarks.—This species is structurally similar to M. georgiae (Nearctic),
and M. flavicoxa (Palearctic).

Biology.—Two females were collected from the cell of the scarab beetle
Peltotrupes youngi Howden, in Florida.

Megaselia (Aphiochaeta) melanderi Borgmeier
Figs. 15, 26

Diagnosis —The male differs from other Nearctic Aphiochaeta species
with 4 scutellars by the character combination: Costa 53% of wing length;
costal bristles long; halter and palpus brown. Terminalia: Epandrium
(Fig. 15) with scattered setulae and 6-8 bristles; hypandrium (Fig. 26)
setulose, bilobed, right lobe longer than left. Female unknown.

Known distribution Florida (Royal Palm Park).

Material examined.—Holotype.

Remarks.—The type-locality for this species is about 35 miles southwest of
Miami, Florida. This region might be considered transitional between the
Nearctic and Neotropical regions.

Megaselia (Aphiochaeta) nigriceps (Loew)
Figs, 1,,2, 3

Diagnosis.—This species differs from other Nearctic Aphiochaeta species
with 4 scutellars by the character combination: Mesanepisternum without
bristles; scutellars long, subequal; costa 50-53% of wing length. Terminalia:
Epandrium (Fig. 1) setulose posterolaterally and with 1-2 long bristles ven-
trally; hypandrium (Fig. 2) bare except for setulose lobe. Oviscapt: Tergum
7 (Fig. 3) long, arched basally.

Known distribution—Widespread in U.S. and western Canada.

Material examined.—Two hundred twelve ¢, 155 2°.

Remarks.—The coloration of this species ranges from yellowish brown to
blackish brown. The darker individuals seem to have a predominantly
northern distribution and the paler individuals have a more southern dis-
tribution. The male palpus and fore basitarsus are generally large, but in
some specimens they are slender. The record of this species feeding on
garbage (Aldrich, 1897) was an error; Aldrich actually had specimens of
M. cavernicola (Brues).

Megaselia (Aphiochaeta) plebia (Malloch)
Figs. 10, 27

Aphiochaeta pallidiventris Malloch, 1919:47. NEW SYNONYMY.
Megaselia (Aphiochaeta) wirthi Borgmeier, 1964:277. NEW SYNONYMY.

224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

NNN
SA

VOLUME 80, NUMBER 2 225

Diagnosis.—This species differs from other Nearctic Aphiochaeta species
by the character combination: Mesanepisternum with 1-2 bristles; fore tibia
with 12-14 strong anterodorsal bristles; halter yellow; costa 45-50% of
wing length. Terminalia: Epandrium (Fig. 10) with 2 strong bristles ven-
trally near base; hypandrium (Fig. 27) setulose. Oviscapt; Terga and
sterna 6 and 7 absent.

Known distribution —Arkansas, Kansas, South Carolina, Virginia.

Material examined.—Fifteen ¢, 12°.

Remarks.—The female of this species is very striking in having tergum
1 dark brown, terga 2-3 entirely or in part pale brown to brownish orange,
and terga 4-6 yellowish orange. Megaselia plebeia was apparently described
from pharate adult males (not 1 male and 2 females as stated by Malloch,
1914). The female of this species (M. pallidiventris) may have been de-
scribed as new because of the striking coloration of the abdomen and in-
adequate description of M. plebeia. Borgmeier (1964) described M. wirthi
before seeing the types of M. plebeia and M. pallidiventris.

Megaselia (Aphiochaeta) proclinata Borgmeier
Figs. 12, 28

Diagnosis—The male can be distinguished from other Nearctic Aphio-
chaeta species by the presence of 6 supra-antennal bristles. Terminalia:
Epandrium (Fig. 12) small, higher than wide, with numerous bristles postero-
laterally, left side with long, unarticulated process directed posteriorly; hy-
pandrium (Fig. 28) setulose laterally, bilobed. Female unknown.

Known distribution.—Virginia.

Material examined.—Holotype.

Remarks.—The presence of 6 supra-antennal bristles is unique for this
group of species. It is possible that one pair are supernumerary. However,
the male of M. proclinata is easily recognized, also, by the unusual shape
of the epandrium.

Megaselia (Aphiochaeta) relicta Borgmeier
iigso oS, los oO

Diagnosis.—This species differs from other Nearctic Aphiochaeta species
with 4 scutellars by the character combination: Mesanepisternum with 1
bristle; halter brown; costa 46-47% of wing length. Terminalia: Epandrium
(Fig. 13) shining, right side with short, unarticulated process posterolat-

<

Figs. 19-30. Megaselia spp. male hypandria. 19. M. capta; 20. M. californiensis;
21. M. georgiae; 22. M. decora; 23. M. laffooni; 24. M. gravis; 25. M. marginalis; 26.
M. melanderi; 27. M. plebia; 28. M. proclinata; 29. M. robusta; 30. M. relicta.

226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

erally; hypandrium (Fig. 30) setulose laterally on right side, lobe without
setulae. Oviscapt: Tergum 7 (Fig. 8) short, an inverted-Y-shape.

Known distribution.—Illinois, Iowa, Kansas, Washington.

Material examined.—Two 4, 3 °.

Megaselia (Aphiochaeta) robusta Schmitz
Figs. 9, 16, 29

Diagnosis—The male differs from other Nearctic Aphiochaeta species
with 4 scutellars by the presence of a row of 10-12 short ventral bristles
on the basal % of the hind femur. Terminalia: Epandrium (Fig. 16) with
4-6 scattered bristles and 1 long bristle; hypandrium (Fig. 29) setulose lat-
erally on right. The female differs by the character combination: Mesan-
episternum without bristles; palpus narrow and with 1 row of ventral
bristles; halter yellowish brown; costa 50-55% of wing length. Oviscapt:
Tergum 7 (Fig. 9) broad, slightly arched basally.

Known distribution —Alaska, western Canada, New Hampshire, northern
Europe.

Material examined.—Nineteen ¢, 21°.

Biology—tThe holotype was reared from larvae in decaying vegetation.

Discussion

The Groups in the genus Megaselia are practical entities and are not neces-
sarily monophyletic. A review of the figures presented in this paper indicates
considerable variation in the male terminalia of Group I Megaselia. There
are some species with primitive, platypezid-like terminalia, and some with
more derived terminalia features.

The Group I Megaselia species (North America) with the most primitive
terminalia features include M. decora, M. marginalis, M. melanderi, M.
plebeia, M. proclinata and M. relicta. The male terminalia of these species
is characterized by one or more unsegmented processes on the epandrium
and/or a bilobed hypandrium.

There are two species in which these conditions can be seen best. In M.
proclinata (Fig. 12) the epandrium has an elongate, unsegmented process
similar to the surstyles of platypezids. The hypandrium of M. marginalis
(Fig. 25) represents the most primitive, unspecialized condition. It is
symmetrical and bilobed, and the lobes are of equal size.

The Group I Megaselia species with the most derived terminalia features
include M. californiensis, M. capta, M. georgiae, M. gravis, M. laffooni, M.
nigriceps and M. robusta. The terminalia of these species is characterized
by a simple epandrium, without lateral processes (except in M. georgiae),
and/or a single-lobed hypandrium.

VOLUME 80, NUMBER 2 227

As knowledge of the male and female terminalia of Megaselia increases,
so will our ability to construct a phylogenetic classification of this genus.

Acknowledgments

I gratefully acknowledge the time and counsel contributed to this re-
search by the late Dr. Jean L. Laffoon. Drs. B. V. Peterson, Biosystematics
Research Institute, Agriculture Canada and W. W. Wirth and R. J. Gagne,
Systematic Entomology Laboratory, HBIII, Agric. Res. Serv., USDA re-
viewed and improved the manuscript.

Literature Cited

Aldrich, J. M. 1897. A collection of Diptera from Indiana caves. In Blatchley, W. S.,
Ed. The fauna of Indiana caves. Indiana Dep. Geol. Nat. Resources, Ann. Rep.
(1896) 21:175-212.

Borgmeier, T. 1964. Revision of the North American Phorid flies. Part II. The

species of the genus Megaselia, subgenus Aphiochaeta. Stud. Entomol. 7:

257-416.

1966. Revision of the North American Phorid flies. Part III. The species
of the genus Megaselia, subgenus Megaselia. Stud. Entomol. 8(1965):1—160.

Lundbeck, W. 1922. Diptera Danica. Genera and species of flies hitherto found in
Denmark. Vol. 6: Pipunculidae, Phoridae. G.E.C. Gad, Copenhagen. 455 pp.

Malloch, J. R. 1914. Notes on Illinois Phoridae (Diptera) with descriptions of three

new species. Bull. Brooklyn Entomol. Soc. 9:56—60.

1919. A new species of Phoridae from Illinois. Can. Entomol. 51:256—

257.

Schmitz, H. 1956. Phoridae. In Linder, E., Ed. Die Fliegen der palearktischen Re-
gion. Vol. 4, Pt. 7. E. Schweizerbart’sche Verlag, Stuttgart. Pp. 369-416
(Lief. 187).

Department of Entomology, Virginia Polytechnic Institute and State
University, Blacksburg, Virginia 24061.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 228-233

TWO NEW SPECIES OF AMERICAN ARADIDAE (HEMIPTERA)
Nicholas A. Kormilev

Abstract—The author proposes two new taxa, Aradus oregonicus new
species and A. usingeri new species, both from Oregon and both belonging
to the “Quilnus group.” A new record and the description of a male is given
for Eunotoplocoris ruckesi Kormilev (1957) from Peru.

By the kind offices of Dr. Harry Brailovsky, Instituto de Biologia
UNAM, Mexico City, Mexico, and Prof. Dr. John D. Lattin, Oregon State
University, Corvalis, Oregon, I have had an opportunity to study two
lots of American Aradidae from collections under their care, for which I
express my sincere gratitude.

Among other common species, two species of Aradus Fabricius, 1803,
from Oregon, belonging to the “Quilnus group,” were new and are de-
scribed in this paper.

The genus Eunotoplocoris Kormilev (1957) was described based on
a single female from Peru. Now I am able to give a description of a
male of the same species, Eunotoplocoris ruckesi Kormilev, 1957.

All measurements in this paper were taken by micromillimeter eyepiece,
25 units = 1 mm. In ratios the first figure represents the length and the
second the width of measured portion. The length of the abdomen in
Aradus F., for convenience, was taken from the tip of the scutellum to the
tip of the genital lobes.

Subfamily ARADINAE
Genus Aradus Fabricius, 1803

“Quilnus group” is characterized by the labium not extending beyond the
fore border of the prosternum and by the straight lateral borders of the
pronotum. It has so far only 10 species, of which three are from the
Nearctic and 7 from the Palaearctic Region. I am now adding two new spe-
cies from Oregon.

Aradus heidemanni Bergroth
Figs. 1-2

Aradus heidemanni Bergroth (1906) is rather common in Oregon and
Northern California.

Measurements of a brachypterous female from Oregon: Head 33:33;
relative length of antennal segments I-IV are 8:20:18:13; pronotum 17:50;
scutellum 35:20; abdomen 125:105. Total length 7.80 mm; width of pro-
notum 2.00 mm; width of abdomen 4.20 mm.

VOLUME 80, NUMBER 2 229

Figs. 1-2. Aradus heidemanni, female. 1. Pronotum, scutellum and hemelytra; 2.
Tip of abdomen from above. Figs. 3-4. Aradus oregonicus, female. 3. Pronotum,
scutellum and hemelytra; 4. Tip of abdomen from above. Figs. 5-6. Aradus usingeri.
5. Pronotum; 6. Tip of abdomen from above. Fig. 7. Eunotoplocoris ruckesi, male,

tip of abdomen from above.

Aradus oregonicus Kormilev, new species
Figs. 3-4

Description—Female. Brachypterous.
Head: As long as its width across eyes (30:30); anterior process com-
pressed laterally, reaching slightly beyond tip of antennal segment I; an-

230 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tenniferous tubercles acute, diverging, lateral tooth absent; postocular tooth
minute, rounded. Antennae 1’ as long as width of head across eyes (45.5:
30); antennal segment I obovate, II and III cylindrical, IV fusiform; rela-
tive length of antennal segments I-IV are 7.5:15:15:8. Labium reaching
hind border of labial groove, which is closed posteriorly.

Pronotum: Trapezoidal, shorter than its maximum width (15:45); an-
terior angles produced forward and rounded; lateral borders straight,
strongly converging and finely serrate; hind angles produced backward and
rounded; hind border evenly sinuate between hind angles. Disc with median
sulcus, flanked by 2 (1 + 1) short carinae, converging posteriorly, and
further laterad by 2 (1 + 1) oblique carinae; depressed along posterior
border medially.

Scutellum: Short, only slightly longer than its maximum width (25:20);
lateral borders raised and rounded, tip narrowly rounded; disc depressed.

Hemelytra: Abbreviated, without membrane; corium reaching tip of
scutellum.

Abdomen: Longer than its maximum width (95:86); postero-exterior
angles of connexiva II-VI not protruding, but lateral border is notched at
sutures between connexiva; postero-exterior angle VII is rounded. Genital
lobes subtriangular, rounded at tip and with a small tubercle on exterior
border.

Color: Black, dark brown on ventral side.

Size: Total length 6.64 mm; width of pronotum 1.80 mm; width of ab-
domen 3.44 mm.

Holotype —Female (Entomological Museum, Oregon State University)
Oregon, Linn Co., Lost Prairie, 3,800’; 38 mi E Sweethome; 29.VII.1958;
J. D. Lattin leg.

Discussion.—Aradus oregonicus is closely related to A. heidemanni but
is smaller; the antennae are relatively shorter, only 1.52 as long as width
of head across eyes (1.78 in A. heidemanni); the lateral tooth is absent; the
postocular tubercles are minute and rounded; the pronotum is less cut
out posteriorly. Scutellum is relatively shorter, almost ovate, with rounded
lateral borders. Lobes of VIII (genital lobes) are much shorter and are
more rounded posteriorly; the incisure between them is shallow with ratio
of depth of incisure to width of head across eyes as 5:30 (12:33 in A. heide-
manni).

Aradus usingeri Kormilev, new species
Figs. 5-6

Description—Female. Macropterous. Head, pronotum, scutellum, baso-
lateral borders of corium and base of veins, connexiva, tergum VII and
genital lobes, covered with short, erect, spiculoid granulation.

VOLUME 80, NUMBER 2 231

Head: Longer than its width across eyes (35:31); anterior process long,
compressed laterally, reaching beyond base of antennal segment II; an-
tenniferous tubercles acute, long, and slightly diverging, reaching ' of
antennal segment I. Lateral tooth absent; preocular tubercles minute, but
distinct; postocular small, rounded. Vertex with 2 (1 + 1) black, callous
spots, connected posteriorly and flanked by erect, spiculoid granulation.
Antennae long, as thick as in A. heidemanni, 1.7X as long as width of
head across eyes. Antennal segment I barrel-shaped, II and II cylindrical,
IV fusiform; relative length of antennal segments I-IV are 7:18:16:12.
Labium thin, reaching hind border of labial groove, which is open pos-
teriorly.

Pronotum: Hexagonal, less than % as long as its maximum width (27:
60); anterior border sinuate; anterolateral angles slightly produced forward,
rounded; lateral borders parallel, converging anteriorly; hind angles rounded
and slightly produced backward. Fore disc with 2 (1 + 1) horseshoe-
shaped black callosities and with 4 (2 + 2) thin ridges, flanked posteriorly
by 2 (1 + 1) short ridges.

Scutellum: Subtriangular, longer than its maximum width (41:30); lat-
eral borders raised and slightly convex, tip rounded, disc depressed.

Hemelytra: Complete, reaching % of tergum VIII; corium reaching
hind border of connexivum IV, clavus reaching tip of scutellum; basolateral
borders of corium expanded, rounded and reflexed; apical angle acute,
apical border convex forming 2 festoons; veins raised.

Abdomen: With subparallel, slightly convex sides, longer than its max-
imum width (100:90); posteroexterior angles of connexiva H-V not pro-
truding, VI protruding, VII forming an obtuse angle. Lobes of VIII sub-
triangular, deeply cleft; ratio length of incisure to width of head across
eyes as 12:31.

Color: Dark brown to black.

Size: Total length 8.28 mm; width of pronotum 2.40 mm; width of
abdomen 3.60 mm.

Holotype.—Female (Entomological Museum, Oregon State University),
Oregon, Corvalis, Benton Co., 14.IV.1958, on side walk; J. D. Lattin leg.

Etymology.—This species is dedicated to the memory of late Prof. Dr.
Robert L. Usinger, who contributed so much to our knowledge of Aradidae.

Discussion.—Aradus usingeri is related to A. heidemanni. It belongs to
the same “Quilnus group” as it has the labium not produced beyond the
hind border of the head. It may be separated from the latter by being
macropterous (females of heidemanni are brachypterous and males stenop-
terous), by having the pronotum hexagonal and not trapezoidal, by pos-
sessing differently shaped genital lobes, and by having curious, spiculoid
granulation.

232 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Subfamily MEZIRINAE
Genus Eunotoplocoris Kormilev
Eunotoplocoris ruckesi Kormileyv

Fig. 7

Description —Male. Head, pronotum and scutellum roughly granulate;
central dorsal plate and connexivum roughly punctured; body covered with
short, curled, rusty hairs; femora and antennal segments I-III with erect
bristles. Micropterous.

Head: Shorter than its width across eyes (61:65); anterior process ro-
bust, slightly constricted at sides, genae produced far beyond clypeus and
cleft, reaching % of antennal segment I. Antenniferous tubercles robust,
dentiform, divaricating, reaching basal % of antennal segment I. Eyes
globose and slightly stalked. Postocular tubercles minute, by far not reach-
ing outer borders of eyes; postocular borders behind them are long and
converging in slightly arcuate line. Infraocular carinae low, crenulate;
vertex raised and with a double row of granules. Antennae relatively
thin, with exception of Ist segment, which is stout and fusiform; II sub-
cylindrical, slightly enlarged apically and tapering at base; III cylindrical;
IV is missing; relative length of antennal segments I-III is 40:25:37.
Labium very short, reaching % of a deep labial groove, which is open
posteriorly.

Pronotum: Much shorter than its maximum width across anterolateral
lobes (26:82). Collar sinuate and granulate; anterior borders laterad of
collar sinuate; anterolateral angles produced into winglike lobes, straight
interiorly and rounded exteriorly, crenulate and produced far beyond collar.
Lateral borders are narrower than anterolateral lobes (80:82), parallel be-
tween themselves; hind border evenly arcuate. Fore disc with a shallow
median depression, flanked by 2 (1 + 1) flat, semicircular callosities, and
further laterad by 2 (1 + 1) high, ovate ridges. Hind disc is greatly reduced
to a narrow strip along hind border of pronotum, separated from fore disc
by a thin, transverse sulcus.

Mesonotum: Subtriangular in the shape of a scutellum, shorter than
its basal width (33:75), truncate posteriorly and raised medially, covered
with very rough granulation.

Metanotum: Consisting of 2 (1 + 1) plates, separated by mesonotum,
raised posteriorly and deeply depressed anteriorly.

Hemelytra: Reduced to small, elongate pads, placed laterad of meso-
notum.

Abdomen: Subquadrate, longer than its maximum width across segment
Il or V (148:135); tergum I raised, convex anteriorly and sinuate pos-
teriorly, separated from metanotum and tergum II by thin sulci. Tergum
II much wider than tergum I, flat, separated from quadrate central dorsal

VOLUME 80, NUMBER 2 233

plate and connexivum by thin sulci. Central dorsal plate consisting of
terga III-VI fused together, it is raised medially and laterally, with 2
(1 + 1) rows of round, callous spots placed in depressions; laterad of them
with 2 (1 + 1) rows of smaller, round, callous spots, also placed in depres-
sions in zigzag pattern. Connexivum roundly produced at segment II;
with parallel, but festooned borders from II-VI; converging at VII. Pos-
teroexterior angles of connexiva from H-VI rounded and protruding; form-
ing small, rounded and diverging lobes on VII. Scars of dorsal scent
gland openings placed on terga IV and V posteriorly. Tergum VII raised for
reception of a large hypopygium (25:33) with a flat, triangular, median
ridge, and semicircular, biheaded ridge surrounding the latter. Paratergites
clavate, reaching % of hypopygium. Spiracles ventral from I-VI and
placed far from border, ventral, but placed near border on VII, lateral and
visible from above on VIII. Metathoracic scent gland openings are large,
gaping, slightly curved and visible from above.

Legs: Unarmed, arolia absent.

Color: Uniformly black.

Size: Total length 11.20 mm; width of pronotum 3.28 mm; width of
abdomen 5.40 mm.

Material—Male (Instituto de Biologia UNAM, Mexico City, Mexico),
Peru, Cuzco; P. Reyes leg.

Literature Cited

Bergroth, E. 1906. Notes on American Hemiptera. Can. Entomol. 38:198—202.
Kormilev, N. A. 1957. Notes on Neotropical Aradidae (Hemiptera), 7. Two new
apterous Aradidae. Am. Mus. Novit. 1860:4.

Research Associate in Entomology, Bishop Museum, Honolulu, Hawaii
96818 (mailing address: 84-05 89th Street, Woodhaven, New York 11421).

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 234-239
DEFENDED HUNTING TERRITORIES AND HUNTING
BEHAVIOR OF FEMALES OF PHILANTHUS GIBBOSUS
(HYMENOPTERA: SPHECIDAE)

Norman Lin

Abstract—Females of the solitary wasp Philanthus gibbosus (F.) de-
fend hunting territories around the particular halictine bee nests they hunt
at, against conspecific females. Halictine bee prey may be taken from
within the nest or outside the bee nest when bees leave or return to
their nests.

Among birds, the females of most species have no territories of their own,
rather they adopt the territory of their mate and aid in the defense of his
territory. Also rare are birds which defend territories solely around specific
food sources such as the hummingbird which may abandon a flowery
shrub, which it had previously defended, when the blossoms fall. Thus
territory is abandoned when food supply fails (Klopfer, 1969). As de-
scribed in the present study, females of the digger wasp, Philanthus gib-
bosus (Fabricius), are at present apparently unique among hymenoptera, if
not territorial organisms in general, in defending individual territories about
the future food supply of their young from conspecific females.

While the life history and behavior of P. gibbosus is in general better
known than any other species of the genus, there are still many major
gaps in our knowledge. Reinhard (1924) reported halictid bee prey being
taken at flowers, and Alcock (1974) obtained indirect evidence of such
hunting by observing females flying from flower to flower and in one case
noted a female carrying a halicitid and a tufted dandelion seed which he
believed indicated the capture was made on a flower. In other areas he
observed bees taken at a large halicitid colony. Peckham and Peckham
(1905) reported that the wasps took guard bees indicating that captures also
occur near or on the ground; Evans and Lin (1959) observed one female
of P. gibbosus, in the course of 10 minutes, enter the nest of an unidentified
halictine bee four times and each time came out with a bee she took to her
own nest. The most common prey species they noted was Lasioglossum
zephyrum (Smith), there being 15 specimens. Augochlora pura (Say) was
one of the least common species, there being three specimens. They con-
ducted their study in Ithaca, New York. Barrows and Snyder (1973) studied
the contents of 15 cells provisioned by Philanthus gibbosus in a vertical bank
containing nest aggregations of Lasioglossum zephyrum located in Douglas
County, Kansas. They found that eight species of halictine bees were used
and L. zephyrum was one of the three species represented by only one speci-
men. This in contrast with the 53 specimens of Augochlora pura, the most

VOLUME 80, NUMBER 2 235

commonly encountered species. They concluded that the wasp did not hunt
near their nest and did not enter Lasioglossum zephyrum burrows to capture
prey. They further suggested that perhaps Philanthus gibbosus hunts pri-
marily around flowers as do P. triangulum (Fabricius) (Tinbergen, 1932)
P. politus Say (Evans and Lin, 1959) and P. bicinctus (Mickel) (Armitage,
1965).

Alcock (1974) studied an aggregation of P. crabroniformis Smith in a lot
in Seattle, Washington. A smaller aggregation of P. gibbosus was inter-
spersed in this aggregation. He reported that P. crabroniformis primarily
captured bees by striking them in flight as they returned to their nest, and he
placed his major emphasis on this method of bee capture which has not
previously been investigated in P. gibbosus and is unknown in the literature
of P. gibbosus.

General Characteristics of Hunting Behavior

Hunting behavior of P. gibbosus was studied intensively during the active
season in the vicinity of a vertical sandbank in which numbers of both
wasps and an unidentified halictine bee nested in number. On 30 Au-
gust between 10:54 AM and 5:36 PM observations were made, barring a few
absences of usually less than an hour, of the attacks by wasps on the 8
marked, and by far most frequently attacked bee nests (A to H) on the
sandhill. Not all attacks were observed due to the temporary absences and
since observations on other aspects of behavior were being studied si-
multaneously. However the representation is an accurate one since similar
observations were made all summer during the active season of the wasp.
The only failing is that on this particular date the wasps were unsuccessful
in capturing even a single bee. However all attacks were made at the nest
entrance whether guarded or not, and wasps, frequently unsuccessful at one
nest, went from nest to nest and occasionally a wasp entered an unguarded
nest and withdrew without prey. Experience has shown that by far the
most successful attacks are made against bees assuming flight in leaving
their nests or on their return to their nests, attacks not observed to occur
on this date. However in the past bees were observed being taken from
within the nest. A total of 63 attacks were observed to have been made
at the entrance of bee nests, whether guarded or not. In attacking, wasps
palpitate the head of the guard or the empty nest entrance with their an-
tenna; and when the guard blocks the entrance with its abdomen, wasps
have been observed attempting to grasp the bee by the abdomen with
their mandibles and to pull it out. They also on occasion vigorously engage
in biting away at the walls of the guarded nest entrance in attempts to gain
entry. There were four distinct episodes in which wasps entered an un-
guarded bee nest. All occurred in nest A which had the largest entrance of
all 8 nests. In one case a wasp entered nest A six times, each time coming

236 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

out in seconds. The same was repeated two times by presumably another
wasp.

On 25 September a rather large halictine bee was caught in midflight by
a female P. gibbosus. The bee was caught in the vicinity of the bee nest
area where other bees were hovering in flight. On capture the pair fell
approximately six inches to the slope below and the wasp was seen to
have its sting pushed up against the bee and definitely appeared to sting
the bee. The wasp then picked up the bee and flew approximately 7 feet
to enter its nest in the slope. Another mid-air attempt at capture by an-
other wasp failed and the bee then took evasive zig-zag flight. Such un-
successful mid-air capture is a common occurrence.

On 3 September a female P. gibbosus was on the slope, and a number of
halictine bees were flying around. One bee flew near the wasp approxi-
mately 4 inches over the ground. The wasp pounced on the bee, malaxated
it and brought the tip of her abdomen up to it in what was a probable
stinging effort; and in about three seconds flew off with the bee. On
3 September in another instance a halictine bee flew about 2 inches over
the slope and a P. gibbosus flew up and pounced on her and the two fell,
locked together, to the slope and rolled down the slope for approximately
4 inches; and after about 4 seconds from the time of capture, the wasp
flew off with the bee. The wasp did appear to malaxite the bee and also
I believe it had the tip of its abdomen in probable stinging behavior pressed
against the bee. On 28 August a number of successful captures of guards
were made at nest A, the nest with the largest entrance which permitted
the entry of all but the larger female wasps.

A wasp probed at a halictine bee nest entrance and palpated the head
of the guard with her antennae. The guard turned to block the entrance
with her abdomen and the wasp soon left. Another wasp suddenly grabbed
a small bee on the outside of her nest and seemed to malaxite it ap-
proximately 10 seconds and to fly off with it; and though it had curved
its abdomen up to the stinging position, it appeared that it did not actually
sting the bee. A few minutes later a wasp was seen to enter nest A. The
female P. gibbosus was small enough to enter. This is only one case of the
individual differences in hunting techniques among female P. gibbosus.
Others involved waiting for returning bees at their nests, pouncing at bees
leaving their nests, making mid-air captures and hunting at flowers or
elsewhere as an individual habit. The territories, to be discussed, briefly
mention that individual wasps have favorite nests around which they
hunt. A wasp entered nest A, and a minute later a wasp entered the
same nest and squeezed approximately half way in and pulled a guard out
and seemed to malaxite it a few seconds and then immediately flew off
with it. I definitely believe this bee was not stung. This nest immediately
had a new guard in its absence. In a few minutes a wasp entered nest A
and in seconds pulled out a bee, malaxited it a few seconds and flew

VOLUME 80, NUMBER 2 237

with it to its nearby nest in the same sand slope. Immediate stinging in
P. gibbosus is often far from the rule. I have; on a number of occasions,
caused a wasp to drop a bee she was carrying in flight to her nest; and, on
being dropped, the bee proved to be uninjured, recovered and flew away.

General Characteristics of Hunting Territories

Females in flight over their vertical bank hunting and nesting area behave
entirely differently toward other females below in the area in comparison
to females located below them in the nesting aggregation of an open field
in Brooklyn where the nests of bees are not close by their prey nests. A fe-
male coming on another female even as much as 2 feet away and the
latter at a bee nest on the slope of her hunting area pounces upon the
latter and either chases her off of they both fly off or the pouncer flies
off. The latter much in the manner of territorial male cicada killer wasps
perching in their territory being pounced on by rival males and almost
invariably pursuing the rival male which pounced on the owner (Lin, 1963)
or the same may be observed among territorial males of Polistes exclamans
Viereck (Lin, 1972). Females even pursue other females in the air. The
following instance represents a case seemingly identical to two territorial
male cicada killer wasps which, both being over their own territory, fly
aggressively at each other and grapple (Lin, 1963). Two females engaged
in hunting behavior were flying inches over the slope and after about 5
seconds came close to each other and when a few inches apart flew at
each other, met in mid-air, grappled, fell to the sand below, continued
to wrestle for about a second or two, separated and both flew off. In another
instance a female wasp was probing nest G when a second female flew
toward G and the first female flew up and made contact with the intruder
when the latter was approximately 2 inches from G. The argument might
be made that females attacking other females are acting toward them as
bee prey rather than territorial rivals. Reasons have already been cited
making this most unlikely; one further reason involves the response of a
female to a male which is nearly identical in appearance to the female.
A male was on the mound of a nest which it had just closed, and it began
flight. A female was engaged in hunting and was at the entrance of nest
A. She spied the slow flying male and flew at the male but she did not hit
him as she obviously would have done to a female; (but apparently recog-
nized him as a male probably by chemical means) she continued to fly
toward him but when approximately one inch from the male she broke off
the pursuit without making contact.

Also recorded on 30 August were the number of aggressive attacks by
one female at another or mutual attacks, the minimum of these were 26 be-
tween 2:06 AM and 5:26 PM with some large time lapses when the ob-
server was not present or when his attention was directed in recording

238 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

other aspects of behavior. At 3:51 PM in about one 3-minute period, four
aggressive female encounters occurred including chasing in the air, pounc-
ing on the ground, and several wrestlings. So much female-female ag-
gressive behavior occurred in that brief span because as many as four fe-
males were hunting at the same time in the small area of the vertical bank.
At 4:06 PM much the same occurred, four females were hunting at the same
time (at 4:12 PM five females were hunting at the same time and at 4:34
PM six females were similarly engaged), and I believe I observed greater
hunting activity than I ever saw at any other time and also far more female
aggressiveness than ever before. I didn’t even record the number of en-
counters but certainly there were in that space of minutes at least five and
probably closer to eight. I believe that during this time of day the highest
frequency of hunting behavior and female P. gibbosus territorial aggres-
sive behavior occurs. This might in fact be what is to be anticipated since
it is probably the time when the bees return to their nest for the day after
gathering pollen; it is also the time of day that the territorial females of
Philanthus gibbosus “wait” for them around their favorite bee nests in their
territories.

One medium or medium large female was marked with paint on 4 Sep-
tember. This was female 2. Female 2 was observed almost daily from
4-14 September in the vicinity of her hunting territory where she par-
ticularly localized around bee nest H which she often sat next to or attacked
the guards in the entrance or pursued some passing bee in flight to or
from the nest. She vigorously defended a territory about 2 square feet
encompassing this area.

Discussion

Long term studies of P. gibbosus have revealed that 1972 was a year
of exceedingly large populations of these wasps in Brooklyn. Halictine
bees were also extremely common during that year. While untrue in other
years, the major source of halictid bee mortality in the Brooklyn areas
studied was due to the predation of P. gibbosus (for a comparison of the
causes of halictine bee mortality in previous years in Brooklyn see Lin,
1964 and 1964-65). Hunting territories among P. gibbosus females was
never observed in the large 40 acre field in Brooklyn (1956-1976) where
years of data were gathered but was observed in the crowded conditions
in a vertical bank in 1972 also in Brooklyn where both wasps and an un-
identified species of halictine bee nested in large numbers.

Hunting territories probably arise under conditions of dense P. gibbosus
populations when they are located in banks which also house the nests
of halictine bees. A safe prediction is that such territories do not exist
in large fields where intensive observation over the years has failed to re-

VOLUME 80, NUMBER 2 239

veal any female-female aggression other than that associated with en-
croachment by one female of another female’s nest where prey species of
bees are probably taken at flowers or dispersed nests. In years when P.
gibbosus is scarce, hunting territories are probably nonexistent over banks
inhabited by both wasps and halictine bees, especially should the latter
be in excessive amount. This may be the explanation for the findings of
Barrows and Synder (1973) that Lasioglossum zephyrum was abundant
in the nesting banks of Philanthus gibbosus and that the wasps apparently
did not hunt near their nest which suggested that competition for prey
was not great. Only prey shortage or predator abundance or both under
these conditions may elicit territorial behavior.

Hunting territories in other digger wasps are here postulated to be
uncommon due to the transitory nature of the location of their food supply
as opposed to a rigidly fixed source in Philanthus provided by the permanent
bee nests located in a small concentrated area of a bank or elsewhere in the
current case.

Literature Cited

Alcock, J. 1974. The behavior of Philanthus crabroniformis Smith (Hymenoptera:
Sphecidae). J. Zool. 173:233-246.

Armitage, K. B. 1965. Notes on the biology of Philanthus bicinctus (Hymenoptera:
Sphecidae). J. Kans. Entomol. Soc. 38:89-100.

Barrows, E. M., and T. P. Snyder. 1973. Halictine bee prey of Philanthus gibbosus
(Hymenoptera: Sphecidae) in Kansas. Entomol. News 84:314—-316.

Evans, H. E., and C. S. Lin. 1959. Biological observations on digger wasps of
the genus Philanthus (Hymenoptera: Sphecidae). Wasmann J. Biol. 17:115-132.

Klopfer, P. H. 1969. Habitats and territories. Basic Books, Inc. New York.

Lin, N. 1963. Territorial behavior in the cicada killer wasp Sphecius speciosus
(Drury) (Hymenoptera: Sphecidae). Behaviour 20:115-133.

—. 1964. Increased parasitic pressure as a major factor in the evolution of

social behavior in halictine bees. Insectes Soc. 11:187—192.

1964-1965. The use of sand grains by the pavement ant Tetramorium
caespitum while attacking halictine bees. Bull. Brooklyn Entomol. Soc. LIX—LX:
30-34.

—. 1972. Territorial behavior among males of the social wasp Polistes exclamans
Vierick (Hymenoptera: Vespidae). Proc. Entomol. Soc. Wash. 74:148-155.

Peckham, G. W., and E. G. Peckham. 1905. Wasps social and solitary. Houghton
Mifflin. New York. 311 pp.

Reinhard, E. G. 1924. The life history and habits of the solitary wasp Philanthus
gibbosus. Smithson. Inst. Annu. Rep. 1922:363-376.

Tinbergen, N. 1932. Ueber die Orientierung des Bienewolfs. (Philanthus triangulum
Fabr.). Z. Vgl. Physiol. 16:305-334.

1487 East 37th Street, Brooklyn, New York 11234.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 240-263
A NEW GENUS AND TWO NEW SPECIES OF ASTEROLECANIID
SCALE INSECTS ON PALM FROM COLOMBIA AND TRINIDAD
(HOMOPTERA: COCCOIDEA: ASTEROLECANIIDAE)

Douglass R. Miller and Paris L. Lambdin

Abstract —A new genus, Grammococcus, and two new species, G. adeto-
corymbus and G. corymbus, are described and illustrated. Affinities and a
diagnosis of the genus are also provided.

A new genus is described for two new species of unusual palm inhabit-
ing asterolecaniid scale insects. Because both species were found in large
populations on their hosts and because palms are frequently grown as
ornamentals in the United States, we felt that it was important to describe
these potential pest species.

Methods and Depositories

Numbers and measurements were taken from 10 specimens and are the
average rounded off to the nearest whole number followed by the range
in parentheses. Measurements are given in microns.

Specimens are deposited in: British Museum (Natural History), London
(BM); California Department of Agriculture, Sacramento (CDA); Florida
State Collection of Arthropods, Gainesville (FSCA); Museo de Historia
Natural de la Ciudad de Mexico, Mexico City (MNC); Museum National
d'Histoire Naturelle, Paris (MNHN); South African National Collection
of Insects, Pretoria (SA); University of California, Davis (UCD); The
University of Tennessee, Knoxville (UT); U.S. National Museum of Natural
History, Washington, D.C. (USNM); Virginia Polytechnic Institute and
State University, Blacksburg (VPI); and Zoological Institute, Academy of
Sciences of USSR, Leningrad (ZI).

Grammococcus Miller and Lambdin, new genus

Type-species.—Grammococcus adetocorymbus Miller and Lambdin, new
species.

Type-locality.—St. Clair, Port-of-Spain, Trinidad.

Affinities —Grammococcus appears to be closely related to Polea Green.
Similarities shared by adult females in the two genera are: Lack of large
8-shaped pores on submargin, presence of submarginal pore clusters, ir-
regularly spaced minute 8-shaped pores on dorsum and bilocular pores
clustered about mouth parts on venter. Grammococcus may be separated
from Polea by the structure of the anal ring, the type of pores in the sub-

VOLUME 80, NUMBER 2 241

marginal pore clusters on the dorsum, the trilocular pores laterad of the
spiracles, the number of labial setae, the reduced number of transverse
rows of multilocular pores, and the lack of a submarginal band of quinque-
locular pores on the venter. For a detailed treatment of Polea see Lambdin
(1977).

Etymology.—The generic name is from the Greek gramme meaning “line”
and coccus (latinized) meaning “seed or scale insect.” The name refers to
the lines of dorsal tubular ducts characteristic of this genus.

Third-Instar Females (Adult)

Diagnosis.—Quinquelocular pores in clusters on dorsosubmargin, large
8-shaped pores absent, minute 8-shaped pores and simple pores present,
tubular ducts in 2 longitudinal lines on each side of body. Venter with
unsegmented antennae, bilocular pores near mouthparts, legs absent, tri-
locular pores near spiracle, multilocular pores in 2 or 3 transverse rows
near vulva and submarginal 8-shaped pores in a submarginal row.

Adult females differ from other instars by the following combination:
With a vulva, unsegmented antennae, ventral multilocular pores, dorsal
tubular ducts, dorsal pore clusters, minute 8-shaped pores, row of tri-
locular pores in each spiracular furrow and bilocular pores; without legs,
marginal 8-shaped pores and dorsal multilocular pores. Adult females are
similar to the 2nd-instar males but are separated by having a vulva, spirac-
ular furrows and multilocular pores.

Second-Instar Females

Diagnosis —Dorsosubmargin with quinquelocular pores in clusters, large
8-shaped pores in medial area, simple disc pores irregularly spaced,
tubular ducts absent. Venter with bilocular pores, minute 8-shaped pores,
and pores near spiracles absent, antennae unsegmented, submarginal 8-
shaped pores extending from cephalothorax to posterior abdominal seg-
ments.

Second-instar females differ from other instars by the following com-
bination of characters: Without a vulva, dorsal and ventral multilocular
pores, dorsal tubular ducts, minute 8-shaped pores, spiracular pores, mar-
ginal 8-shaped pores, legs and bilocular pores; with dorsal pore clusters
and 2 medial rows of large 8-shaped pores. Second-instar females are
similar to 2nd-instar males but differ by lacking dorsal tubular ducts,
minute 8-shaped pores and bilocular pores.

First Instars

Diagnosis—Dorsum with 2 pairs of multilocular pores, 8-shaped pores
in medial and submarginal areas, simple disc pores. Venter with sub-

242 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

marginal tubular ducts, legs with trochanter and femur fused, antennae
segmented.

First instars differ from other instars by the following combination of
characters: With 2 pairs of dorsal multilocular pores, legs, segmented an-
tennae, marginal and submedial rows of large 8-shaped pores and a tri-
locular pore near each spiracle; without a vulva, ventral multilocular pores,
submarginal 8-shaped pores, dorsal tubular ducts, dorsal pore clusters and
bilocular pores. Well-developed legs separate lst instars from all others
except 3rd to 5th-instar males which have either wing buds or wings.

Fifth-Instar Males (Adult)

Diagnosis —Penial sheath short (Fig. 4), with 2 pairs of eyes, antennae
9-segmented, reticulate pattern on several areas of head and thorax.

Fifth-instar males differ from other instars by having a well-developed
penial sheath, sclerotized thorax, 2 pairs of eyes, and 9-segmented an-
tennae; by lacking mouthparts, pores and ducts. Fully developed wings
immediately separate adult males from other instars.

Fourth-Instar Males (Pupae)

Diagnosis.—Pores and ducts absent, antennae 9-segmented, legs 6-seg-
mented, dermal nodules on dorsum.

Fourth-instar males differ from other instars by the following combina-
tion of characters: With 9-segmented antennae, 6-segmented legs, wing
buds and dermal nodules; without pores, ducts and mouthparts. Fourth-
instar males differ from 3rd-instar males by the latter’s having $-segmented
antennae and 4-segmented legs.

Third-Instar Males (Prepupae)

Diagnosis.—Pores and ducts absent, antennae 8-segmented, legs 4-seg-
mented, dermal nodules on dorsum.

Third-instar males can be distinguished from all other instars by having
characters given in diagnosis and by lacking pores, ducts, and mouth-
parts. Third-instar males differ from 4th instar males by the latter’s having
9-segmented antennae and 6-segmented legs.

Second-Instar Males

Diagnosis —Dorsum with clusters of quinquelocular pores on submargin,
S-shaped pores in medial area, minute 8-shaped pores and simple disc
pores. Venter with unsegmented antennae, bilocular pores near mouth-

VOLUME 80, NUMBER 2 243

parts, pores absent near spiracles and 8-shaped pores in submarginal band.
Similar morphologically to adult female but distinguished by absence of
vulva, spiracular furrows and multilocular pores.

Second-instar males differ from other instars by the following combina-
tion of characters: With dorsal tubular ducts, minute 8-shaped pores, un-
segmented antennae and bilocular pores; without a vulva, dorsal and ventral
multilocular pores, spiracular pores, marginal 8-shaped pores and legs.
Second-instar males are similar to 2nd-instar females but differ by having
dorsal tubular ducts, minute 8-shaped pores and bilocular pores.

Grammococcus adetocorymbus Miller and Lambdin, new species
Third-Instar Females (Adult)
Fig. 1

Type material—Holotype adult female on slide with 31 other specimens:
Left label, “Grammococcus adetocorymbus Miller and Lambdin, Holotype,
Paratypes; on palm, Dept. Agr. Grounds, St. Clair, Port-of-Spain A-1035
Trinidad, Nov. 22-18, H. Morrison”; right label gives map of specimens
on slide, locates position of holotype and states “Holotype, Paratype.”
There are 368 paratypes mounted on 30 slides with the same data as the
holotype. Holotype and several paratypes are deposited in USNM; 1 para-
type slide is deposited in each of the following: BM, CDA, FSCA, MNC,
MNHN, SA, UCD, UT, VPI and ZI.

Field features —Occurring on foliage, apparently abundant.

Body measurements.—Holotype mounted, 561 long (paratypes 683 (549-
§30)), 354 wide (paratypes 529 (342-639)).

Dorsum.—Longest anal-lobe seta 33 long (paratypes 31 (23-35)); seta
mesad of longest seta normally touching anal-ring sclerotization, 10 long
(paratypes 11 (9-13)); other dermal setae absent. Large 8-shaped pores
normally absent, present on 8 of 100 randomly selected specimens, presumed
part of exuviae of previous instar. Minute 8-shaped pores irregularly
scattered, about 2 long. Simple disc pores forming 2 pairs of irregular
longitudinal lines, about 30 on each side of body. Paired, simple disc pores
absent. Pore clusters loose, not as compact as on 2nd instar, frequently with
spaces between pores (posterior 2 clusters scattered), pores primarily quin-
quelocular. Number of quinquelocular pores in clusters; 6-7 pores in each
anterior, cephalothoracic cluster (paratypes 6 (2-12)); 7-8 pores in each
posterior, cephalothoracic cluster (paratypes 7 (4-14)); 11 and 12 in each
anterior, abdominal cluster (paratypes 12 (8-18)); and 5 and 6 in each pos-
terior, abdominal cluster (paratypes 9 (4-14)); 32 and 30 quinquelocular
pores on each side of body (paratypes 35 (21-51)). Tubular ducts arranged
in 2 pairs of longitudinal rows, 1 pair submedial and 1 pair mediolateral;

244 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 2 245

40 and 46 on each side of body (paratypes 50 (41-66)); longest duct 33 long
(paratypes 35 (23-43)).

Venter——Antennae 10 long (paratypes 10 (8-13)); each with 1 long,
fleshy seta and 2 short, slender setae. Clypeolabral shield 90 long (paratypes
94 (88-105)), 63 wide (paratypes 68 (63-75)). Labium nearly square, 40 long
(paratypes 41 (38—-48)), 40 wide (paratypes 44 (40-50)); with 2 pairs of minute
setae. Setae in transverse rows on posterior abdominal segments. Spiracles
with associated trilocular pores in spiracular furrow from spiracle to body
margin, 10 and 16 pores in posterior furrows (paratypes 12 (9-14)). Bilocular
pores laterad of mouthparts near junction of labium and clypeolabral shield,
6 and 7 pores on each side of shield (paratypes 7 (5-9)). Submarginal 8-
shaped pores arranged in band extending from anterior of antennae to
abdominal segment VIII or IX, 40 and 52 on each side of body (paratypes
52 (37-62)). Multilocular pores normally 10-locular, rarely 9-, 8-, or 7-
locular; 13 pores (paratypes 12 (9-13)) on abdominal segments VI-VIII.
Anal ring “C” shaped, lateral sclerotized areas without central clear area,
lateral sclerotizations joined by anterior sclerotized bar; 2 pairs of robust
setae of equal length, about 20 long (paratypes 21 (18-23)).

Variation—The paratypes normally have the dorsal clusters of quinque-
locular pores more scattered than on the holotype; the antennal setae
vary, some antennae have 2 fleshy setae and 1 thin seta, some have 1 thin,
long seta, 1 thin, short seta, and 1 fleshy seta; pores in the spiracular
furrows occasionally have 4 or 5 loculi.

Notes.—The above description is based on 182 specimens. The adult fe-
male of G. adetocorymbus differs from G. corymbus by having the lateral
areas of the anal ring connected anteriorly by a thin sclerotized bar, dorsal
clusters of quinquelocular pores, frequently with spaces between the pores,
7 (5-9) bilocular pores, and 52 (37-62) submarginal 8-shaped pores. Gram-
mococcus corymbus differs by having the lateral areas of the anal ring
separate, not connected by a sclerotized bar, dorsal clusters of quinque-
locular pores with pores closely appressed, without spaces between the
pores, 11 (9-14) bilocular pores, and 31 (24-36) submarginal 8-shaped
pores.

Etymology.—The species epithet is from the Greek adetos meaning “un-
bound or loose” and korymbos meaning “cluster of flowers.” The name refers
to the loose clusters of flowerlike pores typical of this taxon.

<

Fig. 1. Grammococcus adetocorymbus, adult female. A, cluster of quinquelocular
pores; B, tubular duct; C, minute 8-shaped pore; D, simple disc pore; E, antenna; F.
bilocular pore; G, trilocular pore; H, submarginal 8-shaped pore; I, multilocular pores;
J, anal ring.

246 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 2 247

Second-Instar Females
Fig. 2

Body measurements.—Mounted, 473 (397-591) long, 286 (226-366) wide.

Dorsum.—Longest anal-lobe seta 29 (25-33) long; seta mesad of longest
seta 7 (5-8) long; other dermal setae absent. Anal ring normally composed
of 2 lateral, sclerotized pieces joined posteriorly and/or anteriorly by
thin sclerotized area, each lateral piece with 2 minute setae 3 (2-4) long.
Large 8-shaped pores, when present, in 2 longitudinal rows on submedial
area of dorsum, each line with about 12 pores; number of pores variable, ap-
parently part or all may slough off during moulting process possibly due
to presence of fungus mycelia; largest pore on each specimen 11 (10-12)
long, 7 (7-8) wide; smallest pore 8 (7-8) long, 5 (5-6) wide. Simple disc
pores forming mediolateral, longitudinal row on each side of body; 11 or
12 in each line. Paired, simple disc pores absent. Pore clusters compact,
clusters of quinquelocular pores normally located as illustrated; 6 (2-10)
pores in each anterior, cephalothoracic cluster; 4 (2-6) pores in each pos-
terior, cephalothoracic cluster; 5 (3-9) in each anterior abdominal cluster;
and 5 (3-7) quinquelocular pores in each posterior abdominal cluster; 20
(11-30) pores on each side of body.

Venter—Antennae 9 (8-12) long; each with 1 long, fleshy seta and
2 short, slender setae. Clypeolabral shield 72 (65-80) long, 56 (53-63)
wide. Labium nearly rectangular, 37 (35-38) long, 34 (33-38) wide; with
2 pairs of minute setae. Setae forming submedial, longitudinal line on pos-
terior 2-4 segments and an occasional marginal line on last 1 or 2 segments.
Submarginal 8-shaped pores represented by 21 (18-24) on each side of
body. Anal ring normally composed of small anterior and/or posterior bar
connecting lateral areas, ring with 2 pairs of setae.

Notes.—The above description is based on 46 specimens. Second-instar
females of G. adetocorymbus and G. corymbus are very similar. The anal
ring of G. adetocorymbus has 2 pairs of setae and normally a small anterior
and/or posterior sclerotized bar connecting the lateral areas. On G.
corymbus the anal ring lacks setae and sclerotized bars connecting the
lateral areas. Also, simple pores are restricted to thoracic region in G.
corymbus while they are arranged in mediolateral longitudinal lines extend-
ing from the anterior area of head to the anal lobes in G. adetocorymbus.

Fig. 2. Grammococcus adetocorymbus, second-instar female. A, cluster of quinque-
locular pores; B, large 8-shaped pore; C, simple disc pore; D, anal ring; E, antenna; F,
submarginal 8-shaped pore; G, seta.

248 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ve

\
8 | A
Y, A" Pica
6 ° 8 \ Ags.
ey

if 8

VOLUME 80, NUMBER 2 249

First Instars

Fig. 3

We have been unable to separate Ist instars of G. adetocorymbus and
G. corymbus and have included only one description.

Body measurements.—Mounted, 329 (293-360) long, 157 (140-183) wide.

Dorsum.—Longest anal-lobe seta 31 (30-33) long, seta mesad of longest
seta 9 (8-13) long. Normally 3 marginal and 1 medial setae on each side
of head. Anal ring variable, normally crescent shaped and without setae
(of 70 specimens, 3 possessed anal-ring setae, all of G. adetocorymbus),
some specimens without crescent-shaped sclerotization. Eyes slightly sclero-
tized. Large 8-shaped pores arranged in longitudinal lines: Marginal lines
each composed of 14 pores, each pore with adjacent sclerotized area,
posterior 3 pores with associated simple pore near adjacent sclerotized
area; submedial rows each composed of 11 (7-13) pores, without adjacent
sclerotized areas. Simple disc pores forming 1 mediolateral line on each
side of body, 11 or 12 pores in each line. One paired simple disc pore
anterior of each eye near body margin. Multilocular pores on mediolateral
area of each side of body; anterior pair of pores near junction of head and
thorax, posterior pair on intersegmental line between segments III and IV;
each pore with 9 or 11 loculi, 9 (8-10) in diameter.

Venter—Antennal segmentation unclear, apparently 6-segmented, 48 (45-
52) long; setae normally as on Fig. 3, occasionally fleshy seta absent on sub-
apical segment. Clypeolabral shield 52 (48-55) long, 39 (37-42) wide. La-
bium nearly rectangular, 29 (26-32) long, 31 (29-36) wide. Legs with tro-
chanter and femur fused; tibia and tarsus fused or separated by weak line;
tarsus with deltoid sensilla near junction of tibia and tarsus; tarsal and
claw digitules capitate, extending beyond tip of claw; claw without denti-
cle. Setae rare, 1 medial seta near antennal base, 6 or 7 minute setae form-
ing submarginal, longitudinal line on abdomen. Spiracle with 1 associated
trilocular pore. Submarginal tubular ducts arranged in a longitudinal
line on each side of body, 8 in each line; submarginal 8-shaped pores repre-
sented by 1 pore near base of each antenna.

Notes.—The above description is based on 68 specimens of G. adeto-
corymbus and 2 of G. corymbus.

<

Fig. 3. Grammococcus adetocorymbus and G. corymbus, first instar. A, simple
disc pore; B, paired simple disc pores; C, seta; D, multilocular pore; E, large 8-shaped
pore; F, large 8-shaped pore with adjacent sclerotized area; G, large 8-shaped pore with
associated simple disc pore; H, anal ring; I, submarginal 8-shaped pore; J, antenna;
K, trilocular pore; L, submarginal tubular duct; M, tarsal claw; N, seta.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

250

t
WY My

aN

VOLUME 80, NUMBER 2 251

Fifth-Instar Males (Adult)
Fig. 4a

Body measurements.——Mounted, 689 (647-738) long, 298 (262-329) wide.

Description—Dorsum with several hairlike setae in each tegular area, sin-
gle seta in sublateral area of segments III, IV, or V-VII, 2 or 3 setae in each
sublateral area of segment VIII. Ventrally setae on ocular sclerite anterior of
ventral eye forward to lateral arm of midcranial ridge, and on mediolateral
area of abdominal segments IV or V-VII. Penial sheath with setae scattered
near ventral slit, setae absent dorsally.

Head subcircular; midcranial ridge with lateral and ventral arms, ventral
arm extending from lateral arms to level of preocular ridge. Preocular
ridge short, extending from dorsal eye to articulatory process of antenna.
Postocular ridge well developed, originating near postoccipital sclerotiza-
tion dorsally, extending to posterior margin of head ventrally. Postoccipital
ridge weakly sclerotized or absent. Ocular sclerites and genae weakly
sclerotized. Dorsal head pores 4 (1-7), near base of each antenna. Pos-
terior tentorial pits present. Dorsal eye 30 (28-33) in diameter, ventral eye 31
(25-33) in diameter. Reticulation on both surfaces of head.

Antenna 435 (421-451) long, about 0.6 as long as body length; apical
segments broader than other segments and 1.5x (1.4-1.6) as long as 3rd seg-
ment. Antennal setae predominately of thin, fleshy type; scape and pedicel
with hairlike setae; apical segment with 1 or 2 antennal bristles and 1 (1-3)
subapical sensory seta. Antennae apparently without placodic or basiconic
sensillae.

Front pair of legs shortest, middle pair normally slightly longer than hind
pair. Total length of trochanter, femur, tibia, tarsus and claw of each
leg as follows: Front 338 (313-368), middle 368 (345-390) and hind 363
(338-375). Each trochanter with 3 pairs of campaniform sensillae. Hind
tibia/tarsus 1.1 (1.1-1.2). Tarsi unsegmented, campaniform sensilla near
base of tibia, and with pair of capitate digtules which extend to tip of
claw. Claws without denticle; digitules of same size and shape as on
tarsus. Legs setae hairlike, without tibial spurs.

Prothorax separated from genae by constriction. Pronotal ridge con-
spicuous, occasionally dorsomedial area lightly sclerotized, nearly touching
proepisternum + cervical sclerite laterally; pronotal sclerites lightly sclero-
tized or absent. Posttergite apparently absent. Proepisternum + cervical
sclerite with anterior ridge articulating with postocular ridge; propleural
ridge well developed. Prosternum conspicuous, heavily sclerotized medially,
less definite laterally.

<

Fig. 4. Grammococcus adetocorymbus, adult male. A, dorsoventral view; B, G.
corymbus n. sp., dorsal view of penial sheath; C, ventral view of penial sheath.

252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Mesothorax with prescutum surrounded by well-developed prescutal
ridges, covered by reticulation pattern. Membranous area posterior of
prescutum. Small area of scutum adjacent of membranous area reticulated.
Scutellum with large internal foramen. Large membranous area between
scutellum and postnotum with noticeable reticulation. Mesopleural ridge
well developed. Episternum incompletely divided by membranous area,
subepisternal ridge weakly sclerotized, not reaching level of membranous
area. Lateropleurite well developed, bounded anteriorly by weakly sclero-
tized extension of marginal ridge of basisternum. Epimeron absent. Basi-
sternum lightly reticulated, divided by well-developed median ridge con-
necting marginal and precoxal ridges; furca large and well developed.

Metathorax without suspensorial sclerites and postnotum. Episternum
and epimeron composed of irregularly sclerotized areas on each side of
short pleural ridge; precoxal ridge absent. Metasternal plate composed of
weakly sclerotized area.

Wings 337 (325-368) long; hamulohalterae absent; without setae, circular
sensoria, or alar lobe.

Abdominal terga inobvious except on segment VIII; sterna irregular,
lightly sclerotized on segments II-VI, well developed on segments VII
and VIII. Dorsal setae on segment VIII on small protuberance, without
glandular pouch.

Genital segment unusually short for asterolecaniid (Giliomee, 1968;
Giliomee and Munting, 1968; Borchsenius, 1960; Russell, 1941); length 60
(58-63), width 63 (60-65); length/width 1.0 (0.9-1.0). Basal rod well de-
veloped. Ventral margin of capsule with conspicuous ridge, dorsal margin
unsclerotized, anus inconspicuous.

Notes.—The above description is based on 29 specimens; of the 29,
7 are in the exuviae of the previous instar. The most conspicuous difference
between G. adetocorymbus and G. corymbus is the shape and chaetotaxy
of the genital segment.

Fourth-Instar Males (Pupae)
Fig. 5

Body measurements.—Mounted, 711 (519-793) long, 422 (268-488) wide.

Dorsum.—Longest anal-lobe seta 25 (18-30) long; seta mesad of longest
seta 9 (5-15) long; other setae representing transverse rows in submarginal
areas of posterior abdominal segments. Dermal nodules near base of wings
and on medial areas of thorax and head. Wing buds 339 (317-354) long.

>

Fig. 5. Grammococcus adetocorymbus, pupa. A, dermal nodules; B, marginal seta;
C, submedial seta.

VOLUME 80, NUMBER 2 253

254. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 2 255

Venter —Antennae 333 (252-370) long. Setae forming mediolateral line
on posterior 3 or 4 abdominal segments on each side. Spiracles without
associated pores. Genital segment lightly sclerotized.

Notes.—The above description is based on 29 specimens.

Third-Instar Males (Prepupae)
Fig. 6

Body measurements——Mounted, 658 (543-732) long, 372 (244-445) wide.

Dorsum.—Longest anal-lobe seta 24 (23-25) long; seta mesad of longest
seta 10 (8-13) long; other setae on submarginal areas of posterior ab-
dominal segments. Dermal nodules near base of wings and on medial
areas of thorax and head. Wing buds about 210 long.

Venter—Antennae 137 (125-145) long. Setae forming mediolateral line
on posterior 3 or 4 abdominal segments on each side of body. Spiracles
without associated pores or furrows. Genital segment lightly sclerotized.

Notes.—The above description is based on 9 specimens.

Second-Instar Males
Fig. 7

Body measurements.—Mounted 489 (390-689) long, 310 (226-445) wide.

Dorsum.—Longest anal-lobe seta 24 (23-28) long; seta mesad of longest
seta 7 (5-9) long; other setae absent. Anal ring composed of 2 lateral,
sclerotized pieces connected by posterior and/or anterior bar, each lateral
piece with 2 minute setae 2 (1-2) long. Large 8-shaped pores, when pres-
ent, arranged in 1 pair of longitudinal lines on medial area of dorsum, each
line with 12 pores; pores variable as on 2nd-instar female; largest pore on
each specimen 13 (11-13) long, 8 (8-9) wide; smallest pore 9 (8-11) long,
7 (6-8) wide. Minute 8-shaped pores scattered over surface, normally
absent near body margin and on mesal area; about 2 long. Simple disc
pores forming sublateral and submedial longitudinal line on each side of
body, submedial line restricted to thorax. Paired, simple disc pores absent.
Pore clusters with pores not closely appressed, normally 4 clusters on each
side of body, rarely 5, extra clusters each with 1 (1-5) pores; 5 (2-8) quin-
quelocular pores in each anterior, cephalothoracic cluster; 4 (1-6) in each
posterior, cephalothoracic cluster; 5 (1-8) in each anterior, abdominal
cluster; and 5 (1-9) in each posterior, abdominal cluster; 20 (6-29) pores on
each side of body. Tubular ducts forming 1 pair of submedial and 1 pair

<

Fig. 6. Grammococcus adetocorymbus, prepupa. A, dermal nodules; B, marginal
seta; C, submedial seta.

256 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 2 257

of submarginal, longitudinal lines on each side of body; 29 (25-32) on
each side of body; longest duct 28 (25-30) long.

Venter—Antennae platelike, difficult to measure; each with 3 small setae.
Clypeolabral shield 76 (70-80) long, 58 (55-63) wide. Labium nearly
rectangular, 36 (33-38) long, 39 (35-43) wide; with 2 pairs of minute setae.
Setae forming submedial and marginal longitudinal line on posterior 3
(2-5) segments. Bilocular pores normally lateral of junction of labium and
clypeolabral shield; 1 (0-2). Submarginal 8-shaped pores represented by
36 (30-43) on each side of body.

Notes——The above description is based on 21 specimens. Second-
instar males of G. adetocorymbus and G. corymbus are very similar. Gram-
mococcus adetocorymbus has the anal ring relatively well developed with
2 pairs of small setae, has 1 (0-2) bilocular pore and has the pore clusters
with the pores loosely arranged. Grammococcus corymbus has a poorly
developed anal ring which lacks setae, has 5 (3-6) bilocular pores and has
the pore clusters with closely appressed pores.

Grammococcus corymbus Miller and Lambdin, new species
Third-instar Female (Adult)
Fig. 8

Type-material—Holotype adult female on slide with 13 other speci-
mens: Left label, “Grammococcus corymbus Miller and Lambdin, Holotype,
Paratypes; on Elaeis quineesis Jackq. (Palmaceae), Melgar (Cund.), Co-
lombia, 29-III-1972. F. Mosquera coll.”; right label gives map of speci-
mens on slide, locates position of holotype and states “Grammococcus
corymbus Miller and Lambdin. Holotype, Paratypes; 14 adult 2.” There
are 146 paratypes mounted on 19 slides with the same data as the holotype.
Holotype and several paratypes are deposited in USNM; 1 paratype slide
is deposited in each of the following: BM, MNHN, SA, UCD, UT and ZI.

Field features —Occurring on foliage.

Body measurements.—Holotype mounted, 573 long (paratypes 526 (445-
604)), 549 wide (paratypes 479 (305-586)).

Dorsum.—Same as G. adetocorymbus except as follows: Longest anal-
lobe seta 28 long (paratypes 28 (25-34)); seta mesad of longest seta not
touching anal-ring sclerotization, 13 long (paratypes 10 (8-15)). Large
8-shaped pores on 1 of 94 specimens. About 26 simple disc pores on each
side of body. Pore clusters compact, without spaces between pores. Num-

<

Fig. 7. Grammococcus adetocorymbus, second-instar male. A, pore cluster; B, tubular
duct; C, large 8-shaped pore; D, minute 8-shaped pore; E, simple disc pore; F, anal
ring; G, antenna; H, bilocular pore; I, submarginal 8-shaped pore; J, seta.

258 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Grammococcus corymbus, adult female. A, pore cluster; B, tubular duct; C,
minute 8-shaped pore; D, simple disc pore; E, anal-lobe seta; F, antenna; G, sub-
marginal 8-shaped pore; H, bilocular pore; I, trilocular pore; J, seta; K, multilocular
pores; L, anal ring.

VOLUME 80, NUMBER 2 259

ber of quinquelocular pores: 8 and 5 in each anterior, cephalothoracic
cluster (paratypes 6 (3-13)); 9 and 5 in each posterior, cephalothoracic
cluster (paratypes 6 (3-9)); 8 and 8 in each anterior, abdominal cluster
(paratypes 7 (3-12)); and 7 and 5 in each posterior, abdominal cluster
(paratypes 6 (3-9)); 32 and 23 quinquelocular pores on each side of body
(paratypes 26 (18-47)). Tubular ducts in 2 pairs of longitudinal lines; 44
and 48 on each side of body (paratypes 42 (34-53)); longest duct 35 long
(paratypes 34 (30-38)).

Venter—Antennae 8 long (paratypes 9 (8-13)). Clypeolabral shield 83
long (paratypes 82 (80-88)), 63 wide (paratypes 61 (58-68)). Labium
40 long (paratypes 39 (38-43)), 35 wide (paratypes 39 (33-48)). Posterior
spiracular furrows with 11 and 13 trilocular pores (paratypes 12 (10-14)).
Bilocular pores represented by 9 and 12 pores on each side of clypeolabral
shield (paratypes 11 (9-14)). Submarginal 8-shaped pores represented by
28 and 33 on each side of body (paratypes 31 (24-36)). Multilocular pores
on posterior abdominal segments near vulva, 11 pores (paratypes 12 (10-
14)). Anal ring in 2 separate pieces, not connected by anterior sclerotized
bar, each half with central clear area; 2 pairs of robust setae, posterior pair
shortest; longest seta 15 (paratypes 14 (8-20)).

Variation—The paratypes occasionally have an additional cluster of
dorsal quinquelocular pores, the anal-ring setae equal in length, ventral
setae on the mediolateral areas of segments 4-9, and the pores in the spirac-
ular furrows with 4 or 5 loculi.

Notes.—The above description is based on 94 specimens. For a com-
parison of G. adetocorymbus and G. corymbus see “Notes” of the former
species.

Etymology.—The species epithet is from the Greek korymbos meaning
“cluster of flowers.” The name refers to the clusters of flowerlike pores
typical of this taxon.

Second-Instar Females
Fig. 9

Same as G. adetocorymbus except as follows:

Body measurements.—Mounted, 435 (323-555) long, 264 (195-348) wide.

Dorsum.—Longest anal-lobe seta about 25 long; seta mesad of longest
seta about 6 long. Anal ring composed of 2 lateral, sclerotized pieces
without anterior or posterior bars, without setae. Largest 8-shaped pores
in medial area about 9 long, 6 wide; smallest about 6 long, 5 wide. Simple
disc pores on thorax only, 3 or 4 submedial pores on each side of body.
Pore clusters compact: 7 (5-10) quinquelocular pores in each anterior,
cephalothoracic cluster; 4 (2-7) pores in each posterior, cephalothoracic

260 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 2 261

cluster; 6 (5-7) in each anterior, abdominal cluster; and 6 (4-7) in each
posterior, abdominal cluster; 23 (18-28) pores on each side of body.

Venter.—Antennae about 8 long. Clypeolabral shield about 64 long, 48
wide. Labium about 34 long, 32 wide. Submarginal 8-shaped pores repre-
sented by 20 (16-23) on each side of body. Anal ring without sclerotized
bars connecting lateral areas and without setae.

Notes.—The above description is based on 3 poor specimens. For a com-
parison of 2nd-instar females of G. adetocorymbus and G. corymbus see
“Notes” of the former species.

Fifth-Instar Males (Adults)

Same as G. adetocorymbus except as follows:

Body measurements.—Mounted, about 610 long, 262 wide.

Body.—Head without postoccipital ridge; area near base of each an-
tenna with 2 (1-3) head pores: Dorsal eye about 30 in diameter; ventral
eye 28 (28-30) in diameter. Antennae about 390 long; apical segment about
equal to length of 3rd segment, not noticeably wider than other segments;
apical segment with 2 subapical sensory setae. Legs with front pair shortest,
hind pair longest. Total lengths of trochanter, femur, tibia, tarsus, claw of
each leg as follows: Front 318 (305-330), middle 336 (335-338), hind
340 (338-343). Hind tibia/tarsus length 1.1. Prosternum smaller than on
G. adetocorymbus. Wings about 350 long. Genital segment 47 (45-50)
long, 60 (58-63) wide; length/width about 0.8; setae of penial sheath
restricted to posterior half of sheath.

Notes.—The above description is based on 3 specimens. For a com-
parison of G. corymbus with G. adetocorymbus see “Notes” of the latter
species. Prepupae and pupae of G. corymbus are unavailable for compari-
son.

Second-Instar Males
Fig. 10

Same as G. adetocorymbus except as follows:

Body measurements.—Mounted, 445 (420-469) long, 280 (238-323) wide.

Dorsum.—Longest anal-lobe seta 24 (23-25) long; seta mesad of longest
seta 4 (3-5). Largest 8-shaped pore on each specimen about 10 long, 6
wide; smallest about 8 long, 5 wide. Minute 8-shaped pores about 3 long.
Simple disc pores fewer than on G. adetocorymbus. Pore clusters with pores

<

Fig. 9. Grammococcus corymbus, second-instar female. A, pore cluster; B, large
8-shaped pore; C, simple disc pore; D, anal ring; E, antenna; F, submarginal 8-
shaped pore; G, seta.

262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

POH |
Fig 10. Grammococcus corymbus, second-instar male. A, tubular duct; B, minute

8-shaped pore; C, simple disc pore; D, pore cluster; E, antenna; F, bilocular pore;
G, submarginal 8-shaped pore; H, seta; I, anal ring.

VOLUME 80, NUMBER 2 263

closely appressed, normally 4 clusters on each side of body, 1 specimen with
1 extra cluster represented by 1 pore; 5 (0-20) quinquelocular pores in
each anterior cephalothoracic cluster; 5 (1-8) in posterior cephalothoracic
cluster; 6 (1-13) in anterior abdominal clusters; and 4 (1-8) in posterior
abdominal clusters; 20 (4-44) pores on each side of body. Tubular ducts
24 (18-27) on each side of body; longest duct about 30 long.

Venter —Clypeolabral shield 76 (75-78) long, 54 (50-55) wide. Labium
38 (35-40) long, 39 (38-40) wide. Bilocular pores near clypeolabral shield,
5 (3-6) pores. Submarginal 8-shaped pores, 34 (31-38) on each side of
body. Anal ring abortive, composed of small, sclerotized spots, without
setae.

Notes—The above description is based on 2 specimens. For a com-
parison of the 2nd-instar males of G. corymbus and G. adetocorymbus see
“Notes” of the latter species.

Acknowledgments

We thank Felipe Mosquera P., Ministerior de Agricultura, Bogota,
Colombia for sending material of G. corymbus. We express our appreciation
to Ronald W. Hodges, Manya B. Stoetzel, Louise M. Russell and Richard
E. White, Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,
USDA, and Michael Kosztarab, Department of Entomology, Virginia Poly-
technic Institute and State University, Blacksburg, for reviewing and criti-
cizing this manuscript. We are also grateful to Pamela Hollyoak for pre-
paring most of the illustrations.

Literature Cited

Borchsenius, N. S. 1960. Fauna of USSR, Homoptera, Kermococcidae, Asterolen-
caniidae, Lecaniodiaspididae, Aclerdidae. (In Russian). Akad. Nauk SSR Zool.
Inst. (n. s. 77) 8, 282 pp.

Giliomee, J. H. 1968. Morphology and relationships of the male of an Asterolecanium
species (Homoptera: Coccoidea: Asterolecaniidae), J. Entomol. Soc. S. Afr.
31:297-308.

Giliomee, J. H., and J. Munting. 1968. A new species of Asterolecanium Targ.
(Homoptera: Coccoidea: Asterolecaniidae) from South Africa. J. Soc. S. Afr. 31:
221-229,

Lambdin, P. L. 1977. A revision of the genus Polea Green. Ann. Entomol. Soc.
Am. 70:911-915.

Russell, L. M. 1941. A classification of the scale insect genus Asterolecanium. Misc.
Pubs., U.S. Dept. Agr. No. 424, 322 pp.

(DRM) Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,
USDA, Beltsville, Maryland 20705; and (PLL) Agricultural Biology De-
partment, The University of Tennessee, Knoxville, Tennessee 37901.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 264-295
LECTOTYPE DESIGNATIONS OF CERTAIN SPECIES OF
THRIPS DESCRIBED BY J. D. HOOD AND
NOTES ON HIS COLLECTION (THYSANOPTERA)

B. R. Pitkin

Abstract.—Corrections to the published lists of J. D. Hood’s papers and
list of new names are given. Lectotypes are designated for 100 species
described by Hood. Lectotypes designated by other authors for Hood’s
species are listed. Notes on Hood’s collections and published papers are
given.

J. Douglas Hood described 1,038 new species and three new varieties of
thrips or Thysanoptera. This represents about 20% of the presently esti-
mated 5,000 described species. Hood also erected 138 new genera and
subgenera and proposed three new specific names including Thrips illicii
for Thrips alysii Hood. Thirteen of the new species and four of the new
genera were described in collaboration with C. B. Williams, (Hood and
Williams, 1915; Hood and Williams in Hood, 1925 (54)).

Bibliographies of Hood’s scientific papers and lists of the new names
proposed in them have been published by Bailey (1949) for 1908-1942 and
ONeill (1974) for 1948-1960. No papers by Hood were published between
1943-1947, due no doubt to the second World War. Hood (1948) also
published a list of the papers he had had published up to that time.

Both Hood (1948) and O'Neill (1974) numbered Hood’s publications and
I have used these numbers in parentheses after date of publication through-
out this note. Unfortunately Hood 1948 (133) gave the same number, Hood
1909 (4), to two different papers (referred to by Bailey (1949) as Hood
1909a and 1909b) and omitted one paper (referred to by Bailey (1949) as
Hood, 1915g). The paper by Hood, 1937 (104), concerns new genera and
species from Africa not America (Hood, 1948 (133)). O'Neill (1977) omitted
Hood, 1958 (174), from her list of Hood’s papers and omitted Helionothrips
compressus Hood, 1954 (153):192-193, from Formosa from the list of
Hood's new names. Unfortunately Hood, 1948 (133), did not publish a list
of names, and Bailey (1949) very occasionally cited the date of publication of
Hood’s papers incorreetly. Thus Hood 1924d was not published until

January 1925 (51); Hood 1927i in January 1928 (71); Hood 1929 in January |
1930 (73); Hood 1933b in January 1934 (86); Hood 1933e in February 1934 |
(87); Hood 1938j in January 1939 (121); and Hood 1941b in January 1942 ©
(132). In addition Cordylothrips peruvianus was described by Hood in |
1937 (110) not in 1927, and Hyidiothrips atomarius was described by Hood |

in 1938 (116) not in 1948. Moreover Bailey misspelled a few specific names

{
{
]
|

[

VOLUME 80, NUMBER 2 265

(cf. Ceuthothrips timuqua Hood, 1938 (116), not ‘timupua’ or ‘timupa’;
Cephalothrips hesperus Hood, 1941 (131), not ‘hexperus’; Atractothrips
bradleyi Hood, 1938 (113), not ‘bardleyi’; Plesiothrips verticalis Hood, 1940
(126), not ‘veticalis’; and Adelothrips macrura Hood, 1941 (131), not
‘macura). In addition Bailey (1949) erroneously included Exophthalomo-
thrips moultoni Hood, 1942 (132) (as ‘1941b’), from Peru in the sub-order
Tubulifera.

A relatively small number of the species Hood described were based on
unique specimens. Some of these were specifically referred to as holotypes
and the others are regarded as such due to their uniqueness. For all but
112 of the remaining species Hood either (a) specifically referred to a holo-
type specimen in the original description or (b) stated in the introduction
to a particular paper that ‘the holotypes and allotypes . . . will remain in
the collection of the author (Hood, 1925 (54); Hood, 1927 (66); and Hood,
1927 (71)) or ‘holotypes and allotypes will remain in the writers collection’
(Hood, 1934 (88)) or ‘holotypes, allotypes and a portion of the paratypes
are in his (Hood’s) collection’ (Hood, 1935 (95)) or ‘holotypes—and the
allotypes in so far as they exist—remain in the authors collection, while
a series of paratypes has been returned . . .. Hood, 1937 (104). By these
criteria I consider that Coremothrips pallidus Hood, 1925 (54); Frankliniella
parvula Hood, 1925 (54); Eupathithrips spectator Hood, 1934 (88); Macro-
phthalmothrips helenae Hood, 1934 (88); Fauriella natalensis Hood, 1937
(104); and Opisthothrips elytropappi Hood, 1937 (104) already have valid
holotype designations (cf. O'Neill, Arnaud and Lee (1971)). Similarly I
consider that Astrothrips angulatus Hood, 1925 (54) (cf. Wilson (1975)) al-
ready has a valid holotype designation.

There are 100 species described by Hood which do not have valid holo-
type or lectotype designations. For all of these species Hood selected a
specimen which he labelled as the holotype. In order to preserve the status
that Hood intended these specimens to have, I am designating them here as
lectotypes. I have also included those lectotypes designated by other
workers in this note.

Hood’s thrips collection was accessioned by the U.S. National Museum
(USNM) in 1965. At that time it was estimated to comprise 60,000 slide-
mounted specimens. These included holotypes or syntypes labelled as
holotypes of all of the species and varieties described by Hood except two.
One of these, Chaeturothrips machadoi Hood, 1954 (154), is in the Dundo
Museum, Angola and the other, Actinothrips monochaetus Hood, 1935 (97),
is in the British Museum (Natural History), London (BMNH). About 1,350
type-specimens of nearly 100 species were accessioned without name and
type-status labels. Some of these specimens also lacked data labels. Each
series of these unlabelled type-specimens was usually identified as such
by an associated paper label written by Hood and bearing the scientific

266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

name of the series and sometimes a note on the characteristics of the spe-
cies. The first slide in each series of unlabelled specimens in all cases ex-
cept that of Glyptothrips arkansanus Hood, 1957 (166), had the word
‘HOLOTYPE’ written directly on the glass slide in black ink. Where Hood
had selected an allotype or ‘morphotype, he labelled those specimens ap-
propriately. None of the specimens, assumed here to be paratypes because
of their collection data, were labelled by Hood as paratypes. I have written
name and type-status labels and where necessary data labels for all of
these unlabelled types. Occasionally Hood changed the spelling slightly in
publication from that on the label. A few times he changed the name com-
pletely. Notes on these changes are included below. I have written the
published name on the slides in the U.S. National Museum. However, it
should be stressed that not all of Hood’s type-specimens are in the USNM.
Hood and the authorities of the U.S. National Museum exchanged a num-
ber of specimens with other institutions and individuals and it is likely
that these specimens are labelled “PARATYPE. I was not able to label
paralectotypes in other locations except the BMNH.

In addition to name and type-status labels I have written data labels for
the 4,000 or more slides in Hood’s collection in the U.S. National Museum
that lacked them. These include numerous type-specimens. Each slide I
labelled was identifiable by a number engraved on the slide, anterior to the
specimen(s). The numbers, referred to as “Hood Nos.,” relate to a
card index in the USNM. I have quoted these numbers in all cases where
possible in the following list.

All names listed below are in their original combinations. For any changes
in nomenclature and synonymy see Jacot-Guillarmod’s Catalogue of the
Thysanoptera of the World (Jacot-Guillarmod, 1970, 1971, 1974, 1975 and
in press). The species are arranged alphabetically within genera and the
genera are arranged alphabetically within families. The families are ar-
ranged systematically from most primitive to most advanced. I have used
the terms macropterous (macr.), micropterous (micr.), brachypterous (brach.)
and apterous (apt.), in the same sense as Hood, to mean fully winged, short
winged, extremely reduced wings and wingless, respectively. The term
dealate indicates that the wings are broken off.

Merothripidae
Merothrips brevisetis Hood, 1954 (149):20-21.

Holotype 2 (macr.)—BRAZIL: Para, Belém, (Instituto Agronomico do
Norte), 2 August 1951 (J. D. Hood). Dead branches of Hevea. USNM Type
71522.

Due to a typographical error there is some ambiguity concerning the
holotype data. The female specimen collected on 23 July is the apterous
‘morphotype.’

VOLUME 80, NUMBER 2 267

Heterothripidae
Aulacothrips dictyotus Hood, 1952 (146):142.

Lectotype ° (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, 25 May
1949 (F. Plaumann). Hood No. 2044. USNM Type 74318. Here designated.

This species was originally described from two females, both in the
USNM. The paralectotype was collected on “‘Compositae: No. 41’ 15 January
1949.

Fauriella natalensis Hood, 1937 (104):98-101.

Holotype ° (macr.)—SOUTH AFRICA: Natal, Zululand, Nolumu, 22
September 1922 (J. C. Faure). In flowers of tree-like Rhus. Faure’s No. T.
50. USNM Type 71232. Designated as a lectotype by O'Neill, Arnaud and
Lee, 1971:25.

This species was originally described from 14 females including a ‘2 para-
type. However in the introduction to the paper Hood refers to ‘holotypes.’
There are 3 female paratypes in addition to the holotype in the USNM.

Heterothrips arisaemae Hood, 1908 (1):362-363.

Lectotype ? (macr.).—USA: Illinois, Urbana, Augerville Woods, 18 May
1907 (F. Gates). In blossoms of Jack-in-the-pulpit. USNM Type 74289.
Here designated. |

This species was originally described from 12 females and 2 males. In
addition to the lectotype there are 6 female and 2 male paralectotypes in
the USNM and all of these have data identical to the lectotype.

Heterothrips sericatus Hood, 1913 (12):66-67.

Lectotype 2 (macr.)—PUERTO RICO: Rio Piedras, 11 June 1912 (T.
H. Jones) [Ex collection H. M. Russell]. “In flowers of guava (Psidium
guajava L.).” Acc. No. 507, 1912. USNM Type 74304. Here designated.

This species was originally described from 33 females and 4 males in-
cluding an ‘allotype’ from Puerto Rico. In addition to the lectotype there
are 13 female and | male paralectotypes in the USNM.

Opisthothrips elytropappi Hood, 1937 (104):102-105.

Holotype ? (macr.)—SOUTH AFRICA: Cape Province, Grahamstown,
21 April 1927 (J. C. Faure). On Elytropappus rhinicerotis. Faure No. T48.
USNM Type 71233. Designated as a lectotype by O'Neill, Arnaud and
Lee, 1971:25.

This species was originally described from 16 females and 5 males includ-
ing a ‘paratype’ of each sex. However in the introduction to the paper
Hood refers to ‘holotypes’ of the species described. There are 3 female
and 3 male paratypes in addition to the holotype in the USNM.

268 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Aeolothripidae
Aeolothrips vehemens Hood, 1927 (67):123-124.

Lectotype ° (macr.)—USA: New Mexico, Chusca Mountains, 1 July
1918 (A. Wetmore). On Populus aurea. Hood No. 397. USNM Type 74281.
Here designated.

This species was originally described from 2 females, 1 of which Hood
referred to as a ‘paratype. This has data identical to the lectotype and
is regarded here as a paralectotype.

Stomatothrips flavus Hood, 1912 (7):64-66.

Lectotype ° (macr.)—MEXICO: Monterey, 5 July 1908 (C. A. Hart).
USNM Type 74266. Here designated.
This species was originally described from “a good series of both sexes.’

Thripidae
Anaphidothrips brasiliensis Hood, 1954 (151):212.

Lectotype 2 (macr.).—BRAZIL: Campo Grande, Distrito Federal, 28
June 1948 (J. D. Hood, A. da Costa Lima and A. Silva). On Andropogon,
probably condensatus Kunth. Hood No. 1667. USNM Type 74357. Here
designated.

This species was originally described from 3 females. Only the lectotype
and 1 female paralectotype are in the USNM, and these have identical
data.

Anaphothrips decolor Hood, 1925 (56):101.

Lectotype 2 (macr.)—USA: Colorado, Golden, (South Table Mountain)
20 June 1918 (L. O. Jackson). Miscellaneous. Hood No. 507. USNM Type
74352. Here designated.

This species was originally described from an unspecified number of
females. In addition to the lectotype there is, in the USNM, a single female
paralectotype, originally labelled as a paratype, which has data identical
to the lectotype.

Arpediothrips mojave Hood, 1927 (69):198.

Lectotype ? (? dealate macr.)—USA: California, Mojave, 14 August
1927 (J. D. Hood). At base of leaves of Joshua tree or Tree Yucca (Yucca
brevifolia Engelm) in the Mojave Desert. Hood No. 804. USNM Type
71234. Designated by O’Neill, Arnaud and Lee (1971:25).

This species was originally described from an unspecified number of
specimens of both sexes from the Mojave Desert, California. In addition to
the lectotype there are 192 female and 9 male paralectotypes in the USNM

VOLUME 80, NUMBER 2 269

and 6 female and 1 male paralectotypes in the BMNH. These are mainly
from Mojave, California but also include specimens from Lancaster, Little-
rock, Victorville, the San Bernardino Mountains and Blythe, California;
Yavapai, Arizona; Deming, New Mexico; and Finlay, Texas collected be-
tween 14 August and 4 September 1927 mainly on Yucca brevifolia but
also on Y. Pbaccata and Y. ?Pelata.

Astrothrips angulatus Hood, 1925 (54):50-51.

Holotype ? (macr.)—BRITISH WEST INDIES: Grenada, 25 March
1915 (C. B. Williams). On leaves of Cacao. Williams No. 599. USNM Type
71516.

This species was originally described from an unspecified number of fe-
males from Grenada, Guadeloupe and Trinidad. Hood (1925) however re-
ferred to ‘holotypes, allotypes and ... paratypes’ of all the species described
in that paper. The designation of a lectotype by Wilson (1975:32-33) is
therefore considered unnecessary. Wilson listed the type-material (as lecto-
type and paratypes) and synonymised the species with Anisopilothrips
venustulus (Priesner).

Bregmatothrips venustus Hood, 1912 (7):67-79.

Lectotype ? (brach.)—USA: Texas, Brownsville, (C. A. Hart). USNM
Type 74358. Here designated.

This species was originally described from ‘several females of both
forms.’

Chirothrips cuneiceps Hood, 1940 (126) :547-550.

Lectotype 2° (macr.)—USA: New York, Oswegatchie, 13 August 1939
(J. D. Hood). Sweeping. USNM Type 74365. Here designated.

This species was originally decribed from 10 females and 2 males in-
cluding 1 female and 1 male paratypes. In addition to the lectotype there
are only 6 female and 2 male paralectotypes in the USNM.

Coremothrips pallidus Hood, 1925 (54):52.

Holotype ° (macr.)—BRITISH WEST INDIES: Trinidad, Evasdale,
near Sangre Grande, 11 October 1916 (C. B. Williams). Cocoa. USNM
type 71235.

This species was originally described from an unspecified number of
macr. females. Hood, 1925 (54), refers to ‘holotypes, allotype and .. . para-
types in the introduction to this paper. This reference is interpreted here
as a holotype designation for this and other species described in that paper.
The designation of a lectotype by O'Neill, Arnaud and Lee (1971) is con-
sidered here unnecessary.

Enneothrips (Enneothripiella) fuscus Hood, 1954 (151):209-210.

270 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Lectotype ? (macr.)—BRAZIL: Para, Corcovado, (Marajo IL, near
Breves), 14 August 1951 (J. D. Hood and F. Camargo). Dead branches
with leaves. Hood No. 2596. USNM Type 74397. Here designated.

This species was originally described from “several females from dead
branches with leaves.” In addition to the lectotype there are 5 macr. female
paralectotypes in the USNM.

Frankliniella achaeta Hood, 1925 (55):81.

Lectotype ? (macr.)—USA: Colorado, 1916 (L. O. Jackson). Sweeping.
USNM Type 72600. Here designated.

This species was originally described from an unspecified number of fe-
males, including a ‘paratype’ from Colorado. In addition to the lectotype
there are 2 female paralectotypes with data similar to the lectotype.

Frankliniella ameliae Hood, 1925 (55):77.

Lectotype 2 (macr.)—PANAMA: Boguete, 28 February 1914 (J. Zetek).
In flower. Hood No. 209. USNM Type 74431. Here designated.

This species was originally described from an unspecified number of
females including a ‘paratype’ from Panama. In addition to the lectotype,
3 female and 2 male paralectotypes were collected with the lectotype; 4
females were collected at the same locality on 29 February 1914; and 1
female was collected at the same locality on 28 February 1914.

Frankliniella auripes Hood, 1915 (27):18-19.

Lectotype 2 (macr.)—PERU: Lima, 13 January 1913 (E. W. Rust). From
‘Jerusalem cherry. USNM Type 74423. Here designated.

The species was originally described from 3 females with identical data.
Only 2 of these, the lectotype and paralectotype, are in the USNM.

Frankliniella difficilis Hood, 1925 (55):73-74.

Lectotype ? (macr.)—FRENCH WEST INDIES: Guadeloupe, 12 March
1915 (C. B. Williams). Flowers of Hydrangea. USNM Type 74434. Here
designated.

This species was originally described from an unspecified number of
specimens of both sexes (including a ‘paratype’) from ‘Guadeloupe and
Martinique. In addition to the lectotype there are 2 female and 7 male
paralectotypes from Guadeloupe and 6 female and 1 male paralectotypes
from Martinique in the USNM.

Frankliniella extremitata Hood, 1937 (106):111-113.

Lectotype 2 (macr.)—PERU: Vicinity of Sani Beni, 840 m, 31 August
1955 (F. Woytkowski). In flowers. Hood No. 1124. USNM Type 74435.
Here designated.

VOLUME 80, NUMBER 2 271

This species was originally described from 12 females including a ‘para-
type. In addition to the lectotype there are 8 female paralectotypes with
data identical to the lectotype.

Frankliniella fuscicauda Hood, 1927 (69):197.

Lectotype ° (macr.)—USA: Arizona, Congress Junction, 22 August
1927 (J. D. Hood). Flowers of Baileya multiradiata. Hood No. 836. USNM
Type 74420. Here designated.

This species was originally described from an unspecified number of fe-
males. In addition to the lectotype female there are 47 females and 7 males
in the USNM, 1 female and 3 males in the BMNH and 1 female and 1
male in the Senckenberg Museum, Frankfurt with the same data as the lecto-
type and a further 5 females and 1 male from Arizona, Aguila, 21 August
1927 (J. D. Hood), on flowers of Baileya multiradiata, Hood No. 847. All
of these are labelled “PARATYPE’ and are regarded here as paralectotypes.

Frankliniella parvula Hood, 1925 (54):49.

Holotype & (macr.)—BRITISH WEST INDIES: Trinidad, Mareval
Valley, 27 March 1915 (C. B. Williams). Flowers of rose. USNM Type
71236.

This species was originally described from an unspecified number of males
and females. However Hood, 1925 (54), refers to ‘holotypes, allotypes and
.. . paratypes’ in the introduction to this paper. This reference is in-
terpreted here as a holotype designation for this and other species de-
scribed in that paper. The designation of a lectotype by O'Neill, Arnaud
and Lee (1971) is considered here unnecessary.

Helionothrips stephaniae Hood, 1937 (104):108-110.

Holotype 2 (macr.)—SOUTH AFRICA: Transvaal, Woodbush, Pieters-
burg, 12 April 1924 (J. C. Faure). On Stephania meyeriana Haw. Hood No.
500. USNM Type 71517. Designated as a lectotype by Wilson (1975:142).

This species was originally described for 11 females including a ‘para-
type. In the introduction to the paper Hood refers to ‘holotypes’ of the
species described.

Heliothrips phaseoli Hood, 1912 (9):113-114.

Lectotype 2 (macr.)—USA: Texas, Brownsville, 26 June 1908 (C. A.
Hart). On bean plants. USNM Type 74325. Here designated.

This species was originally described from ‘many specimens of both
sexes from Brownsville, Texas and Matamoras, Mexico.

Physothrips ventralis Hood, 1918 (42):116.
Lectotype 2 (macr.)—CAMEROON: 23 November 1915 (A. W. Jobbins-

272 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pomeroy). Flowers of monkshood. Hood No. 160. USNM Type 74530.
Here designated.

This species was originally described from ‘numerous specimens of both
sexes from a large variety of flowers’ in Cameroon and southern Nigeria. In
addition to the lectotype there are 70 female and 6 male paralectotypes from
Cameroon and 24 female paralectotypes from southern Nigeria in the
USNM, and 1 female paralectotype from Nigeria in the BMNH.

Plesiopsothrips trinidadensis Hood, 1956 (164):64-66.

Holotype 2 (macr.)—BRITISH WEST INDIES: Trinidad, El Tucuche,
22 June 1951 (J. D. Hood). Dead branches. Hood No. 2529. USNM Type

The unique female holotype was labelled by Hood as ‘trinitatis.’ There
is no doubt that this specimen is the holotype of Plesiopsothrips trinidadensis
Hood.

Pseudodendrothrips alboniger Hood, 1952 (146):145.

Lectotype 2 (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, 26 No-
vember 1949 (F. Plaumann). On Terminalia. Plaumann No. 183. USNM
Type 74449. Here designated.

This species was originally described from 17 females and 6 males from
Terminalia and unidentified plants. In addition to the lectotype there are
11 female and 5 male paralectotypes in the USNM and | female paralecto-
type in the BMNH and all of these are from Nova Teutonia.

Psilothrips pardolatus Hood, 1927 (69):198.

Lectotype ? (macr.)—USA: California, Thermal (elevation, 100 ft) 18
August 1927 (J. D. Hood). Shaken from Atriplex polycarpa Watson. Hood
No. 832. USNM Type 71237. Designated by O'Neill, Arnaud and Lee
(LS7ie25).

This species was originally described from an unspecified number of
specimens of both sexes from California, Arizona and Texas from un-
determined plants. In addition to the lectotype 3 female and 2 male para-
lectotypes were collected with the lectotype at Thermal, California, 6 fe-
male and 1 male paralectotypes from Palm Canyon, California also on A.
polycarpa, 13 female paralectotypes from Ysleta, Texas on A. canescens
(Pursh) Nutt., and 8 female paralectotypes from Quijotoa, Arizona on
A. canescens. All, except 1 female from Palm Canyon which is in the
BMNH, are in the USNM.

Sericothrips occipitalis Hood, 1917 (40):32-34.

Lectotype 2 (macr.)—NIGERIA: Ibadan, 14 January 1915 (A. W. Job-
bins-Pomeroy). On Desmodium lasiocarpum. Hood No. 50. USNM Type
74491. Here designated.

VOLUME 80, NUMBER 2 273

This species was originally described from 3 specimens including a ‘para-
type’ female and an ‘allotype’ male. The paralectotype female was collected
on Bougainvillaea glabra Choisy and the male on Andropogon tectorum
Schum. and Thonn.

Sericothrips opuntiae Hood, 1936 (100):88-91.

Lectotype 2° (macr.)—USA: Arizona, Comobabi, 28 August 1927 (J. D.
Hood). On Opuntia sp. Hood No. 852. USNM Type 74522. Here desig-
nated.

This species was originally described from 12 females and 6 males includ-
ing a ‘paratype’ female. In addition to the lectotype there are 7 female

and 5 male paralectotypes in the USNM and 1 female paralectotype in
the BMNH.

Sericothrips pulchellus Hood, 1908 (1):363-364.

Lectotype ° (macr.)—USA: Ulinois, Muncie, 16 June 1908 (J. D. Hood).
On leaves of Ptelea trifoliata L. USNM Type 74493. Here designated.

This species was originally described from an unspecified number of
specimens of both sexes. In addition to the lectotype there are 7 female
and 3 male paralectotypes in the USNM and 3 female paralectotypes in the
BMNH, and these have data identical to the lectotype.

Sericothrips tiliae Hood, 1931 (78):151-152.

Lectotype 2 (macr.)—USA: New York, Morton, 1 September 1930 (J. D.
Hood and H. M. Hincher). USNM Type 74517. Here designated.

This species was originally described from ‘a large number of specimens
of both sexes from New York.

Taeniothrips aethiops Hood, 1925 (57):2-3.

Holotype 2 (macr.)—CAMEROON: [Kamerun]; 12 February 1916 (A.
W. Jobbins-Pomeroy). Flowers of small shrub. Hood No. 176. USNM
Type 74529.

The type-series in the USNM is labelled “Physothrips aethiops sp. nov.’

Taeniothrips debilis Hood, 1925 (57):5-6.

Holotype 2 (macr.)—NIGERIA: Ibadan, 14 January 1915, (A. W. Job-
bins-Pomeroy). In flowers of Melia azederach. Hood No. 53. USNM Type
75107.

The type-series in the USNM is labelled “Physothrips debilis sp. nov.’
Taeniothrips dilutus Hood, 1925 (57):8-9.

Holotype 2 (macr.)—EAST AFRICA: Rifigi River, 27 July 1917 (A. W.
Jobbins-Pomeroy). Flowers of false ebony. Hood No. 188. USNM Type
74527.

274 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The holotype female is labelled “Physothrips dilutus sp. nov.
Taeniothrips modestus Hood, 1925 (57):3-5.

Holotype ° (macr.)—CAMEROON: [Kamerun]; 16 February 1916 (A.
W. Jobbins-Pomeroy). Flowers of a large bush. Hood No. 177. USNM
Type 74534.

The type-series in the USNM is labelled ‘Physothrips modestus sp. nov.’

Taeniothrips silvestris Hood, 1935 (92):83-84.

Lectotype 2 (macr.)—PANAMA: Panama Canal, Barro Colorado Is-
land, Gatun Lake, October 1933 (S. Aviles). On Dichorisandra hexandra.
Hood No. 1078. USNM Type 74525. Here designated.

This species was originally described from 27 females and 5 males in-
cluding a ‘paratype’ female and an ‘allotype’ male. In addition to the
lectotype there are 20 female and 4 male paralectotypes in the USNM,
and 1 male paralectotype in the BMNH.

Uzelothripidae
Uzelothrips scabrosus Hood, 1952 (146):143-144.

Lectotype 2° (macr.).—BRAZIL: Para, Belém, (Instituto Agronomico do
Norte), 21 August 1951 (J. D. Hood). Dead dry branches of Hevea sp. on
ground. Hood No. 2721. USNM Type 74558. Here designated.

This species was originally described from ‘many specimens’ of both
sexes. In addition to the lectotype there are 2 macr. female, 24 apt. female
and 1 apt. male paralectotypes, all from the type-locality and in the USNM,
and 2 apt. females in the BMNH. There are also 28 apt. females labelled
‘TOPOTYPIC’ in the USNM.

Phlaeothripidae
Actinothrips trichaetus Hood, 1935 (97):248-252.

Lectotype 2° (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
1933 (J. D. Hood). USNM Type 74969. Here designated.

This species was originally described from 46 macr. females, 41 macr.
males and 20 larvae (including a ‘paratype’ of each sex) from Panama and
Ecuador.

Adelothrips eucharis Hood, 1955 (163):84-88.

Holotype ? (macr.)—BRAZIL: Para, Fordlandia, 10 July 1951 (J. D.
Hood). Dead grass. Hood No. 2581. USNM Type 74626.

This species was originally described from 1 macr. female and 1 macr.
male including a holotype female and an allotype male. There are 2 slides

VOLUME 80, NUMBER 2 275

labelled as ‘Adelothrips excellens’ by Hood with data identical to that
published for eucharis. Miss Kellie O'Neill identified these as eucharis,
and I have labelled them as types of that species.

Adraneothrips huachucae Hood, 1927 (69):202-203.

Lectotype ° (macr.)—USA: Arizona, Ramsey Canyon (Huachuca Moun-
tains, 30 mi east of Nogales), 2 September 1927 (J. D. Hood). Beating dead
oak leaves. Hood No. 927. USNM Type 74587. Here designated.

This species was originally described from an unspecified number of
specimens of both sexes. In addition to the lectotype there are 14 female
and 11 male paralectotypes in the USNM, and 2 female and 1 male para-
lectotypes in the BMNH.

Allothrips megacephalus Hood, 1908 (1):373.

Lectotype ? (apt.)—USA: Illinois, Urbana, Augerville Woods, 9 No-
vember 1907 (J. D. Hood). Under bark on living osage-orange tree. USNM
Type 74970. Here designated.

This species was originally described from “several females, one of them
brachypterous.’ In addition to the apt. lectotype there are 4 apt. female
and 1 brach. female paralectotypes in the USNM.

Bagnalliella arizonae Hood, 1927 (69):201.

Lectotype ? (brach.)—USA: Arizona, Fort Huachuca, September 1927
(J. D. Hood). At base of leaves of Yucca elata Engelm. Hood No. 855.
USNM Type 74626. Here designated.

This species was originally described from an unspecified number of speci-
mens of both sexes from Arizona, New Mexico and Texas. In addition to
the lectotype there are 24 macr. female, 111 brach. female and 6 brach. male
paralectotypes in the USNM and 1 brach. female and 1 brach. male para-
lectotypes in the BMNH.

Bagnalliella desertae Hood, 1927 (69):201.

Lectotype 2 (macr.)—USA: California, Victorville, 15 August 1927 (J.
D. Hood). At base of leaves of Yucca baccata Torr. in Mojave Desert. Hood
No. 809. USNM Type 74629. Here designated.

This species was originally described from an unspecified number of
specimens of both sexes. In addition to the lectotype there are 1 macr.
female, 52 brach. female and 16 brach. male paralectotypes in the USNM
and 2 brach. female and 1 brach. male paralectotypes in the BMNH. All
of the type-series are labelled ‘deserti.’

Bagnalliella huachucae Hood, 1927 (69):200.
Lectotype ? (macr.)—USA: Arizona, Ramsey Canyon, (Huachuca Moun-

276 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tains, 30 mi east of Nogales), 2 September 1927 (J. D. Hood). On Yucca sp.
(perhaps Y. buccata Torr.), at base of leaves. Hood No. 928. USNM Type
74631. Here designated.

This species was originally described from an unspecified number of
specimens. In addition to the lectotype there are 7 macr. female, 5 brach.
female and 7 brach. male paralectotypes all bearing the same data as the
lectotype.

Bagnalliella mojave Hood, 1927 (69):200-201.

Lectotype ° (macr.)—USA: California, Mojave, in the Mojave Desert.
14 August 1927 (J. D. Hood). At base of leaves of Joshua tree or Tree
Yucca (Yucca brevifolia Engelm.). Hood No. 804. USNM Type 74630. Here
designated.

This species was originally described from an unspecified number of
specimens of both sexes. In addition to the lectotype there are 1 macr.
female, 85 brach. female and 8 brach. male paralectotypes from Mojave,
19 brach. female paralectotypes from Victorville, and 47 brach. female
paralectotypes from Lancaster, California. All of these, except 3 brach.
females and 1 brach. male from Mojave and which are in the BMNH,
are in the USNM.

Chthonothrips nigrocinctus Hood, 1957 (170):143.

Holotype 2° (apt.)—BRAZIL: Santa Catarina, Nova Teutonia, September
1955 (F. Plaumann). Under fallen leaves. Hood No. 1787. USNM Type
74184.

The last 6 lines of the original description of this species have been
transposed with the first 5 lines of the original description of Chortothrips
gen. nov., Hood, 1957 (170): 143 (O’Neill, 1974). Contrary to O’Neill (1971),
therefore, Hood designated a holotype for nigrocinctus.

Cordylothrips peruvianus Hood, 1937 (110):518-519.

Lectotype 2 (macr.).—PERU: Almirante, Departamento de Amazonas,
(elevation about 1,900 m), 20 December 1936 (F. Woytkowski). Dead
branches in jungle. Hood No. 1143. USNM Type 75002. Here designated.

This species was originally described from 5 females including a “‘para-
type. In addition to the lectotype there are 3 macr. female paralectotypes
in the USNM.

Cryptothrips junctus Hood, 1912 (8):139-142.

Lectotype ? (brach.)—USA: Michigan, Baldwin. USNM Type 74574.
Here designated.

This species was originally described from 18 brach. females, 2 macr. fe-
males and 11 brach. males from Michigan and Illinois.

VOLUME 80, NUMBER 2 277

Cryptothrips rectangularis Hood, 1908 (2):307-309.

Lectotype ° (apt.)—USA: Illinois, Urbana, (near University Forest),
12 May 1908 (J. D. Hood). Under dead bark on peach tree. USNM Type
75001. Here designated.

This species was originally described from ‘four winged individuals.’
These are all in the USNM.

Cryptothrips sordidatus Hood, 1927 (69):199.

Lectotype 2 (macr.)—USA: California, Palo Alto, 4 August 1927 (J. D.
Hood). Beating dead branches of Salix sp. Hood No. 744. USNM Type
75006. Here designated.

This species was originally described from an unspecified number of
specimens of both sexes. In addition to the lectotype, there are 1 brach.
female and 4 brach. male paralectotypes, all with data identical to the
lectotype, in the USNM.

Cyphothrips dorsalis Hood, 1952 (146):172-173.

Lectotype 2 (macr.)—BRAZIL: Para, Belém, (Instituto Agronomico do
Norte), 17 August 1951 (J. D. Hood). Dead leaves of Euterpe oleraceae.
Hood No. 2711. USNM Type 74822. Here designated.

This species was originally described from 5 females and 2 males from
the same locality. In addition to the lectotype there are 2 macr. female
and 2 macr. male paralectotypes in the USNM.

Diceratothrips cubensis Hood, 1941 (131):178-180.

Lectotype ¢ (macr.)—CUBA: Maranzas, San Miguel de los Banos, 18
July 1940 (J. C. Bradley). Dead branches. USNM Type 75013. Here
designated.

This species was originally described from 2 females including a ‘para-
type. Both specimens are in the USNM.

Diceratothrips setigenus Hood, 1941 (131):176-178.

Lectotype ° (macr.)—USA: Texas, Brownsville, 2 March 1939 (J. D.
Hood). Dead branches. USNM Type 75023. Here designated.
This species was originally described from 4 females, including a ‘para-

type. In addition to the lectotype there are 2 female paralectotypes in
the USNM.

Diopsothrips brunneus Hood, 1934 (87):424-425.

Lectotype @ (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
1933 (J. D. Hood and S. Aviles). Dead branches. USNM Type 75026. Here
designated.

278 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

This species was originally described from 6 macr. females from Panama.
Diopsothrips flavus Hood, 1934 (87):423-424.

Lectotype @ (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
1933 (J. D. Hood). USNM Type 75025. Here designated.

This species was originally described from 27 macr. females and 13
macr. males from Panama.

Eupathithrips spectator Hood, 1934 (88):73-76.

Holotype ° (macr.)—PANAMA: Panama Canal, Barro Colorado Island,
Gatun Lake, 25 June 1933 (J. D. Hood). On ripe fruit of Corozo Palm.
Hood No. 947. USNM Type 71238.

This species was originally described from 22 females, 30 males and 3
nymphs taken on ‘Barro Colorado Island, Canal Zone (type-locality) and at
Porto Bello, Panama. This reference and the references to ‘holotypes,
‘allotypes’ and ‘paratypes’ in the introduction of this paper are interpreted
here as a holotype designation. The designation of a lectotype by O'Neill,
Arnaud and Lee (1971) is here considered unnecessary.

Eurythrips citricollis Hood, 1941 (131):240-243.

Lectotype 2 (macr.)—USA: Florida, Winter Park, 11 March 1941 (M.
J. Westfall, Jr.). Fallen pine needles. USNM Type 74662. Here designated.

This species was originally described from 3 macr. and 11 brach. females
including a macr. ‘paratype’ and a brach. ‘paratype.’ In addition to the
lectotype there are 2 macr. female and 9 brach. female paralectotypes in
the USNM and 1 macr. female paralectotype in the BMNH.

Eurythrips nigricornis Hood, 1960 (175):61-63.

Holotype ° (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, June
1957 (F. Plaumann). Dead grasses. Hood No. 2944. USNM Type 72626.

This species was originally described from 2 macr. females (including the
holotype), 19 brach. females and 6 brach. males from dead grasses. There
was an unlabelled series in the USNM, the first specimen of which was
labelled ‘Eury. sculpturus’ and ‘HOLOTYPE [1]. These were identified
by Mound (1976:56) as nigricornis Hood, there is little doubt that they repre-
sent the type-series of that species, and I have labelled them as such. A
brach. paratype of each sex has been deposited in the BMNH.

Eurythrips umbrisetis Hood, 1934 (87):415-416.

Lectotype ° (brach.)—PANAMA: Canal Zone, Frijoles, 18 July 1933
(H. H. Hood and J. J. Hook). Dead leaves of Panicum maximum. Hood
No. 1001. USNM Type 74664. Designated by Mound (1976).

This species was originally described from 2 brach. females from Frijoles,
Canal Zone.

VOLUME 80, NUMBER 2 279

Gastrothrips callipus Hood, 1935 (94):182-186.

Lectotype 2 (macr.).—USA: Texas, Victoria, 1 April 1908 (J. D. Mitchell).
On Chenopodium. USNM Type 72001. Here designated.

This species was originally described from 3 females and 1 male includ-
ing a female ‘paratype’ and male ‘allotype. In addition to the lectotype
there is only the single male paralectotype in the USNM.

Gastrothrips firmus Hood, 1952 (146):162.

Lectotype 2 (macr.)—BRAZIL: Sao Paulo, Itanhaém, 17 June 1948
(J. D. Hood and J. Lane). Dead branches. Hood No. 1640. USNM Type
71995. Here designated.

This species was originally described from 3 females and 2 males. In

addition to the lectotype there are 1 female and 2 male paralectotypes in
the USNM.

Gastrothrips picticornis Hood, 1936 (102):272-275.

Lectotype ? (brach.)—BRAZIL: Rio de Janeiro, (Angra dos Reis),
10 August 1934 (D. Mendes). Dead twigs of Anona squamosa. Hood No.
1208. USNM Type 72004. Here designated.

This species was originally described from 9 brach. females and 2 brach.
males including a ‘paratype’ of each sex. In addition to the lectotype there
are 7 brach. female and 2 brach. male paralectotypes in the USNM.

Gastrothrips proteus Hood, 1934 (87):417-419.

Lectotype 2 (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
1933 (S. J. Hook, H. H. Hood and J. D. Hood). Dead leaves and branches.
USNM Type 71996. Here designated.

This species was originally described from 1 macr. female, 17 apt. females
and 7 apt. males from Panama.

Gastrothrips ruficauda Hood, 1912 (10):156-157.

Lectotype 2 (brach.)—USA: Illinois, Grand Tower/Pulaski, October/
July ?1909 (C. A. Hart and L. M. Smith). On branches of grape/sycamore/
overcup oak (Quercus lyrata Walt.). USNM Type 71997. Here designated.

This species was originally described from 3 females.

Glyptothrips arkansanus Hood, 1957 (166) :59-60.

Lectotype ? (macr.)—USA: Arkansas, Fayetteville, 16 March 1957 (W.
H. Whitcomb). On Andropogon virginicus L. Hood No. 2764. USNM
Type 71993. Designated by Mound and O'Neill (1977).

This species was originally described from ‘29 females (including holo-
type). Mound and O'Neill (1977) were unable to find the type-series in the
systematic collections of the USNM. “However specimens in four series

280 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of unlabelled slides in the numerical collection were identified as ar-
kansanus from the description” (Mound and O’Neill, 1977). Although the
data for these specimens contrasted with the published data “There can
be little doubt that the specimens . . . comprise the type-series of G. ar-
kansanus” (Mound and O'Neill, 1977). Mound & O'Neill (1977) designated
“the female at the head of series 2764” as the lectotype.

Glyptothrips flavescens Hood, 1912 (9):116-117.

Lectotype 2 (macr.)—USA: Illinois, Grand Tower, 1909 (C. A. Hunt).
USNM Type 74679. Here designated.

This species was originally described from 3 macr. females. Two of these
are from the “Type-locality, Grand Tower, and the other is from Pulaski,
Illinois.

Haplothrips graminis Hood, 1912 (7):69-70.

Lectotype 2 (macr.)—USA: Texas, Brownsville (C. A. Hart). USNM
Type 74687. Here designated.
This species was originally described from ‘a good series of both sexes.’

Haplothrips (Anchylothrips) preeri Hood, 1939 (125):565-568.

Lectotype 2 (macr.).—USA: Texas, Palacios, 31 March 1939 (J. D. Hood).
On Spartina alterniflora var. glabra (Muhl.) Fern. USNM Type 74701. Here
designated.

This species was originally described from 10 females and 9 males in-
cluding a ‘paratype’ of each sex, all from Spartina alterniflora at Palacios.
In addition to the lectotype there are 7 female and 6 male paralectotypes
in the USNM and 1 female and 1 male paralectotypes in the BMNH. One
of the females in the USNM has reduced wings.

Holothrips amplus Hood, 1952 (146):160.

Lectotype 2 (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, 31
December 1948 (F. Plaumann). Dry branches. Hood No. 2018-O. USNM
Type 74747. Here designated.

This species was originally described from 3 females. There is only 1
female paralectotype, in addition to the lectotype, in the USNM.

Hoplandrothrips angustatus Hood, 1927 (69):199.

Lectotype 2 (macr.)—USA: Arizona, Nogales. 30 August 1927 (J. D.
Hood). Mesquite-like trees, Acacia or Prosopis. Hood No. 909. USNM
74767. Here designated.

This species was originally described from an unspecified number of
macr. females and brach. males from dead branches of mesquite at
Nogales, Arizona. In addition to the lectotype there are 3 macr. female
and 3 brach. male paralectotypes in the USNM.

VOLUME 80, NUMBER 2 281

Hoplandrothrips longirostris Hood, 1954 (149):46.

Holotype °(macr.)—BRAZIL: Rio de Janeiro, Petropolis, 27 June 1948
(J. D. Hood). Dead and dying branches of various trees. Hood No. 1672.
USNM Type 75090.

This species was originally described from 13 macr. females. None of
the 11 females in the USNM were labelled as Hoplandrothrips longirostris
Hood. The holotype was labelled in ink on the glass slide “Hoplandrothrips
penetralis.. The data for these specimens correspond to the published data
of longirostris. There is no doubt that the specimens are part of the type-
series of this species, and consequently I have labelled them as such.

Hoplandrothrips nigricestus Hood, 1934 (87):429-430.

Lectotype 2 (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
1933 (J. D. Hood). Dead leaves. USNM Type 74766. Here designated.

This species was originally described from 16 macr. females and 7 macr.
males from Panama.

Hoplandrothrips symmetricus Hood, 1942 (132):557-559.

Holotype 2 (macr.)—PERU: Piedras Grandes, Dept. Huanuco (Andes
elevation about 3,000 m), 6 November 1937 (F. Woytkowski). Flowers of a
wild potato. Hood No. 1468. USNM Type 74756.

This species was described from an unspecified number of macr. females
including a holotype. The type-data were omitted from the original de-
scription, and there is only the unique holotype in the USNM.

Idolothrips armatus Hood, 1908 (3):285-287.

Lectotype 2 (macr.)—USA: Illinois, Carbondale, 20 June 1907 (J. D.
Hood). Taken in old dried up Solidago galls. USNM 75030. Here desig-
nated.

This species was originally described from 8 macr. females and 8 macr.
males mainly from galls on Solidago at Carbondale, Illinois. In addition to
the lectotype there are 4 female and 6 male paralectotypes in the USNM
and 1 female paralectotype in the BMNH.

Idolothrips flavipes Hood, 1908 (1):377.

Lectotype 2° (brach.)—USA: Illinois, Dubois, 28 April, 1908 (C. A.
Hart and L. M. Smith). Sifted from dead oak. USNM Type 75034. Here
designated.

This species was originally described from ‘several males and females,
all from Illinois.’ In addition to the lectotype there are 8 brach. female and
4 brach. male paralectotypes in the USNM.

Idolothrips tuberculatus Hood, 1908 (3):287-289.

282 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Lectotype ° (macr.).—USA: Illinois, White Heath, 26 August 1908 (C. A.
Hart). Jarred from branch of white oak (Quercus alba). USNM Type 75043.
Here designated.

This species was originally described from 4 females and 1 male from
white oak at White Heath and Dusky Dell, Illinois. In addition to the lecto-
type there are 1 macr. female and 1 macr. male paralectotypes in the USNM
and 1 macr. female paralectotype in the BMNH.

Lathrobiothrips ramuli Hood, 1934 (87):421-422.

Lectotype ° (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
30 July 1933 (J. D. Hood). USNM Type 74825. Here designated.

This species was originally described from 11 macr. females and 4 macr.
males from Panama.

Leptothrips heliomanes Hood, 1927 (69):202.

Lectotype 2 (macr.)—USA: California, Palm Canyon (near Palm Springs,
elevation 100 ft), 17 August 1927 (J. D. Hood). Shaken from Atriplex poly-
carpa Watson. Hood No. 830. USNM Type 74723. Here designated.

This species was originally described from an unspecified number of
females and males from Palm Canyon, California on an undetermined plant.
In addition to the lectotype there are 5 female and 2 macr. male paralecto-
types in the USNM.

Liothrips avocadis Hood, 1935 (92):97-99.

Lectotype 2 (macr.)—PANAMA: Canal Zone, Ancon, 24 August 1933
(J. Zetek). Avocado. Hood No. 1064. USNM Type 74849. Here designated.

This species was originally described from 16 females and 7 males in-
cluding a ‘paratype’ of each sex. In addition to the lectotype there are 11
female and 6 male paralectotypes in the USNM and 1 female paralectotype
in the BMNH.

Liothrips ordinarius Hood, 1919 (46):101.

Lectotype 2 (macr.)—INDIA: Coimbatore (T. V. Ramakrishna Ayyar).
On shoots of Sesbania grandiflora. Hood No. 368. USNM Type 74655. Here
designated.

This species was originally described from 4 females and 12 males in-
cluding a ‘paratype’ of each sex. In addition to the lectotype there are 1
female and 8 male paralectotypes in the USNM.

Liothrips penetralis Hood, 1935 (92):95-97.

Lectotype ? (macr.)—PANAMA: Panama Canal, Barro Colorado Is-
land, Gatun Lake, 9 August 1933 (J. D. Hood). Leaves, probably of
Trichilia sp. Hood No. 1046. USNM Type 74835. Here designated.

VOLUME 80, NUMBER 2 283

This species was originally described from 9 females and 7 males includ-
ing a ‘paratype’ of each sex. In addition to the lectotype there are 4 female
and 6 male paralectotypes in the USNM and | female paralectotype in the
BMNH.

Liothrips tupac Hood, 1938 (122):414—-417.

Lectotype @ (macr.)—PERU: Vicinity of Celedin, Dept. Cajamarca (in
Andes), 1-3 June 1936 (F. Woytkowski). Beating bush, often containing
dry branches with moss. Hood No. 1187. USNM Type 74836. Here desig-
nated.

This species was originally described from 4 females and 1 male includ-
ing a male ‘allotype’ and female ‘paratype.’ In addition to the lectotype there
are 2 female and | male paralectotypes in the USNM.

Liothrips vigilax Hood, 1938 (122):407-409.

Holotype 2 (macr.)—PERU: Vicinity of Celedin, Dept. Cajamarca (in
Andes), 1-3 June 1936 (F. Woytkowski). Beating bush, often containing
dry branches with moss. Hood No. 1187. USNM Type 74833.

The type-series is labelled ‘vigilans’ rather than ‘vigilax.’

Liothrips xanthocerus Hood, 1927 (69):203.

Lectotype 2° (macr.).—USA: Arizona, Maricopa County, Gillespie Dam,
26 August 1927 (J. D. Hood). Among terminal leaves of Tessaria sericea
(Nutt.) T. & G. Hood No. 888. USNM Type 71239. Designated by O'Neill,
Arnaud and Lee (1971:25).

This species was originally described from an unspecified number of
macr. males and females. In addition to the lectotype there are 8 female
and 17 male paralectotypes in the USNM and 2 female paralectotypes in
the BMNH.

Lissothrips flavidus Hood, 1960 (175):65-66.

Holotype ? (apt.)—BRAZIL: Santa Catarina, Nova Teutonia, May
1957 (F. Plaumann). Litter. Hood No. 2939. USNM Type 72624.

This species was originally described from 6 apt. females. There was an
unlabelled series of 8 apt. females in the USNM (1 apt. female is now in the
BMNH). All of these bear the published type-data and moreover 1 specimen
is labelled “Lisso. flavidus HOLOTYPE.’ I have labelled all 8 apt. females
as type-material.

Lissothrips muscorum Hood, 1908 (1):365.

Lectotype 2 (apt.)—USA: Illinois, Urbana, Augerville Woods, 19 Octo-
ber 1907 (H. E. Ewing and J. D. Hood). Sifted by Mr. Ewing from moss on
stump. USNM Type 74858. Here designated.

284. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

This species was originally described from ‘several apt. females’ from
Illinois. In addition to the lectotype there are 11 apt. female paralectotypes
in the USNM.

Macrophthalmothrips helenae Hood, 1934 (88):79-81.

Holotype 2 (macr.)—PANAMA: Panama Canal, Barro Colorado Island,
29 July 1933 (J. Zetek and J. D. Hood). Dead branches. Hood No. 1019.
USNM Type 71240.

This species was originally described from 34 females and 3 males “all
taken from dead branches on Barro Colorado Island, Canal Zone (type-
locality) and at Porto Bello, Panama. This reference and the reference to
‘holotypes and allotypes’ and ‘paratypes’ in the introduction of this paper
are interpreted here as a holotype designation. The designation of a lecto-
type by O'Neill, Arnaud and Lee (1971) is considered here unnecessary.

Malacothrips fasciatus Hood, 1952 (146):150.

Lectotype 2 (macr.).—BRAZIL: Santa Catarina, Nova Teutonia, 15 May
1949 (F. Plaumann). Grass; Erianthus. Hood No. 2037. USNM Type
74869. Here designated.

This species was originally described from “9? 2 (3 of them macropterous)
and 24 é (brachypterous).’ In addition to the lectotype there are 2 macr. fe-
male, 2 brach. female and 1 brach. male paralectotypes in the USNM.

Malacothrips mediator Hood, 1952 (146):151.

Lectotype 2 (brach.)—BRAZIL: Santa Catarina, Nova Teutonia, 24 May
1949 (F. Plaumann). Grasses. Hood No. 2043. USNM Type 74870. Here
designated.

This species was originally described from 2 females with identical data.
Both are in the USNM.

Megalothrips picticornis Hood, 1927 (69):204.

Lectotype 2 (macr.).—USA: California, 27 July 1927 (J. D. Hood). Beat-
ing Lonicera involucrata and dead Salix. Hood No. 706. USNM Type
75049. Here designated.

This species was originally described from an unspecified number of
females and males from California and Utah. In addition to the lectotype
there are 1 macr. female and 1 macr. male paralectotypes in the USNM.

Megalothrips (?) spinosus Hood, 1908 (2):306-307.

Lectotype ? (macr.)—USA: Pennsylvania, Harrisburg. 10 March (Pa.
State Dept. Agric. Div. Zool.). In burrows of lepidopterous or coleopterous
larvae in dead willow stem. USNM Type 75048. Here designated.

This species was originally described from “2 macr. females.’ How-

VOLUME 80, NUMBER 2 285

ever, in addition to the lectotype there are 2 females labelled ‘paratype’ in
the USNM both with data identical to the lectotype. One of these was
drawn and this is regarded as a paralectotype.

Neothrips corticis Hood, 1908 (1):372.

Lectotype ° (brach.)—USA: Illinois, Urbana, (University forest), 18
January 1908 (J. D. Hood). Under bark on soft maple tree. USNM Type
74874. Here designated.

This species was originally described from ‘several specimens of both
sexes. In addition to the lectotype there are 7 female and 5 male para-
lectotypes in the USNM and 1 female paralectotype in the BMNH.

Neurothrips frontalis Hood, 1952 (146):155.

Lectotype 2 (macr.)—BRAZIL: Sao Paulo, Boracea, Munic de Sale-
sopolis, 5 June 1948 (J. D. Hood). Dead branches with leaves. Hood No.
1578. USNM Type 74876. Here designated.

This species was originally described from 1 female and 4 males from
Boracea, Brazil. In addition to the lectotype there are 3 male paralectotypes
in the USNM.

Oedaleothrips brasiliensis Hood, 1952 (146): 166-167.

Lectotype 2 (apt.)—BRAZIL: Sao Paulo, Serra da Contareira, Franco da
Rocha, 11 June 1948 (J. D. Hood, F. Lane and L. T. Filha). From grasses,
including Andropogon (probably condensatus Kurth.). Hood No. 1606.
USNM Type 75092. Here designated.

This species was originally described from 15 females, 15 males and 6
nymphs with identical data. In addition to the lectotype there are 11
female, 11 male and 6 nymphal paralectotypes in the USNM and 1
female and 1 male paralectotypes in the BMNH.

Oedaleothrips congoensis Hood, 1952 (142):204-209.

Lectotype 2 (apt.)—CONGO BELGE: Nat. Pare Albert, Rwindi Camp,
4 December 1948 (R. S. Bradley). Grass. Hood No. 1722. USNM Type
74999. Here designated.

This species was originally described from 12 females and 7 males in-
cluding a ‘paratype’ of each sex. In addition to the lectotype there are
5 apt. female and 5 apt. male paralectotypes in the USNM and | apt. female
paralectotype in the BMNH.

Orthothrips leptura Hood, 1952 (146):151.

Lectotype 2 (macr.)—BRAZIL: Para, Belém, (Instituto Agronomico do
Norte), 9 August 1951 (J. D. Hood). Dead leaves of Astrocaryum mumbaca.
Hood No. 2682. USNM Type 74882. Here designated.

286 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

This species was originally described from 1 female and 3 males. In ad-
dition to the lectotype there are 2 brach. male paralectotypes in the
USNM.

Palinothrips palustris Hood, 1952 (146):168.

Lectotype ? (macr.).—BRAZIL: Sao Paulo, Sao Carlos, (D. P. de Sauza
Dias), 21 June 1950. Grass. Hood No. 1255. USNM Type 75028. Here
designated.

This species was originally described from 3 females and 6 males. In ad-
dition to the lectotype there are 4 female paralectotypes in the USNM.

Phloeothrips vittatus Hood, 1912 (6):11-12.

Lectotype 4 (macr.)—USA: Michigan, Baldwin, (“Star Lake” near Rain-
bow), 17 August 1908 (J. D. Hood). On rotting poplar stump. USNM Type
74878. Here designated.

This species was described from 2 macr. males. Both specimens are in
the USNM.

Phyllothrips citricornis Hood, 1908 (2):305.

Lectotype ? (macr.).—USA: Illinois, Dubois, 5 May 1908 (L. M. Smith).
On hickory leaves. USNM Type 74842. Here designated.

This species was originally described from an unspecified number of
females from Illinois and Pennsylvania. In addition to the lectotype there
are 8 female paralectotypes and 3 males labelled ‘paratype’ in the USNM
and 1 female paralectotype in the BMNH.

Phyllothrips umbripennis Hood, 1909 (4):30-31.

Lectotype ? (macr.)—USA: Illinois, Carbondale, 12 October 1908
(L. M. Smith). “Jarred from post oak.” USNM Type 71241. Designated
by ONeill, Arnaud and Lee (1971:25).

This species was originally described from ‘many specimens of both
sexes. In addition to the lectotype there are 13 female and 10 male para-
lectotypes in the USNM and 2 female paralectotypes in the BMNH.

Plectrothrips antennatus Hood, 1908 (1):370-371.

Lectotype 2 (macr.).—USA: Illinois, Urbana, 23 June 1908 (J. D. Hood).
On outside of woodshed window. USNM Type 74893. Here designated.

This species was originally described from 2 females and 5 males. In
addition to the lectotype there are 1 female and 3 male paralectotypes in
the USNM and 1 male paralectotype in the BMNH.

Plemmelothrips defectus Hood, 1957 (170):144-145.
Holotype 2 (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, February

VOLUME 80, NUMBER 2 287

1954 (F. Plaumann). Under fallen leaves. Hood No. 2950. USNM Type
74183.

This species was originally described from 4 macr. females, 15 brach. fe-
males and 11 brach. males. None of these were labelled as Plemmelothrips
defectus Hood. The holotype was labelled in ink on the glass slide “Plem-
melothrips deficiens.’ An associated label bears ‘deficiens’ and this has been
struck through and replaced by ‘defectus.’ There is no doubt that the
specimens represent the type-series of defectus Hood, and I have labelled
them as such.

Priesneriella citricauda Hood, 1927 (69):199.

Lectotype 2 (apt.)—USA: California, Palo Alto, 4 August 1927 (J. D.
Hood). Beating dead branches of Salix sp. Hood No. 744. USNM Type
71242. Designated by O’Neill, Arnaud and Lee (1971:25).

This species was originally described from an unspecified number of
apt. males and females. In addition to the lectotype there are 10 apt. fe-
male and 10 apt. male paralectotypes in the USNM and 2 apt. female para-
lectotypes in the BMNH.

Pygothrips albiceps Hood, 1938 (116):401-402.

Lectotype 2 (macr.)—USA: Florida, Homestead, 28 December 1927
(J. C. Bradley and J. D. Hood). Under bark on dead branches. USNM Type
75061. Here designated.

This species was originally described from 5 macr. females and | apt.
male from Florida.

Rhopalothrips bicolor Hood, 1912 (7):73-74.

Lectotype 2 (brach.)—MEXICO: Topo, Chico, 4 July 1908 (C. A. Hart).
On leaves of Opuntia. USNM Type 74622. Here designated.

This species was originally described from 49 brach. females including
a ‘paratype. In addition to the lectotype there are 27 brach. female para-
lectotypes in the USNM and 3 brach. female paralectotypes in the BMNH.

Rhynchothrips capnodes Hood, 1955 (163):106-108.

Holotype ° (macr.)—BRAZIL: Sao Paulo, Boracea, Munic de Sale-
sopolis (elevation 850 m), 8 July 1948 (J. D. Hood). From dead branches
on ground. Hood No. 1598. USNM Type 74239.

This species was originally described from a unique female holotype.
This was originally labelled ‘pullatus’ but this name has been struck through
by someone and ‘capnodes’ has been written below it.

Rhynchothrips rostratus Hood, 1927 (69):203.
Lectotype 2 (macr.)—USA: Arizona, Nogales, (Pajarita Mountains), 31

288 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

August 1927 (J. D. Hood). On oak. Hood No. 912. USNM Type 74920.
Here designated.

This species was originally described from an unspecified number of
females and males. In addition to the lectotype female there are 1 macr.
female and 2 macr. male paralectotypes in the USNM.

Saurothrips assai Hood, 1952 (146):171-172.

Lectotype 2 (macr.)—BRAZIL: Para, Belem, (Instituto Agronomico do
Norte), 11 August 1951 (J. D. Hood). Dead leaves of Euterpe olearacea.
Hood No. 2697. USNM Type 74823. Here designated.

This species was originally described from 11 macr. females and 9 macr.
males. In addition to the lectotype there are 8 macr. female and 8 macr.
male paralectotypes in the USNM.

Scopaeothrips unicolor Hood, 1912 (7):71-72.

Lectotype 2 (brach.)—USA: Texas, Brownsville, (South Texas Garden),
29 June 1908 (C. A. Hart). On Opuntia. USNM Type 74619. Here desig-
nated.

This species was originally described from ‘an excellent series of both
sexes. In addition to the lectotype there are 10 brach. female and § brach.
male paralectotypes in the USNM and 4 brach. female and 12 brach. male
paralectotypes in the BMNH.

Sedulothrips tristis Hood, 1934 (87):434.

Lectotype 2 (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
4 July 1933 (J. D. Hood). Dead leaves and branches of pomarosa (Eugenia
jambos L.). Hood No. 971. USNM Type 71243. Designated by O'Neill,
Arnaud and Lee (1971).

This species was originally described from 24 females and 10 males ‘taken
on Barro Colorado Island (type-locality) Canal Zone and at Porto Bello,
Panama.’

Smicrothrips particula Hood, 1952 (146):173-174.

Lectotype ° (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, 17
February 1949 (F. Plaumann). On Lantana. Hood No. 2036. USNM Type
No. 74818. Here designated.

This species was originally described from an unspecified number of fe-
males. In addition to the lectotype there are 4 macr. female (including 1
dealate female) paralectotypes in the USNM.

Sophiothrips comptus Hood, 1955 (163):69-72.

Holotype ¢° (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, Feb-
ruary 1950 (F. Plaumann). No further data. Hood No. 2084. USNM Type
74254.

VOLUME 80, NUMBER 2 289

This species was originally described from a unique holotype female. This
specimen was originally labelled ‘pictus’ but someone has struck this
name through and has written “‘comptus’ below it.

Sophiothrips panamensis Hood, 1934 (87):428-429.

Lectotype ? (macr.)—PANAMA: Porto Bello, 10 July 1933 (J. D. Hood).
Under bark on dead branches. Hood No. 989. USNM Type 74930. Here
designated.

This species was originally described from 2 macr. females and 1 brach.
female from Panama.

Sophiothrips squamosus Hood, 1934 (87):426-428.

Lectotype 2 (macr.)—PANAMA: Porto Bello, 1933 (J. D. Hood). Under
bark on dead branches. USNM Type 74926. Here designated.

This species was originally described from 3 macr. females, 7 brach. fe-
males, 2 brach. males and 8 brach. males from Panama.

Stephanothrips bradleyi Hood, 1927 (69):204.

Lectotype 2 (apt.)—USA: California, Palo Alto, 4 August 1927 (J. D.
Hood). Beating dead branches of Salix sp. Hood No. 744. USNM Type
74615. Here designated.

This species was originally described from an unspecified number of
apt. males and females. In addition to the lectotype there are 2 apt. female
and 8 apt. male paralectotypes in the USNM and 2 apt. male paralectotypes
in the BMNH.

Stephanothrips carolina Hood, 1938 (119):366-367.

Lectotype ? (apt.)—USA: North Carolina, Rocky Point, (Pender Co.),
21 October 1937 (R. W. Leiby and J. D. Hood). Dead branches of sweet
gum. USNM 74616. Here designated.

This species was originally described from 24 apt. females and 1 apt. male
including a female ‘paratype’ and a male ‘allotype. In addition to the
lectotype there are 18 apt. female and 1 apt. male paralectotypes in the
USNM and 1 apt. female paralectotype in the BMNH.

Strepterothrips conradi Hood, 1934 (87):431-434.

Lectotype 2 (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
9 July 1933 (J. D. Hood and J. Zetek). Dead vegetation. Hood No. 987.
USNM Type 71244. Designated by O’Neill, Amaud and Lee (1971).

This species was originally described from 52 females and 15 males from
‘dead branches of various species of trees and vines.’

Symphyothrips caliginosus Hood, 1952 (146):163-164.
Lectotype ? (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, 13 Jan-

290 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

uary 1949 (F. Plaumann). Dry branches. Hood No. 2018-H. USNM
Type 74735. Here designated.

This species was originally described from 3 females. In addition to the
lectotype there is 1 macr. female paralectotype in the USNM.

Terthrothrips bullifer Hood, 1957 (170):149-150.

Holotype 2° (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, May
1953 (F. Plaumann). Under fallen leaves. Hood No. 2112. USNM Type
74177.

This species was originally described from 2 macr. females, 1 macr. male,
1 brach. male and 1 other male. The type-series in the USNM comprises all
of these except the odd male of unknown morph. The holotype was labelled
‘Terthrothrips calcaratus’ in ink on the glass slide. An associated card how-
ever is labelled “Terthrothrips bullifer, there is no doubt that the specimens
represent the type-series of bullifer Hood, and I have labelled them as such.

Terthrothrips carens Hood, 1957 (170):147-148.

Holotype 2 (macr.)—BRAZIL: Santa Catarina, Nova Teutonia, April
1954 (F. Plaumann). Under fallen leaves. Hood No. 2947. USNM Type
74181.

This species was originally described from 3 macr. females and 5 macr.
males. There are 2 macr. females and 4 macr. males in the USNM and 1
macr. female in the BMNH. None of these were labelled as carens Hood.
The holotype and an associated paper label both bear ‘caritus.’ There is
no doubt however that these specimens represent the type-series of carens
Hood, and I have labelled them as such.

Terthrothrips hebes Hood, 1957 (170):152-153.

Holotype 2? (brach.)—BRAZIL: Santa Catarina, Nova Teutonia, Feb-
ruary 1954 (F. Plaumann). Under fallen leaves. Hood No. 2950. USNM
Type 74173.

This species was originally described from 6 brach. females and 4
brach. males. The type-series in the USNM comprises only 5 brach. females
and 3 brach. males, and none of these were labelled as Terthrothrips hebes
Hood. An associated card is labelled “Terthrothrips hebetatus’ Hood. How-
ever the data on the slides correspond with the published data of hebes and
there is no doubt that these specimens represent that species. Two of the
specimens were labelled ‘HOLOTYPE’ and ‘ALLOTYPE’ respectively.
All of the specimens of the type-series have now been labelled as hebes
Hood. There is also 1 female paratype in the BMNH.

Trachythrips albipes Hood, 1933 (84):214.
Lectotype 2 (apt.)—PANAMA: Panama Canal, Barro Colorado Island,

VOLUME 80, NUMBER 2 291

Gatun Lake, 29 July 1933 (J. D. Hood and J. Zetek). Dead branches. Hood
No. 1018. USNM Type 74617. Here designated.

This species was originally described from ‘many mounted specimens of
which several are males. In addition to the lectotype there are 28 apt.
female and 3 apt. male paralectotypes in the USNM and 1 apt. female para-
lectotype in the BMNH.

Trachythrips deleoni Hood, 1933 (84):213-214.

Lectotype 2 (apt.)—PANAMA: Porto Bello, 9 July 1933 (J. D. Hood).
Dead vegetation. Hood No. 987. USNM Type 71245. Designated by
O'Neill, Arnaud and Lee (1971:26).

This species was originally described from 35 apt. females and 11 apt.
males. In addition to the lectotype there are 27 apt. female and 8 apt. male
paralectotypes in the USNM and 1 apt. female and 1 apt. male paralecto-
types in the BMNH.

Trachythrips frontalis Hood, 1933 (84):214-215.

Lectotype 2 (apt.)—PANAMA: Canal Zone, Frijoles, 7 July 1933 (J. D.
Hood). Dead vine and bush. Hood No. 981. USNM Type 71246. Desig-
nated by O’Neill, Arnaud and Lee (1971:26).

This species was originally described from a ‘large number of . . . speci-
mens of which several are male.’ In addition to the lectotype there are
28 apt. female and 2 apt. male paralectotypes in the USNM and 1 apt.
female paralectotype in the BMNH.

Trachythrips seminole Hood, 1939 (125):613-615.

Holotype 2 (apt.)—USA: Florida, Matheson Hammock, (near Miami),
30 December 1937 (J. D. Hood). Among dead fallen leaves. Hood No. 1377.
USNM Type 74621.

Hood (1939 (125)) described Trachythrips seminole as a new species
and then in the same paper treated it as a new name for an unnamed variety
of watsoni. Bailey (1949) refers to this as watsoni var. fairchildi Watson,
1939.

Trichinothrips latifrons Hood, 1955 (163):83-84.

Holotype 2 (macr.).—BRAZIL: Sao Paulo, Sao Carlos, at Fazenda Salto
(elevation about 837 m), 14 June 1948 (J. D. Hood and D. P. de Souza
Dias). Dead and dying branches of orange trees. Hood No. 1624. USNM
Type 74248.

This species was described from a unique holotype female from “BRAZIL
Sao Carlos, S.P. 14 June 1948, J.D.H. ... from dead and dying branches of
orange trees. There is a single slide, bearing the data cited for the holotype
above, in the USNM. Apart from the data label this slide bears only

292 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the word ‘HOLOTYPE 1. An associated card bears the name “Trichinothrips
frontalis.’ There is little doubt however that this specimen is the unique
holotype of latifrons Hood, and I have labelled it as such.

Trichothrips americanus Hood, 1908 (1):366-367.

Lectotype ? (brach.)—USA: Illinois, Homer, (University forest), 20
March 1907 (J. D. Hood). Under bark on rotten maple stump. USNM Type
74777. Here designated.

This species was originally described from ‘several specimens from Car-
bondale, Homer and Urbana, Illinois.’ In addition to the lectotype there
are 5 macr. female, 25 brach. female and 4 brach. male paralectotypes in
the USNM and 2 brach. female and 1 brach. male paralectotypes in the
BMNH.

Trichothrips angusticeps Hood, 1908 (1):36.

Lectotype ° (brach.)—USA: Illinois, St. Joseph, 4 May 1907 (C. A.
Hart and J. D. Hood). Under bark on rotten oak stump. USNM Type 74779.
Here designated.

This species was originally described from 7 brach. females and 1 brach.
male. In addition to the lectotype there are 3 brach. female and 1 brach.
male paralectotypes in the USNM.

Trichothrips anomocerus Hood, 1912 (8):137-142.

Lectotype 2 (brach.)—USA: Maryland, Plummer’s Island, (near Wash-
ington, D.C.), 18 February 1912 (W. L. McAtee). Under sycamore bark.
USNM Type 71247. Designated by O'Neill, Arnaud and Lee (1971:26).

This species was originally described from 15 brach. females and 7 brach.
males. In addition to the lectotype there are 7 brach. female and 7 brach.
male paralectotypes in the USNM.

Trichothrips buffae Hood, 1908 (1):369.

Lectotype 2 (brach.)—USA: Illinois, Decatur, 22 February 1908 (J. Zetek
and F. C. Gates). Under bark on soft maple tree. USNM Type 74925. Here

designated.
This species was originally described from ‘several brachypterous speci-
mens of both sexes . . . from Homer, Decatur and Urbana, Illinois.’ In

addition to the lectotype. there are 4 brach. female and 2 brach. male para-
lectotypes in the USNM.

Trichothrips graminis Hood, 1934 (87):409-410.

Lectotype ? (macr.)—PANAMA: Canal Zone, (J. D. Hood). Panicum
maximum. USNM Type 74793. Here designated.

This species was originally described from 3 macr. females, 9 brach. fe-
males and 2 brach. males from Panama.

VOLUME 80, NUMBER 2 293

Trichothrips longitubus Hood, 1908 (1):368.

Lectotype ° (macr.)—USA: IUlinois, Carbondale, 19 May 1908 (C. A.
Hart). Sweepings along railroad track. USNM Type 74830. Here desig-
nated.

This species was originally described from 9 macr. females and 1 macr.
male. In addition to the lectotype there are 3 macr. female and | macr. male
paralectotypes in the USNM.

Trichothrips mediamericanus Hood, 1934 (87):138-139.

Lectotype ° (macr.)—PANAMA: Porto Bello, 11 July 1933 (S. J. Hook,
H. H. Hood, J. D. Hood and J. Zetek). Dead branches of cacao. Hood No.
990. USNM Type 71248. Designated by O'Neill, Arnaud and Lee (1971).

This species was originally described from a ‘large number of specimens
of both sexes taken at Porto Bello, Panama (type-locality) on Barro Colorado
Island and at Frijoles.’

Trichothrips militaris Hood, 1935 (87):411.

Lectotype 2° (macr.)—PANAMA: Canal Zone, Barro Colorado Island,
July 1933 (J. D. Hood). Dead branches. USNM Type 74785. Here desig-
nated.

This species was originally described from 5 macr. females from Barro
Colorado Island and Porto Bello, Panama.

Trichothrips smithi Hood, 1909 (4):29-30.

Lectotype 2 (apt.)—USA: HUlinois, Bosky Dell, 20 October 1908 (L. M.
Smith). “Jarred from hard maple branch.” USNM Type 74965. Here desig-
nated.

This species was originally described from 2 apt. females. Both are in the

USNM.
Zeugmatothrips badiipes Hood, 1937 (108):292-295.

Lectotype ? (macr.).—PERU: Quayabamba, Departamento de Amazonas,
(elevation about 1,300 m), 14-19 August 1936 (F. Woytkowski). Dead
branches. Hood No. 1134. USNM Type 75074. Here designated.

This species was originally described from 6 macr. females and 3 macr.
males. In addition to the lectotype there are 3 macr. female and 3 macr.
male paralectotypes in the USNM and 1 macr. female paralectotype in the
BMNH.

Zeugmatothrips cinctus Hood, 1952 (146):170.

Lectotype 2° (macr.)—BRAZIL: Para, Belém, (Instituto Agronomico do
Norte), 14 August 1951 (J. D. Hood and J. M. Pires). Dead leaves of Astro-
caryum murumuru. Hood No. 2703. USNM Type 75082. Here designated.

294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

This species was originally described from 16 females and 11 males from
dead leaves of several genera of palms. In addition to the lectotype there are
11 macr. female and 7 macr. male paralectotypes in the USNM and I macr.
female paralectotype in the BMNH.

Zeugmatothrips mumbaca Hood, 1952 (146): 169-170.

Lectotype ° (macr.)—BRAZIL: Para, Belém, (Instituto Agronomico do
Norte), 11 August 1951 (J. D. Hood). Dead leaves of Asterocaryum
mumbaca. Hood No. 2696. USNM Type 75075. Here designated.

This species was originally described from 9 macr. females and 7 macr.
males. In addition to the lectotype there are 5 macr. female and 5 macr.
male paralectotypes in the USNM and 1 macr. female and 1 macr. male
paralectotypes in the BMNH.

Zeugmatothrips niger Hood, 1952 (146):168—-169.

Lectotype 2 (macr.)—BRAZIL: Para, Belém, (Instituto Agronomico do
Norte), 6 August 1951 (J. D. Hood). Dead leaves of Euterpe oleracea. Hood
No. 2673. USNM Type 75079. Here designated.

This species was originally described from 10 macr. females and 3 macr.
males. In addition to the lectotype there are 5 macr. female and 2 macr.
male paralectotypes in the USNM and 1 macr. female in the BMNH.

Zygothrips americanus Hood, 1912 (9):114—-115.

Lectotype 2 (brach.)—USA: Illinois. USNM Type 74721. Here desig-
nated.

This species was originally described from 50 brach. females, 3 macr. fe-
males and 10 brach. males from various localities in Illinois, Michigan,
Missouri, Nebraska and Maryland.

Acknowledgments

I am greatly indebted to Dr. Laurence Mound and Miss Kellie O'Neill
who instigated my visit to Washington to study Hood’s thrips collection.
Dr. Don Davis and the authorities of the Smithsonian Institution made
my visit. possible. Miss Kellie O’Neill provided much advice and en-
couragement during my visit. To all these I express my sincere thanks.

Literature Cited

Bailey, S. F. 1949. An annotated bibliography of North American Thysanopterists.
Part II. Fl. Entomol. 32(1):11-36.

Hood, J. D. 1948. J. Douglas Hood: Bibliography of Scientific papers. Rev. Entomol.
(Rio de J.) 19(3):499-508.

———. 1958. The terms tergum and sternum, tergite and sternite. Syst. Zool. 7(3):
131-133.

VOLUME 80, NUMBER 2 295

Jacot-Guillarmod, C. F. 1970. Catalogue of the Thysanoptera of the World (Part 1).
Ann. Cape Prov. Mus. Nat. Hist. 7(1):i-216.

——.. 1971. Catalogue of the Thysanoptera of the World (Part 2). Ann. Cape
Prov. Mus. Nat. Hist. 7(2):217-515.

—. 1974. Catalogue of the Thysanoptera of the World (Part 3). Ann. Cape
Prov. Mus. Nat. Hist. 7(3):517-976.

—. 1975. Catalogue of the Thysanoptera of the World (Part 4). Ann. Cape

Prov. Mus. Nat. Hist. 7(4):977—1255.

Catalogue of the Thysanoptera of the World (Parts 5-7). (In press.)

Mound, L. A. 1976. American leaf litter Thysanoptera of the genera Erkosothrips,
Eurythrips and Terthrothrips (Phlaeothripidae: Phlaeothripinae) Bull. Br. Mus.
Nat. Hist. Entomol. 35(2):27-64.

Mound, L. A., and K. O’Neill. 1977. Lectotype designation for Glyptothrips ar-
kansanus Hood (Thysanoptera: Phlaeothripidae) Proc. Entomol. Soc. Wash.
79(2):272.

O'Neill, K. 1971. Chthonothrips nigrocinctus Hood, lectotype designation and de-
scription of male (Thysanoptera: Phlaeothripidae). Proc. Entomol. Soc. Wash.
73(3):336.

——. 1974. J. Douglas Hood: Bibliography of scientific papers, chiefly on
thrips, and index of new names, 1948-1960. Proc. Entomol. Soc. Wash. 76(3):
297-309.

—. 1977. Corrections to “J. Douglas Hood: Bibliography of Scientific papers,
chiefly on thrips, and index of new names, 1948-1960.” Proc. Entomol. Soc.
Wash. 79(3):488.

O'Neill, K., P. H. Arnaud, and V. Lee. 1971. Lectotype designations for certain
species of Thysanoptera described by J. D. Hood. J. Wash. Acad. Sci. 61(1):
24-26.

Wilson, T. 1975. A monograph of the Subfamily Panchaetothripinae (Thysanop-
tera: Thripidae). Mem. Am. Entomol. Inst. (Ann Arbor) 23;1-354.

Department of Entomology, British Museum (Natural History). London,
SW7 5BD, England.

PROC. ENTOMOL. SOC. WASH.
80(2), 1978, pp. 296-308
HABITATS OF LARVAL TABANIDAE (DIPTERA)
IN SOUTH TEXAS!

Patrick H. Thompson, Edward J. Gregg, Donald R. Charanza,
Ralph A. Sauseda and Joseph W. Holmes, Jr.

Abstract—Through the use of physical and chemical methods, the
habitats and interspecific associations of 12 species of Tabanidae in SE
Texas are described. The Navasota River floodplain near College Station was
the most prolific source, producing 298 larvae in 28 soil treatments with
pyrethroid emulsions applied 29 March-12 August 1976. Five species were
represented in the sample population of larvae here: Tabanus atratus F.,
41.8%; T. proximus Walker, 36.8%; T. subsimilis subsimilis Bellardi, 14.4%;
T. trimaculatus Palisot de Beauvois, 5.6%; and T. lineola F., 1.4%. This
total of five species comprised less than one-fifth of the 26 species taken as
adults by Gressitt Traps the previous season (1975); among the 21 species
not collected as larvae in 1976 were numerous forms comprising up to
8.3% of the adult sample population. Conversely, the most abundant
species represented in larval collections was T. atratus, a form including
only 0.1% (14 specimens) of the adults collected in 1975. Larvae of one or
two species predominated at any given site and time, with only several
specimens of a 2nd or 3rd species being found under these conditions.
Low indices of affinity suggest that no specific association existed between
larvae of any two species found together.

The data on larvae reported here were taken in conjunction with those
of adults published for several ecosystems of SE Texas; coastal marshes
(Thompson, 1973a); coastal prairies (Thompson, Blume and Aga, 1977); the
Pine Belt (Thompson, 1973b, 1974b, 1976); and the Post Oak Belt (Thomp-
son, 1974a, 1976, 1977). Because of the kinds of ecosystems selected and
the collecting methods used, these initial studies were not very productive.
Then the discovery of larvae of Tabanus subsimilis subsimilis Bellardi
in varied upland soil situations, and the subsequent successful use of py-
rethrin emulsions for collecting this species for rearing purposes (Thomp-
son, 1975), stimulated further application of this insecticide formulation for
locating larval populations of Tabanidae. Use of pyrethrins improved pro-
ductivity—i.e., more larvae were removed from larger areas in less time—
and surveys, rather than collections, became the major effort.

During the last two years then, larval populations were sampled in eco-
systems which demonstrated large populations of Tabanidae in previous
research with adults, which were located near the laboratory and which
provided soils where intoxicated larvae could be seen after surfacing. The

VOLUME 80, NUMBER 2 297

latter criterion required that surface treatments be made in soils lacking
the dense sod of marshes and grasslands or pasture and lawn turfs. There-
fore, the forest floors of a river floodplain and an upland wooded ridge
some 12 mi away provided the most suitable substrata for application of
pyrethrin emulsions. This paper presents data based upon these collections,
and others using mechanical methods, from south Texas study areas.

The Study Areas

The physiography and vegetation of the two primary study areas near
College Station were described by Thompson (1974a, 1977). Briefly, the
upland locality is a ridge of home properties and adjoining woodlots, the
latter consisting largely of post oaks (Quercus stellata Wang.) on the
ridges proper, and water oaks (Q. nigra L.) in the gullies and washes. The
brushy shrub understory consists mostly of yaupon (Ilex vomitoria Ait.)
and dense lianas. Some 12 miles away, the Navasota River floodplain
forms a 2-mile-wide basin overlain with three natural vegetative cover
types: Upland Forest, like that previously described, on the upper slopes
and flats; Transition Forest on the basal slopes below; and Bottomland
Forest on the floodplain proper. These forests are comprised of woody plant
communities, the nine dominant overstory species of which form a con-
tinuum, gradually replacing one another from the uplands onto the flood-
plain, and then to the river margin, and are: post oak, black hickory
(Carya texana Buckl.), winged elm (Ulmus alata Michx.), overcup oak
(Quercus lyrata Walt.), willow oak (Q. phellos L.), cedar elm (Ulmus
crassifolia Nutt.), water elm (Planera aquatica Gmel.), swamp _ privet
(Forestiera acuminata (Michx.) Poir.), and black willow (Salix nigra L.).
The topography of these lowlands is frequently flooded for short periods
and occasionally, for weeks or months.

Methods

Initial studies employed a variety of mechanical devices and methods of
extraction: A kitchen sieve; a modified Berlese-Tullgren Funnel apparatus;
a hand cultivator for garden use; salt flotation; and a “comb” made from
a wooden dowel and finishing nails (Thompson, 1970a). In the work re-
ported here, the hemispherical kitchen sieve was replaced by one made
from a stainless steel food service tray (12 X 20 X 2.5 in). The floor of the
tray was cut out and covered with % in mesh hardware cloth or 16-mesh wire
screen—the size depending upon the texture of the soil being processed. This
tray-sieve increased the volume and the sieving surface of the soil being
examined and was invaluable for processing sands and alluvium in stream
beds and in river floodplains. In order to find equipment for processing
more soil in less time, several power-driven farm and garden implements

298 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

were also used. A tractor-driven chisel was used to examine floodplain
soils along the Bois d’Arc Creek (where heavy populations of Tabanus s.
subsimilis were then seriously infesting the Red River watershed). Al-
though no larvae were found as the gang of chisel blades cut and turned
the sod to a depth of 1-2 in, the presence of earthworms and white grubs
in the furrows or excavated soil showed that if numerous tabanid larvae
had been present, some would have been exposed in these examinations.
(Under similar circumstances—in plow furrows of cultivated fields—Davis
(1919) and his colleagues found larvae of T. sulcifrons Macquart in Kansas,
Maryland and Mississippi.) In addition to the chisel, a gasoline-driven
rotary tiller was used to turn soils previously shown to contain T. s. sub-
similis, again with negative results.

Two commercial pyrethroid formulations were used as sources of the
active ingredient: An oil solution of pyrethrins (Gulfspray, Gulf Oil
Corp.) which was emulsified with Triton X-100 (Rohm and Haas Co.) be-
fore addition of tap water and an emulsifiable concentrate of resmethrin
(Super Syn 30EC, Redmond Chemical, Inc., Houston, Texas). After the re-
moval of organic debris, 0.002% emulsions were applied as drenches to
measured areas of soil surface. Where rainfall and evaporation allowed,
1 gal of solution per sq yd was usually sufficient to saturate mineral soils. In
most cases, larvae were sought in treated plots for at least 2 h after treat-
ment. Observations, during two treatments yielding 59 and 61 larvae of
T. s. subsimilis (other details of study were presented by Thompson
(1975)), showed that intoxicated larvae can emerge up to 18 h after treatment
and that a large percentage of these, one-third to one-half, can be expected
to surface after the first hour posttreatment. Several important pro-
cedures for posttreatment handling of intoxicated specimens can increase
their survivorship for later rearing purposes. During the present study, up
to half of those larvae emerging within the first several hours, when first
washed with a mild Triton X-100 solution in water and then held in the
lab at 20-25°C, were suitable for rearing in the laboratory.

Considering the temperature, pyrethroids are more active at lower
temperatures down to 5°C, like DDT, whereas these materials become
less effective at temperatures approximating 20-25°C. Toxicological effects,
too, become very significant in affecting behavior and survival after treat-
ment. For example, pyrethrins are more effective irritants upon insects
than synthetic pyrethroids; on the other hand, synthetic pyrethroids, such as
resmethrin, are more potent insecticides. Finally, in addition to their noxious
effects upon insects, the dermatogenic and allergenic properties of these
chemicals should be seriously considered by those persons using them.

Immatures were maintained in 2 oz clear glass jars, the Bakelite lids
of which were center-drilled with 1 in holes and covered by fine-mesh
metal screen. The substratum of washed builder's sand was washed with
tap water weekly and the tabanid larvae were fed those of house flies

VOLUME 80, NUMBER 2 299

and stable flies. A 14% h photoperiod was provided with a timer-controlled
15 W fluorescent bulb.

Results and Discussion

But for the numerous larvae of T. s. subsimilis reported from two grassy
seepage areas near domestic septic tanks (Thompson, 1975), and for lesser
numbers of this species taken here in several subsequent collections, only
one soil treatment in that locality produced larvae of any other species.
In Thompson’s four treatments totaling 81 sq ft of soil, nine larvae were
recovered from the damp and leaf-littered depressions of dry-wash gullies
in the adjacent post oak forest; seven of those larvae were T. s. subsimilis
(instead of one, as previously shown in Thompson's 1975 publication). The
two larvae remaining, included one specimen each of Tabanus atratus F.
and T. trimaculatus Palisot de Beauvois. The only species taken by me-
chanical methods in other collections included one additional specimen of
these two species and four specimens of Leucotabanus annulatus (Say).
The larval habitats of these, and the other species considered here, will
be described in a later section.

Lowland Locale

The most productive collections were made with pyrethrins on litter-
laden soils beside standing water on the Navasota River floodplain near
College Station. Of 28 treatments applied on 18 days from 29 March-12
August 1976, 23 treatments were positive for larvae. These collections
yielded 285 specimens of five species and 13 more individuals of unde-
termined identity. Of the 298 specimens obtained, 1-38 were taken per
positive treatment, with an average of 13.

Relative abundance of different species in the larval sample population
and of species in the larval and adult sample populations —In decreasing
order of abundance, the five species collected as larvae were Tabanus
atratus, T. proximus Walker, T. s. subsimilis, T. trimaculatus, and T. lineola
F. (see Table 1 for numbers). These five tabanid species included only one-
fifth of the 26 species taken by Gressitt Traps with CO, the year before
(Thompson, 1977). Some of these 21 species were abundant or numerous
in the adult sample population; i.e., Tabanus sulcifrons, T. fuscicostatus
Hine and Hybomitra lasiophthalma (Macquart) comprised 8.3, 5.3 and 4.5%
of that catch, respectively. On the other hand, Tabanus proximus, T. s.
subsimilis and T. lineola, the three other species included among the six
most abundant species taken as adults in 1975, were easily observed in
larval collections in 1976 (Table 2). More significantly, the most abundant
species represented in larval collections was T. atratus, a form including only
0.1% of the adults collected in 1975.

Similar results to those above, based upon adult and larval collections

300 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Larval samples of five species of Tabanus collected by pyrethroid treat-
ments of soils on a Navasota River floodplain forest, Brazos Co., Texas, 29 March—12
August 1976 (positive treatments and identified specimens only).

Date atratus — proximus — subsimilis trimaculatus __ lineola Total
Mar. 29 3 4 fi
Apr. 1 14 3 17

2 2 2

9 fi: 2 9

19 3 3 6

19 iL 6 2 (DNs 10

19 8 2 1 11

May 24 1 1

24 27 2 29

24 18 18

25 3 6 4 2 15

25 MI 5 1 1 8

Jun. 11 10 10

14 JME 1 12

18 10 2 12

23 i 7

Aug. 2 13 2 4 19

3 8 14 1 5)

4 19 2 21

4 3 3

4 uf 7

9 35 3 38

12 1 ll

Totals 119 105 4] 16 4 285
Incidence? 52% 52% 48 % 3% 13%

“This figure represents one pupa of this species and is not included in the total of
larvae below, or in any other total presented in this paper.
» The percentage of 23 positive collections producing the species.

in two other Coastal Plain study areas, are presented in Tables 3 and 4.
Larval collections produced approximately one-quarter, or less, of the 40
species taken as adults in each study area. Based on data from the work at
both the Great Swamp National Wildlife Refuge (Thompson, 1967, 1970a)
and the Patuxent Wildlife Research Center (Thompson, 1970b, 1971),
ranking of species demonstrated little correlation between larval and adult
sample populations (Tables 3 and 4).

Larval sample populations in these ecosystems included one or two of
the most abundant dominants and ignored the presence of the others. On
the other hand, the one most abundant species of larva at Great Swamp
and Patuxent was much less numerous in adult collections: Tabanus
marginalis F. included less than 1% of the adult sample population (as for T.

VOLUME 80, NUMBER 2 301

Table 2. Comparison of larval sample population, Navasota River floodplain, 1976
season, with that of adults of Tabanidae from the same locale, 1975 season."

Larvae Adults
Rank % (number) Rank % (number)

Tabanus

atratus 1 41.8 (119) 18 0.1 (14)

proximus 2: 36.8 (105) 2 12.8 (1,648)

subsimilis 3 14.4 (41) 1 55.9 (7,172)

trimaculatus 4 5.6 (16) 8 2.0 (256)

lineola 5 1.4 (4) 3 4.3 (555)
21 spp. left 0 (0) 24.9 (3,195)
Totals 100 (285) 100 (12,840)

* Rank of adults is based upon a total of 26 species collected.

atratus at the Navasota River); and specimens representing either one or
both of T. melanocerus Wiedemann and T. petiolatus Hine (the two are
presently inseparable in the larval stage) dominated larval collections in
a population where the adults were only commonly collected, rather than
very numerous.

Density —From 1-8 sq yds of soil surface (total = 81) were treated in
those applications producing larvae. Densities ranged from 0.4-14.5 larvae
per sq yd, with an average of 3.7. There was no correlation between den-
sity and the presence of standing water or the presence of leaf litter, the
time of collection, the identity of the species collected, or the number of
species associated together.

Interspecific association—As mentioned previously, the most numerous
species in larval collections were also the most frequent species taken

Table 3. Comparison of larval and adult sample populations of Tabanidae, Great
Swamp National Wildlife Refuge, Morris Co., New Jersey 1966—1967.*:?

Larvae Adults
Rank % (number) Rank % (number)
T. marginalis F. 1 80.8 (184) iby 0.3 (20)
C. vittatus Wied. 2 13.1 (30) 2 21.6 (1,394)
T. lineola F. 3 S15) (8) 1 24.4 (aleayis))
C. univittatus Macq. 4 2.6 (6) 14 0.7 (50)
36 spp. left 0 (0) 53.0 (3,434)
Totals 100 (228) 100 (6,471)

* Rank of adults is based upon a total of 40 species collected.
» Several specimens of undetermined species of Atylotus and Hybomitra are not in-
cluded here.

302 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 4. Comparison of larval and adult sample populations of Tabanidae, Patuxent
Wildlife Research Center, Prince Georges Co., Maryland, 1968—-1969.*

Larvae Adults
Rank % (number) Rank % (number)
T. melanocerus Wied.; 1 44.7 (120) 14 0.7 (35)
T. petiolatus Hine? oa = — ll 4.4 (203)
C. vittatus Wied. 2 2527 (69) 1 13.8 (648)
T. trimaculatus Palisot
de Beauvois 3 18.6 (50) 8 6.3 (292)
T. lineola F. 4 3.0 (8) 9 5.6 (259)
T. sulcifrons Macq. 5 OMT (7) 13 1S (64)
T. nigripes Wied. 6 2.2 (6) 23 0.1 (6)
T. marginalis F 7 1.4 (4) 23 0.1 (6)
T. similis Macq. 8 0.7 (2) 32 S08 (1)
C. carbonarius Walk. 8 0.7 (2) SOs Onl (3)
C. dimmocki Hine 9 0.3 (1) 36 — > 0.1 (3)
29 spp. left 0 (0) 67.7 (3,181)
Totals 100 (269) 100 (4,701)

*Rank of adults is based upon a total of 40 species collected.
» These two species are indistinguishable in the larval stage.

then. Yet the incidence of more than two species together in any given
treatment was low (6 of 23 collections or 26%). Furthermore, the use of
Fager’s Index of ae for these treatments illustrates the low frequency
with which any 2 species were found together. This index equals twice the
number of joint occurrences divided by the total occurrences of both species
taken in all samples. Through the use of a table providing the minimum
values of joint occurrence which are significant at the 0.5 level (Fager,
1957), it is possible to recognize quickly pairs of species which are decidedly
associated versus those which show no evidence of affinity at all. At
some later time then, additional and more specialized sampling techniques
can be used to study those associations which are questionable.

With the exception of T. lineola, which was inadequately represented
in the sampling reported here, Fager’s Indices for each association are
shown in Table 5; testing these indices with the table provided by Fager,
no significant association was found between any two of the other four
species considered here.. Therefore, collection of any one of these species
did not increase the chance of finding any other species associated with it.

The low Indices of Affinity found here support the idea that these tabanid
species of the predominantly cannibalistic genus Tabanus compete fiercely
for space in mutually satisfactory habitats. Also, these low Indices for
the large T. atratus and T. proximus species could result more from their
dominant size than from their absolute abundance (as indicated by their

VOLUME 80, NUMBER 2 303

Table 5. Fager’s Indices of Affinity for five species of Tabanus associated to-
gether, Navasota River floodplain, Brazos Co., Texas, 29 March-12 August 1976.

atratus proximus subsimilis trimaculatus __ lineola*
atratus — 0.417 0.522 0.400 —
proximus 0.417 — 0.261 0.400 —
subsimilis 0.522 0.261 — 0.316 =
trimaculatus 0.400 0.400 0.316 == =

lineola*® == = = — =

“This species was inadequately represented for assignment of an index.

related abundance in samples of adult insects—T. proximus) or their habitat
specificity (both species).

Seasonal incidence—Moving down through the chronologically-ordered
list of treatments in Table 1, no striking seasonal differences are apparent
in the incidence of any particular species. But for T. trimaculatus and T.
lineola, the least numerous and least frequent species observed, larvae
were found throughout the entire period of collection—from spring through
summer. With rare exception, larvae of all species were no less than half
an inch long. Although no measurements of larval lengths were made,
the majority of specimens of each species taken in any given collection
were similar in size; this was especially noticeable for the large species,
T. atratus and T. proximus.

The Habitats

The larval stages of 12 species of Tabanidae were obtained by means of
pyrethroid treatments in upland and lowland locales near College Station
and by mechanical means from these and other study areas in SE Texas.

Chlorotabanus crepuscularis (Bequaert)—One larva was taken from the
marginal mud of a slough at the Navasota River Bottoms (NRB), 14 Sep-
tember 1971. Adults of this species were rare from this and the other
locales considered here.

Leucotabanus annulatus (Say).—Large larvae of this species were found
in their typical habitat, rotting deadfall (4 specimens, Upland Locale, Feb-
ruary—May collections; 1 specimen, Huntsville State Park, Walker Co.,
19 June 1972).

Tabanus atratus F.—One of the most common species found; one
specimen was removed from the grassy septic tank seepage area described
by Thompson (1975), where it was associated with 4 larvae of T. subsimilis
—subsimilis (8 June 1976). Some 35 yds away, another larva was found at
the margin of a small, shallow pond of some 500 sq ft of surface, which
was choked with pickerelweed (Pontederia) and duckweed (Lemna) (17
April 1974). Also, on a ridge near this site, Christopher Thompson found

304. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

a larva crawling on the soil surface—a larva which had apparently been
disturbed by his deep excavations several feet away (21 March 1976).
Larvae were found in shoreline situations near other standing waters (drain-
age ditch through a coastal marsh which was described by Thompson
(1973b), 12 July 1971; and a pond on a municipal golf course, College Sta-
tion, 1 April 1974). The most prolific sources were the margins of woodland
pools and sloughs at NRB; four larvae were found here (in the months of
January, February, June and August) before the insecticide treatments
later produced 119 specimens (Table 1). Of the latter samples, 116 speci-
mens were found in samples near standing water; the three specimens
remaining were removed from a leaf-littered wash.

Tabanus cymatophorus Osten Sacken.—One larva of this large, brightly-
marked fly, was taken from the marginal mud of a woodland pond at NRB.
Associated with several T. proximus larvae nearby in this March collection,
the larva was reared to the adult form (female) for determination. The in-
cidence of T. cymatophorus as an immature was low compared with the
commonness of the adults in 1975 (106 females, late June-early August;
Thompson, 1977).

Tabanus lineola F.—This species was found in three NRB collections
made with pyrethrin emulsions (Table 1).

Tabanus molestus Say.—One specimen, identified as a larva, was re-
moved from the shoreline of a slough at NRB, 14 September 1971. Adults
were not common in previous collections (30 specimens, mid-May to mid-
July).

Tabanus petiolatus Hine——Five larvae were taken from a sandy wash
across a firelane in a pine-hardwood forest, Huntsville State Park in Sep-
tember; later in October, three larvae were found in a sandy creek bank of
the same forest. Although three of these eight specimens were reared
for identification, the closely related T. melanocerus is presently un-
known from this study area. Lastly, one larva was taken with T. trimacu-
latus and three other species (see T. trimaculatus) from the Navasota River.
It was later reared for identification.

Tabanus proximus Walker—The most common form collected from
NRB before the 1976 collections using pyrethroids; six collections in March
and one in October produced 24 specimens from the margins of ponds
and pools. As in later collections with pyrethroids, T. proximus was as-
sociated with T. trimaculatus and T. atratus.

Tabanus subsimilis subsimilis BellardiSince the 1975 report, four ad-
ditional collections of T. s. subsimilis were made in the same habitats de-
scribed then: 1 pupa, in a flower bed skirting a patio and 8 larvae in a
grassy alga-covered seepage area (May and June of 1976). In two June
collections, one larva was found at the margin of a grassy pasture pond,

VOLUME 80, NUMBER 2 305

surrounded by very dense willow saplings, and one larva was removed
from the bank of the Navasota.

Tabanus sulicifrons Macquart.—Two specimens, identified as larvae, were
taken from the marginal mud of a drainage ditch through a brackish
coastal marsh near Angleton in July (habitat described in Thompson,
1973a) and from a pile of dry and duff-like manure in the corner of a hay
shed near College Station in September. (Three pupal exuviae, believed to
represent a sibling species, were found several inches above the bottom
of a dry-wash gully in association with three pupal exuviae of T. proximus.)

Tabanus trimaculatus Palisot de Beauvois.—This species was represented
in collections from all major locales studied: In a grassy seepage area (1
specimen) together with large numbers of T. s. swbsimilis (Thompson, 1975);
at the margin of a 2-acre impoundment with Ludwigia submerged along
the littoral zone nearby and smartweed (Polygonum) at the shoreline above
it (1 larva); in the sandy-creek bed of a pine-hardwood forest, the site pre-
viously described for Tabanus petiolatus, but on a different occasion (2
larvae); as the predominating species in marginal mud of NRB sloughs in
association with Chlorotabanus crepuscularis, Tabanus atratus, T. molestus
and T. petiolatus (7 specimens); at this slough and at woodland pools nearby
in March of the following year, in association with 11 specimens of T.
proximus and 1 of T. atratus (7 specimens); and dominating the latter spe-
cies here the next October (17 specimens to 4 of T. atratus); and finally, again
in March of the following year, from a woodland pond margin here, where
1 specimen was a small minority among the 13 T. proximus specimens
found.

Tabanus venustus Osten Sacken.—One specimen, later reared to the
adult form (female), was found in March at the margin of a small pond
near the College Station municipal airport.

Conclusions

If the adults of Coastal Plain Tabanidae are ubiquitously obvious to the
observer, the larvae are patently obscure. Paradoxically, this situation is
occasionally reversed, with larval numbers in samples greatly exceeding
those of adults: Tabanus reinwardtii Wiedemann (Cameron, 1926; Philip,
1931; Schwardt, 1936; Stone, 1930; and Pechuman, 1972); T. marginalis
(Philip, 1931; Teskey, 1969; and Thompson, 1970a (q.v., Table 3)); Mery-
comyia brunnea Stone (Jones and Anthony, 1964); and Merycomyia whitneyi
(Johnson) (Pechuman, 1964; Goodwin, 1973; and Philip, Weems, and
Fairchild, 1973). In either case of adult or larval preponderance, this dis-
parity between numbers of the two forms has complicated population esti-
mation and retarded understanding of larval ecology.

306 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Comprehension of tabanid larval ecology has also been obscured by the
relative ease of finding larvae in marginal habitats, by the relative difficulty
of finding them in deeper soils in habitats where they have been previously
established and by the consequent understandable biases of scientists
focusing their attentions on marginal niches and surficial soils.

This information, based upon accumulated field experience, has been
reenforced by several notions originated and perpetuated in the literature—
notions categorizing the immatures of Tabanidae, collectively, as aquatic
or semiaquatic insects, rather than as soil insects; as specialized forms in
choice of habitat niches, instead of as generalized ones; and as air-breathing
insects, rather than as those depending very largely upon cutaneous respira-
tion.

Thus biased collecting and published misconceptions regarding the larvae
of the family have led to arbitrary classification of groups of species ac-
cording to habitat. Usage of such terms as aquatic, littoral, submerged
and terrestrial have been applied to certain species rather than to the habi-
tats where those species were observed in specific instances. (The larvae of
the so-called terrestrial species have been so described because they were
found away from habitats near any accumulations of free water. Al-
though some Tabanidae are obviously adapted morphologically for aquatic
existence and others are commonly found submerged, proximity to free
water probably has little causal relationship in habitat selection for most
species of the Atlantic Coastal Plain of North America.)

In conclusion, the cumulative effect of this experience and these ideas in
North American research has been to encourage the search for habitats
which are as distinctive as the species inhabiting them and to discourage
recognition and study of the critical factors limiting the distribution of
tabanid larvae in general. For it is the general body of knowledge about
larval habitat selection—the critical environmental characteristics many spe-
cies of Tabanidae hold common—that reveals most about the biology of
most of these species as individuals. To wit, the life of the generalized larval
type is unknown. The large body of published larval collection data for
Coastal Plain species, the most obvious habitat descriptions of which have
been conveniently summarized in tabular form for 109 species and sub-
species by Goodwin (1967), documents the generalized and nonspecific
nature of this habitat selection.

Acknowledgments

We gratefully acknowledge the help of Mr. Lee W. Bailey for allowing
us the use of his land on the Navasota River, in this and in past research; of
Mr. Mason C. Cloud, Jr., Texas Forest Service, for providing literature
sources; of Mr. Jeffrey B. Tucker in aiding the search for larvae of T. s.
subsimilis on the Bois d’Arc; of Dr. F. W. Plapp for technical advice in

VOLUME 80, NUMBER 2 307

the pertinent toxicology; of the Drs. L. E. Ehler, L. L. Pechuman, and H. J.
Teskey for their comments on the manuscript; and of Christopher L. Thomp-
son for his astute observation and collection of a passing T. atratus larva.

Literature Cited

Cameron, A. E. 1926. Bionomics of the Tabanidae (Diptera) of the Canadian Prairie.
Bull. Entomol. Res. 17:1—42.
Davis, J. J. 1919. Contributions to a knowledge of the natural enemies of Phyllophaga.
Ill. Nat. Hist. Surv. Bull. 13:53-138 (Tabanidae, pp. 97-100).
Fager, E. W. 1957. Determination and analysis of recurrent groups. Ecology 38:
586-595.
Goodwin, J. T. 1967. Contributions to the taxonomy of immature horse and deer
flies (Tabanidae, Diptera). PhD dissertation, Univ. Tenn., Knoxville, 125 pp.
——. 1973. Immature stages of some Eastern Nearctic Tabanidae (Diptera). IV.
The Genus Merycomyia. J. Tenn. Acad. Sci. 48:115-118.
Jones, C. M., and D. W. Anthony. 1964. The Tabanidae (Diptera) of Florida.
USDA Tech. Bull. No. 1295, 85 pp.
Pechuman, L. L. 1964. A synopsis of Merycomyia (Diptera: Tabanidae). Proc.
Entomol. Soc. Ont. 94:62-67.
—. 1972. The horse flies and deer flies of New York (Diptera, Tabanidae).
Search 2:1-72.
Philip, C. B. 1931. The Tabanidae (Horseflies) of Minnesota. Minn. Agr. Expt.
Stn. Tech. Bull. No. 80, 132 pp.
Philip, C. B., H. V. Weems, Jr., and G. B. Fairchild. 1973. Notes on Eastern
Nearctic Haematopota, Merycomyia, and Chrysops, and description of male of
C. zinalus. Fla. Entomol. 56:339-346.
Schwardt, H. H. 1936. Horseflies of Arkansas. Univ. Ark. Agr. Expt. Stn. Bull. No.
332, 66 pp.
Stone, A. 1930. The bionomics of some Tabanidae (Diptera). Ann. Entomol. Soc.
Amer. 23:361—304.
Teskey, H. J. 1969. Larvae and pupae of some eastern North American Tabanidae
(Diptera). Mem. Entomol. Soc. Can. No. 63, 147 pp.
Thompson, P. H. 1967. Abundance and seasonal distrubution of the Tabanidae
(Diptera) of the Great Swamp, New Jersey. Ann. Entomol. Soc. Am. 60:1255—
1260.
—. 1970a. Larval Tabanidae (Diptera) of the Great Swamp, New Jersey. Ibid.
63 :343-344.
—. 1970b. Tabanidae (Diptera) of Patuxent Wildlife Research Center, Laurel,
Maryland. Ibid. 63:572-576.
1971. Larval Tabanidae (Diptera) of the Patuxent Wildlife Research Center,
Laurel, Maryland. Ibid. 64:956-957.
1973a. Tabanidae (Diptera) of Texas. I. Coastal marsh species, West Gal-
veston Bay; incidence, frequency, abundance, and seasonal distribution. Proc.
Entomol. Soc. Wash. 75:359-364.
1973b. Tabanidae (Diptera) of Texas. II. Pine Belt species, Huntsville
State Park; incidence, frequency, abundance, and seasonal distribution. Ibid.
75:430-435.
1974a. Tabanidae (Diptera) of Texas. HII. Post Oak Belt species, Navasota
River Watershed; incidence, frequency, abundance, and seasonal distribution.

Ibid. 76:35-38.

308 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

——. 1974b. Tabanidae (Diptera) of Texas. IV. Pine Belt species, the Big Thicket;
incidence, frequency, abundance, and seasonal distribution. Ibid. 76:315-321.

—. 1975. Larval habitats of Tabanus subsimilis subsimilis Bellardi in southeast
Texas (Diptera, Tabanidae). Ibid. 77:494—500.

—. 1976. Tabanidae (Diptera) of Texas. V. Second-year collections from
Huntsville State Park and the Navasota River Watershed. Ibid. 78:376-381.

—. 1977. Tabanidae (Diptera) of Texas. VII. Comparisons of upland and
lowland tabanid populations in southeast Texas. Ibid. 79:564—574.

Thompson, P. H., R. R. Blume, and A. Aga. 1977. Tabanidae (Diptera) of Texas.
VI. Coastal Prairie species, Victoria County. Ibid. 79:266—-269.

Veterinary Toxicology and Entomology Research Laboratory, Agric. Res.
Serv., USDA, College Station, Texas 77840.

Footnote

‘This paper reports the results of research only. Mention of a pesticide in this paper
does not constitute a recommendation for use by the U.S. Department of Agriculture
nor does it imply registration under FIFRA as amended.

VOLUME 80, NUMBER 2 309
NOTE

WILLIAM DWIGHT PIERCE, BIOGRAPHICAL NOTES AND A
REVIEW OF HIS BOOK THE DEADLY TRIANGLE

W. Dwight Pierce, as he was often known, died nearly 11 years ago, but
no obituary has come to my attention. During his long life (16 November
1881-29 April 1967) he was productive in several aspects of our science
for more than 60 years. The recent appearance of his unusual book on en-
tomological history (The Deadly Triangle. A brief history of medical and
sanitary entomology. 1975. 138 pp.) makes timely both a review of the
book and brief biographical notes. This small, soft-bound, posthumously and
privately published book is distributed in accordance with Dr. Pierce’s
will by the executor of his estate, Dr. Richard B. Loomis, Department of
Biology, California State University, Long Beach, California 90840. The
supply is limited, but there may be a second printing.

Largely in language understood by the layman, for Dr. Pierce often
lectured before general groups and felt that entomologists should do a better
job explaining science to the public, he has attempted to give a broad
view of the early history of science, especially the background of those
diseases of man that later were found to be transmitted by arthropods. A
typical “triangle,” in terms of the title, is represented by a man, a mosquito,
and the causative organism (Plasmodium) of human malaria. Much of
Dr. Pierce’s experience in the field of medical entomology dates from
the World War I period when he headed a committee responsible for making
information on entomology available to the armed forces. However, he
refers (page 99) to an unpublished compilation of over 2,800 pages covering
more than 500 diseases of man and animals, so the subject was of long-time
interest to him.

The book lacks illustrations except for a diagram on the cover and, un-
fortunately, the offset type of printing is noticeably small. The writing is
rather anecdotal in style, probably unsuitable for beginning students, but
much background information of potential interest to advanced entomol-
ogists and sanitarians is included. About 45 pages deal with the nature of
diseases in early times; then there are brief accounts of malaria and nu-
merous Other arthropod-borne diseases, with particular reference to the
period when proofs of transmission were first obtained. Later, the urgent
problems brought about by World War I are discussed. World War II,
with its more global problems, is discussed in less detail than it deserves; for
example, there is a decidedly short account of dengue, which was a serious
disease on Saipan. Finally, there is an expanded discussion of 12 principles
governing insect transmission of disease, a list of literature cited, index to
authors and other persons mentioned, and a short subject index. A few

310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Dr. W. Dwight Pierce (right) shown with Dr. Paul H. Arnaud on a balcony of the
Los Angeles County Museum in late 1953 (from a kodachrome by Gurney).

of the publications cited appeared as recently as the late 1950’s and 1960,
but a hasty check shows that Dalmat’s important volume (Dalmat, H. T.
1955. The Black Flies (Diptera, Simuliidae) of Guatemala and Their Role as
Vectors of Onchocerciasis. 425 pp.), Cushing’s short history (Cushing, E. C.

VOLUME 80, NUMBER 2 311

1957. History of Entomology in World War II. 117 pp.), and Zinsser’s
classic work (Zinsser, H. 1934. Rats, Lice and History. 301 pp.) are
omitted. More recently, several other books that cover some of the same
historical aspects as Pierce’s, have appeared, one of the most recent being
a very attractive and readable book (Cloudsley-Thompson, J. L. 1976.
Insects and History. 242 pp.).

Dwight Pierce was born in Illinois and studied entomology at the Uni-
versity of Nebraska where he obtained bachelor’s and master’s degrees. In
1917 he received the doctorate in entomology from George Washington
University, Washington, D.C. He mentions that one of his first jobs was
packing chinch bug fungus in Professor Lawrence Bruner’s laboratory in
1901 for distribution to farmers. He was employed in 1904-1919 by the
U.S. Department of Agriculture and worked on insects attacking field crops,
especially cotton, as well as those affecting the health of man and animals.
A 1912 paper by Pierce et al. on the relationships of a large number of
insects (pests, parasites, hyperparasites, predators) directly or indirectly
associated with the cotton plant was an outstanding early example of the
interdependent associations of insects and plants (Pierce, W. D., R. A.
Cushman and C. E. Hood. 1912. The Insect Enemies of the Cotton Boll
Weevil, U.S. Dept. Agr., Bur. Entomol. Bull. 100:1-99). An instructive
diagram from the paper, which illustrates the relationships of this boll
weevil complex, was reproduced in a standard text (Allee, W. C., O. Park,
A. E. Emerson, T. Park and K. P. Schmidt. 1949. Principles of Animal
Ecology. 837 pp.) as a notable example of interspecies relationships.
Pierce also became an active systematist on weevils and Strepsiptera. He
was located in Washington, D.C. a portion of the time but at other times
chiefly in the South. In 1917, with the assistance of about a dozen col-
leagues, Pierce published a book on potential harmful insect introductions
(Pierce, W. D. (Ed.) 1917. A Manual of Dangerous Insects Likely to be In-
troduced in the United States through Importations. 256 pp.). During World
War I he was a leader in several activities of the Bureau of Entomology that
were planned to give full assistance in solving military problems that in-
volved insects. He describes on page 69 how a special class of about 25
entomologists was formed, and 500 copies of the proceedings were mailed
weekly to entomologists and military representatives throughout the U.S.
This was the genesis of another book (Pierce, W. D. (Ed.) 1921. Sanitary
Entomology. The entomology of disease, hygiene and sanitation. 518 pp.).
About 10 collaborators assisted him in the book’s preparation.

After leaving USDA employment, Dr. Pierce worked as a consultant,
during which period he had assignments in the Philippine Islands and
elsewhere, before he began serving on the staff of Biological Abstracts in
1931-1936. From 1937 he was associated with the Los Angeles County
Museum in California, where he again did systematic work on a variety
of insects including fossils from calcareous nodules of Miocene age found in

312 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the Calico Mountains of southern California and others from Pleistocene
or more recent deposits of the Rancho La Brea asphalt pits. A second
posthumous publication was issued privately (Pierce, W. D. 1975. The
Sand Dune Weevils of the genus Trigonoscuta with a correlation of their
anatomy to the Geological History of our Coast Lines. 160 pp.). A few
details of Dr. Pierce’s study habits during his last years are contained
in the Editor's Comments of the Trigonoscuta paper. He continued to
do some teaching and to provide consulting service. He and his wife,
who also has died, had no children.

Dwight Pierce seems to have been a consistently imaginative, studious
entomologist, glad to tackle new and difficult problems. Even if not al-
ways successful in an outstanding way, he made important and unforgettable
contributions. In the historical account reviewed here, he tried to plant
new ideas for further investigation, though some may not appear orthodox
now. During his career, medical entomology developed from its infancy
as a discipline into a major speciality with outstanding accomplishments of
international scope. From the time of his earliest participation in this spe-
cialty, during which he was associated with L. O. Howard, W. D. Hunter,
and F. C. Bishopp, he appears to have been constantly recording the back-
ground events, trying to place developments in the context of fundamental
principles, and explaining the subject to both the lay public and pro-
fessional workers seeking more information. Throughout his career, he
was a scholarly naturalist of the old school. At the time of his death he
was one of the oldest members, in both age and length of membership,
of the Entomological Society of Washington.

Ashley B. Gurney, Cooperating Scientist, Systematic Entomology Labora-
tory, IIBIII, Agric. Res. Serv., USDA, c/o U.S. National Museum, Washing-
ton, D.C. 20560.

NOTE

NEW DISTRIBUTIONAL RECORDS FOR TWO SPECIES OF
NERTHRA SAY FROM MEXICO
(HEMIPTERA: GELASTOCORIDAE)

A small collection of Gelastocoridae from México recently identified for
Dr. Harry Brailovsky A., Instituto de Biologia, México, D. F. included
the following new distributional records.

Nerthra spangleri Polhemus, 1972. Proc. Entomol. Soc. Wash. 74(3):306.
26 and 28, Isla Isabel, Nayarit, México, 31 Jan. 1976. J. Palacios.

VOLUME 80, NUMBER 2 313

Polhemus listed Sinaloa, Sonora and Colima in the original description, so
the occurrence in Nayarit is not surprising. The species will undoubtedly
eventually be found in Jalisco as well.

Nerthra usingeri Todd, 1954. Pan-Pac. Entomol. 30(2):116. A_ single
6 collected on the Estacion de Biologia, Chamela, Jalisco, México, 5
Dec 1976, H. Brailovsky. This is first Mexican record for the species. It
was previously known only from California.

E. L. Todd, Systematic Entomology Laboratory, IIBIII, Agric. Res. Serv.,
USDA c/o NHB-127, U.S. National Museum, Washington, D.C. 20560.

NOTE

A “NOCTURNAL” FORAGING RECORD FOR
DIOGMITES NEOTERNATUS (DIPTERA: ASILIDAE)

Robber flies are generally considered to be diurnal insects usually be-
ing active between about 9:30 AM and 7:30 PM. However, some species,
in particular those in the desert, have been observed to mate during the
night (Lehr, 1959, Proc. Fourth Congr. All Union Entomol. Soc. 1:76-78;
Lehr, 1964, Proc. Sci. Res. Inst. Protection Plants, Alma-Ata. 8:213-244 (In
Russian); Newkirk, 1963, Ann. Entomol. Soc. Am. 56:234-236). In addition,
Rau (1938, Ann. Entomol. Soc. Am. 31:540-556) reported Deromyia ternatus
Loew foraging and capturing house-flies on a screen-door of a city shop
at dusk. Presumably, this species was using the light from the shop to
see and capture its prey. Since such behavior among Asilidae is in-
frequently reported, we would like to report the following similar “noc-
turnal” foraging behavior for Diogmites neoternatus (Bromley). It is also
of interest to note that to our knowledge, this is the first published record
of this asilid occurring in Virginia.

While collecting insects in the vicinity of Fairfax, Virginia, we frequently
found D. neoternatus in open weedy fields, on the edges of forested areas
and less frequently in the forests. We also found numerous specimens
of this species trapped in the staircase of our three-story apartment build-
ing. Diogmites neoternatus have been found by other investigators in
open areas of dry fields (Scarbrough, 1974, Proc. Entomol. Soc. Wash.
76:385-396), in clover fields (Artigas, 1966, Ohio J. Sci. 66:401-421), and
moist bushy woods or fields (Bromley, 1931, Ohio State Univ. Mus., Sci.
Bull. 1:3-19; Bromley, 1950, Ann. Entomol. Soc. Am. 43:227-239).

On 24 July 1976 at 12:15 PM, we observed several D. neoternatus resting
on the walls of the aforementioned well-lit apartment building staircase,

314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

as well as flying about the staircase (according to Bromley, 1950, Ibid.,
D. neoternatus “has been taken at light,” which may mean that this species
was collected at night near a light). Much to our surprise one male captured
a prey (Diptera: Xylomyidae, Xylomya sp.) in flight by the light of the
staircase. The male then flew around with his fore legs held above his
thorax in a posture similar to that exhibited by male D. angustipennis Loew
during courtship (see Lavigne and Holland, 1969, Univ. Wyo. Agr. Exp. Stn.
Sci. Monogr., No. 18, 61 pp.), held the prey between his middle and hind legs
and tried to “grab” onto the walls. He eventually held onto a crevice in the
wall with his left fore leg, inserted his hypopharynx in the prey and rested
on the wall with all six legs. As the male fed he frequently held onto the
wall with one of his fore legs, manipulated the prey with the rest of his
legs and reinserted his hypopharynx. The male would then either hang
onto the wall for a short period of time with one fore leg or immediately
grab onto the wall with all six legs. Similar methods of manipulating prey
have been observed for other species of Diogmites (Bromley, 1946, Bull.
Conn. Geol. Nat. Hist. Surv. No. 69, 48 pp.; Lavigne and Holland, Ibid.).
We continued to watch the male until we disturbed him and he dropped
the prey in flight. In addition to feeding on a dipteran, as reported here,
Artigas (Ibid.) states that D. neoternatus feeds principally on bees and
wasps, and occasionally flies and “bugs.”

We would like to thank R. J. Lavigne (Entomology Section, University
of Wyoming, Laramie) for his confirmation of the identity of Diogmites
neoternatus.

D. Steve Dennis and Jeanne A. Gowen, 3065 South Cathay Circle, Aurora,
Colorado 80013.

VOLUME 80, NUMBER 2 315
BOOK REVIEW

A Bibliography of Quantitative Ecology. 1977. V. Schultz, L. L. Eber-
hardt, J. M. Thomas, M. I. Cochran. 361 pages. Dowden, Hutchinson &
Ross, Inc., Stroudsberg, Pennsylvania. Cost $18.50.

This bibliography includes over 2,000 references arranged in 28 topics
relating to quantitative ecology including age, frequency distribution,
models, sampling, and taxonomy. References are through 1974, and there
are some titles from 1975. The format of the book is such that entries are
arranged in two columns per page with the pages consisting of offset re-
production of camera-ready copy prepared from computer printout or what
would appear to be computer printout. Within each topic the papers are
arranged alphabetically by the surname of the first author. The index
presented at the end includes the names of all authors. Thus, second and
third authors of joint publications can be identified.

Information retrieval from the bibliography is a problem. If one wants
to know something about negative binomial distributions, he will have to
scan all of the titles under frequency distribution or know that C. I. Bliss,
R. A. Fisher, and a few other statisticans work in the research field. A sub-
ject index would have been useful.

Some of the papers have pertinence in more than one field, but each
paper is listed only once under the 28 topics. Because of the way the
bibliography is arranged, it is impossible to identify a paper which tangen-
tially approaches a topic in which the user wants more information. An
in-depth cross index would have added substantially to the value of this
book.

Another source of improvement would be uniform application of key
words following each citation. Some references have key words added, but
others do not.

Considering a topic as broad as quantitative ecology, it would be nearly
impossible to prepare a complete bibliography. I suspect that the 2,000
plus references are comparable to the emergent part of an iceberg: seven-
eighths of its mass remains submerged. For instance, nearly all of the
references are English language papers, but there are innumerable non-
English language papers on quantitative ecology.

All things considered, this is a reasonable first attempt at collecting the
literature about quantitative ecology. However, in thumbing through the
book I had the feeling one has when he goes to an unfamiliar library: the
information is there, but how does one find it?

Gordon Gordh, Systematic Entomology Laboratory, IIBIII, Agric. Res.
Serv., USDA (present address: Division of Biological Control, Department
of Entomology, University of California, Riverside, California 92502).

316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
SUMMARY REPORTS OF SOCIETY OFFICERS FOR 1977

Treasurer

(1 November 1976 to 31 October 1977)

Special
General publication
SUMMARY: fund fund Total
On hand, 1 Nov. 1976 1,194.51 18,386.72 19,581.23
Total receipts 31,996.40 7,429.07 39,425.47
Total disbursements 29,607.49 887.50 30,494.99
On hand, 31 Oct. 1977 3,583.42 24,928.29 28,511.71

Editor
(Calendar Year 1977)

Four numbers of the Proceedings were published in 1977. The 662 pages
printed represented 84 scientific articles, 16 notes, 3 book reviews, 3 an-
nouncements and the minutes for 7 meetings of the Society. Page charges
were waived for 14 articles totaling approximately 74 pages. Full page
charges were paid for immediate publication for 7 articles totaling 63
pages.

The January 1977 issue of the Proceedings was dedicated as The Alan
Stone Commemorative Issue on the occasion of Dr. Stone’s 73rd birthday.
A new format for articles was instituted with the July 1977 issue. Adoption
of the new format will mean that larger print will be used throughout the
articles, that there will be more type/page, and that, with several minor
editorial changes, the Society will realize a reduction in its printing costs.
Page charges were raised to their present level in December 1975. No in-
crease in page charges is expected at this time. Costs of reprints were raised
in March 1977.

In September 1977, the Society sponsored the publication of “Pictorial
Key to Species of the Genus Anastrepha (Diptera: Tephritidae)” by George
C. Steyskal. This article is now available from the Custodian for $1.50.

Publications Committee: Earlene Armstrong, Barnard D. Burks, Ashley
B. Gurney, George C. Steyskal, Manya B. Stoetzel (Editor).

PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

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Memnnsneers (On (Cymapid alls) 0k Fk 6.00
Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman __ 35
Unusual Scalp Dermatitis in Humans Caused by the Mite Dermato-
UII Ly UR. WANT a .25
A Short History of the Entomological Society of Washington, by Ashley
CURES a SR ae el ae 50

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(EE SVEN CHGS C75 | ee tg a 1.50

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
No. 1. The North American Bees of the Genus Osmia, by Grace

Sandhouse. 1939 tte dai aE ee ere Sesh $15.00
No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga,

by amamG. Boving. 1942 pee oe ee! 15.00
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figs roo bat VWalson (Oman. 1949 . 15,00
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Back issues of the Proceedings of the Entomological Society of Washington are
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the Society.

Prices quoted are U. S. currency. Dealers are allowed a discount of 10 per cent
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orders should be placed with the Custodian, Entomological Society of Wash-
ington, c/o Department of Entomology, Smithsonian Institution, Washington,
D.C. 20560

CONTENTS

(Continued from front cover)

Lectotype designations of certain species of thrips described by J. D. Hood and
notes on his collection (Thysanoptera) B. R. PITKIN

Terminalia of some North American species of Megaselia (Aphiochaeta) and de-
scriptions of two new species (Diptera: Phoridae) W. H ROBINSON

Ethology of Cerotainia albipilosa Curran (Diptera: Asilidae) in Maryland: Court-
ship, mating and oviposition A. G. SCARBROUGH

Synopsis of the North American Pyrgotidae (Diptera) G. C. STEYSKAL

Habitats of larval Tabanidae (Diptera) in south Texas
P. H. THOMPSON, E. J. GREGG, D. R. CHARANZA,
R. A. SAUSEDA and J. W. HOLMES, JR.

NOTES:

A “Nocturnal” foraging record for Diogmites neoternatus (Diptera: Asilidae)
D. S. DENNIS and J. A. GOWEN

William Dwight Pierce, biographical notes and a review of his book The Deadly
Triangle A. B. GURNEY

New distributional records for two species of Nerthra Say from México (Hemiptera:
Gelastocoridae) E: L. 2ODD

BOOK REVIEW:
A Bibliography of Quantitative Ecology G. GORDH
SUMMARY REPORTS OF SOCIETY OFFICERS FOR 1977 .

216

179 5

149

296

313

309

312

315
316

VOL. 80 JULY 1978 a NO. 3
m 70673 NURSON 4a

“PROCEEDINGS.

of the Tr higee

INTOMOLOGICAL SOCIETY
ot WASHINGTON

DEPARTMENT OF ENTOMOLOGY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY

CONTENTS

Morning glory bees and the Ipomoea pandurata complex (Hymenoptera: An-
thophoridae) D: FE. AUSTIN: © 397

Oviposition and host feeding behavior of Aphelinus asychis (Hymenoptera: Chal-
cidoidea: Aphelinidae) on Schizaphis graminum (Homoptera: Aphididae)
and some reactions of aphids to this parasite H. BOYLE and E.M.BARROWS 441

Taxonomic notes on Zagrammosoma, a key to the Nearctic species and descriptions
of new species from California (Hymenoptera: Eulophidae) G. GORDH 344

A new stonefly from West Virginia (Plecoptera: Chloroperlidae)
P. P. HARPER and R. F. KIRCHNER 403

Two new Ceratocapsus Reuter 1876, from the eastern United States (Hemiptera:
Miridae) T. J. HENRY 383

Description and life cycle of a new species of Histiostoma (Acari: Histiostomi-

dae) associated with commercial mushroom production
A. HILL and K. L. DEAHL 317

Two new species of Tarsonemus (Acari: Tarsonemidae) associated with commercial
mushroom production A. HILL and K. L. DEAHL 330

A new species of Pygmephorus (Acari: Pygmephoridae) associated with commercial
mushroom production A. HILL and K. L. DEAHL 335

The immature stages and biology of Mallota posticata (Fabricius) (Diptera:
Syrphidae) C. T. MAIER 424

(Continued on back cover)

en,

ENTOMOLOGICAL SOCIETY

OF WASHINGTON
ORGANIZED Marcu 12, 1884

OFFICERS FOR 1978 |

Doucias W. S. SUTHERLAND, President HELEN SOLLERS-RIEDEL, Hospitality Chairwoman
Donatp R. Davis, President-Elect Victor E. ADLER, Program Chairman
Wayne N. Maruis, Recording Secretary Joyce A. Urmar, Membership Chairwoman;
DonaLp R. WHITEHEAD, Corresponding Secretary Sueo NakaHara, Custodian}
F. CurisTIAN THOMPSON, Treasurer D. W. S. SurHERLAND, Delegate, Wash. Acad. Sei.

Manya B. StoerzeE., Editor
Publications Committee
EARLENE ARMSTRONG ASHLEY B. GURNEY
WayneE E. CLark GrEorGE C. STEYSKAL

Honorary President
C. F. W. MuESEBECK

Honorary Members
FREDERICK W. Poos Ernest N. Cory RayMonp A. St. GEORGE

All correspondence concerning Society business should be mailed to the appropriate officer at the following
address: Entomological Society of Washington, c/o Department of Entomology, Smithsonian Institution,’
Washington, D.C. 20560.

MEETINGS.—Regular meetings of the Society are held in the Ecology Theater, Natural History Building,
Smithsonian Institution, on the first Thursday of each month from October to June, inclusive, at 8 P.M.
Minutes of meetings are published regularly in the Proceedings.

MEMBERSHIP.—Members shall be persons who have demonstrated interest in the science of entomology,
Annual dues for members are $9.00 (U.S. currency) of which $8.00 is for a subscription to the Proceedings
of the Entomological Society of Washington for one year.

PROCEEDINGS.—Published quarterly beginning with January by the Society at Washington, D.C. Members

in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember sub-
scriptions are $15.00 per year, domestic, and $17.00 per year, foreign (U.S. currency), payable in advance,
All remittances should be made payable to The Entomological Society of Washington.

The Society does not exchange its publications for those of other societies.

Please see inside back cover of any 1972—1975 issue for instructions regarding preparation of manuscripts

STATEMENT OF OWNERSHIP }
Title of Publication: Proceedings of the Entomological Society of Washington. "
Frequency of Issue: Quarterly (January, April, July, October).

Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society ol
Washington, c/o Department of Entomology, Smithsonian Institution, 10th and Constitution NW,
Washington, D.C. 20560.

Editor: Dr. Manya B. Stoetzel, Systematic Entomology Laboratory, BARC-W, Beltsville, Maryland 20705)
Managing Editor and Known Bondholders or other Security Holders: none. 4

i ;
This issue was mailed 18 July 1978 qT
Second Class Postage Paid at Washington, D.C. and additional mailing office.

ALLEN PRESS, INC. eereD LAWRENCE, KANSAS

use [

PROG. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 317-329
DESCRIPTION AND LIFE CYCLE OF A NEW SPECIES OF
HISTIOSTOMA (ACARI: HISTIOSTOMIDAE) ASSOCIATED
WITH COMMERCIAL MUSHROOM PRODUCTION

Aagje Hill and Kenneth L. Deahl

Abstract—The immature and adult stages of a new species of Histiostomi-
dae, Histiostoma heinemanni, associated with commercial mushroom pro-
duction, are described and compared with those of H. feroniarum (Dufour).
Methods for rearing H. heinemanni, data on its biology and evidence that
it may spread mushroom pathogens are presented.

While collecting mites from commercial mushroom beds and compost piles
in Chester County, Pennsylvania, we frequently found adult and immature
stages of new species of Histiostoma, almost identical with H. feroniarum
(Dufour). Histiostoma feroniarum, which has also been found on mushrooms
as well as in many other habitats, has been redescribed by Scheucher (1957),
Hughes and Jackson (1958), and Jary and Stapeley (1936) under the name
Histiostoma rostro-serratum (Megnin). Hughes (1976) synonymized Anoetus
under Histiostoma. We are using her nomenclature for the idisomal struc-
tures.

This work was done in collaboration with the Systematic Entomology
Laboratory, IIBIII, Agricultural Research Service, U.S. Department ot
Agriculture, Beltsville, Maryland 20705. We undertook laboratory breed-
ing experiments to study the life-cycle of the new species of Histiostoma
and to collect all stages so that they could be described and figured.

Methods of Rearing and Course of Development

Histiostoma species have been associated with environments with high
relative humidity. Rearing experiments were started by transferring groups
of hypopi to petri dishes containing potato-dextrose agar and some decay-
ing potato that served as food. The dishes were sealed with tape to prevent
the hypopi from escaping and then placed in an incubator at 27°C. Some
hypopi entered the quiescent stage within a few hours after being transferred
to the dishes. The first tritonymphs were seen on the 2nd day and the
first eggs were seen on the 4th day. The females selected dry and com-
pacted material on which to lay eggs, either singly or in small clusters. The

first larvae were seen on the 5th day and the first protonymphs were
seen on the 6th day. Without overpopulation or food depletion the hypopus
stage was omitted and the first tritonymphs were seen on the 7th day. In-
creasing numbers of hypopi were found in dishes kept from 4-6 weeks. Like
Scheucher (1957), we saw hypopi standing on legs III and IV and making

|
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|
}

318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

searching movements with legs I and II, apparently ready to attach to the
moving objects, usually insects by which they are dispersed.

Parthenogenesis experiments were like the rearing experiments except that
only one hypopus was put in each petri dish. Only two of these hypopi
out of approximately 30 specimens developed into adult females that
laid eggs. Cultures became infested with fungi, which probably prevented
the development of these stages. Eggs of one of these females gave rise to
three tritonymphs, fourteen heteromorphic males and two homeomorphic
males; eggs of the other gave rise to one tritonymph, twenty-two hetero-
morphic males and twelve homeomorphic males.

Histiostoma heinemanni Hill and Deahl, new species

Diagnosis—The adults and immatures of Histiostoma heinemanni can
easily be distinguished from those of other Histiostoma spp. by the platelet
(pl) located between scapular internal setae (sci) and scapular setae (sce)
(Fig. 1). Gnathosoma (Figs. 36-37): Gnathosoma identical in immatures
and adults, except for size. Chelicerae (Fig. 36) bifurcate distally with 6
small teeth and laterally with 9 larger teeth; flagellum present; proximal
part of digitus fix us (df) with 3 projections; digitus mobilus (dm) smaller
than df. Pedipalps (p) (Fig. 37) with 2 flagella, subequal in length;
membranous structure present on venter of pedipalps.

Female

Figs. 1-6

Holotype.—Idiosoma pyriform, 440 u long and 227 mw wide; length of
paratypes (10) averages 363 (279-467) uw leng and 203 (147-267) mw wide;
cuticle with fine projections.

Dorsum (Fig. 1).—Anterior of propodosoma sculptured; vertical internal
setae (vi) in front of vertical external (ve); vi not enlarged basally; sci in
front of sce; sci and sce border platelet (pl). All prodosomal setae sub-
equal in length. Dorsal propodosoma and hysterosoma separated by trans-

verse groove. Setae dj, d;, l-l; and h smooth with enlarged bases and |

slender distally. Oil gland between d; and 1; ; 3 pores in same region.
Bursa copulatrix dorsal, located about '% the length of the hysterosoma from
caudal end.

Venter (Fig. 2)—Epimera I Y-shaped. Transverse genital opening be-
tween coxae II and II. Two pairs of almost circular rings, anterior pair
between coxae II and III, posterior pair at level of coxae IV. Setae
cx III slightly shorter and finer than cx I; setae ga and gp subequal in length
and somewhat longer than % the length of cx I; setae gp and subequal in
length; setae ay and a; slightly shorter than 2x the length of a,; ay (Fig. 1)
slightly shorter than 3X the length of aj.

VOLUME 80, NUMBER 3 319

Figs. 1-6. Histiostoma heinemanni, female. 1. Dorsum; 2. Venter; 3-6. Respectively,
legs I-IV.

320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Legs (Figs. 3-6).—With 5 free segments; tarsi relatively long, with stout
claw originating in short pretarsus. All setae spinelike. Chaetotaxy: Tarsi,
13-12-10-10; tibiae, 2-2-1-1; genua, 2-2-0-0; femora, 1-1-0-1; trochanters
1-1-1-0. Solenidiotaxy: Tarsi, 3-1-0-0; tibiae, 1-1-1-1; genua, 2-1-0-0. For
trochanters and femora see Figs. 1 and 2.

Male
Figs. 7-12

Two types of males are present in the cultures: Homeomorphic males with
normal size spines on tarsus II and heteromorphic males with two enlarged
spines on tarsus II. Intermediate stages are found. Idiosoma of 8 hetero-
morphic males averages 298 (255-319) mw long and 147 (128-160) pw wide.
Idiosoma of 7 homeomorphic males averages 262 (230-287) mw long and
127 (121-134) mw wide. Idiosoma more rectangular and smaller than in
females; cuticle covered with small projections.

Dorsum (Fig. 7).—Anterior of propodosoma sculptured; setae vi in front
of ve; sculptured platelet (pl) bordered by setae sci and sce. All propodo-
somal setae subequal in length, enlarged basally except vi slender distally.
Hysterosomal setae d,-d;, l-l; and h correspond with those of female
in relative size and distribution, but setae dy are absent. Oil gland between
setae d and l;; pore at level of |. |

Venter (Fig. 8)—Four chitinous ringlike structures at level of coxae IV.
Arrangement of setae and epimera as in female. Genitalia located about |
46 the length of the hysterosoma from caudal end; 2 pairs of internal setae, |
the shorter located more dorsally, and 4 pairs of anal setae.

Legs (Figs. 9-12).—Tarsi broader and shorter than in female. Chaeto- |
taxy and solenidiotaxy as in female, except for tarsus I, which has 10 setae |
and 2 solenidia. |

Tritonymph
Figs. 13-18

Idiosoma and gnathosoma of 10 specimens average 223 (121-294) pw long |
and 133 (70-185) j wide. The rather large variation in measurements
probably reflects the presence of both male- and female-forming trito- _
nymphs.

Dorsum (Fig. 13).—Anterior of propodosoma and platelet (pl) connected, —
both sculptured. Type, arrangement and relative size of dorsal setae similar |
to those of female; setae dy missing; dorsal setae located on distinct bosses. |
Dorsal pores missing.

Venter (Fig. 14)—Two pairs of sclerotic ringlike structures at level of |
coxae IV. Setae ga above anterior ringlike structure. Chaetotaxy similar to
that of female, but gp lacking. No sexual structures.

Legs (Figs. 15-18)—Chaetotaxy and solenidiotaxy as in female.

VOLUME 80, NUMBER 3 321

Figs. 7-12. Histiostoma heinemanni, male. 7. Dorsum; 8. Venter; 9-12. Respectively,
legs I-IV.

322 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 13-18. Histiostoma heinemanni, tritonymph. 13. Dorsum; 14. Venter; 15-18.
Respectively, legs I-IV.

VOLUME 80, NUMBER 3 323

Hypopus
Figs. 19-24

Idiosoma of 10 specimens averages 81 (103-217) long and 138 (74-160) uw
wide.

Gnathosoma (Fig. 20)—Elongate; ratio correlation of length; width is
1:4; 1 pair of flagellum longer than gnathosoma; anterolateral pair of
short setae present.

Dorsum (Fig. 19).—Propodosoma broad; anterior of propodosoma with
interrupted groove (as figured); setae vi in front of ve. Transverse groove
complete, posterior with broken-line area. Hysterosoma oval shaped; pos-
terior broadly rounded; surface smooth. Dorsal setae short, setiform, sub-
equal in length.

Venter (Fig. 20)—Epimera I fused medially; sternum free posteriorly,
epimera II connected with epimera III by a fine sclerotized extension from
latter; epimera III free; posterior and anterior of posterior sternum free.
Suckers on epimera IT on coxal plate III, and laterad on genital area. Setae
sh, genital setae, and “anal setae” small and setiform. Anal opening located
about 7% the length of the suctorial plate from caudal end.

Chaetotaxy of legs (Figs. 21-24).—Tarsi, 8-9; tibiae, 2-2; tibiotarsi, 7-7;
genua, 3-2-2; femora, 1-1-0-1; trochanters 0-0-1-0. Solenidiotaxy: Tarsi, 3-1;
tibae, 0-1; genua, 0-1-0.

Protonymph
Figs. 25-30

Average length of idiosoma of 7 specimens: 221 (160-262) mw long and
124 (108-134) px wide; cuticle punctate.

Dorsum (Fig. 25)—Anterior of propodosoma sculptured; platelet (pl)
bordered by setae sci and sce. Dorsal chaetotaxy as in female, but setae
relatively smaller.

Venter (Fig. 26)—One pair of sclerotic ringlike structures at level of
coxae IV. No sexual structures present. Chaetotaxy as in female, but setae
ga and gp lacking.

Leg (Figs. 27-30).—Legs differ from those of female in lack of spines on
trochanters I-III; leg IV differs also in absence of spine and famulus on
tibia and absence of 3 spines around pretarsus. Chaetotaxy: Tarsi, 13-12-
10-7; tibiae, 2-2-1-0; genua, 2-2-0-0; femora, 1-1-0-0; trochanters 0-0-0-0.
Solenidiotaxy: Tarsi, 2-1-0-0; tibiae, 1-1-1-0; genua, 2-1-0-0.

Larva
Figs. 31-35

Idiosoma of 8 specimens avergaes 160 (109-185) long and 93 (76-109)
wide.

;

324 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 19-24. Histiostoma heinemanni, hypopus. 19. Dorsum; 20. Venter; 21-24. Re- |
spectively, legs I-IV.

VOLUME 80, NUMBER 3 325

Figs. 25-30. Histiostoma heinemanni, protonymph. 25. Dorsum; 26. Venter; 27-30.
Respectively, legs I-IV.

326 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 31-35. Histiostoma heinemanni, larva. 31. Dorsum; 32. Venter; 33-35. Re-
spectively, legs I-III. Figs. 36-37. Histiostoma heinemanni, gnathosoma of all stages.

VOLUME 80, NUMBER 3 327

Dorsum (Fig. 31).—Anterior of propodosoma sculptured and contiguous
with sculptured platelet (pl). Dorsal setae on distinct sculptured bosses.
Chaetotaxy as in female, but d; and 1; absent.

Venter (Fig. 32)—One pair of chitinous ringlike structures anterior to
coxae II. Setae cx I, cx III and a, present; setae a;—az, ga and gp absent.

Legs (Figs. 33-35).—Leg I differs from that of female in absence of spine
on trochanter, 1 solenidion on tarsus, and famulus on pretarsus; legs II and
III differ in absence of spine on trochanter. Chaetotaxy: Tarsi, 13-12-10;
tibiae, 2-2-1; genua, 2-2-0; femora, 1-1-0; trochanters, 0-0-0. Solenidiotaxy:
Tarsi, 1-1-0; tibiae, 1-1-1; genua, 2-1-0.

Deposition of type material—Holotype: Female, USNM No. 3836 from
mushroom compost from the mushroom farm of R. Dowell, Toughkenamon,
Pennsylvania, 8 August 1976, by A. Hill and K. L. Deahl. Paratypes were
collected from horse manure, first-stage mushroom and mushroom compost
in Chester County, Pennsylvania as follows: R. Dowell, Toughkenamon,
8 August 1976; P. Yeatman, Avondale, 13 October 1976; Keystone Com-
pany, Coatesville, 31 August 1976; G. Guizetti, Toughkenamon, 15 Septem-
ber 1976; C. Nigro, Cochranville, 8 August 1976. Hypopi were also collected
from phorid flies caught in the mushroom houses. We also obtained sev-
eral stages from the Experimental Mushroom House, BARC-West, USDA,
Beltsville, Maryland. Paratypes are deposited with the following: U.S. Na-
tional Museum of Natural History, Washington, D.C.; Institute de Mede-
cine Tropicale, Antwerp, Belgium; Ryksmuseum van Natuurlijke Historie,
Leiden, Netherlands; Dr. R. L. Heinemann, Longwood College, Farm-
ville, Virginia; Acarology Laboratory, Ohio State University, Columbus,
Ohio; Catholic University, Nijmegen, Netherlands; Hungarian Natural His-
tory Museum, Budapest, Hungary.

Etymology.—This species is named for Dr. R. L. Heinemann, Long-
wood College, Farmville, Virginia.

Discussion

In the family Histiostomidae, the generic classification has been based on
the hypopus stage. Differences between hypopi can be minute and older
descriptions were sometimes incomplete, so identification has been difficult.
Except for the absence of the broken-line area posterior to the transverse
groove and contiguity of the epimera IV, the hypopus of H. feroniarum
described by Scheucher (1957) is identical with hypopus of H. heinemanni
in that the former has an extra pair of dorsal setae and one sensory seta
on tarsus I instead of two.
A significant difference between the two species is the presence of a
platelet on the dorsal propodosoma of adults and developmental stages of
the species only, this platelet has not been described previously. The form
of the platelet is probably growth dependent because the platelet is some-

328 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

times medially split into two or nearly so. In some specimens (Fig. 13)
the platelet is connected with the anterior sculptured part of the pro-
podosoma, but usually it is separate and bordered by setae sci and sce.
Compatible with previous descriptions are dorsal bosses, obvious in the de-
velopment stages but also readily visible in the adults.

For H. feroniarum, Scheucher (1957) shows 11 pairs of dorsal setae and
3 pairs of anal setae in the female and the male; Hughes and Jackson (1958)
show 12 pairs of dorsal setae in the female and 11 in the male, and 3 pairs
of anal setae for both. Histiostoma heinemanni has 11 pairs of dorsal and
4 pairs of anal setae in the female and 10 pairs of dorsal and 4 pairs of anal
setae in the male. Also, Scheucher (1957) figures only 1 sensory seta on
tarsus I of the female of H. feroniarum, whereas 2 sensory setae are present
on tarsus I of the female of H. feroniarum, as described by Hughes and
Jackson (1958), and on tarsus I of the female of H. heinemanni, as described
here. Furthermore, Scheucher (1957) describes a second claw lacking in
other descriptions on tarsi I and II of the male of H. feroniarum. The life
cycle of H. heinemanni seems very similar to that described for H. fero-
niarum except that we found both homeomorphic and heteromorphic males
in the culture, whereas Scheucher (1957) found only heteromorphic males of
H. feroniarum. The difference could be inherent in the species or could
be a result of environmental conditions. In the mushroom houses we fre-
quently saw many H. heinemanni on mushrooms that were decaying as a
result of infection with Verticillium malthousei Ware, Pseudomonas tolaasii
Paine (Deahl, unpublished) and various secondary saprophytic invaders.
Histiostoma heinemanni apparently does not feed on the mushroom tissue
but is probably attracted by the presence of the microorganisms. How-
ever, by feeding on the microorganisms, this mite may be important in the
spread of diseases. Only one species, Histiostoma gracilipes (Banks) is known
to injure mushrooms by feeding on the spawn (Compton, 1933).

Acknowledgments

We thank Edward W. Baker, Systematic Entomology Laboratory, I[BIII, —
Agricultural Research Service, USDA, and R. L. Heinemann, Longwood ;
College, Farmville, Virginia, for their help with writing this paper. |

Literature Cited |
Compton, C. C. 1933. Successful mite control saves costly mushroom loss. 46. Ann. |
Rept. Ill. Agric. Expt. St. for 1932-1933. Pp. 156-158.
Jary, S. G., and J. H. Stapeley. 1936. Investigations on the insect and allied pests |
of cultivated mushrooms—IV. Observations upon the tyroglyphid mite Histio-
stoma rostro-serratum Megnin. T.S.—E. Agric. Coll. Wye. 38:67-74.
Hughes, A. M. 1976. The mites of stored food and houses. Ministry of Agriculture, |
Fisheries and Food Technical Bulletin 9. 100 pp.

VOLUME 80, NUMBER 3 329

Hughes, R. D., and C. G. Jackson. 1958. A review of the Anoetidae (Acari). Virginia
J. Sci. 9:5-198.

Scheucher, R. 1957. Systematik und Okologie der deutschen Anoetinen. In Beitrage
zur Systematik und Okologie Mitteleuropaischer Acarina. Band I, Teil 1 Aka-
demische Verlagsgesellschaft, Geest & Portig. K.-G, Leipzig. pp. 283-384.

Vegetable Laboratory, PGGI, BARC-West, USDA, Beltsville, Maryland
20705 (A. Hill, former research assistant and graduate student from Botany
Department, Catholic University, Nijmegen, Netherlands).

PROC. ENTOMOL. SOC. WASH.

80(3), 1978, pp. 330-334

TWO NEW SPECIES OF TARSONEMUS (ACARI: TARSONEMIDAE)
ASSOCIATED WITH COMMERCIAL MUSHROOM PRODUCTION

Aagje Hill and Kenneth L. Deahl

Abstract—Females of two new species of Tarsonemidae, Tarsonemus
mercedesae and T. lukoschusi, associated with commercial mushroom pro-
duction in Pennsylvania are described and figured.

A survey of mites associated with commercial mushroom production
was made near Kennett Square, Pennsylvania. Two new species in the
Tarsonemidae were found. The specimens described in this paper were col-
lected from first-stage mushroom compost and horse manure outside mush-
room houses. The terminology of Lindquist (1969) is used for the idiosomal
and gnathosomal structures. The terminology of Suski (1966) is used for the
chaetotaxy and solenidiotaxy of the legs. This work was done in collabora-
tion with the Systematic Entomology Laboratory, IIBIII, Agricultural
Research Service, U. S. Department of Agriculture, Beltsville, Maryland
20705.

Tarsonemus mercedesae Hill and Deahl, new species
Figs. 1-4

Tarsonemus mercedesae can be distinguished from other species in the
genus by the presence of a dark spot on the membrane between genu and
tibiotarsus III. Only the female is known.

Gnathosoma (Fig. 2)—Pharynx short and slender, with a pair of dis-
tinct glandular structures posteriorly. Ventral and dorsal marginal setae
subequal in length.

Dorsum (Fig. 1).—Body oval, broadest at metapodosomal area. Propo-
dosomal shield subtriangular and partially covering gnathosoma. Shield
somewhat less than 1.5x as broad posteriorly as it is long medially. Sensilla
spherical, finely spiculate and mostly covered by the propodosoma. Vertical
setae (V) longer than distance between bases; scapular setae (Sc) almost 2
as long_as V. Marginal setae on tergum I subequal in length to V and
longer than other hysterosomal setae; median setae on terga IL and III
equal in length, ’% as long as marginal setae on tergum I and stronger
than other dorsal setae; marginal setae on terga III and IV subequal in
length and shorter and more slender than other dorsal setae. Pores on
terga I, II and IV.

Venter (Fig. 2)—Anteromedian apodeme interrupted between coxae
I and II and not connected with apodemes II; apodemes II slightly curved;
posterior portion of anteromedian apodeme trifurcate anteriorly, with

VOLUME 80, NUMBER 3 331

Figs. 1-2. Tarsonemus mercedesae, female. 1. Dorsal view; 2. Ventral view.

sclerotized area posteriorly. Coxal setae I less than ‘2 as long as II, which
are relatively long. Transverse apodeme complete. Apodemes III extend
medially to posterior of coxal setae III and laterally to anterior extremities
of coxae III; medial parts of apodemes III curved posteriorly; apodemes
IV slender and connected with posteromedian apodeme, with median
nodule; posteromedian apodeme slender, complete, anterior bifurcate and
not strongly sclerotized. Posteromedial lobe between coxae IV_ broader
than long and distally rounded. Genital-anal plate distinct and with 1
pair of short, slender setae.
Chaetotaxy and solenidiotaxy of legs—Leg I (Fig. 3): Femur, genu, tibio-
- tarsus: 4-4-8 + 4 solenidia + 4 eupathidia. Leg II (Fig. 4): Femur, genu,
tibia, tarsus: 3-3-4-5 + 1 solenidion. Leg III (Fig. 2): Femorogenu, tibia,
tarsus: 2-3-4. Leg IV (Fig. 2): Femorogenu 2, tibiotarsus 2.
Measurements of type-material—Holotype: Idiosoma plus gnathosoma
179 mw long; idiosoma 92 ww wide. Idiosoma of paratypes (3) averages 169
(163-173) w long and 93 (85-101) p wide.
Deposition of type-material—Holotype: Female, USNM, No. 3761 from

332 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

6

Figs. 3-4. Tarsonemus mercedesae, female. 3. Leg I; 4. Leg II. Figs. 5-6. Tarso-
nemus lukoschusi, female. 5. Leg I; 6. Leg II.

first-stage mushroom compost, P. Yeatman, Avondale, Pennsylvania, 13
October 1976, by A. Hill and K. L. Deahl. Paratypes: Three from horse
manure, Keystone Mushroom Company, Coatesville, Pennsylvania 31
August 1976, by A. Hill, in U.S. National Museum of Natural History.

Etymology.—This species is named for Dr. Mercedes Delfinado, New
York State Museum, Albany, New York.

Tarsonemus lukoschusi Hill and Deahl, new species
Figs. 5-8

Tarsonemus lukoschusi can be distinguished from other species in the
genus by the presence of broadly lanceolate sensilla, previously found
only in some species of Steneotarsonemus, such as S. phyllophorus (Ewing)
and S. laticeps (Halbert) (R. Smiley, personal communication). Only the
female of is known.

Dorsum (Fig. 7).—Body oval. Propodosomal shield sub-triangular, not
covering gnathosoma. Vertical setae (V) shorter than distance between
bases. Scapular setae (Sc) fine and % longer than V. Sensilla broadly
lanceolate with conspicuous spicules and partially covered by propodosomal
shield. Dorsal hysterosomal setae simple. Setae on tergum I short and sub-
equal in length; marginal setae on terga III and IV subequal in length.
Three pairs of glandular openings on dorsum.

Venter (Fig. 8)—Apodemes I fused, Y-like, and connected with antero-
median apodeme; apodemes II not connected with anteromedian apodeme

VOLUME 80, NUMBER 3 333

ai 8

Figs. 7-8. Tarsonemus lukoschusi, female. 7. Dorsal view; 8. Ventral view.

and each with a medial and a distal nodule. Anteromedian apodeme weak-
ened between apodemes I and II; posteriorly with diffuse sclerotization.
Transverse apodeme is complete and irregular (as figured). Apodemes HI
extend medially to posterior of coxal setae III and laterally beyond coxal
III; posteromedian apodeme bifurcate anteriorly; apodemes IV_ finer
than III, with median nodule; coxal setae I, II, and III finer and shorter
than IV. Posteromedial lobe between coxae IV about as broad as long.
Genital-anal plate distinct and with 1| pair of short slender setae.

Chaetotaxy and solenidiotaxy of legs.—Leg I (Fig. 5): Femur, genu, tibio-
tarsus: 4-4-1] + 3 solenidia + 3 eupathidia. Leg II (Fig. 6): Femur, genu,
tibia, tarsus: 3-3-4-6 + 1 solenidion. Leg III (Fig. 8): Femorogenu, tibia,
tarsus; 1-3-4. Leg IV (Fig. 8): Femorogenu 2, tibiotarsus 2. Femur II with
large flange. On tibia II of holotype, protuberance appears spine-like; it
appears as a protrusion in paratypes.

Measurements of type-material—Holotype: Idiosoma plus gnathosoma
182 uw long and 101 pw wide. Idiosoma of paratypes (6) averages 163 (142-
196) uw long and 92 (81-106) x wide.

Deposition of type-material—Holotype: Female, USNM No. 3760 from

Best stage mushroom compost, P. Yeatman, Avondale, Pennsylvania, 13

334 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

October 1976, by A. Hill and K. L. Deahl. Paratypes: Two are in Acarol-
ogy Laboratory, Ohio State University, Columbus, Ohio; 2 in Biosystem-
atics Research Institute, Ottawa, Ontario, Canada; and 4 in U.S. Na-
tional Museum of Natural History, Washington, D.C.

Etymology.—This species is named for Dr. F. S. Lukoschus, Catholic
University, Nijmegen, Netherlands.

Acknowledgments

We thank Edward W. Baker and Robert L. Smiley, Systematic Entomol-
ogy Laboratory, IIBIII, Agricultural Research Service, USDA, and Mer-
cedes D. Delfinado, New York State Museum and New York State Educa-
tion Department, Albany, New York, for their help with writing this paper.

Literature Cited

Lindquist, E. E. 1969. Review of Holarctic tarsonemid mites (Acarina: Prostigmata)
parasitizing eggs of pine bark beetles. Mem. Entomol. Soc. Can. 60:111.

Suski, Z. W. 1966. Nomenclature of leg setation in the mite family Tarsonemidae
(Acarina: Heterostigmata). Bull. Acad. Pol. Sci. Cl. II Ser. Sci. Biol. 14:635-

638.

Vegetable Laboratory, PGGI, BARC-West, USDA, Beltsville, Maryland
20705 (A. Hill, former research assistant and graduate student from Botany
Department, Catholic University, Nijmegen, Netherlands).

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 335-343
A NEW SPECIES OF PSEUDOPYGMEPHORUS (ACARI:
PYGMEPHORIDAE) ASSOCIATED WITH
COMMERCIAL MUSHROOM PRODUCTION

Aagje Hill and Kenneth L. Deahl

Abstract—Both sexes of a new species of Pygmephoridae, Pseudo-
pygmephorus smileyi, from horse manure around commercial mushroom
production houses in Pennsylvania are described and figured.

Mites associated with commercial mushroom production were studied at
the Beltsville Agricultural Research Center in 1976. Most of the mites were
collected from compost and horse manure around and in the mushroom
houses near Kennett Square, Pennsylvania. This work was done in collabora-
tion with the Systematic Entomology Laboratory, IIBIII, Agricultural Re-
search Service, U.S. Department of Agriculture, Beltsville, Maryland 20705.

Several species of mites belonging to the family Pygmephoridae and
Tarsonemidae were found. Among them were males and females of an
undescribed species of Pseudopygmephorus, and these are described here.
As far as we know, this is the third species in this genus for which males are
known. The terminologies of Mahunka (1973) and Norton and Ide (1974)
are used for the idiosomal structures, leg chaetotaxy and _ solenidiotaxy.

Pseudopygmephorus smileyi Hill and Deahl, new species

The female of this species resembles the female of Psewdopygmephorus
sellnicki (Krezal, 1959) in having similarly curved dorsal terga. It can be
separated from the latter species by the absence of strong spinelike setae
on tarsi If and III. Smiley (1978) describes the male of P. sellnicki. The
male of P. smileyi differs from the male of P. sellnicki in having slender
setae rather than strong spines on tarsi II and III.

Female
Figs. 1-6

Gnathosoma (Fig. 1).—Slightly elongate with 2 pairs of simple dorsal
setae, anteromedial pair longest. Palpus with 1 segment, dorsally with 2
pairs of simple setae, ventrally with 1 small solenidion and a larger sucking-
like apparatus.

Dorsum (Fig. 1).—Propodosoma convex anteriorly, with concave margins,
rather small in comparison to body. Exobothridial setae (exo) longer than
distance between bases. Interbothridial setae (in) very small, without
spicules. Rostral setae lacking. Setae c. on tergum I longer than exo setae;

336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

exo

Fig. 1. Pseudopygmephorus smileyi, female with gnathosoma enlarged, dorsal view.

VOLUME 80, NUMBER 3 Son

Fig. 2. Pseudopygmephorus smileyi, female with gnathosoma enlarged, ventral view.

338 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

4 D

Figs. 3-6. Pseudopygmephorus smileyi, legs I-IV of female.

setae c; on tergum I, d; on tergum II, and e; on tergum III about equal
and '% the length of c2; setae f; somewhat longer than % the length of setae
c, and f, longer than c;. All setae spiculate, unless otherwise stated.

Venter (Fig. 2)—Apodemes II longer than I, both connecting with antero-
median apodeme; anteromedian apodeme connecting with transverse apo-
deme; posteromedian apodeme connecting with apodemes III and IV; 5th
apodeme lacking. Propodosomal setae la spiculate, somewhat shorter than
distance between their bases; setae 1b bifurcate and spiculate; setae 2a
spiculate and longer than 1b; setae 2a twice as long as 2b. Hysterosomal
setae 3a and 3c not as long as 3b; setae 3b thicker than 3a and 3c; setae
4a and 4c subequal in length; setae 4b longer than other hysterosomal
setae. Caudal setae 3h longest and spiculate; setae 1h and 2h short and
slender. All hysterosomal setae smooth, unless otherwise stated.

Chaetotaxy and solenidiotaxy of legs—lLeg I (Fig. 3): Femur, genu,
tibiotarsus: 3-4-12 + 4 solenidia + 4 eupathidia. Leg II (Fig. 4): Femur,
genu, tibia, tarsus: 3-3-4 + 1 solenidion, -6 + 1 solenidion. Leg HI (Fig.
5): Femur, genu, tibia, tarsus: 2-2-4 + 1 solenidion, -6. Leg IV (Fig. 6):
Femur, genu, tibia, tarsus: 2-1-4 + 1 solenidion, -6.

VOLUME 80, NUMBER 3 339

Fig. 7. Pseudopygmephorus smileyi, male with gnathosoma enlarged, dorsal view.

340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Measurements of type-material—Holotype: Idiosoma plus gnathosoma

275 mw long and 126 pw wide. Idiosoma of female paratypes (10) averages
272 (225-320) uw long and 136 (167-113) pw wide.
Male
Figs. 7-12

The males vary in body size and in the size of the leg setae. Venter and
dorsum are drawn from different specimens.

Gnathosoma (Fig. 7)—Small, elongate, with 2 pairs of simple dorsal
setae, 1 pair of solenidia, and 1 pair of simple ventral setae; anteriorly
with oblong sucking-like apparatus. Palpi absent, chelicerae not visible.

Dorsum (Fig. 7).—Propodosomal shield rounded anteriorly; wider pos-
teriorly than long; medially with 3 pairs of spiculate setae. Prodorsal
setae pd; and pdy, % the length of pd; setae pd; more than 2 as long
as pd; and pd. combined. Hysterosoma with 3 pairs of setae; c; short; cs
more than 2x as long as c;; d % the length of cz. All hysterosomal setae
spiculate. Opisthosomal setae e; and e2 spiculate; e2 located anteriorly.
Solenidia and microsetae (ms) present on sculptured genital area.

Venter (Fig. 8)—Apodemes I short and connecting with anteromedian
apodeme; apodemes II not connected with anteromedian apodeme; antero-
median apodeme connecting with transverse apodeme; posteromedian apo-
deme connecting with apodemes IV and V; apodemes III not connecting
medially. Propodosomal setae la smooth and longer than weakly spiculate
setae lb. Coxal setae 2a longer and stronger than 2 b; both smooth. Setae
2a and la subequal in length; coxal setae 3a and 3c subequal in thickness
and length; setae 3b longer than 3a and 3b; coxal setae 4a and 4c similar
to 3a and 3c; setae 4b longer than 4a and 4c. Aedeagus (aed) as figured.

Chaetotaxy and solenidiotaxy of legs—Leg I (Fig. 9): Femur (not fig-
ured), genu, tibia, tarsus: 3-4-6 + 2 solenidia, -9 + 2 solenidia + 4
eupathidia. Leg II (Fig. 10): Femur (not figured), genu, tibia, tarsus:
3-3-4 + 1 solenidion, -7 + 1 solenidion. Leg HI (Fig. 11): Femur (not
figured), genu, tibia, tarsus: 1-2-4 + 1 solenidion, -7. Leg IV (Fig. 12):
Femur, genu, tibia, tarsus: 1-1-3 + 2 solenidia, —4.

Measurements of type-material—Average of paratypes (5): length: 222
(184-242) w; width 108 (74-127) p.

Deposition of type-material—Holotype: Female, USNM No. 3759 from
horse manure, Keystone Mushroom Company, Coatesville, Pennsylvania, 31
August 1976, A. Hill and K. L. Deahl. Paratypes: Five ¢ and 182 in U.S.
National Museum of Natural History, Washington, D.C.; 12 at Univer-

Fig. 8. Pseudopygmephorus smileyi, male, ventral view.

1

VOLUME 80, NUMBER 3

342 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 9-12. Pseudopygmephorus smileyi, legs I-IV of male (femora not figured).

sity of Alabama, Tuscaloosa, Alabama; 1? in Rijksmuseum van Natuurlijke
Historie, Leiden, Netherlands; 12 in Acarology Laboratory, Ohio State
University, Columbus, Ohio; 12 at Catholic University, Nijmegen, Nether-
lands; 12 in Hungarian Natural History Museum, Budapest, Hungary; 1?
with U.S. Forest Service, Pineville, Louisiana; 12 in Zoologisches Institut
and Zoologisches Museum, Universitat Hamburg, Hamburg, Germany; para-
types with the same data as holotype.

Etymology.— This species is named for Mr. Robert L. Smiley, Systematic
Entomology Laboratory, IIBIII, USDA, Beltsville, Maryland.

Acknowledgments

We thank Edward W. Baker and Robert L. Smiley, Systematic Entomol-
ogy Laboratory, IIBIII, Sci., USDA, for their help with writing this paper.

Literature Cited

Krezal, H. 1959. Systematik und Okologie der Pyemotiden. Beitrage zur Syste-
matik und Okologie mitteleuropaischer Acarina. Band I: Tyroglyphidae und
Tarsonemini. Teil. 2:385-625.

Mahunka, J. 1973. Neue Tarsonemiden (Acari) aus der Mongolei. Ann. Hist.
-nat. Mus. Nat. Hung. 65:309-315.

Norton, R. A., and G. I. Ide. 1974. Scutacarus baculitarsus agaricus, n. subsp.
(Acarina: Scutacaridae) from commercial mushroom houses, with notes on
phoretic behavior. J. Kans. Entomol. Soc. 47:527-534.

VOLUME 80, NUMBER 3 343

Smiley, R. L. 1978. Taxonomic studies on Pygmephorus species from the western
hemisphere, with a key to females and an overview of the current problems
for classification (Acari: Pyemotidae and Pygmephoridae). Int. J. Acarol. 4(2):
125-160.

Vegetable Laboratory, PGGI, BARC-West, USDA, Beltsville, Maryland
20705 (A. Hill, former research assistant and graduate student from Botany
Department, Catholic University, Nijmegen, Netherlands).

NOTE

A NEW NAME FOR COLPOCEPHALUM ABBOTTI PRICE
(MALLOPHAGA: MENOPONIDAE)

Price (1976. Syst. Entomol. 1:63) described Colpocephalum abbotti, and
based the specific name on that of the ibis type-host, Threskiornis aethiopica
abbotti (Ridgway).

Unfortunately, Kellogg (1899. Occas. Pap. Calif. Acad. Sci. 6:36) had
previously described Colpocephalum abbotti for a louse taken from a gull,
Larus sp. I had overlooked this earlier use of C. abbotti due to the fact
that this name is now associated with the menoponid genus Actornithophilus
Ferris and is further a junior synonym of A. piceus (Denny). A shortcoming
in my card file failed to bring this to my attention. I thank Dr. R. C.
Dalgleish, Rensselaerville, New York, for being more efficient and for
pointing this homonymy out to me.

So, with C. abbotti Price a junior primary homonym of C. abbotti Kellogg,
I hereby provide Colpocephalum tandani nomen novum to replace C.
abbotti Price. This name is in recognition of the numerous excellent con-
tributions of Dr. B. K. Tandan to Mallophaga taxonomy.

Roger D. Price, Department of Entomology, Fisheries and Wildlife, Uni-
versity of Minnesota, St. Paul, Minnesota 55108.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 344-359
TAXONOMIC NOTES ON ZAGRAMMOSOMA,
A KEY TO THE NEARCTIC SPECIES AND
DESCRIPTIONS OF NEW SPECIES FROM CALIFORNIA
(HYMENOPTERA: EULOPHIDAE)

Gordon Gordh

Abstract.—Zagrammosoma intermedium, new species, and Z. melinum,
new species, are described from California; Z. intermedium parasitizes
Lithocolletis nemoris, and Z. melinum parasitizes Bucculatrix sp. The fe-
male of Z. nigrolineatum Crawford is described. Zagrammosoma. inter-
lineatum Girault is synonymized with Z. multilineatum (Ashmead) (NEW
SYNONYMY); Z. sanguineum Girault is synonymized with Z. nigrolineatum
Crawford (NEW SYNONYMY). A key to North American species of Zagram-
mosoma is provided; and distribution, host associations and taxonomic notes
are given for each North American species.

The name Zagrammosoma was proposed by Ashmead (1904) as a replace-
ment name for Hippocephalus Ashmead, 1888, which was preoccupied by
Hippocephalus Swainson, 1839, in fishes. Catalogs of North American
Zagrammosoma have summarized published taxonomic and biological in-
formation about the species (Muesebeck et al., 1951; Peck, 1963; Burks, in
press). Presently eight species are included in Zagrammosoma from North
America. Two new species are described in this paper, and two species
are synonymized.

European and American workers differ in opinion regarding the generic
position of Zagrammosoma and Cirrospilus Westwood. Boucéek and Askew
(1968) regard Zagrammosoma as a subgenus of Cirrospilus. This position
was followed by Kerrich (1969). American workers consistently have
maintained Zagrammosoma and Cirrospilus as generically distinct. Twenty
species of Cirrospilus (sensu American authors) are recognized in North
America. These will be considered in another paper.

The biologies of both genera are poorly studied but host relationships
seem to be slightly different. Representatives of both genera are parasites
of leafmining Lepidoptera and Diptera, but some species of Cirrospilus

also parasitize leafmining Hymenoptera. Cirrospilus frequently acts as a |
hyperparasite of braconids and ichneumonids that attack leafminers, but |
only rarely have species of Zagrammosoma been found acting in a hyper-—

parasitic role. When the biological associations of species in both genera
are studied, more subtle differences may be found.

Zagrammosoma is abundant in the western states of North America and |
is especially well represented in California. Cirrospilus appears more com-

mon in the central and eastern states.

VOLUME 80, NUMBER 3 345

Some morphological characters may be of importance in separating
these genera. There is a difference in the shape of the head: Zagram-
mosoma species have the vertex vaulted between the compound eyes, and
the head is elongate; Cirrospilus species do not have the vertex vaulted be-
tween the compound eyes and the head shape is usually oval. Most
specimens in both genera shrivel after death so this character is not always
visible. Specimens of Cirrospilus have a well-developed median pro-
podeal carina (except one undescribed species). Specimens of Zagram-
mosoma do not have a median propodeal carina, or it is weakly developed.

Generic concepts in the Eulophinae show that differences between genera
are often slight and qualitative. If Zagrammosoma and Cirrospilus are
considered congeneric, then a strong argument could be made for synony-
mizing Microlycus Thomson with Necremnus Thomson and Hemiptarsenus
Westwood with Notanisomorpha Ashmead because the differences between
these genera are qualitative and slight.

Thus it seems that we do not know enough about the biology, distribution
and morphological variation of Zagrammosoma, Cirrospilus and related
genera of Eulophinae. For the present it seems more appropriate to main-
tain them as generically distinct until they are better known.

Genus Zagrammosoma Ashmead

Hippocephalus Ashmead, 1888:App. VIII. Type species: Hippocephalus
multilineatus Ashmead. Monotypic.

Zagrammosoma Ashmead, 1904:354, 393. Replacement name for Hippo-
cephalus Ashmead, not Hippocephalus Swainson.)

Zagrammatosoma Schulz, 1906. Spolia Hym., pg. 142. Unjustified emend.

Atoposoma Masi, 1907. Bol. Lab. Zool. Gen. Agric., Portici. 1:276.
Atoposoma variegatum Masi. Monotypic.

Key to North American Zagrammosoma Based on Females

1. Metasomal terga predominantly dark reddish, at least mesally, and

with ornate color pattern (Figs. 1, 3, 7) 2
— Metasomal terga predominantly pale or without ornate color pattern

(Figs. 4, 6) 6
2. Forewing hyaline; dorsal surface of adstigmal area densely setose

(Fig. 16) nigrolineatum Crawford
— Forewing infuscated; dorsal surface of adstigmal area asetose (Figs.

10, 13, 15) 3

3. Forewing infuscation extending parallel to marginal vein from
_ stigmal vein to basal cell (Figs. 11, 15); area posterior to junction of
submarginal vein and marginal vein asetose; postmarginal vein pale,
but as long as stigmal vein 4

346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

— Forewing infuscation “U” shaped, extending from stigmal vein

to junction of submarginal vein and marginal vein (Fig. 10); area pos-

terior to juction of submarginal vein and marginal vein with coarse
dark setae; postmarginal vein about % as long as stigmal vein 5

4. Dark brown mesosomal stripe broad (Fig. 4), as wide as distance

between parallel longitudinal grooves on scutellum; notaulices

pale; area posterior to junction of marginal vein and stigmal vein
asetose (Fig. 15) centrolineatum Crawford

— Dark brown mesosomal stripe narrow (Fig. 5); notaulices dark

brown; area posterior to junction of marginal vein and _ stigmal
vein setose (Fig. 11) intermedium, new species

5. Metasoma entirely reddish brown; propodeal callus uniformly red-

dish brown; apex of hind femur pale, remainder reddish brown
mirum Girault

— Metasoma dusky reddish brown ventrally with pale spots laterally;

propodeal callus pale yellow, remainder of propodeum reddish

brown; apex and basal % of hind femur pale, remainder dusky
flavolineatum Crawford

6. Forewing with several infuscated spots or if infuscated spots faint,

then junction of submarginal vein and marginal vein, marginal vein,

and stigmal vein dusky or darkened; dorsal surface of adstigmal

area asetose (Figs. 10-15) i
— Forewing hyaline; dorsal surface of adstigmal area densely setose
(Fig. 16) nigrolineatum Crawford

7. Anterior margin of mesoscutum with dark transverse stripe that is
enlarged laterally forming a spot (sometimes concealed beneath
posterior margin of pronotum) (Fig. 1) americanum Girault

— Anterior margin of mesoscutum without transverse stripe 8

8. Scutellum with longitudinal, medial stripe (Fig. 3); hind femur with
dorsal stripe and apical spot; forewing blade with numerous setae
(Fig. 12) multilineatum (Ashmead)

- Scutellum without longitudinal, medial stripe (Fig. 2); hind femur
pale yellow, without stripe or spot; forewing blade with fewer setae
(Fig. 14) melinum, new species

Zagrammosoma americanum Girault
:
Figs. 1, 13

Zagrammosoma americanum Girault, 1916:126-127.

Type-locality—Boulder, Colorado.
Girault described this species from one female. Girault separated Z. —
americanum from Z. multilineatum based on the conspicuous, round, black
dot near the apex of the hind femur. Some specimens of Z. americanum

VOLUME 80, NUMBER 3 347

Figs. 1-3. Dorsal view of Zagrammosoma species. 1. Z. americanum; 2. Z. melinum;
3. Z. multilineatum.

have this dot misshapen, and in others it is almost a stripe. The dorsal
i i
longitudinal stripe on the hind femur is also variable.
A more reliable character to distinguish these species is a transverse,
g is
dark stripe along the exposed margin of the mesoscutum (which also
i gS i gS
projects beneath the posterior margin of the pronotum). Laterally this
stripe becomes two enlarged spots. This character is present only in
Z. americanum.
The propodeal pigmentation of Z. americanum is variable and resembles

348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Z. multilineatum (the anterior and posterior margins are dark). In most
specimens of both species the pigmentation on the meson extends postero-
laterally and forms a “W.” In some specimens it forms a median dot.

This species is abundant in California and is often found in association
with pine. It has been reared from Coleotechnites milleri (Busck), Coleoph-
ora laricella (Hubner) and Asphondylia galls.

Zagrammosoma centrolineatum Crawford
Figs. 4, 15

Zagrammosoma centrolineatum Crawford, 1913:256.

Type-locality.— California.

Crawford described this species based on two females taken in Los
Angeles County and one female taken in Sonoma, California. The specimen
from Sonoma is conspecific with the specimens from Los Angeles. The
paratype from Los Angeles is missing the metasoma.

Crawford’s original description is accurate; supplementary illustrations
of the habitus and forewing (Figs. 4, 15) will make recognition of this dis-
tinctive species relatively easy. Based on forewing characters this species
is closely related to Z. mirum, Z. flavolineatum and Z. intermedium. It can
be distinguished from these species based on characters given in the
key.

Little information has been gathered on Z. centrolineatum. It has been
recovered from Oregon, California and Utah. Hosts include Caloptilia
alnivorella (Chambers), Lithocolletes mediodorsella Braun, Lithocolletes
sp. on Populus spp. and Quercus dumosa, “leaf blotch mine” on Q. wislizenii
and Tischeria sp. on Q. dumosa englemanii.

Zagrammosoma flavolineatum Crawford
Fig. 10

Zagrammosoma flavolineatum Crawford, 1913:255-256.

Type-locality —Boulder Co., Colorado.

Crawford described this species from one female specimen, and the
original description is accurate. For a discussion of this species see com-
ments under Z. mirum.

Zagrammosoma intermedium Gordh, new species
Figs. 5, 11

Type-locality—Palo Alto, California.

Female.—1.8 mm long. Body coloration as illustrated (Fig. 5); anterior
aspect of head pale yellow except dark stripe extending from dorsomedial
margin of compound eye to anterior ocellus; gonostylus dark brown; an-

VOLUME 80, NUMBER 3 349

Figs. 4-6. Dorsal view of Zagrammosoma species. 4. Z. centrolineatum; 5. Z. inter-
medium; 6. Z. mirum.

tennal pedicel with dusky spot on dorsal surface; funicular segments red-
dish brown, club dark brown. Forewing color pattern and setation as il-
lustrated (Fig. 11). Legs pale yellow.

Head similar in shape and proportions to Z. centrolineatum.

Mesosoma with uniform alutaceous sculpture except on lateron of
-metanotum, pattern somewhat larger and not as deeply incised as in Z.
centrolineatum. Pronotum with numerous scattered, short, dark setae
and a row of larger setae along posterior margin; mesoscutum with 12 dark

|
|

350 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

setae which become progressively larger posteriorly; scutellum with 2 pairs
of large, dark setae with the posterior pair larger than the anterior pair.
Propodeum with weak but complete median carina; callus with long,
pale setae; spiracle round, about 1 diameter from anterior margin of
propodeum.

Metasomal tergum with alutaceous sculpture, but pattern not as strongly
incised as on mesosoma. Terga 1-3 with short, pale setae along postero-
lateral margin; terga 4-6 more densely setose and pattern complete
transversely; tergum 7 uniformly setose along apical 4%. Gonostylus densely
setose.

Male.—Unknown.

Described from one female taken at Stanford University, Palo Alto, Cali-
fornia during 1947 from parasitized Cameraria nemoris (Walsingham) by
J. W. Tilden. Holotype deposited in the U.S. National Museum of Natural
History (USNM Type 75663).

This species is similar to Z. centrolineatum but can be distinguished from
that species on the forewing characters given in the key and by the
width of the mesosomal stripe.

After the description of this species the head was inadvertently lost. I
have decided to describe the species on the basis of an imperfect speci-
men because the host is known and the species has a distinctive habitus.

This species is the same as that referred to by Tilden (1949) in his short
note on leafminer parasites.

Etymology.—The specific name is a Latin adjective (intermedius) and
means intermediate.

Zagrammosoma melinum Gordh, new species
Figsae2 nos.

Type-locality.—Coalinga, California.

Female.—2.3 mm long. Body pale yellow except the following (Fig. 2);
dark stripes under compound eye extending from medial margin to posterior
ocellus; pronotum with 2 parallel, longitudinal, narrow stripes, 1 lat-
eral and 1 dorsomesal; mesoscutum with lateral, longitudinal mesal stripe
and a short, dark line on notaulices; scutellum with 2 short stripes halfway
between median carina and spiracles; posteriomesal margin dark; basal
gastral tergum with lateral spot; pygostylus brown; distal % of gonostylus

dark brown. Antennal scape and pedicel with dark dorsal stripes (Fig. 9).
Forewing hyaline except faint dusky cloud beneath stigma and junction of |

submarginal vein and marginal vein (Fig. 14). Legs pale yellow; pretarsi
brown apically.

Head in frontal aspect 1.2x wider than high; compound eyes protuberant, |
asetose, 1.4 taller than length of malar space. Head surface alutaceous; |
vertex, medial margin of compound eye, face, and clypeal margin with |

:

VOLUME 80, NUMBER 3 351

Figs. 7-8. Dorsal view of Zagrammosoma nigrolineatum.

sparse vestiture of pale, fine setae; clypeal margin straight. Torulus sit-
uated halfway between imaginary transverse line connecting ventral mar-
gins of compound eyes and eye midline.

Antenna (Fig. 9) 9-segmented (1, 1, 2, 2, 3); scape setose, alutaceous, 5.0
longer than wide; pedicel 1.6 longer than wide, setae more robust than
setae on scape; alutaceous; anelli transverse, setose, smooth; funiculars
subequal in length, setose and bearing rhinaria; club compact, 1.7x longer
than wide, wider than funiculars, setose, each subsegment with rhinaria.
Mandible 6-toothed. Maxillary palpus 1-segmented; labial palpus 1-seg-
mented.

Mesosoma except metanotum alutaceous; meson of metanotum smooth,
lateron striate. Posterior margin of pronotum with a line of fine, pale
setae; scapula laterally with pale, fine setae; mesoscutum with 5 pairs of

352 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

fine setae; scutellum with 2 pairs of fine, pale, long setae; metanotum
asetose; propodeal callus with long, fine, pale setae, medial carina weakly
developed but complete.

Metasoma oblong-ovate from above, 1.16 longer than mesosoma, aluta-
ceous; posterior margin of terga with lateral line of setae incomplete on terga
1-3 but progressively increasing in number mesally such that line is
complete on segments 4-7; apical 12 of tergum 1 uniformly setose; sterna
mesally setose; ovipositor extending from base to apex of metasoma,
1.96 longer than hind tibia, 2.1< longer than middle tibia, 5.19x longer
than gonostylus. Pygostylus well developed, apparently with 4 long and
1 short setae.

Forewing moderately setose distal to junction of submarginal vein and
marginal vein; admarginal area asetose on dorsal surface of wing; costal
cell with a line of setae; marginal fringe short.

Male.—Unknown.

Described from eight females taken at Coalinga, California during
August 1939 from parasitized Bucculatrix sp. on cottonwood by F. P.
Roullard. Holotype and female paratypes deposited in the U.S. National
Museum of Natural History (USNM Type 75665).

This species is similar to Z. multilineatum but can be distinguished from
that species based on the following characters: Z. melinum lacks a medial
longitudinal stripe on the scutellum and apical hind tibial spot, and the in-
tensiveness of setation on the forewing is considerably less.

Variation.—Although the type-series is not extensive, there does appear
to be some color variation. The propodeum may be pigmented, the meso-
scutal stripe may be complete and the mesal portion of the metasomal
tergum may be pigmented.

Etymology——tThe specific name is from Latin (melinus) and means
yellow-colored.

Zagrammosoma mirum Girault
Figs. 6, 10

Zagrammosoma mirum Girault, 1916:119-120.

Type-locality —Claremont, California.

This species was described from a single specimen. It is similar to Z.
flavolineatum, and topotypical material of both species should be collected
to determine whether they are conspecific. Only the type-specimen of Z.
flavolineatum exists, but several specimens of Z. mirum have been accumu-
lated in the U.S. National Museum, Natural History, collection. The char-
acters that distinguish these species are the pale propodeal callus and pale
basal ‘2 of the hind femur on Z. flavolineatum, and the uniformly dark

353

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VOLUME 80, NUMBER 3

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Antenna of Zagrammosoma melinum.

Fig. 9.
mosomad species.

354. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

coloration of the propodeum and dark hind femur with pale apex on Z.
mirum.

Graf (1917) discussed a species he called Z. flavolineatum which attacked
potato tuber moth, Phthorimaea operculella (Zeller), in Southern Cali-
fornia. The illustrations of Graf's parasite lead me to conclude that he
was dealing with Z. mirum because the coloration of the hind femur and
propodeal callus is identical with that species.

Hosts of Z. mirum include Lithocolletis sp., Tischeria sp. and Liriomyza
pictella (Thomson).

Zagrammosoma multilineatum (Ashmead)
Figs: 3) 512

Hippocephalus multilineatum Ashmead, 1888: VII.

Type-locality.—Riley Co., Kansas.
Zagrammosoma multilineata var. punicea Girault, 1911:123.
Zagrammosoma interlineatum Girault, 1916:125-126. NEW SYNONYMY.

Type-locality.—District of Columbia.

In Girault’s original description of Z. interlineatum he compared the
species to Z. multilineatum. I find no structural characters to differentiate
the two, and the color pattern of interlineatum is within the range of
variation exhibited by Z. multilineatum. Therefore, the synonymy is pro-
posed.

Ashmead (1888) described multilineatum from two specimens and wrote
that the species was characterized by a longitudinal stripe extending from
the base of the torulus to the clypeal margin and two lines extended be-
neath the compound eye. The type-specimens also have a longitudinal
stripe along the central third of the hind femur and a dark apical spot on
the outer surface only. Since Ashmead’s description this species has been
recovered from Florida west to Idaho, Puerto Rico, and South America.
Kerrich (1969) has provided supplementary descriptive notes on this species
based on two specimens.

Not all of the material identified as Z. multilineatum in the U.S. Na-
tional Museum, Natural History, collection conforms to the type-specimens.
I have noted four phenotypes: (1) presence of a “nose” stripe, two stripes
beneath the compound eye, and a stripe on the hind femur; (2) absence of —
a “nose” stripe, spotless and stripeless hind femur and convergent stripes |
beneath the compound eye; (3) absence of the “nose” stripe, stripes be- |
neath the compound eye and spotless and stripeless hind femur; and (4)
absence of the “nose” stripe, presence of two convergent stripes beneath |
the compound eye and a stripe and spot on the hind femur. There is _
geographical overlap among the phenotypes. Careful biological study |
of this species is necessary to determine whether sibling species are in-_
volved.

355

VOLUME 80, NUMBER 3

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356 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Girault (1911) named a “variety” of Z. multilineatum called puniceum
from four specimens parasitizing Tischeria malifoliella Clemens taken
by Quaintance at Washington, D.C. Girault reported that the distinguish-
ing character was coloration, Z. multilineatum “variety” puniceum was
pink and Z. m. multilineatum was yellow. When comparing Girault’s type-
material with other specimens of Z. multilineatum, I noted that the stripes
beneath the compound eye converge, there is no “nose” stripe and the hind
femur has a dorsal stripe and an apical stripe. Although Girault emphasized
the difference in color, it is probably a cyanide induced reaction and I do
not consider puniceum a subspecies.

The problem is complicated further because the specimens are labeled
“Zagrammosoma multilineatum var amoverta MS, Girault,” but apparently
not in Girault’s handwriting. The specimens are card-point mounted, and
the type number is 9641. The U.S. National Museum, Natural History,
type-catalog holds this name, but apparently Girault changed the name after
it was entered in the catalog and before the manuscript was published.

This species is the most abundantly collected Zagrammosoma in North
America and has been recovered from many hosts including Diptera and
Lepidoptera. The most common hosts include: Bucculatrix canadensisella
Chambers, Coleotechnites milleri (Busck), Lithocolletis ostensackenella
(Fitch), Antispila nyssaefoliella Clemens, Phyllonorycter craetaegella
(Clemens), Agromyza pusilla (auct., nec Meigen) and Liriomyza_ sativae
Blanchard.

Zagrammosoma nigrolineatum Crawford
Figs. 7, 16

Zagrammosoma nigrolineatum Crawford, 1913:257.
Type-locality —Compton, California.

Zagrammosoma sanguineum Girault, 1916:133. NEW SYNONYMY.
Type-locality.—Colorado.

Crawford’s original description indicates that this species was based on
two females, but examination of the type-series shows that they are both
males. Several females have been acquired over the past 60 years; con-
sequently, the following description can be provided.

Female 1.8 mm long. Head yellow except for 2 dark brown longitudinal
stripes on frons, 3 spots surrounding ocelli, 2 occipital stripes, 2 large spots
originating near oral fossa extending dorsally on either side of hypostomal
bridge toward occipital foramen then diverging toward posterolateral mar-
gin of compound eyes; prementum dark; dark spot between toruli. Thorax |
yellow except metallic green lateral longitudinal and medial longitudinal
stripes on pronotum, most of mesoscutum, scapulae along notauli; mesal_
% of scutellum and meson of metanotum. Entire propodeum except supra-_ |
coxal flange metallic green. Posterior margin of proepisternum, ventral

VOLUME 80, NUMBER 3 357

i}
yl
1!

!

SSS =

Fig. 16. Forewing of Zagrammosoma _ nigrolineatum.

0.33 of prepectus, ventral 0.50 of mesepisternum, mesepimeron dark.
Metasomal terga predominantly dark reddish brown; sterna mesally dusky,
laterally yellow; gonostyli dusky. Coxae yellow (except base of hind
coxa), trochanters yellow; femora yellow except dorsolongitudinal stripe
on front femur, base of hind femur; tibiae and tarsi dusky. Antennal
scape with dorsal stripe; pedicel, anelli dark brown, funiculars and club
slightly lighter.

Head in frontal aspect 1.33 wider than tall; compound eyes not strongly
protuberant. Vertex and frons alutaceous; face, malar space smooth; com-
pound eye 1.13x taller than malar space length. Head surface setose,
compound eye setose, setae moderately long, pale; clypeal margin straight;
toruli beneath imaginary transverse line extending between compound
eyes, separated by 1.5 torular diameters. Antenna 9-segmented (1, 1, 2,
2, 3); scape reaching vertex, with reticulate striae, 5.70 longer than wide,
with a few pale, thin setae; pedicel 2.0x longer than wide, moderately
setose, usually about 1.5x longer than Ist funicular segment; anelli trans-
verse with small, pale setae; funicular segments subequal in size, bearing
setae and rhinaria; club 2.23x longer than wide with setae, rhinaria. Man-
dible 5-toothed. Maxillary palpus 2-segmented; labial palpus 1-segmented.

Mesosoma with alutaceous sculpture; pronotum with a row of moderately
large setae on posterior margin; mesoscutum with 2 or 3 pairs and scutel-
lum with 2 pairs of large setae; scapula with 5 setae; axilla and metanotum
asetose; metanotal apex mesally pointed, displacing anterior portion of pro-
podeum; propodeal median carina not reaching posterior margin; callus
with moderate vestiture of pale, long, thin setae.

Metasoma smooth, 1.16 as long mesosoma; terga 1-3 setose laterally,
4-5 with transverse line of setae, 6-7 with uniform vestiture of pale, thin
setae; sterna sparsely setose mesally. Ovipositor 1.11 as long as hind

358 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tibia, 1.19 as long as middle tibia, 4.0x as long as gonostylus; ovipositor
extending from basal % of metasoma to apex. Pygostylus as long as wide,
with 2 long and 3 short setae.

Forewing submarginal vein 1.17 longer than marginal vein; stigmal
vein 1.3x longer than postmarginal vein; costal cell with a line of setae and
a few setae along anterior apical margin; marginal fringe 0.07X maximal
wing width.

Described from numerous specimens collected in Canada and the western
United States. Hosts include Coleotechnites milleri (Busck) on Pinus con-
torta and Jacaranda acutifolia infested with Phytoliriomyza jacarandae
Steyskal and Spencer M.S., Argyresthia pilatella Braun, and Ocnerostoma
strobivorum Freeman.

Variation —The color pattern of this species varies considerably. A long
series of specimens taken at Targhee, Idaho by J. H. McLeod from cotton
leafminer shows that the body coloration varies from pale yellow to a
metallic blue-green mesosoma and reddish-brown metasoma. Ratio of
pedicel length to first funicular segment length sometimes has been used
as a taxonomic character. However, in this species the ratio is variable;
the pedicel may be longer or shorter than the first funicular segment. Oc-
casionally there are supernumerary setae on the scutellum. The ocelli al-
ways have dark spots surrounding them. Pale specimens have small spots;
extensively metallic specimens have the entire interocellar area pigmented.
This species has been recovered from Apanteles spp. attacking leafminers
in Canada.

Girault (1916) described Z. sanguineum based on one female taken in
Colorado. The type-specimen has the head crushed on a slide and mounted
in Canada balsam and the body point-mounted. The body is red, but again
I suspect that this is a cyanide-induced reaction. Although the antennae are
broken, the pedicel is definitely longer than the first funicular segment.
Morphologically this specimen falls within the range of variation exhibited
by Z. nigrolineatum and thus the synonymy is proposed.

Acknowledgments

I wish to thank Ms. Deborah Green (Univ. Calif., Berkeley), Mr. J. Hall
(Univ. - Calif., Riverside), Drs. R. Luck (Univ. Calif., Riverside) and
C. M. Yoshimoto (Biosystematics Research Institute, Ottawa, Can.) for
providing material at their disposal. The illustrations were prepared by
Kate Conway, Biruta Ackerberg and Linda Heath.

Literature Cited

Ashmead, W. H. 1888. Descriptions of some unknown parasitic Hymenoptera in
the collection of the Kansas State Agricultural College received from Prof.
E. A. Popenoe. Kans. Agric. Exp. Stn. Bull. 3:App., I—-VIII.

VOLUME 80, NUMBER 3 359

1904. Classification of the chalcid flies. Mem. Carnegie Mus. 1(4):225-551.

Boutek, Z., and R. R. Askew. 1968. Index of Entomophagus Insects. Hym. Pale-
arctic Eulophidae (excl. Tetrastichinae). Le Francois, Paris. 254 pp.

Burks, B. D. (In press). Family Eulophidae. In Krombein, K. V., P. D. Hurd, Jr., D. R.
Smith, and B. D. Burks, Eds. Catalog of Hymenoptera in America north of
Mexico. Smithsonian Institution Press. Washington, D.C.

Crawford, J. C. 1913. Descriptions of new Hymenoptera, No. 6. Proc. U.S. Nat.
Mus. 45:241-260.

Girault, A. A. 1911. Synonymic and descriptive notes on the Hymenoptera Chal-
cidoidea with descriptions of several new genera and species. Arch. Nat. Jahrg.
77(1): 119-140.

—. 1916. New North American Hymenoptera of the family. Eulophidae. Proc.
U.S. Nat. Mus. 51(2148):125-133.

Graf, J. E. 1917. The potato tuber moth. U.S. Dep. Agric. Bull. 427, 56 pp.

Kerrich, J. G. 1969. Systematic studies of eulophid parasites (Hym., Chalcidoidea),
mostly of coffee leafminers in Africa. Bull. Entomol. Res. 59(2):195-228.
Muesebeck, C. W. F., K. V. Krombein, and H. K. Townes. 1951. Hymenoptera of
America North of Mexico. Synoptic Catalog. USDA Agric. Monogr. No. 2,

1420 pp.

Peck, O. 1963. A catalog of the Nearctic Chalcidoidea (Insecta: Hymenoptera).
Can. Entomol. Supl. 30, 1092 pp.

Tilden, J. W. 1949. Notes on parsites of certain microlepidoptera. Pan-Pac. Entomol.

25(1):27-28.

Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. Educ. Admin.,
USDA, c/o U.S. National Museum, Washington, D.C. 20560 (now at:
Division of Biological Control, Department of Entomology, University of
California, Riverside, California 92521).

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 360-372
NEOTYPE DESIGNATION OF CULEX QUINQUEFASCIATUS
SAY (DIPTERA: CULICIDAE)

Sunthorn Sirivanakarn and Graham B. White

Abstract—To promote nomenclatural stability concerning the inter-
pretation and use of the name Culex quinquefasciatus Say for the southern
(tropical) house mosquito, a neotype male is here designated. It was
chosen from a series of specimens reared from an egg raft, collected in
1969 at New Orleans, Louisiana, USA. Description and illustrations of the
quinquefasciatus male, female and associated pupa and larva are pro-
vided. Under the Law of Priority, the name quinquefasciatus Say 1823
takes precedence over all accepted junior synonyms, notably fatigans
Wiedemann 1828.

In the intervening two decades since the name Culex quinquefasciatus
Say (1823:10) was discussed by Stone (1956 [1957]:342-343) and adopted
by Stone et al. (1959) as the valid name for the southern (tropical) house
mosquito, some significant contributions have been made towards an
objective resolution of the nomenclatural arguments concerning this well
known taxon. Fundamental to the solution of this problem is the identity
and nomenclatural status of the only surviving mosquito specimens that
were collected by Thomas Say. These were sent by Say to Wiedemann
between 1523 and 1828 and later were deposited in the Naturhistorisches
Museum in Vienna. Among this material are specimens which Wiedemann
(1828:12-13) described as Anopheles ferruginosus. In a footnote he stated
that the description was based on “original” specimens of Culex quinque-
fasciatus (i.e., material from Say but not necessarily type-material, see
Belkin, 1977:44).

In 1905, L. O. Howard (in Coquillett, 1906:7) examined four specimens
labelled as ferruginosus and reported that they were Culex, not Anopheles.
This discrepancy has led subsequent culicidologists to suspect or speculate
that some of the specimens in the type-series of ferruginosus may repre-
sent the original material from which Say (1823:10-11) drew his descrip-
tion of quinquefasciatus. If this were the case it would be possible to select
and designate one of these specimens as a lectotype of quinquefasciatus,
thus removing all doubts about the identity of the species to which this
name has been applied.

In an attempt to clarify and to resolve the above and other intimately re-

lated problems, Belkin (1977:45-52) critically reexamined all existing |

Say material of mosquito species described by Wiedemann (ferruginosus,
crucians and pungens) at the Naturhistorisches Museum in Vienna in the
summer of 1966. Of the 4 so-called ferruginosus specimens mentioned by

VOLUME 80, NUMBER 3 361

Coquillett (1906), Belkin found only 3 with determination labels from
Wiedemann. These 3 specimens represent an Anopheles species conform-
ing to Wiedemann’s description of ferruginosus but not to Say’s descrip-
tion of quinquefasciatus. The fourth specimen lacks a definite determina-
tion label, and was identified by Belkin as Culex. As discussed by Belkin,
Howard probably saw this specimen; but, as it bears no Wiedemann labels,
it cannot be taken as type-material of any species described by Wiedemann,
particularly Culex pungens to which it apparently belongs. This informa-
tion rules out any possibility of designating a lectotype of quinquefasciatus
from the existing ferruginosus syntype series.

The type-specimens of Culex pungens, and Wiedemann’s description
of this species, agree perfectly with Say’s description of quinquefasciatus
and it appears possible that pungens might have been described from
original specimens of quinquefasciatus. However, as the pungens type-
specimens cannot be proven to have come from Say, their standing in rela-
tion to quinquefasciatus is equivocal. Based on these lines of argument,
derived from his examination of the ferruginosus and pungens material,
Belkin (1977) concluded that the ferruginosus specimens are unacceptable
as the original material (type) of quinquefasciatus and that Wiedemann’s
description of pungens was probably based on the specimens of quin-
quefasciatus. Other information from the description and labels of Wiede-
mann’s species indicates New Orleans as the origin of the Say material.
Although the exact locality of quinquefasciatus cannot be determined
from Say’s notes, it is safe to assume that some of the original material may
have come from somewhere in the vicinity of New Orleans to where the
type-locality was restricted by Belkin, Schick and Heinemann (1966:4-5).

From a careful consideration of the involved problems fully discussed
by Belkin (1977) we are satisfied that none of the material from Say, as
used for the description of ferruginosus Wiedemann, is eligible for designa-
tion as lectotype of quinquefasciatus. The rest of Say’s original material
is no longer existent in the United States. Harris, who studied the Thomas
Say collection shortly after Say’s death, reported that the Diptera were
entirely destroyed (Weiss and Ziegler, 1931). Thus there seems to be no
possibility that other original type-material of quinquefasciatus will be
found for proper lectotype designation.

We also concur with Belkin (1977) that, since none of the Anopheles fer-
ruginosus specimens can be considered as the original material (type) of
Culex quinquefasciatus, a suitable neotype from New Orleans should
be designated in order to clarify and to stabilize the nomenclature.
In accord with the interpretation by Stone (1956 [1957]), as adopted in both
editions of the World Catalog of mosquitoes (Stone et al., 1959:254; Knight

and Stone 1977:217) and as analyzed further by Belkin (1968b:47; 1977:

45-52), we recognize that the original description of quinquefasciatus by
Say (1823:10-11) applies to the Culex species commonly known as the

362 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tropical or southern house mosquito. Furthermore, it seems highly un-
likely that Say, who described 6 North American Culicidae in the years
1823-1827, would have failed to name this familiar pest, and none of his
other descriptions could readily be confused with it. In accordance with the
Law of Priority (Article 23, International Code of Zoological Nomenclature,
1964:23; 1974:79-81), therefore, the name quinquefasciatus Say 1823 takes
precedence over all accepted junior synonyms, notably fatigans Wiedemann
1828 (see Knight and Stone, 1977:217-219 for complete synonymy).

In support of previous and present interpretations of the name quinque-
fasciatus, the original description given by Thomas Say is reproduced in
Fig, 1.

Neotype Designation and Depository

Neotype é (No. 9) with associated pupal and larval skins and slide of
genitalia (No. 691013-1), reared from an egg raft collected on 18 Septem-
ber 1969 in New Orleans, Louisiana, U.S.A., by personnel of the New Orleans
Parish Mosquito Control (George T. Carmichael, director); to be deposited
in the U.S. National Museum, Washington, D.C. (USNM).

Other specimens reared from the same egg raft as the neotype have been
deposited in the following institutions:

(1) British Museum (Natural History), London, Great Britain: 14 (No.
11) with associated pupal and larval skins and genitalia slide (No. 691013-2),
12 (No. 2) with associated pupal and larval skins and 2 whole larvae.

(2) Services Scientifiques Centraux, O.R.S.T.O.M., Bondy, France: 1é
(No. 17) with associated pupal and larval skins and genitalia slide (No.
691013-3), 12 (No. 5) with associated pupal and larval skins and 2 whole
larvae.

(3) Australian National Insect Collection, C.S.I.R.O., Canberra, Aus-
tralia: 1¢, with slide of genitalia (No. 760318-1), 12 (No. 8) with as-
sociated pupal and larval skins and 2 whole larvae.

(4) Department of Entomology, National Science Museum, Tokyo, Japan:
146 with slide of genitalia (No. 760318-4), 12 with associated pupal and
larval skins and 2 whole larvae.

The rest of the material in this series, which consists of 84 (3 with genitalia
slides No. 760329-2, 3, 5), 42 with associated pupal and larval skins
(No. 1, 15, 16, 18), 122 (2 with slides of cibarial armature No. 760329-1, 2)
and several whole larvae are placed in the collection of the USNM. These
specimens are available for deposition in other museums upon request.

Description and Illustrations

The description and illustrations of quinquefasciatus presented here are
composite and comprehensive, based on a detailed study of the neotype and

VOLUME 80, NUMBER 3 363

2. ©. 5-fasciatus. Body cloathed with cinereous
hair; abdomen annulate with blackish.

Inhabits the western states.

fyes deep black ; antennee fuscous, region of the
base paler; proboscis black; thorax with a dilated
dorsal fuscous vitta; pectus cach side varied with
blackish ; halteres entirely whitish ; scutel glabrous:
wings with dusky nervures, immaculate ; feet mode-
rate, fuscous; thighs whitish; abdomen cinereous:
tergum with five black, broad, fascie: tail black
above.

Length about one-fifth of an inch; proboscis one-
tenth of an inch.

This is an exceedingly numerous and troublesome
species. We found them in great numbers on the
Mississippi in May and June. The hairy covering
is very deciduous, and when an individual is caught
by hand, the back of the thorax, in consequence of
being denuded by the touch, exhibits the dorsal
vittz of a blackish colour confluent at the base, with
an oval black spot on each side. The abdominal an-
nuli are sometimes fuscous or even light brown.

Legs much shorter than those of the preceding spe-
cles, but like them in not being annulated.

Fig. 1. Reproduction of the original description of “Culex 5-fasciatus” as published
by Say, 1823:10-11.

all other specimens in this series. Altogether, 43 specimens (13 males, 20
females, 10 fourth instar larvae) and 11 associated pupal and larval skins
have been examined. The descriptive terminology used follows Belkin (1962,
1968a) and Belkin et al. (1970). For a brief diagnosis of the adults and im-

364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

legs

T
1-0
fore —
cibarial armature
mid —
al ac
Wo“) )
4 | VC })
hind — : LS /
<
2 J
9 o
genitalia
Fig. 2.

Culex quinquefasciatus. A, female head and thorax, lateral view; B, male
head, lateral view; C, female cibarial armature; D, female thorax, dorsal view; E, legs

>
anterodorsal views; F, male, female tarsal claws; G, wing, dorsal view; H, female ab-
domen, dorsal view; I, female genitalia.

.
:

VOLUME 80, NUMBER 3 365

matures of quinquefasciatus, consult Belkin (1962, 1968a), Bram (1967) and
Sirivanakarn (1976).

Male (Fig. 2)—Measurements based on neotype. Wing 3.6 mm. Fore-
femur 1.8 mm. Proboscis 2.7 mm. In general as described for female ex-
cept for the following. Head: Palpus exceeding proboscis by full length
of segment 5; segments 2 and 3 entirely dark scaled; segment 3 sometimes
with a few pale scales on lateral surface in middle, apical 0.25-0.40 with a
ventrolateral tuft of 10-12 dark bristles, ventral surface with a row of sev-
eral short, pale hairlike setae extending from base to apex; segments 4 and
5 entirely dark scaled on dorsal surface, lateral, ventral and mesal sur-
faces with numerous bristles; ventral surface of segment 4 with a pale
scaled line from base to about 0.75 of total length; ventral surface of segment
5 with a distinct pale scaled spot at base. Proboscis entirely dark scaled or
sometimes with a poorly defined pale ring at false joint which is located
at about 0.75 of the length from base. Antenna shorter than proboscis,
flagellar whorl long, densely plumose. Legs: Claws of fore- and midlegs
enlarged, external claw larger than internal, both with a distinct subbasal
denticle; claws of hindleg small, equal and simple. Wing: Scales on
branches of veins R, M and Cu less dense than those in the female.
Abdomen: Tergites II-VII with complete, evenly broad basal pale bands,
all of which are connected with basolateral pale spots laterosternad; length
of basal band about '% of segment width.

Male genitalia (Fig. 3A)—Segment IX: Tergal lobe poorly developed,
with 1-2 irregular rows of 10-12 strong setae; sternum broad, finely spic-
ulate, without setae or scales. Sidepiece: Slender, conical, about 0.35 mm in
length; inner tergal surface with 1-2 irregular rows of about 15 subequally
strong setae extending from basal % to slightly beyond level of subapical
lobe; lateral tergal surface with about 20 heavy bristles and several weaker
bristles; apex with a row of 6-7 setae on sternal surface. Subapical lobe:
Broad; specialized setae of proximal and distal divisions clearly divided;
proximal divisions with 3 strong rodlike setae (a—c) of subequal length;
rod a straight with abruptly pointed apex; rods b and c gently curved with
hooked apices; rod c thinner than a and ), its base more or less separated
from the latter distad; distal division with 3 slender bladelike or rodlike
setae in group d-f on mesal surface and 1 broad leaflet (g) and 1 strong
flattened seta (h) on lateral surface. Clasper: simple, typically sickle-
shaped, about 0.75 of length of sidepiece; outer subapical margin without
distinct annulation or crest of spicules; 2 ventral tiny setae present distad of
median curvature on ventral surface, dorsal seta absent; spiniform subapical,
short, flattened and apically blunt. Phallosome: Apical portion of lateral
plate with outer and inner divisions; median portion of outer division with
a prominent apically pointed tergal mesal spine (or tergal arms of several
authors) which is straight so that both spines on each lateral plate are nearly

366 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

*— ventral arm
(inner division)

Fig. 3. Culex quinquefasciatus. A, male genitalia, dorsal view; B, pupa, cephalo-
thorax, C, pupa, cephalothorax, abdomen and paddle.

—
——

VOLUME 80, NUMBER 3 367

parallel; lateral portion of outer division with a small or weakly sclerotized,
divergent lateral spine and a small, apically rounded lateral basal process;
inner division represented by a simple, broad, leaflike ventral arm which is
sharply pointed and strongly divergent laterad; DV/D ratio [or distance
between apices of tergal mesal spine and ventral arm (DV)/distance be-
tween apices of tergal mesal spines (D)] usually 1, varies from 0.7-1.
Proctiger: Apical crown large, dark, composed of 4-5 flat and blunt spicules
laterally and numerous spinelike spicules laterally and mesally; paraproct
well sclerotized; basal sternal process rudimentary or poorly developed,
at most 0.03 mm in length; cercal sclerite poorly sclerotized; cercal setae
34.

Female (Fig. 2)—Wing 4.2 mm. Forefemur 1.98 mm. Proboscis 2.3
mm. Abdomen 3.24 mm. General coloration light brownish. Head: Eyes
contiguous above antennal pedicels; decumbent scales on dorsum of
vertex narrow, crescent-shaped, rather coarse and predominantly pale beige
in center, fine and whitish on orbital line; erect scales numerous, evenly
spread, largely dark brownish except for a few pale ones in center; lateral
patch of broad appressed scales whitish; frontal bristles strong, yellowish
or golden; upper orbital bristles weaker, dark brownish; suborbital bristles
weak, pale yellowish to dark brownish. Clypeus bare, integument dark
brownish. Palpus 4-segmented, about 0.2 of proboscis length, largely dark
scaled, apex of segment 4 usually tipped with some pale scales on inner
dorsal surface. Proboscis completely dark scaled on labium; labial basal
setae 4 with 2 lateral ones strong and as long as palpus and 2 median ones
weaker and shorter. Antenna slightly shorter or as long as proboscis;
pedicel with a distinct patch of semi-erect scales and setae on inner dorsal
surface; flagellum 13-segmented; flagellar segment 1 with or without a few
pale scales; 5-6 flagellar bristles, very weak and sparse, their length about
2x as long as one flagellar segment. Cibarial armature: Cibarial dome
oval, strongly imbricate; cibarial bar evenly concave except for slight pro-
jection at middle; about 30 teeth, all short, apices blunt, truncate or abruptly
pointed; 3-4 median teeth weakly developed and lightly pigmented, lateral
teeth stronger and dark pigmented. Thorax: Mesonotal integument brown-
ish or lighter, but not blackish; mesonotal scales narrow, crescent-shaped
and dense, more or less uniformly pale beige or dull yellowish on disc, pale
whitish on extreme anterior promontory, lateral margin of supra-alar, mid-
dle of prescutellar space and scutellar lobes; acrostichal bristles well de-
veloped in a double row from anterior promontory to near prescutellar
space; dorsocentral and supra-alar bristles strong; mid-scutellar lobe with
7-8 bristles, lateral scutellar lobe with 6-7 bristles. Integument of pronotum
same color as mesonotum,; anterior pronotal lobe (apn) with 6-8 strong
bristles and several pale scales on dorsal surface. Posterior pronotum
(ppn) with a broad patch of narrow, pale beige scales on anterior upper
surface; 5-6 strong, dark posterior bristles. Pleural integument paler than

368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

A
em, i
49 te ;
Nin ; i
AN} ae §
ARN vi i
= WA Vs +
WAN
rs yh \ ‘
wy \
5

Fig. 4. Culex quinquefasciatus, larva. A, head; B, mental plate; C, thorax and abdo-
men I-VI; D, abdomen VII, VIII, siphon and saddle; E, comb scale; F, pecten tooth.

VOLUME 80, NUMBER 3 369

mesonotum and without definite pattern of darkened areas; whitish scale
patches present, distinct, restricted to propleuron (ppl), sternopleuron (stp)
and mesepimeron (mep); ppl with a small scale patch at base on lateral
surface; stp with a broad scale patch on uppermost corner and a separate
vertical scale patch along posterior border; mep with a broad scale patch at
same level as upper corner of stp and several loosely packed scales among
upper mesepimeral bristles; ppl with 5-7 bristles and 5 other weak setae;
lower mep bristles 1-2 and sometimes 3; upper mep bristles about 10.
Legs: Anterior surface of forecoxa with several strong, curved bristles
and a broad scale patch, latter largely dark on lower surface, pale whitish,
forming a distinct spot on upper lateral surface; anterior surface of mid-
and hindcoxae with a narrow whitish scale patch; trochanters and bases
of femora pale scaled; anterior surface of fore- and midfemora dark
scaled, apex tipped with pale scales, ventral surface whitish scaled; an-
terior surface of hindfemur with a broad longitudinal pale stripe from base
to apex, dorsal surface dark scaled, ventral surface whitish scaled; all tibiae
dark on dorsal surface, apex tipped with pale scales, ventral surface pale;
all tarsi completely dark or blackish scaled; claws of all legs small, equal
and simple. Wing: Scales on all wing veins dark and dense; plume scales
on Rs, Rs and Ry,; narrow, linear; cell R2 about 3x as long as length of
Rs,3; furcation of cell M. at same level as or slightly distad of furcation of
cell Rz; alula fringed with a row of 12-14 dark, narrow scales; upper and
lower calypters fringed with numerous long, yellow, hairlike setae. Halter:
Peduncle pale and bare; knob cupshaped, covered with several pale
scales. Abdomen: Tergites II-VII with distinct basal pale bands and baso-
lateral pale spots, latter on tergites II-V not distinct from above; tergum I
with dark caudal scale patch, basal bands on tergites II-V broadened in
middle, narrow laterally and not connected with basolateral pale spots;
basal bands on tergites VI-VIII evenly broad and comnected with basolateral
pale spots or streaks which are visible from above; sternites predominantly
yellowish. Genitalia: Sternite VIII with distinct median emargination; lat-
eral caudal margin with a row of 7-8 strong, curved bristles, median caudal
margin with several weaker bristles. Tergite IX narrow with an irregular
row of about 10 bristles on lateral caudal margin, median portion bare. Cerci
short, thumblike, about 0.15 mm in length, with numerous setae largely
restricted to apical lateral surface. Postgenital plate rounded on posterior
caudal margin, apical 0.5 with a double lateral row of 6-7 bristles, with
1-2 of most distal bristles strongest. Posterior cowl narrow, ribbonlike,
with numerous spicules. Vaginal sclerite horseshoe-shaped or in form of
a U. Sigma with a dense tuft of 8-9 strong setae.

Pupa (Fig. 3B, C).—Abdomen 3.6 mm. Paddle 0.90 mm. Trumpet 0.72

mm; index 5. Detailed chaetotaxy as figured. Cephalothorax: Yellowish
| white with indefinite darkened areas along margin of posterior middorsal
| ridge, leg and wing cases; setae 1- to 3-C triple; 5-C 4-5 branched; 8-C usu-

|
|

370 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ally 3-4 branched (2-4); 9-C 2-3 branched. Trumpet: Meatus narrow and
dark in basal 0.25, apical 0.75 gradually broadened or more or less cylindrical
and pale; apical margin truncate or slightly emarginated; pinna oblique and
long, 0.30-0.38 of total length. Metanotum: Darkened in middle, pale
laterad; seta 10-C 8-10 branched; 11-C double; 12-C 3-4 branched (2-5).
Abdomen: Segments I-IV darkened in middle, pale towards lateral margin;
segments V-VIII uniformly pale yellowish to whitish; setae 3-I to 3-III
double, 3-III sometimes single; 5-II and 5-III 4-5 branched; 6-I and 6-II
single; 7-I and 7-II double; 1-II small, brushlike, dendritic, with 15-16
distal branches; 1-III to 1-VI subequal, 4-5 branched, 0.50-0.75 of length
of segment following; 1-VII shorter, usually 4 branched (3-4); 5-IV to 5-
VI strong, as long as or slightly longer than segment following; 5-IV usually
triple, sometimes double; 5-V and 5-VI double; 6-III to 6-V subequal,
usually triple (2-4); 6-VI stronger, usually 4 branched (3-4); 4-VII double;
9-VII usually 4 branched (3-4); 9-VIII 5-8 branched. Paddle: Very broad,
hemispherical; color whitish to almost transparent; external buttress and
midrib distinct, but not infuscate; outer margin smooth or minutely spic-
ulate; setae 1-P and 2-P minute, single.

Larva (Fig. 4)—Head 0.78 mm. Siphon 1.3 mm; index 4. Saddle 0.38
mm; siphon/saddle ratio 3.3. Detailed chaetotaxy and general features as
figured. Head: Broader than long; integument pale yellowish from
level of ocular bulge to anterior margin of frontoclypeus, darker posteriorly,
collar brownish; ocular bulge prominent; labrum narrow; seta 1-C pale,
proximally flattened, distally filamentous, its length about 0.5 of the distance
between bases of the pair; 4-C single, as long as or slightly longer than the
distance between bases of the pair; 5-C and 6-C usually 5 branched (4-6),
strong, subequal, their apices reaching slightly beyond mouthbrush; 7-C
8-10 branched, slightly shorter than 5-C and 6-C; 13-C 4 branched; 14-C
single; 16-C and 17-C not developed. Antennal shaft 0.50-0.75 of head
length, straight or weakly curved outward in middle; proximal portion with
numerous strong spicules, distal portion beyond base of setae 1-A with
or without a few spicules; pigmentation same as head capsule; 1-A large,
fan-shaped, with about 22 strongly pectinate branches; 2-A and 3-A single,
bristlelike and pale, both situated subapically. Mental plate brownish, with
10-13 lateral teeth on each side of a median tooth. Mouthbrush composed of
numerous long, yellowish filaments. Thorax: Integument glabrous; setae
1-P to 8-P strong, subequal, 1-P to 3-P single; 4-P double; 7-P usually
double, sometimes triple or 4 branched; 8-P usually double, sometimes
triple or 4 branched; 14-P single; 3-M single; 4-M double; 8-M 6-8 |
branched; 9-M and 9-T 5-6 branched; 7-T 7-10 branched; 12-T single; |
13-T 3-7 branched. Abdomen: Segment I-IV: Integument glabrous; setae |
6-I and 6-II usually 4 branched, sometimes 3; 7-I double, sometimes triple; |

VOLUME 80, NUMBER 3 aia

1-III to 1-VI strong, 0.50-0.75 of seta 6-III to 6-VI, 1-III and 1-IV usually
single, sometimes double; 1-V and 1-VI double; 6-III to 6-VI all double.
Segment VII: Seta 1-VII 3-4 branched; 3-, 7-, 10- and 12-VII single; 4-VII
single or double. Segment VIII: Lightly spiculate; comb scales 30-40, all
broad, short, subequal, apical fringe rounded, composed of evenly fine
spicules; seta 1-VIII 5-6 branched; 2-VIII and 4-VIII single; 3-VIIT 7-8
branched; 5-VII 4 branched. Saddle complete, pigmentation whitish or light
yellowish; spiculation and sculpture practically absent or poorly developed;
posterior caudal margin weakly spiculate; seta 1-X single, very distinct; 2-X
with 1 short and 1 long branch; 3-X single; 4-X (ventral brush) consists of
6 pairs of setae, all inserted within grid; anal gills stout, apex pointed, as
long as or slightly longer than saddle length. Siphon: Rather stout and
thick, somewhat fusiform; acus present, blackish, tube yellowish with
variable amount of brownish tinge; pecten teeth developed, 6-12 in a ventral
lateral row from base to about 0.3 of total length of siphon; 3-4 distal
teeth with 3 graded strong basal denticles and 1 spinelike apical denticle;
siphonal tufts 4 pairs (total 8), placed beyond pecten; 2 proximal pairs
strong, subequal, 6-8 branched, as long as siphonal width at point of at-
tachment; 2 distal pairs reduced, 4-6 branched; most distal pair placed
subventrally, the other more proximal, laterally; seta 2-S pale, single, spini-
form; median caudal filament of spiracular apparatus developed and dis-
tinct.

Acknowledgments

We are most grateful to Professor John N. Belkin, Department of Biology,
University of California, Los Angeles, for making available to us his topo-
typic material of quinquefasciatus for description and neotype designation,
for encouraging us to fulfil these tasks and for reading the final draft.
Our thanks for innumerable favors and valuable suggestions for improv-
ing this paper are due to: Drs. Curtis W. Sabrosky, F. Christian Thomp-
son and Alan Stone (retired), Systematic Entomology Laboratory, IBIII,
Federal Research, Sci. Educ. Admin., Agriculture Research Service, USDA;
Drs. Paul Freeman and Roger W. Crosskey, Department of Entomology,
British Museum (Natural History); Dr. Oliver S. Flint, Jr., Department of
Entomology, Smithsonian Institution; Dr. Ronald A. Ward, Department of
Entomology, Walter Reed Army Institute of Research; and Dr. Richard V.
Melville, Secretary to the International Commission on Zoological Nomen-
clature. Finally, we thank Thelma Ford Smith and Vichai Malikul, Medical
Entomology Project, for preparing the illustrations.

This work was supported by Research Contract No. DAMD-17-74-C-
4086 from the U.S. Army Medical Research and Development Command,
Office of the Surgeon General, Washington, D.C.

372 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Literature Cited

Amendments to the Internationol Code of Zoological Nomenclature Adopted since
the XVI Intemational Congress of Zoology, Washington. 1963. 1974. Bull.
Zool. Nomencl. 31:77-101.

Belkin, J. N. 1962. The mosquitoes of the South Pacific (Diptera, Culicidae). Univ.
Calif. Press, Berkeley and Los Angeles. 2 vols. 606 and 412 pp.

———. 1968a. Mosquito studies (Diptera, Culicidae) VII. The Culicidae of New
Zealand. Contrib. Am. Entomol. Inst. (Ann Arbor) 3(1):1-182.

——. 1968b. Mosquito studies (Diptera, Culicidae) IX. The type specimens of
New World mosquitoes in European museums. Contrib. Am. Entomol. Inst.
(Ann Arbor) 3(4): 1-69.

———. 1977. Quinquefasciatus or fatigans for the tropical (southern) house mos-
quito (Diptera: Culicidae). Proc. Entomol. Soc. Wash. 79:45-52.

Belkin, J. N., S. J. Heinemann, and W. A. Page. 1970. Mosquito studies (Diptera,

Culicidae) XXI. The Culicidae of Jamaica. Contrib. Am. Entomol. Inst. (Ann —

Arbor) 6(1):1—458.
Belkin, J. N., R. X. Schick, and S. J. Heinemann. 1966. Mosquito studies (Diptera,

Culicidae) VI. Mosquitoes originally described from North America. Contrib.

Am. Entomol. Inst. (Ann Arbor) 1(6):1-39.

Bram, R. A. 1967. Contributions to the mosquito fauna of Southeast Asia.—Il.
The genus Culex in Thailand (Diptera: Culicidae). Contrib. Ann. Entomol.
Inst. (Ann Arbor) 2(1):1—296.

Coquillett, D. W. 1906. A classification of the mosquitoes of North and Middle
America. U.S. Bur. Entomol., Tech. Ser. 11. 31 pp.

International Code of Zoological Nomenclature. 1964. 2nd ed. London, International
Trust for Zoological Nomenclature. xix + 176 pp.

Knight, K. L., and A. Stone. 1977. A catalog of the mosquitoes of the world (Diptera:
Culicidae). 2nd ed. The Thomas Say Foundation, Entomol. Soc. Am. Vol. 6:
x - 611 pp.

Say, T. 1823. Descriptions of dipterous insects of the United States. J. Acad. Nat.
Sci. Phila. 3:9-54.

Sirivanakarn, S. 1976. Medical entomology studies—III. A revision of the sub-
genus Culex in the Oriental region (Diptera: Culicidae). Contrib. Am. Entomol.
Inst. (Ann Arbor) 12(2):1—272.

Stone, A. 1956 (1957). Corrections in the taxonomy and nomenclature of mosquitoes _

(Diptera, Culicidae), Proc. Entomol. Soc. Wash. 58:333-344.
Stone, A., K. L. Knight, and H. Starcke. 1959. <A synoptic catalog of the mosquitoes

of the world (Diptera, Culicidae). Thomas Say Foundation, Entomol. Soc. Am. |

Vol. 6. 258 pp.
Weiss, H. B., and G. M. Ziegler. 1931. Thomas Say, early American naturalist.
Charles C. Thomas, Springfield, Ill. 260 pp.

Wiedemann, C. R. W. 1828. Aussereuropaische zweiflugelige Insekten. Vol. 1. |

Hamm. 608 pp.

(SS) Medical Entomology Project, Smithsonian Institution, Washington, _
D.C. 20560; and (GBW) Department of Entomology, British Museum (Nat-

ural History), London SW7 5BD, Great Britain.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 373-379
AN INFESTATION OF MILTOGRAMMINE SARCOPHAGIDAE
(DIPTERA: SARCOPHAGIDAE) IN A POPULATION OF
HYBOMITRA LASIOPHTHALMA (MACQUART)
(DIPTERA: TABANIDAE)

Patrick H. Thompson

Abstract—A second isolation of Macronychia sp. near aurata (Coquillett)
was made from adult specimens of Tabanidae in south-central Texas.
Fifty-nine sarcophagid larvae were obtained from 270 adult females of
Hybomitra lasiophthalma (Macquart) which were taken 7-14 April 1977, in-
clusive, during their peak emergence on the Navasota River floodplain
near College Station, Texas. The apparent high incidence of parasites in
this tabanid population (22%) is qualified by the observed frequency of
multiple parasitism; of 4 parasitized females dissected, 3 suffered multiple
infestations of 2, 3 and 5 larvae, respectively. The chronology of para-
sitization of H. lasiophthalma by this sarcophagid suggests that the tabanid
serves as a first host species for the parasite, while species of Tabanus, such
as Tabanus subsimilis subsimilis Bellardi, act as hosts of subsequent genera-
tions of parasites. Once again, the parasite was found infesting T. s. sub-
similis.

The isolation of miltogrammine sarcophagid larvae from adult females of
Tabanus subsimilis subsimilis Bellardi was reported by Thompson (1978).
That particular infestation involved a population of this tabanid collected
near Easterwood Airport and the sewage treatment plant, College Station,
Texas 28 May through 14 July 1976. Subsequently, the sarcophagid para-
site was identified as Macronychia sp. near aurata (Coquillett), hereafter re-
ferred to as Macronychia sp. The 1978 report considered the chronological
and ecological details coincident with the parasitization; methods of rearing
the parasite, and its larval behavior; relevant biology for the host and for
the parasite; and the implications of these findings for the known life
histories of miltogrammine sarcophagids and for the management of noxious
populations of Tabanidae.

During a study of Hybomitra lasiophthalma (Macquart) the following
year, Macronychia sp. was again observed commonly infesting females of

_ the host species. The details of this sarcophagid-tabanid association will

"
:

:

be compared with those previously recorded for Macronychia sp. and T. s.

_ subsimilis (Thompson, 1978).

Studies of reproductive physiology of Tabanidae, preparatory to rearing
_ work, were begun with the first spring dominant found in the vicinity of
_ College Station. All females of Hybomitra lasiophthalma collected during

|
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

374

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VOLUME 80, NUMBER 3 375

Table 1. Daily catches of Hybomitra lasiophthalma females taken 7-14 April 1977,
inclusive, and numbers of infesting Macronychia sp. larvae emerging from them."

Day of April
i 8 9 10 11 12 13 14 Totals
No. tabanids 22 34 28 22 42 AT 40 35 270
No. sarcophagids 1 6 5 1 5 32 6 3 59
Percent infested 4 18 18 4 12 68 15 9 22

*Eleven larvae were excepted; these were found in the abdominal cavities of dis-
sected females.

the period 7-14 April, inclusive, from two Manitoba Traps located on the
Navasota River, were held in the lab to observe additional instances of
parasitism. Catches were augmented with CO. generated from dry ice.

Methods

All dead H. lasiophthalma females removed from holding cages in the lab-
oratory were retained together by date in 10-dram glass vials. The centers
of the polyethylene lids of these containers were cut out with a heated
No. 12 cork borer; and the lids were subsequently lined with a 25-mm disc
of 70-mesh brass screen (Brass Strainer Cloth, C. O. Jellif Corp., Southport,
Connecticut 06490). These vials of dead flies were examined daily for the
presence of parasite larvae and the first larvae were found on 8 April, when
they emerged from dead H. lasiophthalma females trapped the day before.
Sarcophagid specimens were reared on decapitated crickets in glass Petri
dishes. Parasites removed from flies taken on the same date were reared
together. Further details of rearing were previously described (Thompson,

1978).

Observations

A total of 59 sarcophagid larvae was removed from vials containing 270
adult females of H. lasiophthalma. More than half of these (32) emerged
from host specimens taken on 12 April, the 6th day of collection (Table 1).

Mortality of the larvae was very high (81.4%) in contrast to that suffered

by the lesser number of specimens reared from Tabanus subsimilis sub-
similis (17.7%; Table 2).

Development and behavior—The periods of parasite larval develop-
ment and of pupal development, under the same physical conditions as for
T. s. subsimilis, were considerably longer (Table 2). The high mortalities
and attenuated development of parasites found in H. lasiophthalma could re-
flect microbial infection. Again, larval behavior was scavenger like; but

376 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Summary data of Macronychia sp. infestations in Hybomitra lasiophthalma
and Tabanus subsimilis subsimilis.*

H. lasiophthalma T. s. subsimilis

No. larvae collected 59 27
No. larvae reared ll 14

no. females 7 4

no. males 4 10
Mortality rate (%) 81.4 ae
Larval period (days) 8-14, avg. 10.3 2-8, median 6
Pupal period (days) 18-20, avg. 18.8 15-17, avg. 15.9
No. tabanid adults 270 3,000 (est.)
Parasitism rate (%) 3-22, 1 (est.)
Parasite loads BONS eS (A)) 7,8 (2)

“Data on Tabanus subsimilis subsimilis presented in detail in Thompson (1977).
» Rate based upon 17 larvae reared, rather than 27, because 10 were preserved for
taxonomic purposes.

also again, parasites attacked dead hosts via intact cervical membranes
(4 instances).

Discussion

Briefly, miltogrammine Sarcophagidae are cleptoparasites of wasps and
bees. The two North American species of Macronychia, in particular, have
been reared from nests of two sphecid wasp genera. The findings relating
to tabanid parasitism described here, conflict with those supporting a
cleptoparasitic, and thus secondarily parasitic life history for this species.

Multiple parasitism.—First, as with Tabanus subsimilis subsimilis, the
occurence of multiple infestations in one host suggests that such larvae
were specimens that originated together from the same females foraging
near the host. (In 3 of 4 host infestations examined in vivo, 2, 3 and 5
larvae were found together in the abdominal cavity.) Secondly, these
larvae were the same size.

If these assumptions regarding multiple infestations by sibling larvae
are correct, the mode of life of Macronychia sp. could be that of a primary
parasite. Although another order of insects was involved, Dr. Paul Arnaud
of the California Academy of Sciences, San Francisco, called my atten-
tion to a documented instance of such primary parasitism by a miltogram-
mine species. From 2 camel crickets, he isolated 7 larval specimens of
Hilarella hilarella (Zetterstedt)—specimens which subsequently pupated
and emerged (Arnaud, 1954). Dr. K. V. Krombein, Smithsonian Institution,
suggested that Macronychia species could normally parasitize Diptera and
that apparent instances of these flies parasitizing the prey of sphecid wasps

|

VOLUME 80, NUMBER 3 3t7

could, instead, represent primary parasitism of other species of Diptera be-
fore they fall prey to the wasps (Krombein, personal communication).

Rate of parasitism.—The parasite-host ratio was estimated because host
females of Hybomitra lasiophthalma were not isolated individually. As-
suming that 59 larvae infested host flies singly, this rate would be 22%;
conversely, if 8 parasites infested each host (8 was the most observed), this
rate would be 3%. Obviously, the actual rate lies between 3% and 22%.

One factor would surely have increased this rate. Dissections of only a
few H. lasiophthalma females produced 4 specimens with dead or dying
larval parasites. Therefore, unless this small sample is atypical, many in-
festing parasites never completed development.

The host.—(A review of this tabanid, currently in preparation, will detail
and document information in the following summary.) Presently, H.
lasiophthalma is one of the most widespread and abundant species of its
genus in North America, extending from British Columbia to Nova Scotia,
and south to Texas and Georgia. In southern Canada and the northern
United States, where the larvae breed extensively in sphagnum bogs and
shrub-sedge marshes, the adults emerge in June and July. As this form
extends southward through the eastern states, its distribution is largely con-
fined to rivers where it breeds predominantly in heavy organic soils of
floodplain forests and where its seasonal distribution constricts and _ re-
cedes, the adults emerging over a shorter period in spring, rather than
through midsummer, as farther north. In such riverine localities, and their
surrounding ridges, this horse fly is often a dominant species and a major
pest of cattle and horses. Parity in H. lasiophthalma has been described and
females oviposit a blackish, tar-like mass of eggs on the leaves of reeds,
sedges, cattails and shrubs growing in low areas which locally characterize
the larval habitat. As is also typical of other North American species of
Hybomitra and Tabanus, the adults commonly feed on nectars (this is
often evidenced by pollen grains on the mouth parts of trapped specimens):
often pursue rapidly-moving vehicles; fly near roadside puddles and wood-
land pools where they frequently touch the water surface, or “dip,” as they
circle above it; and hover in sunlighted woodland openings (males), prob-
ably in frequent association with mating. This tabanid was one of the
first North American species reared in the laboratory from eggs collected in
the field; and as with the eggs, the larvae and the pupae have been ob-
served and collected from lowland habitats in nature. In addition, the im-
matures and both sexes of the imago have been described. Hybomitra
lasiophthalma has been the subject of other studies involving physiology
and disease transmission.

The relationship.—Most significant to the association described here,
are the chronological facts causally relating populations of the parasite to
those of the two known host species of the Navasota River floodplain and its

378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

adjacent uplands. Macronychia sp. was first observed in Tabanus sub-
similis subsimilis during the period 28 May through 14 July 1976; and
the following year, in Hybomitra lasiophthalma, from 7 through 14 April.
Although these infestations were observed in different years, their chronol-
ogy suggest that H. lasiophthalma could serve as an important host for
the first generation of sarcophagids; then the 2nd and subsequent gen-
erations of the parasite could infest populations of Tabanus s. subsimilis
or other numerous forms found during the summer months (Fig. 1).

The curve for T. s. subsimilis presented here is based upon collections
made in the uplands during the previous year (1975; Thompson, 1977);
this curve closely corresponds with that taken in the lowlands the same
year, but for the following reservation. Catches at the river declined in
August, 1975 because of flooding, whereas catches in the uplands con-
tinued to increase for 3 weeks after that.

The 6-week interim between infestations of the two host populations,
as depicted in this figure, could account for a projected trough between
generations, as based upon development times observed in the laboratory
(Table 2).

Concerning the seasonal distribution of Tabanus s. subsimilis, intensive
collections of adults through most of the year in several Texas ecosystems
showed that this species, in contrast to Hybomitra lasiophthalma, is asea-_
sonal; i.e., it is found continuously throughout 9 months of the year and -
peaks anytime during most of that period—from late March to late May |
(Thompson, 1975) or in late summer (Thompson, 1977). Therefore, this.
horse fly is abundantly accessible as a potential host insect during most of |
the season. |

A second infestation of Tabanus subsimilis subsimilis—Macronychia sp. |
was again isolated from this species, in this case, from bottomland popu-
lations collected with catches of Hybomitra lasiophthalma. Two females
which had been held in the laboratory for 3 and 7 days, respectively, con-
tained 2 small 2nd-instar larvae (1 female, 12 April); and 1 large Ist-instar
larva (1 female, 8 April). Of these specimens observed in the abdominal |
cavity, one larva in each fly was alive when it was found.

Conclusions

As with Tabanus s. subsimilis, the circumstances surrounding collec-
tion and retention of Hybomitra lasiophthalma preclude any reasonable pos-
sibility of unnatural parasitism. With both species, specimens were re-
moved from trap collection containers within the laboratory and were
then held alive in the lab with small numbers of their own species (H.
lasiophthalma) or of other Tabanus species (T. s. subsimilis). No other in-
sects or extraneous materials of any kind were contained in any cages of

VOLUME 80, NUMBER 3 379

either species. In addition, no miltogrammine sarcophagids were found in
Manitoba Trap catches containing infested specimens of these tabanid
species. (Several miltogrammine species of other genera have been taken
in small numbers from Gressitt Traps.) In fact, the Manitoba Traps used
during these and similar studies of the past 13 years, have taken very few
numbers of few species of other insects. Finally, infested horse fly speci-
mens were found in numerous samples of the 2 species—S in the case of
H. lasiophthalma and 7 for T. s. subsimilis; and these samples were isolated
from one another in containers inaccessible to other insects the size of
Macronychia.

The incidence of Macronychia parasitism in Tabanidae is now well
documented. The extent of this association with the 2 host tabanids named
here, and of others locally abundant, remains to be defined by further
survey. Parasite-host specificity, and consequent high rates of parasitism,
could offer a potential means of managing pest populations of species which
otherwise remain unmanaged.

Acknowledgment

I gratefully acknowledge the aid of Mr. Joseph W. Holmes, Jr. in
servicing traps; of the Drs. Raymond J. Gagné and Curtis W. Sabrosky,
Systematic Entomology Laboratory, IBII, Fed. Res., Sci. Educ. Admin.,
USDA, for their determinations of Macronychia; and of the Drs. B. J.
Cook, L. E. Ehler, G. B. Fairchild, R. J. Gagné, K. V. Krombein, L. L.
Pechuman, C. B. Philip, and F. E. Wood for their comments on the manu-
script.

Literature Cited

Amaud, P. H. 1954. Hilarella hilarella (Zetterstedt) (Diptera: Sarcophagidae) para-
site upon a rhaphidophorid (Orthoptera: Gryllacrididae). Can. Entomol. 86:135-
136.

Thompson, P. H. 1975. Larval habitats of Tabanus subsimilis subsimilis Bellardi in

southeast Texas (Diptera: Tabanidae). Proc. Entomol. Soc. Wash. 77:494—500.

1977. Comparisons of upland and lowland tabanid populations in south-

east Texas. Proc. Entomol. Soc. Wash. 79:564—574.

—. 1978. Parasitism of adult Tabanus subsimilis subsimilis Bellardi (Diptera:
Tabanidae) by a miltogrammine sarcophagid (Diptera: Sarcophagidae). Proc.
Entomol. Soc. Wash. 80:69-74.

Veterinary Toxicology and Entomology Research Laboratory, Fed. Res.,
Sci. Educ. Admin., USDA, College Station, Texas 77840.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 380-382
THE ALLOTYPE OF ARENASELLA MALDONADOI AND CHANGE
OF DEPOSITORY FOR TWO OF FENNAH’S HOLOTYPES
(HOMOPTERA: TROPIDUCHIDAE AND ISSIDAE)

J. Maldonado Capriles and Angel Berrios

Abstract.—The male allotype of Arenasella maldonadoi Caldwell is de-
scribed from specimens collected from the type-locality. The holotypes of
Tangella pustulifrons Fennah and Colpoptera galatea Fennah are trans-
ferred to the collection of the U.S. National Museum of National History
(USNM), Washington, D.C.

Caldwell (1951) described the second species of Arenasella from female
specimens collected by the senior author near the Recreation Area at El
Yunque National Forest, Luquillo, Puerto Rico. We now describe the male
allotype from a series of specimens collected from almost the same place.
The specimens were collected from a bromeliad (Guzmania sp.). This ful-
goroid protects itself during the day by living under the sheaths of the
leaves that form the base of the plant.

Fennah (1965) described Tangella pustulifrons from specimens collected
by the senior author at Christiana, Jamaica and Colpoptera galatea from
St. John, Barbados, and he deposited the holotypes in Maldonado’s (JMC)
collection. These types now have been transferred to the collection of the
USNM.

Tropiduchidae
Arenasella maldonadoi Caldwell, 1951:224.

The characters of the male agree closely with those of the female. Head
as illustrated in Figs. 1, 2 and 3. Forewing 2.5x as long as broad, Fig.
7. The apex of the clavus is as described by Caldwell for the female, but
the correct venation of the apical half of the forewing is as in Fig. 7. Some
of the longitudinal veins are missing in Caldwell’s drawing. The meso-
pleurite has a black spot. We think Caldwell meant the last thoracic
sternite instead of “abdominal sternite” as there is a conspicuous black
spot on the posterolateral corner of the metapleura almost hidden by the

=
Figs. 1-7. Arenasella maldonadoi Caldwell, male. 1. Head, frontal; 2. Head and |

tp)

pronotum, dorsal; 3. Head and pronotum, lateral; 4. Genital capsule and aedeagus, —
lateral; 5. Aedeagus, dorsal; 6. Aedeagus, ventral; 7. Forewing.

VOLUME 80, NUMBER 3 381

382 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

abdomen. Ventrally near base of the anterior coxa there is a small
black spot. This last spot is inconspicuous in the females at hand and
not mentioned by Caldwell. Looking at the insect from the ventral side
the spots on the procoxa, mesopleura, and prebasally on the costa of the
forewing are distributed arc-like. These spots are also visible in lateral
view. The last abdominal sternum has a conspicuous spot at the lateral
extremity. Length 4.9-5.0 mm.

Genital segments as in Figs. 4-6.

Allotype, male, Puerto Rico, El Yunque National Forest, 20-22 March
1954, J. Maldonado Capriles and S. Medina collectors, in the USNM. Three
paratypes in the USNM, two in the collection of the Department of En-
tomology of the Agriculture Experiment Station of the University of Puerto
Rico, Rio Piedras, and one in JMC.

Tangella pustulifrons Fennah
Tangella pustulifrons Fennah, 1965:99.
Male holotype formerly in JMC, now in the collection of the USNM, Type
No. 75328.
Issidae
Colpoptera galatea Fennah
Colpoptera galatea Fennah, 1965:103.
Male holotype formerly in JMC, now in the collection of the USNM,
Type No. 75329.
Literature Cited

Caldwell, J. S., and L. F. Martorell. 1951. Review of the Auchenorhynchous Homop-
tera of Puerto Rico. Part II. The Fulgoroidea except Kinnaridae. J. Agric.
Univ. P.R. 34(2):133-269.

Fennah, R. G. 1965. New species of Fulgoroidea (Homoptera) from the West
Indies. Trans. R. Entomol. Soc. London. 117(4):95-126.

Department of Biology, University of Puerto Rico, Mayaguez, Puerto
Rico 00708 (JMC now at: Department of Anatomy, School of Medicine,
Catholic University of Puerto Rico, Ponce, Puerto Rico 00731).

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 383-387
TWO NEW CERATOCAPSUS REUTER 1876, FROM
THE EASTERN UNITED STATES (HEMIPTERA: MIRIDAE)!

Thomas J. Henry

Abstract.—Two new species of Ceratocapsus are described: Ceratocapsus
spinosus from Pennsylvania and C. aurantiacus from Florida. The adult of C.
spinosus and the male genitalia of both species are illustrated.

While examining light trap material from the Frost Entomological Mu-
seum, Pennsylvania State University, I discovered two new species of
Ceratocapsus from the United States. One species, taken at the Archbold
Biological Station in Florida is the first truly orange-red Ceratocapsus and
the other species, taken in western Pennsylvania is the largest member of this
genus. They are described and illustrated here.

Ceratocapsus spinosus Henry, new species

Figs. 1-5

Male.—Length 5.83 mm, width 2.0 mm; impunctate, generally fuscous
to black, clothed with fine, short setae, intermixed with erect, pilose setae.
Head: Length 0.82 mm, width 0.96 mm, vertex 0.34 mm, dorsal width
of eye 0.30 mm; black, shiny, finely granulate, not punctate, median line
shallowly grooved, basal carina distinct. Rostrum: Length 1.80 mm,
brown, reaching middle of mesocoxae. Antennae: I, length 0.62 mm, light
brown, with several erect, stout setae; I, length 2.04 mm, weakly thickened
to apex, brown to dark brown, especially on apical '2, clothed with fine,
recumbent, brown to black setae, fewer at base; III, length, 0.92 mm, dark
brown; IV, length, 0.80 mm, dark brown. Pronotum: Length 1.00 mm,
width at base 1.60 mm, finely granulate, shiny, black becoming lighter
or more brown at base, void of pubescence; mesoscutum black; scutellum
black, apex brown, transversely rugose, sparsely set with brown, pilose
setae. Hemelytra: Brown, somewhat alutaceous, clothed with short, re-
cumbent, pale setae intermixed throughout (except on embolium and
cuneus) with brown, pilose setae; cuneus fuscous to black; corium appearing
darker due to fuscous abdomen beneath. Membrane: Translucent, more
fumate on apical '%, veins colored as membrane. Venter: Shiny fuscous
to black, ostiolar peritreme pale, abdomen clothed with recumbent, pale
setae, those setae on genital segment longer. Legs: Uniformly brown,
hind tibiae somewhat darker; setae and tibial spines colored like segments.
Genitalia: Left paramere three pronged (Figs. 2, 3); basal prong at
right angle to middle prong, with 2 distinct subapical spines; dorsal margin

384

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

perc,
Renesas
eS

Fig. 1. Ceratocapsus spinosus, habitus.

VOLUME 80, NUMBER 3 385

of middle prong with 1 spine; right paramere simple, relatively straight
(Fig. 4) basal process simple and blunt; theca (Fig. 5).

Female —Unknown.

Holotype.—é, Pennsylvania: Beaver Co., Pa., Rt. 168, 6 mi SW of Dar-
lington, 4 August 1975, Mrs. M. A. Carter collector (USNM type #74022).
Paratype: 4, 1 August 1975, same data as holotype (PSU collection).

Remarks.—Ceratocapsus spinosus keys to nigrocephalus Knight in Knight
(1923, 1941), but differs from it by the larger size, the uniformly black
cuneus, the brown corium, the relative lengths of the antennal segments
and the structure of the male genitalia.

This species may be separated from all other Ceratocapsus by the large
size, the black head, pronotum, scutellum and cuneus, the brown hemelytra
clothed with pilose setae and the male genitalia. The basal prong of the
left paramere bears two subapical barbs or spines which easily char-
acterize this species and are the basis for the name spinosus.

Ceratocapsus aurantiacus Henry, new species
Figs. 6-S

Male.—Length 3.36 mm, width 1.48 mm, impunctate, general color
orange to orange red, clothed with pale to silvery, sericeous pubescence,
intermixed with more erect, pale setae. Head: Length 0.68 mm, width
0.78 mm, vertex 0.30 mm, orange, sparsely set with erect, pale setae;
eyes reddish. Rostrum: Length about 1.30 mm (imbedded in glue), reach-
ing posterior margin of metacoxae. Antennae: I, length 0.38 mm, pale
testaceous; II, length 1.06 mm, pale testaceous, apex lightly tinged with
orange; III, length 0.64 mm, testaceous, tinged with orange; IV, length 0.52
mm, pale orange or reddish. Pronotum: Length 0.70 mm, width at base
1.20 mm, shiny orange to reddish orange, sparsely set with erect and semi-
erect, simple setae; mesoscutum orange; scutellum orange, more reddish
just before apex, apex pale orange, weakly but distinctly rugose. Hemelytra:
Generally orange to orange red; outer margin of corium, area just before
cuneal fracture and cuneus distinctly orange red; sparsely set with semi-
erect, pale setae, clavus and corium clothed with silvery, sericeous pu-
bescence. Membrane: Fumate, veins colored like membrane. Venter:
Orange to red, abdomen red, genital segment more orange and clothed with
long, pale setae. Legs: Pale testaceous; tibiae weakly spined; claws and
tarsi testaceous. Genitalia: Left paramere three pronged; basal prong
short, appressed along dorsal margin of middle prong, apex strongly re-
curved (Fig. 6); right paramere sickle-shaped with basal process notched
apically (Fig. 7); theca (Fig. 8).

Female.—Length 3.48 mm, width 1.60 mm. Head: Length 0.68 mm, width
0.78 mm, vertex 0.36 mm. Rostrum: Length about 1.10 mm (imbedded in

386 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 2-5. Ceratocapsus spinosus. 2. Left paramere, lateral view; 3. Left paramere,
inside lateral view; 4. Right paramere; 5. Theca of aedeagus. Figs. 6-8. Ceratocapsus
aurantiacus. 6. Left paramere, lateral view; 7. Right paramere, lateral view; 8. Theca
of aedeagus.

glue), reaching middle of mesocoxae. Antennae: Testaceous; I, 0.32 mm, |
red dash on inside margin; I, 1.14 mm, apex tinged with orange; III, 0.64
mm, apical '2 tinged with orange; IV, 0.46 mm, reddish orange, pale at base.
Pronotum: Length 0.70 mm, width at base 1.24 mm. The females are sim-
ilar to the male in color and form except for minor variation noted in
the antennae.

Holotype.—¢, Archbold Biological Station, Highlands Co., Fla., 5-3-1967,
S. W. Frost collector (USNM Type #75435). Allotype: ¢, same data as
holotype, 5-5-1967 (USNM). Paratypes: 22, same data as other types
(PSU collection).

Remarks.—Ceratocapsus aurantiacus keys to taxodii Knight or vicinus |
Knight (Knight, 1941) and to vicinus Knight (Blatchley, 1926), but differs
from both species in color and the structure of the male genitalia. |

Ceratocapsus aurantiacus is characterized most easily by the orange to

VOLUME 80, NUMBER 3 387

orange-red color and the structure of the male genitalia. The basal prong
of the left paramere is reduced but strongly recurved apically; the basal pro-
cess of the right paramere is rather broad and apically notched. The name
aurantiacus is derived from the Latin adjective aurantium, meaning orange.

Acknowledgments

I thank Dr. K. C. Kim of the Frost Entomological Museum, Pennsylvania
State University for the loan of specimens and Drs. Kim, and K. R. Valley
and A. G. Wheeler, Jr., Bureau of Plant Industry, Department of Agri-
culture for reviewing the manuscript, and Mr. G. C. Steyskal, Systematic
Entomology Laboratory, IIBIII, Fed. Res., Sci. Educ. Admin., USDA for
his suggestion and explanation of the name aurantiacus.

Literature Cited

Blatchley, W. S. 1926. Heteroptera or true bugs of eastern North America. The
Nature Publishing Co., Indianapolis. 1,116 pp.

Knight, H. H. 1923. Family Miridae (Capsidae). Pp. 422-658. In Britton, W. E.
(ed.) The Hemiptera or sucking insects of Connecticut. Conn. State Geol.
Nat. Hist. Surv. Bull. No. 34.

——. 1941. The plant bugs or Miridae, of Illinois. II]. Nat. Hist. Surv. Bull. 22,
234 pp.

The Frost Entomological Museum, Department of Entomology and the
Pennsylvania State University, University Park, Pennsylvania 16802 (mail
address: Bureau of Plant Industry, Pennsylvania Department of Agri-
culture, Harrisburg, Pennsylvania 17120).

Footnote

* Authorized for publication on 15 August 1977 as paper no. 5349 in the Journal
Series of the Pennsylvania Agricultural Experiment Station.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 388-396
BIOLOGY AND IMMATURE STAGES OF
ANTICHAETA BOREALIS (DIPTERA: SCIOMYZIDAE),
A PREDATOR OF SNAIL EGGS

William H Robinson and B. A. Foote

Abstract.—Larvae of all investigated species of Antichaeta feed on snail
eggs. Females of the North American A. borealis Foote oviposit directly
onto egg masses of the snail Oxyloma sp., and all three larval instars feed
solely on the eggs. The mature larva usually leaves the egg mass before
forming a puparium. Antichaeta borealis is multivoltine in northeastern
Ohio. Three species of Ichneumonidae, Phygadeuon sp. A, Phygadeuon
sp. B and Mesoleptus sp., were reared from puparia collected in the study
area. The egg, three larval instars and puparium are described and il-
lustrated. The life histories of the reared species of Antichaeta are reviewed
and compared.

The genus Antichaeta is one of the smaller genera in the family Sciomyzi-
dae, consisting of 5 Palearctic and 8 Nearctic species. An undescribed spe-
cies is known from Michoacan, Mexico (K. Valley, personal communica-
tion). Biological information is available for 7 of the 13 described species.
Investigations of the biology and immature stages of Antichaeta species were
conducted by Fisher and Orth (1964), Knutson (1966) and Knutson and
Abercrombie (1977). Larvae of all investigated species feed on eggs
of snails.

This paper outlines the life history and describes the immature stages
of A. borealis Foote, a widespread North American species whose larvae
attack eggs of terrestrial snails of the genus Oxyloma (Succineidae). Spe-
cific morphological features of the immature stages are illustrated in de-
tail. Laboratory rearings and observations reported here are based on
material collected during the spring and summer of 1965 at an extensive
marsh located about 6 kilometers east of Kent, Ohio.

Biology of Antichaeta borealis

In northeastern Ohio, the only area where the biology was studied, adults —
and larvae were found most commonly in open, permanent marshes and —
frequently along the shaded borders of such areas. The habitat distribution |

of the fly is similar to that of the principal food snail, Oxyloma sp.
Recently emerged females of A. borealis had a premating period (from
emergence to first copulation) of 24-36 hours. No specific courtship be-
havior was observed before mating. The mating position was similar
to that of other species of the genus (Fisher and Orth, 1964; Knutson, 1966).

VOLUME 80, NUMBER 3 389

The male was situated dorsally and faced in the same direction as the
female, with the anterior portion of his head (not including the antennae)
extending to about the mid-length of the thorax of the female. The male’s
front tarsal segments were laid along her parafrontal areas, laterad to the
fronto-orbital bristles and bending them slightly inward. The tarsi ex-
tended over the female’s fronto-facial ridge, just laterad to her antennae.
His middle legs remained free or occasionally rested on the basicostal
margins of the female’s half-outstretched wings. The hind tarsi of the
male were placed on the abdomen of the female. Copulation was not re-
stricted to any particular part of the day, and often lasted an hour. Adults
mated frequently before and after oviposition.

Egg masses of Oxyloma sp. (probably O. retusa (Lea)) apparently provide
the only natural oviposition site for A. borealis. Although laboratory-reared
females also readily oviposited onto egg masses of Physa sp., Helisoma sp.,
Catinella vagans (Pilsbry) and other species of Oxyloma, they did not
lay eggs on living or dead snails; and they oviposited on vegetation only
when egg masses were not available. Females confined with males had a
pre-oviposition period of 3-4 days. Before laying eggs, females exhibited
a curious behavior that apparently was a prerequisite for oviposition. They
first searched the bottom of the breeding jar for a suitable oviposition site
(i.e., snail egg masses). Upon locating a mass, the female would walk onto
it and spend several minutes exploring its surface with her front tarsi. After
this brief routine she would begin laying eggs. Eggs were scattered over
the surface of the masses, with 1-30 eggs deposited on each egg mass in
the laboratory. Eggs deposited on Oxyloma sp. egg masses were so oriented
that the upturned anterior and posterior ends projected above the surface
of the egg mass. Soon after the eggs were laid they appeared to be en-
circled by a colorless liquid which gave the appearance of a small halo
around each egg. Eggs that were placed onto egg masses of other snail
species did not show this phenomenon. Several females each laid over
150 eggs within 40 days, and one adult deposited 207 eggs within a 35-day
period. Hatching occurred within 24-36 hours.

After hatching, each first-stage larva penetrated the egg mass and began
:attacking the contained eggs. Because of the viscosity of the gelatinous
‘matrix surrounding Oxyloma eggs and the position of the eggs within the
|matrix, it was necessary for the larva to expend considerable energy and
‘time in locating individual eggs. The larva traveled through the matrix
in an undulatory manner. After a larva located an egg it began rasping at
the vitelline membrane with its mouthhooks. After puncturing the mem-
brane, the larva extended its anterior end into the egg and began feeding
on the developing snail embryo. First-stage larvae occasionally entered
the egg cell completely, in the process of ingesting its contents. A larva
consumed the contents of one egg in a few hours then crawled through the

390 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

csp

Rist
%
eee

Figs. 1-8. Antichaeta borealis, immature stages. 1. Anterior segments, third-stage
larva; 2. Lateral view, third-stage larva; 3. Posterior spiracular disc, posterior view,
first-stage larva; 4. Posterior spiracular disc, posterior view, second-stage larva; 5.
Posterior spiracular disc, posterior view, third-stage larva; 6. Anterior spiracle, second-
stage larva; 7. Anterior spiracle, third-stage larva; 8. Posterior spiracular plate, third-
stage larva. Abbreviations: asp, anterior spiracle; csp, sclerotized sensillum; fh, float
hair; mh, mouthhook; psp, posterior spiracle; ss, spiracular slit; va, ventral arch.

gelatinous matrix in search of other eggs. During this search for eggs and
also during feeding, the posterior spiracles of the first-stage larva were
frequently out of contact with the ambient air. The first stadium lasted
3-4 days, and by the 3rd day the larva had developed to such a size
that the posterior spiracles could be extended above the surface of the
egg mass. First-stage larvae in Oxyloma egg masses each consumed from
4-7 eggs. Molting took place within the egg mass.

First-stage larvae developing in Physa sp. and Helisoma sp. egg masses
exhibited, for the most part, the same feeding behavior as those utilizing
Oxyloma egg masses. Larvae in Helisoma egg masses fed on the underside
of the mass (the side without a hardened protective covering). Relatively

|

VOLUME 80, NUMBER 3 391

few of the first-stage larvae feeding on Physa sp. and Helisoma sp. egg
masses reached the second instar.

Second-stage larvae remained in the Oxyloma egg masses and actively fed
on the living snail embryos. Although second instars were restricted pri-
marily to feeding on snail eggs, one larva that had not fed for 24 hours
was observed to kill and eat a recently hatched Oxyloma snail. When several
eggs had been laid on one egg mass and the larvae subsequently became
crowded, several left the original egg mass and searched for others. Second-
stage larvae fed with the posterior spiracles in contact with the surface of
the egg mass. The second stadium lasted 2-3 days. By the 3rd day most of
the larvae had acquired a faint red color in their digestive tracts, and their
Malpighian tubules were dark maroon. The reddish coloring in the gut
persisted during feeding and molting but disappeared when the larva det-
ecated. The number of Oxyloma eggs consumed by each second-stage
larva varied from 3-11, the usual number being 7. Like the preceding in-
star, molting took place within the egg mass. Second-stage larvae feeding
on Physa and Helisoma egg masses failed to develop fully and died before
the end of the second stadium.

Third-stage larvae continued to feed on eggs within the masses. The
reddish color of the intestine and Malpighian tubules became more pro-
nounced, and the tracheal system became dark gray. During the 7-10
days of the third stadium, a larva consumed considerably more eggs than
the other 2 instars combined. It was not unusual for a third instar to
destroy 105-110 Oxyloma eggs.

When third-stage larvae became fully developed, they stopped feeding,
frequently abandoned the egg mass and burrowed into the cotton or moist
filter paper in the rearing dishes where pupariation occurred. Hardening
of the puparium required about 5 hours. The prepupal period lasted ap-
proximately | day; the pupal period, about 15 days.

The earliest collection record for adults in northeastern Ohio was 16
May (1965); the latest, 10 December (1965). Adult populations peaked
around the end of June or the first part of July. Larvae and pupae were
discovered in nature between May and September. Antichaeta borealis is
multivoltine and overwinters as diapausing pupae. In northeastern Ohio
first-generation adults emerged in early May.

_ Puparia collected in January (1966) and retained at temperatures be-
»tween 5-7°C for at least 60 days produced adults after 8-10 days after
being returned to room temperature. Adult males were usually the first
to emerge and were followed by females in 24-48 hours. Reared adults
lived 7-44 days. Field collected adults lived 3-36 days in the laboratory.
Three species of Ichneumonidae, Phygadeuon sp. A, Phygadeuon sp. B
and Mesoleptus sp. (determined by Dr. William Mason) were reared from
puparia collected in marshes near Kent. Each infested puparium pro-
duced only 1 wasp.

392 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 9-13. Antichaeta borealis, immature stages. 9. Cephalopharyngeal skeleton,
third-stage larva; 10. Cephalopharyngeal skeleton, first-stage larva; 11. Cephalopharyn-
geal skeleton, second-stage larva; 12. Lateral view, puparium; 13. Lateral view, egg.
Abbreviations: asp, anterior spiracle; at, accessory teeth; db, dorsal bridge; dw,
dorsal window; es, epistomal sclerite; hs, hypostomal sclerite; m, mycropyle; psp,
posterior spiracle; va, ventral arch; vw, ventral window.

Description of Immature Stages
Egg (Fig. 13)—Length 0.62-0.71 mm, greatest width 0.21-0.24 mm. White.
Elongate-ovoid, ventral surface more curved than dorsal surface. Micropylar
end slightly truncate (in dorsal view), posterior end rounded. Contents of

egg white, developing.embryo yellowish white. Chorion distinctly retic- |

ulated, with 4- to 7-sided cells. Posterior end without reticulations.
First-stage larva (Figs. 3, 10)—Length 1.10-2.16 mm, greatest width
0.32-0.51 mm. Translucent. Integument slightly papillose, not pubescent.

Metapneustic. Posterior spiracular disc (Fig. 3) at apex of elongate postanal |
if if 8 I it
portion of segment 12; disc with 2 pairs of weak, rounded lobes (ventro- |

VOLUME 80, NUMBER 3 393

lateral lobes slightly larger than ventrals) and 2 spiracular plates, each
with 2 spiracular slits. Postoral spine patch reduced to a few rows of dark
spinules around anterior margin of segment. Dorsal patches on segments
6-11 reduced to few scattered spinules. Venters of segments 6-11 with
a few dark spinules on anterior creeping welt; middle and posterior creeping
welts with patch of dark spinules, 4-6 spinules wide; spinules few or lacking
laterally. Anterior portion of segment 12 ringlike, bearing continuous band
of dark spinules; postanal portion greatly extended dorsally. Perianal pad
transverse, lobes enlarged. Cephalopharyngeal skeleton (Fig. 10) lightly
pigmented, length 0.18-0.23 mm, indentation index approximately 66.
Mouthhooks lightly pigmented, triangular with 2 small openings dorsally;
darkly pigmented bifid hooks on anterodorsal margin; no accessory teeth.
Ventral arch (Fig. 10) curved with 14-16 teeth. Cornua of pharyngeal
sclerite without distinct openings; dorsal cornu ’% longer than ventral cornu.
(Based on 10 larvae.)

Second-stage larva (Figs. 4, 6, 11).—Length 1.87-2.74 mm, greatest
width 0.58-0.74 mm. Unicolorous white. Integument papillose, slightly
pubescent. Anterior spiracles (Fig. 6) rounded, with 20-23 marginal papillae;
papillae close together and not as elongate as those of 3rd instar. Pos-
terior spiracular disc (Fig. 4) bearing 2 spiracular plates each with 3 spirac-
ular slits and 4 branching, hairlike, interspiracular processes. Spinule distri-
bution similar to that of Ist instar. Sparse patch of dark spinules on anterior
creeping welt on venters of segments 6-11 reduced to small patch along
meson. Cephalopharyngeal skeleton (Fig. 11) pigmented, length 0.31-
0.46 mm, indentation index approximately 55. Mouthhooks deeply pig-
mented, each with single, lightly pigmented accessory tooth. Ventral
arch with winged process laterally; anterior margin with 16-18 variously
sized teeth. Ventral cornua slightly longer than dorsal cornua, both with
unpigmented windows posteriorly. (Based on 10 larvae.)

Third-stage larva (Figs. 1, 2, 5, 7, 8, 9)—Length 3.60-5.51 mm, greatest
width 0.77-1.10 mm. Yellowish tan. Integument papillose, appearing pu-
bescent due to dense covering of small, 0.04-0.06 mm spinules. Body
elongate, subcylindrical, tapering anteriorly, somewhat truncate posteriorly.
Segment 1 weakly bilobed apically (in dorsal view), each lobe bearing
lightly sclerotized sensillum (csp) anterodorsally; postoral segment bearing
spine patch posteriorly, with band extending halfway-up on each side of
segment (Fig. 1). Anterior spiracles tan, (Fig. 7) dorsolateral on posterior
margin of segment 2, subcircular with 22-23 marginal papillae; outer lateral
surface of each papilla with small, irregular, clear area. Posterior spiracles
at tip of elongate stigmatic tubes on posterior spiracular disc. Spiracular
disc (Fig. 5) with 2 pairs of weak lobes; ventrolateral lobes somewhat
larger than ventrals; 2 spiracular plates (Fig. 8), each with 3 elongate spirac-
ular slits and dark stigmatic scar; 4 transparent, branched interspiracular

394 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

processes or float hairs on each plate. Segments 3-11 with numerous trans-
parent spinules; segments 1 and 2 without spinules. Each segment (3-11)
divided ventrally and dorsally by secondary integumentary folds to form
anterior, middle and posterior welts. Segment 3 with row of dark spinules
on anterodorsal margin, venter of segment with sparse spinule band an-
teriorly. Segment 4 without dark spinules dorsally; venter of segment with
sparse spinule band, 3-4 spinules wide, on anterior margin. Segments
5-6 with band of a few dark spinules on anterior and middle welts. Seg-
ments 6-11 without distinct dark spinules dorsally or ventrally. Segment
12 with a few pre-anal spinules, located along meson. Perianal pad bi-
lobed, anus longitudinal on mid-line. Cephalopharyngeal skeleton (Fig. 9)
deeply pigmented, length 0.77-0.79 mm, indentation index approximately
51. Mouthhooks (in dorsal view) converging anteriorly, separated pos-
teriorly; 3 lightly pigmented accessory teeth (at) beneath each hook part.
Anterior margin of ventral arch (va) with 22-23 anteriorly directed, variously
sized teeth. Epistomal sclerite (es) fused to parastomal bars of pharyngeal
sclerite. Pharyngeal sclerite with dorsal and ventral cornu separated by
sinus extending slightly more than halfway to anterior edge of sclerite,
ventral cornua with elongate window (vw), dorsal cornua with small, taper-
ing window (dw); narrow parastomal bars arising from anterior margin of
sclerite. (Based on 10 larvae.)

Puparium (Fig. 12)—Length 3.71-4.82 mm, greatest width 1.70-2.12
mm. Dark brown. Elongate, subcylindrical, anterior end flattened dorso-
ventrally, posterior end tapered. Ventral surface slightly more flattened
than dorsal. Integument dull, opaque, with pubescent appearance due
to tanned spinules of 3rd instar. Intersegmental and secondary intraseg-
mental folds distinct. Segment 1 invaginated. Anterior spiracles (asp) tan,
fan-shaped. Postanal portion of segment 12 narrower than pre-anal por-
tion: Perianal pad bare; lobes on posterior spiracular disc reduced; stig-
matic tubes uplifted and divergent; spiracular plates yellow. (Based on
10 puparia.)

Discussion

Information available on the studied species of Antichaeta indicates that
the species segregate primarily by habitat and species of food utilized. In
North America, A. testacea Melander and A. borealis are commonly found
in permanent marshes and are associated with snails that produce eggs
more or less continuously throughout the year. Both of these species of
Antichaeta are multivoltine. The habitats of A. melanosoma Melander are
vernal pools, temporary woodland pools and areas in large marshes
where there is a seasonal drop in water level. In such habitats the host
snail, Aplexa hypnorum (L.), is most active at 1 or 2 times a year. As might
be expected, this species is univoltine (Knutson and Abercrombie, 1977).

VOLUME 8C, NUMBER 3 395

The 4 European species, A. analis (Meigen), A. atriseta (Loew), A. obliviosa
Enderlein and A. brevipennis (Zetterstedt), all seem to be associated with
vernal marshes (Knutson, 1966).

A wide variety of foods is utilized by different Antichaeta species. Ac-
cording to Fisher and Orth (1964), first- and second-stage larvae of A.
testacea can feed on the eggs of several snail species, including those of
Oxyloma, Physa, Succinea, Radix, Stagnicola, Pseudosuccinea and Helisoma.
Third-stage larvae continue to feed on snail eggs but also can kill and eat
juvenile and adult snails. Larvae of A. testacea show the most generalized
feeding habits of all the studied species of Antichaeta. In the laboratory,
first- and second-stage larvae of A. melanosoma and A. borealis feed on
the eggs of several species of snails, but unlike A. testacea, the larvae
failed to continue development on eggs other than those of the host species.
Third-stage larvae of A. melanosoma killed and ate adult snails of the host
species (Knutson and Abercrombie, 1977), but third-stage A. borealis larvae
limited their feeding to Oxyloma eggs and refused juvenile or adult snails.
Under laboratory conditions larvae of A. analis and A. brevipennis fed on
eggs of Lymnaea and Succinea snails, but neither attacked juvenile or adult
snails.

In areas where species overlap in habitat distribution, they are segregated
by differential food utilization. For example, in northeastern Ohio, 3 spe-
cies of Antichaeta, A. borealis, A. fulva Steyskal, and A. melanosoma, occur
in the same marsh habitats at approximately the same time of year (late
April-June). The first species utilizes Oxyloma eggs; the second, Lymnaea
sp. eggs; and the third, eggs of Aplexa and Physa. There may be some
slight habitat segregation (especially with A. melanosoma) among the 3
species, but it does not seem to be as significant an isolating factor as
the differences in food utilization.

Acknowledgments

We are grateful to Dr. William Mason, Agriculture Canada for identifying
the Ichneumonidae. We wish to thank Sandra Evanson and Karen
Humphries for the illustrations. Drs. K. R. Valley, Bureau of Plant Industry,
Harrisburg, Pennsylvania; and L. Knutson, Systematic Entomology Lab-
oratory, IIBII, Fed. Res., Sci. Educ. Admin., USDA, Beltsville, Maryland,
reviewed and improved the manuscript.

Literature Cited

Fisher, T. W., and R. E. Orth. 1964. Biology and immature stages of Antichaeta
testacea Melander (Diptera: Sciomyzidae). Hilgardia 36: 1-29.

Knutson, L. V. 1966. Biology and immature stages of malacophagous flies: Anti-

chaeta analis, A. atriseta, A. brevipennis, and A. obliviosa (Diptera: Sciomyzidae).

Trans. Am. Entomol. Soc. 92(1):67-101.

396 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Knutson, L., and J. Abercrombie. 1977. Biology of Antichaeta melanosoma (Diptera:
Sciomyzidae), with notes on parasitoid Braconidae and Ichneumonidae (Hymenop-
tera). Proc. Entomol. Soc. Wash. 79:111—125.

(WHR) Department of Entomology, Virginia Polytechnic Institute and
State University, Blacksburg, Virginia 24061; and (BAF) Department of
Biological Sciences, Kent State University, Kent, Ohio 44242.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 397-402
MORNING GLORY BEES AND THE IPOMOEA PANDURATA
COMPLEX (HYMENOPTERA: ANTHOPHORIDAE)

Daniel F. Austin

Abstract.—Oligolectic bees and their association with Ipomoea pandurata
and its allied species were studied in North America. Cemolobus ipomoeae
was found to be closely related to Ipomoea pandurata; Melitoma grisella
has been shown to be linked with Ipomoea leptophylla; Ptilothrix sumichrasti
was found with Ipomoea longifolia. The exact relationships between each
bee species and each plant species are different.

According to the literature three bee genera in the Anthophoridae, An-
cyloscelis, Cemolobus and Melitoma, are oligolectic to the genus Ipomoea
(Michener in Muesebeck et al., 1951; Michener, 1954). Two of these, Cemo-
lobus and Melitoma, occur in the eastern United States. Moreover, Robert-
son (1929) suggested that these bees were probably associated with
Ipomoea pandurata (L.) Meyer, one of the few endemic species of
Ipomoea in the eastern United States. My study began in the summer of
1971 largely as an attempt to confirm the existence of close relationships
between these bees and plants. Once oligolecty'! was documented, the fol-
lowing two summers were used for field study of widely separated popula-
tions of I. pandurata. Patterns of insect behavior at the flowers were found
to be essentially constant throughout the range of bees and plants.

Materials and Methods

Field studies were made in Florida, Georgia, South Carolina, North
Carolina, Virginia, Pennsylvania, Ohio, Indiana, West Virginia, Illinois,
Missouri, Kentucky and Tennessee. In addition to the field studies, her-
barium material of Ipomoea has been examined in A, FAU, FLAS, GH,
UNC, US, MO, and USF (abbreviations follow Lanjouw and Stafleu,
1959). The herbarium and field studies were augmented with various floras
to provide accurate distributions (Britton & Brown, 1898; Small, 1903, 1933;
Wooten & Standley, 1915; Deam, 1940; Tidestrom & Kittell, 1941; Fernald,
1950; Kearney & Peebles, 1951, 1961; Jones et al., 1955; Mohlenbrock &
Voight, 1959; Gleason & Cronquist, 1963; Steyermark, 1963; Justice & Bell,
1968; Radford et al., 1968; Conard, 1969; Correll & Johnston, 1970; Wharton
& Barbour, 1971; Brown, 1972; Ellis & Chester, 1973). Insects were deter-
mined with the help of Paul D. Hurd, Jr., and several other members of
the Entomology Department, Smithsonian Institution as well as Mitchell
(1960, 1962).

398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ipomoea pandurata, Intrafloral Ecology

Robertson (1891, 1925a, 1925b, 1928, 1929) reported a series of insect
visitors to I. pandurata. Of these only two show oligolecty to these plants:
Cemolobus ipomoeae Robertson and Melitoma taurea (Say).

Oligolectic bees——While few studies have been made of the life history
of the monotypic Cemolobus (Robertson, 1891, 1925a, 1925b), all past and
present data indicate that it utilizes pollen only from I. pandurata.

Cemolobus is a matinal species, although cool, foggy, or rainy mornings
may cause the bees to shift their activities later in the day. Under normal
weather conditions the first bees arrive at I. pandurata flowers between 6:30
and 7:00 AM. For about the first two hours the majority of individuals visit-
ing the flowers are females. Males arrive later and may remain active until
12:00 or 1:00 PM.

Females enter the flowers head-first to sip nectar; pollen is collected
with the legs during this time. After nectar is taken they may either back
out of the tube onto the inner edge of the limb to collect pollen with the
front legs, or they may turn around within the tube facing out and collect
pollen. Usually there is a shift from backing out early in the day to turn-
ing around within the tube later. On a few occasions these females
were seen “drumming” with the wings while sitting on the inner edge of
the limb and collecting pollen with the front legs. Pollen is transferred to
the scopae of the posterior legs either while on the flower limb or in
the air. These normally brown females are conspicuously white with
pollen as they leave the plants to return to their nests.

The scopae of each female are capable of carrying large amounts of
pollen and this requires the bees to visit several flowers before they are
filled. Flights from flower to flower are rapid and direct. Although no
male bees collect pollen it is common for them to visit several flowers and
then to perch on the corolla limb or nearby plants to clean themselves
of pollen.

When the flowers open the stigmas have no pollen on them, but grains
are left on the stigma by each visit. Bagged or emasculated flowers or
solitary plants on the fringe of the range that are not pollinated produce
no fruits, while pollinated flowers fruit abundantly.

Melitoma taurea is a member of a small American genus of bees that
is known to utilize only Ipomoea pollen in their nests (Michener in
Muesebeck et al., 1951; Michener, 1954, 1975). The species M. taurea is
not specific to a single species of Ipomoea in the eastern United States,
but has been collected at the flowers of I. sagittata Lam. and I. aquatica
Forsk. (Austin, unpublished), I. purpurea (L.) Roth (Robertson, 1929) and
Calystegia sepium (L.) R. Brown (Robertson, 1891). Robertson indicated
that the bees were only taking nectar from Calystegia and not collecting
pollen.

VOLUME 80, NUMBER 3 399

Melitoma taurea is also a matinal bee, and females often appear at the
flowers of I. pandurata before Cemolobus. As with Cemolobus females,
visits to the plants begin between 6:30 and 7:00 AM. Melitoma enters the
floral tube head-first to sip nectar. While doing this they actively brush
pollen from the anthers with their legs. Unlike Cemolobus, they almost in-
variably turn around within the corolla tube to face outward. Once they
have turned around they collect more pollen and transfer it to their scopae.
When the scopae are full, the females leave the plants; they later return
with empty scopae.

Melitoma uses the flowers of I. pandurata as trysting sites. Once the
females have collected pollen for about two hours, the number of males
visiting the plants increases markedly. Males enter flowers much less
often than do females. The most common behavior is for the males to
rapidly fly from flower to flower, pausing on the limb only long enough
to determine if a female is inside the tube. If no female is present, the
males rapidly visit other blossoms. Once a flower is found containing
a female, the male enters and copulation occurs. About one flower visit in
ten is used by the males to take nectar.

Other insects.—A series of insects has been recorded by Robertson (1891,
1925a, 1925b, 1928, 1929) as visitors to I. pandurata; my own collections
include most of these. Several of these visitors are oligolectic bees with
close pollen relationships to other plants, e.g. Peponapis and Xenoglossa to
Curcurbita (Hurd et al., 1971). Most common among the other insects
visiting I. pandurata is Bombus.

Bombus auricomus (Robertson), B. griseocollis (Degeer), B. vagans F.
Smith and B. pennsylvanicus (Degeer) (Robertson, op. cit.) visit throughout
the range of the plants. I found B. pennsylvanicus to be the predominate
species. These bees are often found taking nectar when no other insects
are active, particularly on foggy or drizzly mornings. Bombus spp. transfer
pollen frequently although they have never been seen storing it in their
scopae. They do often stop to clean pollen from their body and leg hairs.
Members of this genus are the only visitors seen visiting the flowers on
the extreme northern limits of J. pandurata range.

Cemolobus, Melitoma and Bombus are considered the primary polli-
nators of I. pandurata because they transfer pollen to the stigmas with each
flower visit, and usually have two hours to visit flowers and transfer pollen
before other visitors appear. Other visitors serve as occasional pollinators.

Ipomoea pandurata, Relatives and Other Bees

The closest relative of I. pandurata in the United States is I. leptophylla
Torrey. Ranges of these two plants overlap in eastern Oklahoma and north-
eastern Texas. In the area of overlap is a rare population, I. shumardiana
(Torrey) Shinners. Robert Pearce has proposed that this population is the

400 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

result of hybridization between I. pandurata and I. leptophylla (Shinners
in Correll and Johnston, 1970).

Ipomoea leptophylla is a prairie species known to range from south-
eastern Montana south into Texas. This plant was studied by Linsley
(1960) near Los Montoyas, New Mexico. Linsley found Melitoma grisella
(Cockerell and Porter), a matinal pollen collector, to be the major visitor
and pollinator of this species. Three other genera of bees visit I. leptophylla,
but their visits were for nectar only and no pollen was collected. Melitoma
grisella is the only Ipomoea oligolege found in the northern part of the range
of the plants.

In the small area of range overlap of I. pandurata and I. leptophylla in
Oklahoma and Texas, there is also an overlap of Melitoma taurea and
M. grisella. If hybridization is the origin for the plants named I. shumar-
diana, either of these bees or Bombus could be responsible for the crossing.

Ipomoea longifolia Bentham is known from two border counties in Ari-
zona south to the state of Queretaro, Mexico. Linsley et al. (1956), Butler
(1967) and Cazier and Linsley (1974) studied this species and found two
oligolectic bees associated with the plants. Most closely allied with I.
longifolia is the bee Ptilothrix sumichrasti (Cresson); also gathering pollen
was Melitoma segmentaria (Fabricius) (= Melitoma euglossoides Lepeltier
and Serville). Exhaustive study of the biology of this Ptilothrix indicates
that the species is oligolectic to I. longifolia as long as the plant is flowering
(Linsley et al., 1956; Linsley and MacSwain, 1958; Michener, 1974; Torchio,
1974). The Ptilothrix is not as species-specific as Cemolobus but takes
pollen from other Ipomoea species when I. longifolia has finished its bloom-
ing season. Ipomoea pringlei Gray is the species noted to be visited by
both Ptilothrix and Melitoma as the flowering season of I. longifolia ends.

Summary.—The data show that two species of bees have specific re-
quirements for Ipomoea pollen in the eastern United States. Cemolobus
ipomoeae is specific in pollen collecting habits and uses only I. pandurata
pollen. Both the plants and the bees are endemic to the deciduous forest
areas of the eastern United States. Melitoma taurea is found in the same
region, but is less specific in pollen collecting as it takes pollen from
several species of Ipomoea.

Related to I. pandurata is the prairie species I. leptophylla. Present
data indicate that this plant is closely linked with Melitoma grisella. At
present it is not possible to determine whether I. shumardiana is a hybrid
between I. pandurata and I. leptophylla.

The closest relative to I. leptophylla and I. pandurata is the Arizonan |
and Mexican species I. longifolia. Ptilothrix sumichrasti is oligolectic to
I. longifolia, however, the relationship is somewhat different from the other |
United States species. As long as I. longifolia is in flower, this Ptilothrix
uses pollen only from that species. When this Ipomoea stops flowering,
the bees shift to another species.

VOLUME 80, NUMBER 3 401

Acknowledgments

My sincere appreciation goes to Paul D. Hurd, Jr. (Smithsonian Institu-
tion) who turned my attention in this direction during the 1971 Summer
Institute in Systematics. Curators of the herbaria mentioned were gracious
in allowing me to examine their collections and in answering my ques-
tions. C. D. Michener (University of Kansas) checked my state distribution
map for Cemolobus and Melitoma. E. G. Linsley (University of Calli-
fornia) and P. D. Hurd, Jr., offered several useful comments on the original
manuscript.

Literature Cited

Baker, H., and P. D. Hurd, Jr. 1968. Intrafloral ecology. Annu. Rev. Entomol.
13:385-414.

Britton, N. L., and A. Brown. 1898. An illustrated flora of the northern United States,
Canada and the British possessions. Vol. III. Scribner's Sons, New York.
Brown, C. A. 1972. Wildflowers of Louisiana and adjoining states. Louisiana State

Univ. Press, Baton Rouge.

Butler, G. D., Jr. 1967. Biological observations on Ptilothrix sumichrasti (Cresson)
in southern Arizona. Pan-Pac. Entomol. 43:8—14.

Cazier, M. A., and E. G. Linsley. 1974. Foraging behavior of some bees and wasps
at Kallstroemia grandiflora flowers in southern Arizona and New Mexico. Am.
Mus. Novit. 2546:1-20.

Conard, H. S. 1969. Plants of central Florida. Ridge Audubon Soc., Lake Wales.

Correll, D. M., and M. C. Johnston. 1970. Manual of the vascular flora of Texas.
Texas Research Foundation, Renner.

Deam, C. C. 1940. Flora of Indiana. Indianapolis.

Ellis, W. H., and E. W. Chester. 1973. Summer and fall wildflowers of land be-
tween the lakes. Austin Peay State Univ., Clarksville.

Faegri, K., and L. van der Pijl. 1966. The principles of pollination ecology.
Pergamon Press, Oxford.

Fernald, M. L. 1950. Gray’s manual of botany, ed. 8. Boston.

Gleason, H. A., and A. Cronquist. 1963. Manual of the vascular plants of north-

eastern United States and adjacent Canada. D. van Nostrand, Princeton.

Hurd, P. D. Jr., E. G. Linsley, and T. W. Whitaker. 1971. Squash and gourd bees
(Peponapis, Xenoglossa) and the origin of the cultivated Cucurbita. Evolution
25:218-234.

Jones, G. N., G. D. Fuller, H. E. Ahles, G. S. Winterringer, and A. Flynn. 1955.
Vascular plants of Illinois. Mus. Sci. Ser. Vol. VI., Univ. Illinois Press, Urbana.

Justice, W. S., and C. R. Bell. 1968. Wildflowers of North Carolina. Univ. North
Carolina Press, Chapel Hill.

Kearney, T. H., and R. H. Peebles. 1951. Arizona flora. Univ. California Press,

Berkeley.

1961. Arizona flora, ed. 2. Univ. California Press, Berkeley.

Lanjouw, J., and F. A. Stafleu. 1959. Index herbariorum. Regnum Veg. 15:1-249.

Linsley, E. G. 1960. Observations on some matinal bees at flowers of Cucurbita,
Ipomoea and Datura in desert areas of New Mexico and southeastern Arizona.

| J. N.Y. Entomol. Soc. 68: 13-20.

_ Linsley, E. G., and J. W. MacSwain. 1958. The significance of floral constancy

among bees of the genus Diadasia (Hymenoptera, Anthophoridae). Evolution

12:219-223.

402 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Linsley, E. G., J. W. MacSwain, and R. F. Smith. 1956. Biological observations on
Ptilothrix sumichrasti (Cresson) and related groups of Emphorine bees. Bull.
South. Calif. Acad. Sci. 55:83-101.

Michener, C. B. 1954. Bees of Panama. Bull. Am. Mus. Nat. Hist. 104:1—-175.

1974. Further notes on nests of Ancyloscelis. J. Kans. Entomol. Soc. 47:

19-22.

——. 1975. Nests of Paranthidium jugatorium in association with Melitoma taurea
(Hymenoptera: Megachilidae and Anthophoridae). J. Kans. Entomol. Soc. 48:
194-200.

Mitchell, T. B. 1960. Bees of the eastern United States. Vol. I. N.C. Agric.
Exp. Stn. Tech. Bull. 141:1-538.

—. 1962. Bees of the eastern United States. Vol. II. N.C. Agric. Exp. Stn.
Tech. Bull. 152:1-557.

Mohlenbrock, R. H., and J. W. Voight. 1959. <A flora of southern Illinois. S.
Illinois Univ. Press, Carbondale.

Muesebeck, C. F. W., K. V. Krombein, and H. K. Townes. 1951. Hymenoptera of
America north of Mexico. Synoptic Catalog. USDA Agric. Monogr. No. 2.

Radford, A. E., H. E. Ahles, and C. R. Bell. 1968. Manual of the vascular flora
of the Carolinas. Univ. North Carolina Press, Chapel Hill.

Robertson, C. 1891. Flowers and insects, Asclepiadaceae to Scrophulariaceae. Trans.
St. Louis Acad. Sci. 5:569-598.

—. 1925a. Heterotropic bees. Ecology 6:412—436.

———. 1925b. Habits of the Hibiscus bee, Emphor bombiformis. Psyche 32:278—

282.
1928. Flowers and insects. Published privately, Carlinville, Illinois.
1929. Phenology of oligolectic bees and favorite flowers. Psyche 36:112-—

118.

Small, J. K. 1903. Flora of the southeastern United States. Published by the author,
New York.

—. 1933. Manual of the southeastern flora. Univ. North Carolina Press, Chapel
Hill.

Steyermark, J. A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames.

Tidestrom, I., and Sister T. Kittell. 1941. Flora of Arizona and New Mexico. Catholic
Univ. of America Press.

Torchio, P. F. 1974. Notes on the biology of Ancyloscelis armata Smith and com-
parisons with other Anthophorine bees (Hymenoptera: Anthophoridae). J.
Kans. Entomol. Soc. 47:54-63.

Wharton, M. E., and R. W. Barbour. 1971. <A guide to the wildflowers and ferns
of Kentucky. Univ. Press of Kentucky, Lexington.

Wooten, E. O., and P. C. Standley. 1915. Flora of New Mexico. Contrib. U.S. Nat.
Herb. 19:514—-519.

Department of Biological Sciences, Florida Atlantic University, Boca
Raton, Florida 33431.

Footnote

1 Oligolectic, as used here, refers to bees whose females utilize pollen from one or
a few species of plants as food for their young. This usage conforms to that by
Robertson (1929), Michener (1954), Linsley & MacSwain (1958), Faegri and van der
Pijl (1966) and Baker and Hurd (1968) among others. Some biologists object to such
usage since it ignores adult male activities.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 403-406
A NEW STONEFLY FROM WEST VIRGINIA
(PLECOPTERA: CHLOROPERLIDAE)

P. P. Harper and R. F. Kirchner

Abstract.—Alloperla aracoma is described from the Guyandotte River
system in West Virginia; it is closely related to Alloperla leonarda Ricker.
Both species are illustrated.

During an investigation of the stonefly fauna of West Virginia, the
junior author collected a series of specimens which at first appeared to be
a form of Alloperla leonarda Ricker 1952; on closer examination of the male
genitalia, however, it became apparent that the series represented a new
species; its description follows.

Alloperla aracoma Harper and Kirchner, new species

Figs. 1-3

Body length.—Six (4) to 7.5 (2) mm; wing length: 6.5-8 mm. General
habitus similar to other Alloperla; color uniformly pale green (yellowish
white in alcohol-preserved specimens), wings with a light greenish tinge;
an obscure series of brownish marks on the mid-dorsum of abdominal terga
2-7, not forming a definite mid-dorsal stripe.

Male genitalia —Genitalia resemble those of other Alloperla. Abdominal
segments 8 and 9 with rows of dark spines on their lateral margins. Tergum
10 cleft with a median depression into which fits the epiproct. Epiproct
fixed to the anterior border of tergum 10 by an anchor-like structure; it
is prolonged posteriorly as a thin rod and produced as a forward directed
process (Fig. 1); the process (Figs. 2 and 3) gradually expanded distally and
its apex truncate with 2 lateral knobs; process covered dorsally and _ lat-
erally with setae except at the tip and it bears on its ventral surface 2 rows
of a dozen or so thorn-like spines; length of the process of the epiproct at
least 7 that of basal rod.

Female genitalia—Hind margin of the 8th abdominal sternum produced
into triangular subgenital plate which is somewhat thickened in its mid-
dle; the plate is very similar to that described and illustrated by Baumann
(1974) for Alloperla imbecilla (Say).

Holotype.—é, West Virginia, Logan County, Holden, Frogtown Hollow
of the Copperas Mine Fork of Island Creek in the Guyandotte River system;
_7 May 1975; R. F. Kirchner. Allotype 2°, same data. Paratypes: 154, 42, 7
May 1975; 92, 152, 12 May 1975; 8¢, 12, 2 May 1976; 9¢, 152, 8-15 May
1976; 8¢, 102, 8 May 1977, all from Frogtown Hollow. One ¢, 22, West
Virginia, Mingo County, Laurel Fork of Pigeon Creek; 30 May 1976; R. F.
Kirchner and T. Mayberry Jr.

404 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-3. Male genitalia of Alloperla aracoma. 1, dorsal view of terminal ab-

9

dominal segments. 2, epiproct, side view. 3, process of epiproct, dorsal and ventral
aspects.

The holotype and the allotype will be deposited in the U.S. National
Museum. Paratypes will be in the USNM, the Entomological Collection
of the Université de Montréal and in the authors’ collections.

Derivation of the name.—From Princess Aracoma, daughter. of Chief
Cornstalk of the Shawnee Indians, who had a village on Middleburg
Island on the Guyandotte River.

T ype-locality Frogtown Hollow is an intermittent stream, 3-4 feet in
width, at an elevation of 800-1,200 feet above sea level. Riparian vegeta-
tion includes yellow-poplar, American beech, sugar and red maples, black
birch, yellow buckeye, hemlock, cucumber and umbrella magnolias, white
ash, red oak, shagbark hickory, spicebush, wild hydrangea, witch hazel and
hornbean. Associate Chloroperlidae are Hastaperla brevis (Banks), Alloperla
imbecilla (Say), Alloperla usa Ricker and Sweltsa mediana (Needham &
Claassen).

Diagnosis.—The male of Alloperla aracoma bears much superficial re-
semblance to A. leonarda (see Ricker, 1952, Fig. 132 and Harden and
Mickel, 1952, Pl. X, Figs. 7-8 as A. sylvia). The distinguishing features
are the following: In A. aracoma the process of the epiproct is long, about
%s as long as the basal rod (Fig. 1); in A. leonarda the process is less than
% as long as the rod (Fig. 5). Though both species bear lateral knobs
on the tip of their epiproct (Figs. 3 and 4), the shapes of the processes are
markedly different, and only A. aracoma possesses the rows of ventral
spines (Figs. 2 and 3). Furthermore only A. leonarda has large fleshy lobes

VOLUME 80, NUMBER 3 405

Figs. 4-5. Male genitalia of Alloperla leonarda. 4, process of epiproct, dorsal
aspect. 5, epiproct, side view.

adjacent to the lobes of the epiproct (Fig. 4). In Hitchcock’s (1974) key to
the northeastern species of Alloperla (s. 1.), the female of A. aracoma will
key out to A. imbecilla from which it can be distinguished at present only

by its smaller size; A. atlantica Baumann, a species long confused with A.
-imbecilla, will also key there and again the smaller size of A. aracoma will
_be the main diagnostic feature.

Literature Cited

Baumann, R. W. 1974. What is Alloperla imbecilla (Say)? Designation of a neo-
type, and a new Alloperla from Eastern North America (Plecoptera: Chloroperli-
dae). Proc. Biol. Soc. Wash. 87:257-264.

406 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Harden, P. H., and C. E. Mickel. 1952. The stoneflies of Minnesota (Plecoptera).
Univ. Minne. Agr. Exp. Sta., Techn. Bull. 201:1-84.

Hitchcock, S. W. 1974. Guide to the insects of Connecticut. VII. The Plecoptera
or stoneflies of Connecticut. St. Geol. Nat. Hist. Surv. Bull. 107:1-262.

Ricker, W. E. 1952. Systematic studies in Plecoptera. Ind. Univ. Publ., Sci. Ser.
18: 1-250.

(PPH) Département de Sciences biologiques, Université de Montréal,
C.P. 6128, Succursale “A,” Montréal, Québec H38C 3J7, Canada; and
(RFK) Rt. 1, Box 412-A, Barboursville, West Virginia 25504.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 407-423
TAXONOMY AND DESCRIPTION OF TWO PRIONINE
CERAMBYCIDAE FROM SOUTHERN ARIZONA:
A NEW SPECIES OF STENODONTES AND NEW STATUS
FOR NEOMALLODON ARIZONICUS (COLEOPTERA)

Durward D. Skiles

Abstract.—Recent studies of the prionine Cerambycidae of southern
Arizona have revealed two new forms, a new species, Stenodontes (Mal-
lodon) madericus, and the previously unrecognized male of Stenodontes
(Neomallodon) arizonicus (Casey). Both species inhabit oak woodlands
from central Arizona to northwestern Mexico and the larvae infest the
heartwood of living evergreen oak trees. Discovery of these new forms
necessitates revisions of Linsley’s (1962) keys to the North American
genera of the prionine tribe Macrotomini and the North American species
of the genus Stenodontes. Clearly generically distinct from other Mexican
and North American Macrotomini, S. arizonicus is redescribed and the sub-
genus Neomallodon Linsley is elevated to generic status.

Recent studies of the southern Arizona prionine Cerambycidae presently
assigned to the genus Stenodontes (Linsley, 1962) have revealed two pre-
viously unrecognized forms, a new species described below as Stenodontes
(Mallodon) madericus, and the heretofore unknown male of Stenodontes
(Neomallodon) arizonicus (Casey).

Although infrequently collected, S$. madericus ranges from north central
Arizona to northwestern Mexico and can be found among other species of
Macrotomini in almost any large collection of southern Arizona Cerambyci-
dae. It is indeed remarkable that this distinctive prionine has remained
undescribed for so long, but the small number of specimens collected
prior to the last few years and the similarity of habitus of the females
and minor males of the western species of Stenodontes are undoubtedly
responsible. As a case in point, the author identified his first specimens
of S. madericus as females of S. lobigenis (Bates), and it was not until
the holotype male was collected that it was clear a distinct species was
at hand. Realization that the species was undescribed was further delayed
by the fact that the strong, basally retracted mandibles suggested that the
specimen was a male of S. arizonicus.

The latter species, originally described over sixty years ago (Casey,
1912), remains rare in collections despite the fact that collecting has per-
sisted within its range for almost a century. Less than ten specimens
of S. arizonicus were known to Linsley (1962) when he revised Stenodontes.
The lack of material, the short mandibles, and the rather similar appearance

408 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of the sexes led Linsley to include only the female in his revision, although
Linsley et al. (1961) refer to a male taken in Cave Creek Canyon in the
Chiricahua Mountains of southeastern Arizona. The latter specimen, how-
ever, was known only to Knull (Linsley, personal communication).

Intensive collecting in southern Arizona by several individuals in re-
cent years has enabled the author to examine a substantial series of S. ari-
zonicus and, thereby, to distinguish the male and female and to significantly
extend the known range of the species. A revised description of the species is
presented below, and on the basis of the rather subdued sexual dimorphism
and other obvious structural differences from all closely related taxa, it is
proposed that the monotypic subgenus Neomallodon Linsley be elevated to
generic status.

Description of these new forms necessitates revisions of Linsley’s (1962)
keys to the North American genera of the prionine tribe Macrotomini
and the North American species of the genus Stenodontes. To ensure
proper separation of the taxa and to remove the confusion of species al-
most invariably encountered in collections, the keys and descriptions are
rather detailed. Because all but one species (S. chevrolati Gahan) of the
North American Macrotomini occur in Mexico', and since Linsley (1934)
originally described Neomallodon arizonicus (Casey) as Aplagiognathus
remotus Linsley, the Mexican genus Aplagiognathus is included in the
key to the Macrotomini.

Key to the North American Genera of Macrotomini

L Antenna with 3rd segment shorter than scape; scutellum con-
cave 2

- Antenna with 3rd segment longer than scape; scutellum very
convex Strongylaspis |

2(1). Antennal tubercle obtuse; mandible nearly vertical, almost gla-
brous, neither carinate nor tuberculate above, prominently uni-
dentate internally at about middle, not excavated internally;
lateral margin of pronotum crenulate or denticulate, never spinose

Archodontes
= Antennal tubercle obtuse or acute; mandible only moderately de-
flexed or nearly horizontal, basal % in profile tumid or broadly
triangular above, or more or less carinate above and excavated
internally, usually bi- or tridentate internally; lateral margin

of pronotum denticulate to prominently spinose 3

3(2). Sexual dimorphism evident but rather subdued; mandible shorter
than head, either robust or extremely short, dorsal and/or outer
margin strongly retracted at base (except occasional females),
always possessing numerous long setae on external margin; sub-
mentum unsculptured, broadly, shallowly, transversely impressed,

VOLUME 80, NUMBER 3 409

usually finely to moderately, transversely rugose, occasionally
polished and shallowly punctate; metepisternum broad 4
- Sexual dimorphism pronounced; mandible usually neither robust
nor strongly retracted at base, in males as long or longer than
head (except some minor males); if mandible strongly retracted at
base, metepisternum narrow; submentum not as above, coarsely
granulate-punctate and/or coarsely, usually longitudinally, ru-
gose, posterior and anterior regions distinctly contrasting, an-
terior region usually recessed, occasionally only more finely
punctate; metepisternum broad or narrow Stenodontes
4(3). Mandible in profile evenly arcuate dorsally, rather carinate above,
distinctly excavated internally, a stout, broadly emarginate tooth
before apex; antennal tubercle acute or scarcely obtuse; pro-
notum with lateral region obsoletely punctate, lateral margin
denticulate, distinctly reflexed; metepisternum broad, inner mar-
gin straight or feebly convex Neomallodon
- Mandible in profile with basal % tumid or subtriangular dorsally,
not or scarcely excavated internally, a strong, broadly rounded
tooth before apex; antennal tubercle obtuse; pronotum with
lateral region densely granulate-punctate, lateral margin spinose,
not reflexed; metepisternum very broad, inner margin dis-
tinctly convex Aplagiognathus

Genus Neomallodon Linsley, NEW STATUS
Stenodontes (Neomallodon) Linsley, 1957:2; Linsley, 1962:18.

Sexual dimorphism not pronounced. Body large, robust, rather depressed.
Head well developed; antenna robust, attaining basal % of elytron, scape
longer than 3rd segment, 3rd segment robust, subequal to 4th; antennal tu-
bercle acute or scarcely obtuse; mandible deflexed, robust, distinctly shorter

than head, in profile rather evenly arcuate above, excavated internally, a
stout, broadly emarginate tooth at apical %, outer margin strongly retracted
-at base; submentum shallowly, transversely impressed. Pronotum very trans-
verse, disk polished, almost impunctate, lateral margins reflexed, parallel
in male, converging anteriorly in female; metepisternum broad, inner mar-
}gin straight or feebly convex; elytral apices rounded, sutural angle
‘rounded or subangulate.

Type-species.—Paramallus arizonicus Casey (by original designation).
Range.—Southeastern Arizona, and presumably northwestern Mexico.

|

Neomallodon arizonicus (Casey), NEW STATUS
rigs lar?

Paramallus arizonicus Casey, 1912:228.

410 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Neomallodon arizonicus (Casey). Left, male; right, female. To afford a
full view of the mandibles, the heads are only slightly deflexed.

Aplagiognathus remotus Linsley, 1934:161.

Stenodontes (Neomallodon) arizonicus (Casey), Linsley, 1957:3 (synonymy
of P. arizonicus and A. remotus); Linsley, 1962:19; Linsley et al., 1961:3
(record).

Male.—Form elongate-robust, sides subparallel. Integument shining,
glabrous, medium to dark red brown. Head deflexed, bisected by a longi-
tudinal groove, fine basally, becoming broader and deeper on frons,
abruptly bifurcating to form clypeal suture; dorsum coarsely, moderately
to densely punctate, punctures usually less dense in broad median region
extending from occiput to frons; antennal tubercle acute or scarcely
obtuse; antenna attaining basal %

’s of elytron, rather sparsely punctate,
flagellar segments carinulate, basal segments distinctly broader than apical
segments, 2nd segment transverse, 3rd 7 as long as scape, subequal to

VOLUME 80, NUMBER 3 411

4th and 5th, thence to 10th scarcely increasing in length, 11th distinctly
longer than 10th, shorter than scape; mandible deflexed about 45°, tumid,
very robust, distinctly shorter than head, above broadly carinate and rather
evenly arcuate in profile, excavated internally with a stout, broadly
emarginate tooth at apical % and a strong tooth at base, outer margin
arcuate, strongly retracted at base, shallowly, sparsely punctate, a re-
curved seta arising from each puncture, inner margin more densely
punctate, pubescence erect; submentum shallowly, transversely impressed,
rather finely, transversely rugose; genal process broadly emarginate, sub-
angulate below, rounded above. Pronotum shining, twice as broad as long,
narrower than base of elytra, anterior margin broadly sinuate or feebly
trisinuate, basal margin bisinuate, lateral margins subparallel, denticulate,
distinctly reflexed; disk polished, impunctate or very finely, sparsely punc-
tate, lateral regions vaguely rugose, punctures, large, shallow, obsolete ex-
cept near disk; prosternum feebly impressed, very finely, sparsely punc-
tate in front of coxae, finely to moderately rugose anteriorly, pubescence
sparse, erect; metepisternum rather broad, finely punctate, inner margin
straight or feebly convex; mesosternum, metasternum, and metepisternum
covered with long, dense, erect, golden pubescence; scutellum broadly
rounded, glabrous, nearly impunctate. Elytron 4.4-4.6 times as long as
basal width, widest at or behind middle; surface shining, finely, sparsely
punctate basally, more densely and vaguely so apically; apices separately,
broadly rounded, suture rounded or subangulate. Legs short; femora
shining, impunctate, with few scattered setae; tibiae sparsely pubescent,
2 thin lines of dense, golden pile on lower margins, punctures sparse, large,
vague. Abdomen shining, glabrous beneath, thinly pubescent laterally,
densely pubescent on lateral margins, 5th sternum densely fringed with
long, golden setae, broadly subtruncate, feebly emarginate, usually very
transverse and shorter than 4th. Length, apex of elytron to base of
mandible, 40-47 mm.

Female.—Head distinctly smaller than in male. Mandible shorter and less
robust than in male, moderately retracted at base, internal teeth less pro-
nounced, preapical tooth truncate or feebly emarginate. Lateral margins of
pronotum converging anteriorly. Elytron 4.5-4.7 times as long as basal
width. Fifth abdominal sternum less transverse than in male, apex rounded
or subtruncate, often feebly emarginate at middle. Length, 37-47 mm.
| Type-locality—Of arizonicus, Arizona; of remotus, Mt. Washington, al-
titude 6,000 feet, Santa Cruz Co., near Nogales, Arizona.

Known range.—Oak woodlands of southeastern Arizona (Fig. 2).
Flight period—Late June to late July.

Host.—Quercus.

Habits.—The habits of Neomallodon arizonicus remain poorly known, and

412 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

specimens, particularly males, are rare in collections. Adults are oc-
casionally attracted to light during the early hours of darkness, but their
flight is also crepuscular, for I have taken a male from the porch of the
Santa Rita Lodge in Madera Canyon, Arizona during the evening twilight.
Although observations are not sufficient to permit a detailed comparison,
the larval habits and host preferences of N. arizonicus are apparently
rather similar to those of its sympatric relative Stenodontes madericus, as
larvae of each species have been found working in the heartwood of a
living evergreen oak tree.

The only host record for Neomallodon arizonicus has been provided by
C. E. Langston, who reared several adults (only one of which I have seen)
from oak (either Quercus emoryi Torrey, Q. oblongifolia Torrey or Q. ari-
zonicus Sargent) collected at Texas Pass in the Dragoon Mountains, Co-
chise County, Arizona. The larvae were found mining the heartwood of a
large, living limb which had been seriously weakened by their extensive
work. Living wood, however, is apparently not essential to larval develop-
ment, since adults emerged from the limb over a period of two or three
years. Moreover, in the trunks and larger limbs of dead oak trees in
southeastern Arizona one occasionally finds extremely large emergence
holes which are probably those of Neomallodon arizonicus. The holes
are much larger than those characteristic of Stenodontes madericus, and
are not readily attributed to Prionus californicus Motschulsky or P. heroicus
Semenov, the only other sufficiently large cerambycids which are known
or presumed to infest oak trunks in southeastern Arizona. Like many other
members of the genus Prionus, P. californicus feeds primarily on the
roots of living trees and pupates in the soil, and only occasionally feeds
and pupates in dead logs or stumps (Linsley, 1962). The larval habits of
P. heroicus have not been determined, but they are probably similar to
those of P. californicus, since females have been found in leaf litter at the
bases of both living oak trees (D. Skiles) and oak stumps (Hovore and
Giesbert, 1976).

Diagnosis.—This species has clear affinities with both Archodontes and
Aplagiognathus. Casey (1912) erected the genus Paramallus to include
both his new species P. arizonicus and what is now known (Linsley, 1962) as
Archodontes melanopus (.). The distinctly deflexed head, rather stout,
externally arcuate mandibles, plane, unsculptured submentum, broad met-
episternum, and general facies of the pronotum indeed give the females of
Neomallodon a very archodontine appearance, and I have often found the
two species confused in collections. However, as Linsley (1957, 1962)
recognized, Neomallodon is readily generically separated from Archodontes
by the rather flattened body, acute antennal tubercles and basally retracted,
internally excavated, bidentate mandibles.

Linsley (1934) originally placed his species remotus in Aplagiognathus,

VOLUME 80, NUMBER 3 413

but upon examining the type of Paramallus arizonicus Casey, concluded
that the two species were identical and placed them in synonymy in a
new stenodontine subgenus, Neomallodon (Linsley, 1957). Unfortunately,
Linsley was unable to examine the types side by side (Linsley, personal
communication), and the rather subdued sexual dimorphism—the sexual
dimorphism apparent in Fig. 1 is extreme for the species, the female being
minor, the male very major—coupled with the fact that the Casey type
is a large, robust, but obviously female, specimen, led him to conclude that
the type and paratype of Aplagiognathus remotus were also female. The
acute antennal tubercles and internally excavated and dorsally carinate
mandibles then seemed to place the species in Stenodontes.

I have examined the genitalia of both the Casey type and the Linsley
type and paratype. The former is female, and the latter, as originally
stated by Linsley (1934), are male. The stout, basally retracted mandibles,
unsculptured submentum and minimal sexual dimorphism indicate that
Neomallodon is quite distinct from Stenodontes and more closely related
to Aplagiognathus. From the latter, however, Neomallodon is distin-
guished by the dorsally arcuate, internally excavated mandibles, the acute
antennal tubercles, and the distinctive structure of the pronotum.

Material examined.—Arizona. Holotype 2 of Paramallus arizonicus Casey
(USNM type 36405. No collection data with specimen. Casey (1912) gives
the locality and collector as “Arizona-Levette.”); holotype ¢ (CAS type
3822) and paratype ¢ (CAS) of Aplagiognathus remotus Linsley, Mt.
Washington, near Nogales, Santa Cruz Co., alt. 6,000 ft., VII-20-1919
(J. A. Kusche). Santa Cruz Co.: Sycamore Canyon, 12 mi SE Ruby, 12,
VII-22-71 (D. G. Marqua); Nogales, 12, VII-10-57 (Stange and Hard-
ing); Madera Canyon, 1é (flying at evening twilight), VI-28-VII-4-73,
12 (at light), VII-15-75 (D. D. Skiles); 12, VII-3-66, 12, VII-23-24-71 (F.
T. Hovore, at light); 12, VII-10-70, 12, VII-12-73 (E. F. Giesbert, at light);
16, VII-11-57, 24 8, VII-12-57 (Stange and Harding); 12, VII-19-25-57
(R. L. Westcott); 12, VII-22-65, 12°, VII-8-71, 12, VII-19-71 (D. G. Marqua);
12, VII-16-72 (A. E. Lewis). Pima Co.: Madera Canyon, 1°, VII-11-70
(E. F. Giesbert, drowned in creek); Molino Basin, NE Tucson, 12,
VII-10-73 (D. G. Marqua, at light). Gila Co.: Sierra Ancha Mts., 14
(D. K. Duncan, labeled Archodontes cilipes Say). Cochise Co.: South-
_western Research Station, 5 mi SW Portal, 12, VII-20-71 (E. F. Giesbert, at
light); 3.5 mi SW Portal, 12, VII-11-66 (R. G. Beard); Miller Canyon,
Huachuca Mts., 12, VII-14-69 (A. E. Lewis); Texas Pass, 1¢, VI-1-73
(C. E. Langston, reared ex Quercus); Cochise Stronghold, Dragoon Mts.,
12, VII-29-57 (C. W. O’Brien); 12, VII-18-77 (D. G. Marqua, at light).
Santa Rita Mts., 12, VH-11-50 (J. G. Rozen). Additional Arizona localities
known to me are: Cochise Co.: Huachuca Mts. and Miller Canyon,
Huachuca Mts. (Linsley, 1962:19).

414

Key to North American Species and Subgenera of Stenodontes

Antennae not or scarcely surpassing middle of elytra in male, not
attaining middle in female; mandible of male at most but little
longer than head, distinctly pilose internally

Antennae attaining at least apical % of elytra in male, middle in
female; mandible of male narrow, usually very much longer than

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

bo

head, almost glabrous (Stenodontes s. str.) chevrolati Gahan

Metepisternum broad, inner margin straight or feebly convex;
mandible without strong preapical tooth on inner margin (Ortho-

mallodon) dasytomus (Say)

Metepisternum narrow, inner margin slightly concave in female,
strongly concave in male; mandible with a strong preapical tooth
on inner margin (Mallodon)

Integument medium to dark red brown; mandible tumid and re-
tracted at base, particularly in profile (Fig. 4), outer margin rather
strongly, evenly arcuate from base to tip, never but slightly longer
than head; lateral margin of pronotum finely spinose, the subbasal
spine most prominent, often 2 or more as long as remaining
spines; anterior '2 of submentum deeply recessed, recession not
attaining lateral sutures; genae not produced over submentum,
but lateral and usually basal portions of submentum distinctly
produced over recession in males, scarcely so in females

madericus, new species

Integument very dark brown or piceous; mandible not tumid or
retracted at base, outer margin straight or slightly arcuate over
basal 7%, then rather abruptly inflexed to tip, in males (except
some minor males) distinctly longer than head; lateral margin
of pronotum generally finely denticulate, subbasal dentule oc-
casionally prominent, not spinose; anterior '2 or % of submentum
deeply recessed over entire width; genae distinctly produced over

recession in males, less so in females lobigenis (Bates)

Stenodontes madericus Skiles, new species
Figs. 2-6

Stenodontes arizonicus Hovore and Giesbert, (not Casey), 1976:350 (habits).

Male.—Form elongate, flattened, sides sub-parallel. Integument shining,
glabrous, medium to dark red brown. Head very coarsely, rugosely punctate
around eyes, punctures becoming sparse and separate in broad, polished
median region extending from occiput to frons and continuing laterally
onto antennal tubercle; dorsum with a longitudinal impression, very fine
basally, becoming broad and deep on frons, abruptly bifurcating to form

VOLUME 80, NUMBER 3 415

Fig. 2. Known geographical ranges of: Neomallodon arizonicus (Casey), +;
Stenodontes madericus Skiles, O. Heavy circle indicates the type locality of S$. madericus.
Small solid dots show the locations of the cities of Phoenix, Tucson, Chihuahua.

clypeal suture; occiput finely granulate-punctate, a few punctures ex-
tending onto vertex; antennal tubercle acute; antenna attaining middle
of elytron, scape coarsely, closely punctate externally, pedicel and basal
flagellar segments finely, sparsely punctate, segments beginning with

416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Stenodontes madericus Skiles. Left, holotype male; right, allotype female.

3rd or 4th carinulate externally, carinulae becoming more numerous dis-
tally, apical segments completely carinulate, 3rd segment shorter than

scape, longer than 4th, 5th through 10th subequal or gradually increasing
I 8 S | 8 om

in length, 11th distinctly longer than 10th, subequal to scape, feebly ap-
pendiculate; mandible longer than head, tumid, strongly retracted at base,

basal '2 bulbous in profile, subangulate above, about 3x as thick as base
1

of apical ‘2, abruptly declivous at middle of mandible; outer margin —

coarsely, densely punctate, a long recurved seta arising from each punc-

ture, inner margin more densely punctate, setae very long, erect, a strong

blunt tooth before apex, a strong triangulate tooth at or before middle;
1

submentum coarsely, rugosely punctate, anterior 12 deeply recessed, re- |

cession not attaining lateral suture, sides and base distinctly, usually dra-
matically, produced over recession, genal-submental suture carinate, not
produced. Pronotum 2x as wide as long, anterior margin broadly tri-

sinuate, basal margin broadly bisinuate, often with a 3rd vague sinus at

middle, lateral margin finely spinose, subbasal spine most prominent,

VOLUME 80, NUMBER 3 417

oe

ea se
OY eat eee Ce,

*

Fig. 4. Profile of male S. madericus, showing bulbous, basically retracted mandible.

usually 2 as long as others; disk polished, very finely, sparsely punctate,
punctures becoming much coarser and denser on sides which are very
broadly, longitudinally rugose, base of disk with 2 transverse, oval im-
pressions clearly visible to the naked eye; prosternum densely, usually
finely, punctate and thinly pubescent, sparsely punctate between coxae;
metepisternum very narrow, finely, densely punctate and pubescent, inner
margin strongly concave; mesosternum and metasternum covered with
dense golden pile, except shining, rather sparsely pubescent midline re-
gion which is narrow basally and broadly flaring apically; scutellum
glabrous, very finely, very sparsely punciate. Elytra at least 2* as long
_as basal width, widest at about middle, base wider than pronotum; surface
smooth, somewhat dull, often vaguely rugulose basally near suture, often
finely, sparsely punctate apically; apices broadly, separately rounded, suture
angulate or minutely dentate. Legs short; femora somewhat slender, shining,
very sparsely, finely punctate, glabrous except for sparse, recurved setae
near lower posterior margins; tibiae less shining, more coarsely, densely
punctate than femora, glabrous except for scattered, erect, golden setae
and 2 thin lines of dense, golden pubescence covering at least apical halves
of lower margins. Abdomen with sternites polished, glabrous and im-
punctate at middle, sides sparsely punctate, each puncture with a reddish-

418 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Presumed mid-instar larva of S$. madericus found with an adult male in
trunk of living Quercus hypoleucoides. a, lateral view. b, dorsal view. c, ventral view.

golden recumbent seta, setae becoming denser on lateral and apical mar-
gins; 5th sternum transverse, shorter than, or subequal to, 4th, apex
broadly emarginate, densely fringed with long, reddish-golden setae.
Length, apex of elytron to base of mandible, 26-40 mm.

VOLUME 80, NUMBER 3 419

Fig. 6. Fragment of trunk of Q. hypoleucoides containing three emergence holes
of S. madericus.

Female.—Mandible shorter than head, less strongly retracted at base than
in male; preapical tooth less prominent; antenna attaining basal % of
elytron, 3rd segment 7 as long as scape, 4th through 6th scarcely decreas-
ing, 7th through 10th scarcely increasing in length, 11th subequal to scape,
not appendiculate; sides of submentum not or slightly produced over apical
recession. Pronotal spines more pronounced, subbasal spine 2-4 as
long as remaining spines; prosternum usually more shining and sparsely
punctate than in male. Metepisternum less narrow than in male, inner
margin only slightly concave. Fifth abdominal sternum longer than 4th,
apex truncate, notched, or broadly emarginate. Length, 30-43 mm.

Type-locality—Roundup Picnic Ground, elevation 5,400 feet, Madera
Canyon, Santa Cruz Co., Arizona.

Known range.—Oak woodland of north central Arizona to northwestern
Mexico (Fig. 2).

Flight period—tLate June to early August.

Habits.—Stenodontes madericus is much less common and widespread
than its sympatric congeners S. logigenis and S. dasytomus, but adults are
attracted to light in oak woodland throughout much of southeastern Ari-
zona. The only confirmed host for S. madericus is living silver leaf oak
(Quercus hypoleucoides A. Camus). The latter species is characteristically
rather small, the diameter of the trunk seldom exceeding 10 or 12 inches.

An adult male (abdomen still distended) and three presumed mid-instar
larvae of S. madericus were found at the type-locality on 21 June 1977 in
the six to eight inch diameter trunk of a living Q. hypoleucoides (E. F.
Giesbert), near where several adults had been taken at night in July of
a previous year, walking on living trunks of the same species (J. S. Cope
and D. G. Marqua). (Here it should be noted that the remarks of Hovore

420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

and Giesbert (1976), concerning the oak infesting habits of S. arizonicus,
in fact refer to S. madericus.)

The larvae were mining the heartwood and had excavated several inter-
mingling galleries which were parallel to the grain and almost entirely
filled with a rather coarse, tightly packed mixture of granular frass and
feculae. The extent of the galleries and the simultaneous presence of three
larvae and an adult in the same gallery system indicate that the larvae re-
quire at least two years to develop and often reinfest the same tree. The
larvae construct oval emergence holes, often through scars devoid of bark
(Fig. 6), and leave both the holes and the outer two or three inches of the
exit galleries open, thus creating the impression the adults have already
emerged.

Emergence holes which appear to be those of S. madericus are not un-
common in Madera Canyon and other oak woodland regions of south-
eastern Arizona. Infested trees seldom seem to suffer from the initial at-
tack. However, trees which have been repeatedly infested sometimes
appear to be in decline, and on occasion I have seen a large, living oak limb
seriously weakened or even broken off as a result of extensive larval work.

Given that most oak infesting cerambycids, including prionines, are not
confined to a single host species, it is possible that S. madericus attacks
several species of evergreen oaks, and perhaps even deciduous oaks. Quercus
hypoleucoides is certainly not the only host species, since S. madericus has
been taken in both Oak Creek Canyon and the Superstition Mountains
of Arizona where Q. hypoleucoides does not occur (Kearney and Peebles,
1960). There are also indications that S. madericus does not attack only
living trees. Three specimens have been taken from beneath slabs of
bark on standing dead oaks—a pair in copulo (end-to-end and facing apart)
in the Huachuca Mountains of Arizona (F. T. Hovore) and a male in Cave
Creek Canyon near Portal, Arizona (A. E. Lewis). In addition, one oc-
casionally finds in southeastern Arizona a standing dead oak thoroughly
riddled with emergence holes which appear to be those of S. madericus.

Diagnosis.—This species, like Archodontes melanopus and the other
North American species of Stenodontes (but apparently unlike Neomallodon
arizonicus) is quite variable in size and exhibits significant mandibular
allometry, characteristics which undoubtedly have been largely responsible
for the confusion of S. madericus with other species of Macrotomini, par-
ticularly S. lobigenis. Despite the fact that S. madericus has until now
remained unrecognized, most specimens are readily separated from sim-
ilar forms by the characters given in the preceding keys. The tumid,
basally retracted mandibles (Fig. 4) and deeply excavated submentum im-
mediately separate males and major females from other species of Steno-
dontes, and the sculptured submentum and narrow metepisternum readily
distinguish all specimens from Neomallodon and Archodontes. However,

VOLUME 80, NUMBER 3 421

minor females of S. madericus and S. lobigenis are often difficult to
separate, primarily because the submental sculpture and mandibular pro-
files of minor females are not distinctive. Minor females of S. lobigenis
from the United States, mainland Mexico and northern Baja California
are generally separable from specimens of S. madericus by their dark brown
color and short, denticulate pronotal spines, but specimens of S. lobigenis
from the cape region of Baja California and from various islands in the
Gulf of California are often red brown and have spinose pronotal mar-
gins. The latter, however, can be distinguished from S. madericus by the
inner margin of the mandible, which in S. madericus is bidentate and in
S. lobigenis is vaguely crenate and lacking a distinct subbasal tooth.
Stenodontus madericus clearly belongs to the subgenus Mallodon Audinet-
Serville, and the description of Mallodon given by Linsley (1962) must be
modified to accommodate tumid, basally, retracted mandibles. Within
the subgenus, S$. madericus is a member of what Lameere (1902) referred
to as the Mexican group of Nothopleurus Lacordaire, i.e., S. lobigenis and
S. subsulcatus (Dalman). The essential morphological differences between
these species are expressed almost exclusively in the structures of the
mandibles and submenta. Stenodontes subsulcatus is quite possibly known
from only three specimens, all types, none of which I have seen. However,
the original description of Mallodon gnatho White (1853) and the detailed
description of the type of Nothopleurus ebeninus Lacordaire by Lameere
(1902), who synonymized both with Prionus subsulcatus Dalman and
placed them in Stenodontes, are sufficient to draw the following con-
clusions. Stenodontes madericus is probably most closely related to
S. subsulcatus, but differs from the latter in the more dramatically recessed
submentum, less pronounced mandibular structure and smaller size. The
latter, to be sure, is a poor species character but nevertheless, the White
type, a male, is 42 mm long and the Lacordaire type, also a male, is 45 mm
long. Both are thus larger than the largest male (40 mm) of the rather sub-
stantial type-series of S. madericus. The mandibles of S. subsulcatus are
-apparently similar to those of S. madericus, but differ in having two con-
tiguous preapical teeth on the inner margin and the basal swelling dis-
tinctly triangular in profile and terminating apically in a tooth-like pro-
cess. Stenodontes subsulcatus is known only from Honduras and _ the
Yucatan and is therefore considerably separated geographically from
S. madericus. On the other hand, S. lobigenis and S. madericus are sym-
patric throughout most of the range of the latter, and their specific differ-
entiation presumably arose via exploitation of distinct, but contiguous,
biomes—evergreen oak woodland and desert scrub.
Remarks.—This species is named to commemorate the type-locality,
Madera Canyon, Arizona, long noted for its rich and unusual avifauna and
entomofauna by professional scientist and amateur naturalist alike.

422 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Material examined.—Holotype ¢ (U.S. National Museum of Natural
History, Washington, D. C., USNM type #75481) from Roundup Picnic
Ground, elevation 5,400 feet, VII-14-75, and allotype from Santa Rita
Lodge, elevation 4,960 feet, VII-15-75, Madera Canyon, Santa Cruz County,
Arizona (D. D. Skiles, at light). Fifty-seven paratypes. Arizona. Coconino
County: Indian Gardens, 6 mi N Sedona, 1? VIII-7-67 (S. M. Anderson,
labelled Archodontes melanopus serrulatus é). Santa Cruz County, Madera
Canyon: 49, VI-28-VII-4-73, 246, 192 VII-15-75 (D. D. Skiles, at light); 13,
42, VII-18-65, 32, VII-19-65, 16, 12, VII-20-65, 16, 292, VII-5-75, 42,
12, VII-20-75 (J. S. Cope, at light and walking on living oak trunks); 1¢,
12, VII-11-57( Stange and Harding); 22°, VI-25-76 (at light), 1¢ (in living
oak trunk), VI-21-77 (E. F. Giesbert); 19, VII-25-73, 16, VII-26-76 (F. T.
Hovore, at light); 26, VII-10-15-75 (A. E. Lewis, at light); 12, VII-29-71
(C. E. Langston, at light); 12, VII-11-63, 12, VI-28-75 (G. C. Walters, Jr., at
light); 16, VII-2-65 (D. N. Harrington, 4,880 feet); 12, VII-23-65, 14, 32,
VII-8-71, 12, VIII-4-72, 16, VII-14-75, 12, VII-18-75 (D. G. Marqua, at
light and walking on living oak trunks). Gila County: Globe, 1¢, July
(D. K. Duncan). Cochise County: Carr Canyon Road, elevation 6,000
feet, Huachuca Mts., 16, 12, (in copulo under bark of oak), VII-24-76
(F. T. Hovore); Southwestern Research Station, 5 mi SW Portal, 14
(dead about one year), 1¢é, 12, VII-3-76 (D. D. Skiles, at light); 3 mi SW
Portal, 146, VII-25-66 (A. E. Lewis, under bark of dead oak). No county
given (very old label): Superstition Mts., 12, July (D. K. Duncan). Mexico.
Chihuahua: 8 mi W Matachic, elevation 7,200 feet, 14, VH-8-47 (D. Rocke-
feller Exp., Schramel, AMNH); 15 mi E Cuauhtemoc, elevation 6,600 feet,
12, VII-11-64 (J. A. Chemsak and J. Powell, at light).

Also seen but not included as paratypical owing to the questionable
validity of the locality (A. E. Lewis, personal communication), 12, labeled
Gila Bend, Ariz., 5 Aug 1954, (A. E. Lewis).

Paratypes reside in the collections of the California Academy of Sciences,
the American Museum of Natural History, the Essig Museum of En-
tomology, University of California, Berkeley, and the collections of the
author, J. S. Cope, E. F. Giesbert, F. T. Hovore, C. E. Langston, A. E. Lewis,
D. G. Marqua, and G. C. Walters, Jr.

Acknowledgments

I am indebted to E. G. Linsley and J. N. Belkin for many fruitful discus-
sions and for reviewing the manuscript.

The following institutions and individuals are gratefully acknowledged
for the loan of specimens and type-material: U.S. National Museum of
Natural History (USNM), Smithsonian Institution, Washington, D.C. (T. J.
Spilman); California Academy of Sciences (CAS), San Francisco, Cali-

VOLUME 80, NUMBER 3 423

fornia (D. H. Kavanaugh); Essig Museum of Entomology, University of
California, Berkeley (J. A. Chemsak); American Museum of Natural His-
tory (AMNH), New York, New York.

The major portion of the collection and specimen data were obtained
through the generous cooperation of several private collectors: J. S. Cope,
E. F. Giesbert, F. T. Hovore, C. E. Langston, A. E. Lewis, D. G. Marqua,
and G. C. Walters, Jr. I am particularly grateful to C. E. Langston and
F. T. Hovore for information on the rearing of N. arizonicus and to E. F.
Giesbert for information on the larval habits of $. madericus and for compar-
ing specimens of S. madericus with material in the Casey collection at
the USNM.

I also wish to thank V. Roth and the Southwestern Research Station of
the American Museum of Natural History for assistance and collecting
privileges, L. D. Skiles for collecting assistance, S. R. Skiles for typing and
editing the manuscript, and J. Sells for preparing Fig. 2. The photo-
graphic work was done by the author.

Literature Cited

Casey, T. L. 1912. Memoirs on the Coleoptera. 3:228.

Hovore, F. T., and E. F. Giesbert. 1976. Notes on the ecology and distribution
of western Cerambycidae (Coleoptera). Coleop. Bull. 30:350.

Kearney, T. H., and R. H. Peebles. 1960. Arizona Flora. 2nd ed., Univ. Calif. Press,
Berkeley and Los Angeles.

Lameere, A. 1902. Revision des Prionides. Quatriéme Mémoire—Sténodontines. Mem.
Soc. Entomol. Belg. 9:63—110.

Linsley, E. G. 1934. Notes and descriptions of west American Cerambycidae
(Coleoptera). Entomol. News. 45:161.

—. 1957. Descriptive and synonymical notes on some North American Cerambyc-
idae (Coleoptera). Amer. Mus. Nov. 1828:1—21.

——. 1962. Cerambycidae of North America, Part II], Taxonomy and _ Classifica-
tion of the Parandrinae, Prioninae, Spondylinae, and Aseminae. Univ. Calif.
Berkeley Publ. Entomol. 19:1—102.

Linsley, E. G., J. N. Knull, and M. Statham. 1961. A List of Cerambycidae from
the Chiricahua Mountain Area, Cochise County, Arizona (Coleoptera). Amer.
Mus. Nov. 2050:3.

White, A. 1853. Catalogue of Coleopterous Insects in the Collection of the British
Museum, Part VII, Longicornia I, p. 45.

Institute of Geophysics and Planetary Physics, University of California,
Los Angeles, California 90024.

Footnote
*While Neomallodon arizonicus has apparently not been collected in Mexico (see

Fig. 2), it is unlikely that this is a result of the species’ observance of a_ political
boundary.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 424-440
THE IMMATURE STAGES AND BIOLOGY OF MALLOTA
POSTICATA (FABRICIUS) (DIPTERA: SYRPHIDAE)

Chris T. Maier

Abstract—The immature stages of Mallota posticata (Fabricius), which
are described and illustrated, occurred in or near wet treeholes in deciduous
trees. Larvae developed in the wet treehole detritus upon which they
fed. In central Illinois, larvae of univoltine M. posticata attained full size
by late summer, 2’2-3'% months after hatching, and then they entered dia-
pause. In the spring, they usually crawled out of treeholes and pupariated
in the soil at the base of the tree. Laboratory-reared larvae required
several months of chilling to terminate diapause and subsequently to
pupariate. Males generally pupariated before females, but both remained
in the puparium about 13-14 days when they were maintained at 22°C
and on a 17 hour photophase and a 7 hour scotophase. Adults exposed
to a 0400-2100 photophase eclosed mostly between 0600 and 1000—the
equivalent of morning hours in nature.

In the field, adults fed upon open, actinomorphic blossoms which pro-
duced substantial quantities of pollen—a requisite for normal ovarian de-
velopment. Mating, which is described, occurred near flowering plants and
near treeholes. Females readily oviposited (17.7 + 1.30 eggs per clutch) in
artificial oviposition containers placed in the forest. Oviposition in con-
tainers occurred principally in June between 1100 and 1800 CDT.

The numerous investigations on syrphids in the tribe Eristalini deal
mostly with representatives of the genus Eristalis, especially E. tenax (L.),
and only rarely with members of other genera. Eristaline larvae, “rat-tailed
maggots, are adapted for an aquatic existence. In particular, they have
an extensible, caudal respiratory tube for obtaining atmospheric oxygen,
anal papillae for facilitating ionic exchange and an elaborate filtering
apparatus for feeding on detritus (Hartley, 1963; Hase, 1926; Krogh, 1943;
Kruger, 1926; Roberts, 1970; Wahl, 1900; Wichard and Komnick, 1974).
These structures and the external sensory organs have provided most of
the characters used in taxonomic descriptions of larvae (Dixon, 1960;
Hartley, 1961; Hennig, 1952). Dixon (1960), Dolezil (1972), Hartley (1961),
Hennig (1952), Johannsen (1935) and others published partial keys to
third-instar eristaline larvae.

The larvae of Mallota bautias (Walker), M. cimbiciformis (Fallen) and
M. posticata (Fabricius) develop in detritus-containing rot pockets, usually
wet treeholes in upright deciduous trees (Becher, 1882; Britten, 1917; Coe,
1953; Johannsen, 1935; Lintner, 1882; Lundbeck, 1916; Snow, 1949). Johann-

VOLUME 80, NUMBER 3 425

sen (1935), Lintner (1882), Morse (1910) and Snow (1949) described the
gross external morphology of one or more of the immature stages of M.
posticata but largely ignored its life history. The taxonomic descriptions
are not sufficiently detailed for use in comparative studies. Furthermore,
I shall show that Johannsen’s (1935) description of the anal papillae of
the larva is erroneous. Unfortunately, Hennig (1952) and Snow (1949)
incorporated the spurious information into their taxonomic keys.

Adults of M. posticata, apparent Batesian mimics of bumblebees, visit
open, actinomorphic blossoms from April to July in the eastern United
States (Graenicher, 1910; Robertson, 1928; Waldbauer and Sheldon, 1971;
Waldbauer et al., 1977; Maier, unpublished data). Nothing else is published
about the behavior of adults except for Curran’s (1925) description of
the oviposition behavior of an unidentified species of Mallota. Fashing
(1973, 1974, 1975 and 1976), however, reported that M. posticata females
are dispersal agents for two species of treehole-inhabiting mites.

Materials and Methods

Descriptions are based on specimens collected in nature and on others
reared in the laboratory. Live eggs, larvae and puparia were used for most
measurements of external structures. Measurements that were made on im-
matures or on biological events are usually reported as the mean + standard
error.

The principal research site, Sand Ridge State Forest, is in an extensive
sand area in central Illinois near the Illinois River. The vegetation at
Sand Ridge consists mostly of oak-hickory forest, disturbed sand prairie,
and pine plantations (Maier, 1977a). The forest is dominated by the oaks
Quercus marilandica Muenchh., and Q. velutina Lam. and by the hickories
Carya ovalis (Wang.) Sarg. and C. tomentosa (Poir.) Nutt.

Laboratory colonies were started with the eggs of wild females and main-
tained for three generations without noticeable deleterious effects. The
standard diet for rearing larvae was a 1:1 mixture of Purina Fly Larvae
Media (Ralston Purina Co., St. Louis, Missouri) and homogenized treehole
detritus. The mixture was saturated with water and usually supplemented
with 3-4 grams of Vitamin Diet Fortification Mixture (ICN Pharmaceuticals,
Inc., Cleveland, Ohio) per liter of wet diet. Larvae reared in crispers at
22°C and on a 17 hour photophase and a 7 hour scotophase reached full

size in approximately three months and then entered diapause. One month

later, they were placed in crispers containing moist, coarse treehole detritus
and kept at 2°C and on a 10 hour photophase and a 14 hour scotophase.
After 4-5 months, the larvae were transferred to 6.5 < 12.5 cm crystalliz-
ing dishes which were placed in crispers partially filled with sand. The
dishes were filled with the semi-artificial diet and exposed to the initial

426 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

temperature and photoperiod regimes. In 1-3 weeks, the larvae crawled out
of the rearing dishes and pupariated in the sand.

Adults were usually kept in plastic-covered 0.6 x 0.6 X 0.6 meter cages
and maintained at 75% relative humidity, at 22°C, and on a 17 hour
photophase and a 7 hour scotophase. Each cage contained two Petri
dishes of pollen (collected in a pollen trap on a honeybee hive), two dishes
with a solution of 10% glucose and 10% sucrose, and a 4 liter plastic jug
for oviposition. The jug had a 10 cm circular opening on the side and
held 1.5 liters of homogenized, water-soaked treehole detritus and strips of
bark. The experiment monitoring ovarian development was carried out in
0.3 x 0.3 x 0.3 meter cages.

The oviposition containers described above were also used to record
egg-laying activity in nature. Seasonal and diurnal patterns of oviposi-
tion were measured with 20 and 30 containers, respectively.

Descriptions of the Immature Stages

Egg.—Length 1.5 + 0.01 mm, maximum width 0.6 + 0.02 mm (N = 10).
Elongate oval in outline, ends bluntly rounded (Fig. 1). White; chorionic
surface microscopically sculptured, pattern under light microscope as in
Fig. 1.

Mature third-instar larva—Body length 20.8 + 0.72 mm, maximum
width 6.4 + 0.12 mm, length of retracted caudal respiratory tube 13.6 +
0.58 mm (N = 14); ratio of body length to retracted respiratory tube
length 1.5:1.0.

Body slightly fusiform, truncate anteriorly, abruptly tapered posteriorly
to base of caudal respiratory tube (Fig. 2). Cuticle translucent in life,
cream to dirty white after fixation. Segmentation indistinct; longitudinal
plicae anteriorly, numerous transverse plicae posteriorly. Short, unpig-
mented hairs sparsely cover body; longer hairs laterally and posteriorly,
in longitudinal bands laterally.

Antenno-maxillary sense organ (AM) and mouth circled by brown spinules
(Fig. 3); inflatable, pilose lobes lateral to mouth; verricule between an-

tenno-maxillary sense organ and mouth. Two anterior spiracles (AS),

brown, with 17-18 elliptical to circular openings on anteroventrally di-
rected. face and its margins (Figs. 2, 4). Ventral prolegs well developed
(Figs. 2, 3, 5), 1 thoracic, 6 abdominal pairs. Crochets brown apically,

= |

Fig. 1. Mallota posticata, egg. Figs. 2-8. Mallota posticata, mature third-instar
larva. 2, lateral view. 3, ventral view, anterior end. 4, anteroventral view, left an-
terior spiracle. 5, lateral view, left fourth abdominal proleg. 6, cross section, arrange-
ment of sensilla on abdominal segments 1-7. 7, ventral view, outline of protrusible

VOLUME 80, NUMBER 3 427

ae / : ‘

| v

|

| 4 4 7

‘ \ I A At

usr: t ; 4 no yy » \ ¥
Se yy /

eed \
fH Gti hi ——— N |
My, wy ) Sony
xr YS }
=
5 mm AX
=~ ye = if
Ne

1] fi
Ni \ ff
\\ Vr Hy | }
\Y | V | /
Sh 4 ay
S /
NN aR
/, / f / i \ \
// | | ! HY \\\
|| \
[/ I /} I] uta
SV HOGA LOIN ON
Y I ra \| \| \) ~
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anal papillae. 8, spiracular plate of caudal respiratory tube. Abbreviations: AS,
anterior spiracle; AM, antenno-maxillary sense organ.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Imm

i

13

TN
w

VOLUME 80, NUMBER 3 429

multiserial, in 4-5 rows becoming indistinct dorsally (Fig. 5); crochet ar-
rangement posterolateral at anterior end of body, nearly lateral at pos-
terior end. Abdominal segments 1-7 with 20 sensilla each (Fig. 6); ar-
rangement of sensilla on other segments similar to that described by
Hartley (1961) for Mallota cimbiciformis. Anal papillae protrusible, 6 an-
terior ones, 8 posterior ones (Fig. 7). Caudal respiratory tube immediately
anterior to brown, sclerotized distal portion, bearing anteriorly directed,
unpigmented spines on longitudinal cuticular ridges. Spiracular plate of
caudal respiratory tube convex, with 2 prominent spiracular scars cen-
trally, encircled by 8 laterally fringed setae (Fig. 8).

Cephalopharyngeal skeleton as in Figs. 9-13; length 4.1 + 0.14 mm (N =
7). Mandibular lobes (ML) well developed, ribbed (Figs. 9, 11), with
comb of filaments on inner ridges (Fig. 12). Pharyngeal sclerite with pos-
teriorly projecting dorsal cornuae (DC) connected by dorsal bridge (DB)
(Fig. 11), ribbed cibarial filter ventrally (Figs. 9-10). Pharyngeal sclerite
black to amber anteriorly, translucent posteriorly except for light brown,
posteriorly tapering strip at each margin of cibarial filter (Figs. 9-10).
Two pairs of bars projecting anteriorly from pharyngeal sclerite (Fig. 9),
black to amber; dorsal pair unconnected, at lateral margins of brownish
epipharyngeal plate (EP) (Fig. 11); ventral pair joined by concave, black
to brown, sclerotized bridge (Figs. 10-11); labial sclerite (LS) supported
posteriorly by bridge. Floor of pharyngeal sclerite with cibarial filter of
9 filament-lined ridges, converging posteriorly (Fig. 10); inner 7 ridges with
single row of filaments on each side (Fig. 13), outer 2 ridges with 1
row of filaments.

Puparium.—Length excluding caudal respiratory tube 14.9 + 0.25 mm,
maximum width 8.1 + 0.14 mm (N = 15).

Inflated, tear-shaped in dorsal view (Fig. 14), fusiform in lateral view
(Fig. 15), strongly convex dorsally, nearly flat ventrally. Light brown, en-
tirely rigid. Longitudinal swelling at each dorsolateral margin; indistinct
transverse plicae, increasing posteriorly. Sparsely pubescent, larval cuticular
ornamentation present but indistinct. Two dehiscent plates at anterior
end (Fig. 16); anterior plate trapezoidal, evenly convex, with diverging
spiracles at anterior margin. Posterior plate abruptly turned anteroventrally

Figs. 9-13. Mallota posticata, cephalopharyngeal skeleton. 9, lateral view. 10, ventral
view, excluding mandibular lobes. 11, anteroventral view, junction between mandibular
lobes and pharyngeal sclerite. 12, lateral projections on cuticular ridges of mandibular
_ lobes. 13, filament-lined cuticular ridge of cibarial filter. Abbreviations: DB, dorsal
bridge of pharyngeal sclerite; DC, dorsal cornua of same; EP, epipharyngeal plate;
LP, labial plate; LS, labial sclerite; ML, mandibular lobe.

430 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2mm

VOLUME 80, NUMBER 3 431

at lateral margins; dorsum nearly rectangular, slightly convex, with nearly
straight, diverging, anterodorsally directed pupal respiratory horns. Horns
with numerous tubercles except at base and on lower surface (Fig. 17); each
tubercle with spiracular opening at apex.

Material studied —Descriptions based on material from Sand Ridge State
Forest, Mason County, Illinois; voucher specimens deposited in the Il-
linois Natural History Survey.

Johannsen’s (1935) brief description of M. posticata immatures (preceded
by a question mark) agrees with mine except for the number and the ar-
rangement of anal papillae in the larva. He found 13 anal papillae, 6 an-
terior and 7 posterior, rather than the 14 anal papillae, 6 anterior and 8
posterior, reported here. He did not associate the larva with an adult
and, thus, apparently described either another eristaline species, probably
a Mallota, or a damaged specimen of M. posticata. Unfortunately, Lintner’s
(1882) description is too fragmentary to permit comparison.

The immature stages of the North American M. posticata differ from
those of the sympatric M. bautias (Snow, 1949) and the european M. cimbici-
formis (Coe, 1953; Hartley, 1961). Although Snow (1949) described only
the puparium of M. bautias, he noted in a key that the larva has 7 anterior
and 7 posterior anal papillae. Furthermore, in contrast to the M. posticata
puparium, the M. bautias puparium described by Snow (1949), and others
examined by me, have a nearly flat anterior dehiscent plate, the distal por-
tion of the pupal respiratory horns bent dorsally, and distinct transverse
plicae on the abdominal dorsum.

The M. cimbiciformis larva has no pubescence on the posterior abdominal
dorsum, 25 openings on each anterior spiracle, and 12 anal papillae
(Hartley, 1961). Any one of these characteristics separates M. cimbiciformis
from M. posticata. In addition, the absence of dorsal abdominal pubescence
and the greater number of spiracular openings on the anterior spiracle of
the M. cimbiciformis puparium distinguishes it from the M. posticata
puparium.

The general morphology of the cephalopharyngeal skeleton of M. posti-
cata resembles that of several Eristalis species (Hartley, 1963; Wahl, 1900)
and Myjiatropa florea (L.) (Roberts, 1970), taxonomically related spe-
cies with similar feeding habits. Therefore, Roberts’ (1970) account of the

function of the feeding structures in M. florea probably applies well to
'M. posticata and most other eristaline larvae.

Figs. 14-17. Mallota posticata, puparium. 14, dorsal view. 15, lateral view, ex-
cluding caudal respiratory tube. 16, dehiscent plates. 17, posterodorsal view, left pupal
| respiratory horn.

432 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Biology

Larvae were collected from treeholes in upright, living Carya spp.,
Liquidambar styraciflua L., Populus deltoides Marsh, Quercus alba L.,
Q. marilandica, Q. velutina Lam., and Ulmus sp. in Illinois, Indiana or
Michigan. The openings to these treeholes occurred at various heights
but were mostly near ground level. The cavities usually contained more
than 3 liters of detritus and held standing water for more than 2 months
out of the year. There were 1-51 larvae per treehole; the number generally
increased with the quantity of detritus. The 51 larvae were in a treehole
in Populus deltoides that contained approximately 60 liters of detritus. In
the summer, larvae were usually near the bottom of treeholes where the
particles of detritus were the smallest and the moisture was the greatest.
Other organisms commonly found in the same treeholes included Acarina
(Fashing, 1973, 1974, 1975 and 1976), helodids (Elodes, Prionocyphon), cera-
topogonids, psychodids (Telmatoscopus) and rarely other syrphids (e.g.,
Somula decora Macquart and Spilomyia longicornis Loew). Mallota bautias
larvae never occurred in treeholes containing M. posticata larvae.

The filter-feeding larvae of M. posticata consumed fine-grained detritus
and other material trapped in treeholes. Most treehole detritus originates
from trees, for example, from leaves, wood and acorns. Wind and the
activities of ants facilitate the deposition of detritus in treeholes. In the
laboratory, larvae often protruded and then pulsated their anal papillae
while feeding. Thus, in addition to an osmoregulatory function (Krogh,
1943; Wichard and Komnick, 1974), the anal papillae may also be used
to put food particles into suspension and to circulate them.

In nature, larvae attained full size and weight (0.35-0.55 grams) and
entered diapause by late summer, approximately 22-3’ months after
hatching. Diapausing larvae were inactive, relatively inflexible and pink
(due to particles of unknown origin suspended in the hemolymph). These
larvae also had an empty gut and contracted cephalothoracic region.

Larvae overwintered in diapause in treeholes. In early spring, they
commonly congregated against the inner treehole wall in moist but not
water-soaked detritus. Exposure to cold was necessary for the termina-
tion of diapause and for the synchronous emergence of adults, an important
consideration in any univoltine species. In the laboratory, full grown
larvae that were subjected to 2°C and a 10 hour photophase and 14 hour
scotophase for 4-5 months and then to 22°C and a 17 hour photophase and
7 hour scotophase pupariated over a 1-3 week period. By contrast, larvae
that were continuously maintained at 22°C and on a 17 hour photophase
and a 7 hour scotophase failed to pupariate within 14 months after hatch-
ing.

At Sand Ridge State Forest, larvae usually left treeholes from April
to June and pupariated in the surface of the sandy soil within 1-2 m of

VOLUME 80, NUMBER 3 433

20
C1) @ (N=56)
E} o (N=60)

Number of Puparia

Dy a Se Salonen SG 26 D2 24 26 28 30
Days After Cold Treatment

Fig. 18. Pattern of Mallota posticata pupariation after exposure to 2°C and a 10
hour photophase and a 14 hour scotophase.

the tree trunk. They occasionally remained in the treeholes and pupariated
near the surface of the detritus in the cavity. Pupariation usually occurred
over a 2 week period at any one treehole. Although puparia were frequently
discovered, the actual departure of larvae from treeholes was witnessed only
twice, at 1400 and 1515 CDT on 19 May 1975. Ants foraging in the vicinity
attacked both larvae. One escaped and pupariated in the soil, and the
other climbed the tree until out of sight. At 1100 on the following day,
ants successfully captured and killed a larva at the same treehole.

In the laboratory, most larvae crawled out of rearing dishes between
1100 and 1900 of a 0400-2100 photophase, wandered on the sand up to
several hours and finally pupariated in the sand, often under the dishes.
The pupal respiratory horns protruded through the puparium 54-72 hours
later. Larvae that were exposed to 2°C and a 10 hour photophase for 140
days began to pupariate only 9 days after they were switched to 22°C
and a 17 hour photophase (Fig. 18). The main time from the termination
of the cold treatment to pupariation was 15.2 + 0.38 days for males and
20.4 + 0.41 days for females. Males and females remained in the puparium
12-15 days and 13-15 days, respectively (Table 1). Most males (48.72%)
eclosed on day 13, and most females (60.00%) eclosed on day 14. Similarly,

434 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Time spent in the puparium by both sexes of Mallota posticata at 22°C and
on a 17 hour photophase and a 7 hour scotophase.

J Males Females
Days in
puparium’ Number % of total Number % of total
12 2 5), 118) 0 0.00
13 19 48.72 8 22.86
14 16 41.02 21 60.00
15 2 alls 6 17.14
Total 39 35

*Puparia were collected and observed at 1500 each day.

Lintner (1882) reported that a male and a female, respectively, spent 12
and 14 days in the puparium.

Eclosion was concentrated between 0600 and 1000, with a distinct peak
at 0700, when the photophase extended from 0400-2100 (Fig. 19). In na-
ture, individual adults were seen to eclose at approximately 0800, 1000,
and 1100 CDT. Akre et al. (1973) also noted a morning peak in eclosion
for the syrphid Microdon cothurnatus Bigot.

Adults frequented open, actinomorphic blossoms of species which pro-
duce substantial quantities of pollen per flower or inflorescence. In cen-
tral Hlinois, they fed on the pollen, nectar, or both of Ceanothus americanus
L., Celastrus scandens L., Cornus drummondi Meyer, C. racemosa Lam.,
Heracleum lanatum Michx., Osmorhiza sp., Pastinaca sativa L., Prunus
serotina Ehrh., Ptelea trifoliata L., Rhus glabra L., Rosa carolina L., Rubus
allegheniensis Porter, R. occidentalis L., Sambucus canadensis L., and
Viburnum sp. In other parts of their range, they visited Maianthemum
canadense Dest. (Graenicher, 1910) and Rosa setigera Michx. (Robertson,
1928). At Sand Ridge State Forest, adults foraged primarily on Cornus
drummondi, C. racemosa, Rosa carolina and Sambucus canadensis and
demonstrated considerable flower constancy (Maier, 1977b).

Table 2 shows that virgin females required pollen, a protein source,
for normal ovarian development. In the absence of pollen, little growth
was evident after 8 days; but with pollen some eggs were fully formed
after 2-4 days. Schneider (1948) also found that pollen consumption was
a prerequisite for rapid ovarian development in the syrphid Episyrphus
balteatus (De Geer).

Maier (1977b) noted that the mate-seeking activities of Mallota posticata
males were coordinated temporally and spatially with female activity. In |
the morning, males searched blossoms to find feeding females. In the |
afternoon, most males defended territories around wet treeholes and at-_
tempted to mate females arriving to oviposit. Matings occurred at both |
sites. |

|

VOLUME 80, NUMBER 3 435

40

30

20

Number Eclosed

0400 0600 0800 1000 1200 1400 1600 1800 2000

Hour of Eclosion (CDT)

Fig. 19. Periodicity in the eclosion of Mallota posticata adults at 22°C and on a
17 hour photophase (0400-2100 CDT).

No overt courtship was evident prior to copulation in M. posticata.
Males responded almost instantaneously to flying or stationary females.
In less than 2 seconds, a male grasped a female and achieved genital con-
tact. Often a brief flight followed with the male carrying the female.
Pairs alighted within 1-10 m of the site of encounter, generally on foliage,
the trunk of a tree or occasionally on the forest floor. The male usually
hooked his prothoracic and mesothoracic tarsal claws onto the costal wing
margins of the female. He clasped the abdomen of the female with his
metathoracic tibiae and enlarged femora. A shrill sound, apparently created
by the vibrating thoracic sclerites of one or both flies (Aubin, 1914), fre-

Table 2. The average weights (mg) of the ovaries of unmated Mallota posticata
females that were fed pollen or pollen-free diets for 2, 4, 6 or 8 days after eclosion.

Days
after
eclosion Number Without pollen Number With pollen
2, 4 3:0 O26 5 PAI) Se sh PAS,
4 3 3\5) a= (O18) 4 26.5 + 4.38
6 3 Bhi S= OPAS 4 32.6 + 1.47
8 3 3.8 + 0.06 5 46.7 + 5.66

436 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

400

00
Ss
Wy
8

© 7200
sc)
Q
S
=

100

O

oa 6° Vs / V5 ; % 1h “3 ‘Ns We “fo2 Vag

pe eee ete ee
MAY JUNE

Sampling Period

Fig. 20. Number of Mallota posticata eggs laid in 20 artificial oviposition sites
during 2-day periods between 23 May and 29 June 1976.

quently accompanied the initiation of mating. If a female was not initially
receptive, the male moved anteriorly, maintained a firm grip on the fe-
male and vigorously stroked the eyes of the female with his prothoracic
legs. A male repeatedly stroked an unreceptive female at brief intervals
until one or both departed.

In the laboratory, both sexes mated frequently during the first 20 days
of their life. One male captured at a flower, marked and released was
resighted 11 days later while mating near a treehole. Most pairs copulated
for 1-3 hours in cages, but the duration of actual sperm transfer is unknown.

Gravid females flew upwind to treeholes. They circled all trees within
10 m of a tree containing a treehole and, after locating the treehole, usu-
ally hovered at the entrance for 0.5-1.0 minute before landing. Hovering
often included intermittent 0.5-1.0 second bobbing or down-and-up flights
before the opening of a treehole. After landing, females meticulously probed
crevices in the bark near the opening or in the wood of the inner wall of the
treehole with their ovipositor. Once they selected an oviposition site, they
rapidly laid a group of eggs. Eggs were depusited in one or rarely two
areas of a treehole, typically under bark at the entrance, above the water
line on the inner treehole wall or on the water surface. The mean egg
clutch, which was laid in detritus-containing artificial oviposition sites

VOLUME 80, NUMBER 3 437

250

~ 200

S

|

YH

(50

uy

™~

aS)

100
50

0800 !000 1200 1400 !1600 1800 2000
Sampling Hour (CDT)

Fig. 21. Total number of Mallota posticata eggs laid per hour in 30 artificial ovi-
position sites uniformly distributed over 0.5 hectare of forest floor and monitored for
10 days in June 1976.

placed in the forest understory, was 17.7 + 1.30 eggs (N = 103). The
mean time from hovering to departure at a natural treehole was 20.3 + 3.63
minutes (N = 6). Thirty females placed in the same laboratory cage laid
11,474 eggs over 30 days, an average of 382.4 eggs each.

In 1976 at Sand Ridge State Forest, egg-laying in artificial oviposition
sites was concentrated in June, with a peak on 14-15 June (Fig. 20). Little
Oviposition occurred prior to mid-May or after early July. The oviposition
peak on 14-15 June probably reflects a peak in female emergence in
early June. In the laboratory, females commenced ovipositing 3-6 days
after eclosion and deposited most of their eggs in the first 15-20 days of
their life. In nature, females oviposited primarily between 1100 and 1800
CDT in artificial oviposition sites (Fig. 21). The relatively low egg-laying
activity between 0800 and 1000 corresponds to the period when females
are feeding at flowers growing in fields adjacent to forests (Maier, 1977b).

In laboratory cages, the mean longevity of adults was 19.2 + 2.23 days
(N = 23) for males and 21.4 + 2.33 days (N = 25) for females. Two males

438 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of unknown age, which were captured at flowers and marked, were resighted
11 and 15 days later.

In conclusion, several of the significant developmental events in the
life of M. posticata have apparently been selected to occur in the spring
when environmental conditions are most favorable. After the termination
of diapause, larvae pupariate and adults subsequently eclose as numerous
species of flowers began to blossom near suitable larval habitats in deciduous
forests. Most spring flowers have an unspecialized floral morphology which
permits the adult syrphids to feed easily on the pollen, a requisite for rapid
ovarian development, and on the nectar, an energy source. Furthermore,
fragile, young larvae probably have a better opportunity for survival in
the spring than they do later because the water level in treeholes is highest
and the quantity of detritus is greatest due to the accumulation during the
previous two seasons.

Acknowledgments

I express my sincere appreciation to W. E. LaBerge, P. W. Price, G. P.
Waldbauer and D. W. Webb for their critical appraisals of earlier drafts
of this paper. The Illinois Department of Conservation and especially Olen
Bortel, Caretaker of Sand Ridge State Forest, willingly gave assistance
throughout this project. A special thanks is due to E. R. Jaycox who pro-
vided the pollen.

This investigation was completed as part of a dissertation submitted to
the University of Lllinois, Urbana-Champaign, in partial fulfillment for
the Ph.D. degree. The Department of Entomology and the University Re-
search Board, University of Illinois, Urbana provided financial assistance.

Literature Cited

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Becher, E. 1882. Ueber die ersten Stinde einiger Syrphiden und eine neue
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Britten, H. 1917. Mallota cimbiciformis, Fln., bred from rotten wood. Trans.
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Coe, R. L. 1953. Mallota cimbiciformis Fallen (Diptera: Syrphidae) breeding in
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VOLUME 80, NUMBER 3 439

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and Son, London. 591 pp.

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Wahl, B. 1900. Ueber das Tracheensystem und die Imaginalscheiben der Larve
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Waldbauer, G. P., and J. K. Sheldon. 1971. Phenological relationships of some

440 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

aculeate Hymenoptera, their dipteran mimics, and insectivorous birds. Evolution
25:371-382.

Waldbauer, G. P., J. G. Sternburg, and C. T. Maier. 1977. Phenological relationships
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Wichard, W., and H. Komnick. 1974. Feinstruktur und Funktion der Analpapillen

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1-10.

Department of Entomology, University of Illinois, Urbana, Illinois
61801.

Present address—Department of Entomology, Connecticut Agricultural
Experiment Station, P. O. Box 1106, New Haven, Connecticut 06504.

PROC. ENTOMOL. SOC. WASH.
80(3), 1978, pp. 441-455
OVIPOSITION AND HOST FEEDING BEHAVIOR OF
APHELINUS ASYCHIS (HYMENOPTERA: CHALCIDOIDEA:
APHELINIDAE) ON SCHIZAPHIS GRAMINUM
(HOMOPTERA: APHIDIDAE) AND SOME
REACTIONS OF APHIDS TO THIS PARASITE

Hillary Boyle and Edward M. Barrows

Abstract.—Fifty-four Aphelinus asychis (Walker) were observed during
their first encounters and ovipositional attacks with unparasitized Schizaphis
graminum (Rondani). Oviposition occurred when wasps inserted their ovi-
positors in aphids for at least 60 s, and aphids were not paralyzed prior
to or during ovipositor insertion. Before leaving aphids, 65% of the wasps
made single ovipositor eversions (SE’s), 35% made multiple ovipositor
eversions (ME’s). Everted ovipositors did not always pierce aphids. Single
eversions (as opposed to ME’s) preceded oviposition in 75.7% of 33 ob-
servations. Durations of ovipositor eversions which preceded oviposi-
tion were significantly longer than durations of eversions that did not
precede oviposition. Twenty wasps were observed host feeding. Prior
to feeding, wasps stung and paralyzed aphids. Oviposition and host feeding
were discrete activities always carried out on separate aphids. Wasps’ initial
approaches and attacks on aphids, however, were similar in both types of
attack. Wasps took significantly longer to sting aphids than to oviposit.
Aphid reactions to wasps included body jerking, antennating, walking
away and kicking with one leg. Aphids reacted violently to stinging and
some ovipositional attacks. Alarm behavior in some aphid groups oc-
curred.

This study was conducted in an attempt to identify behaviors involved
in oviposition and host feeding of a hymenopterous parasite, Aphelinus
asychis (Walker), on the greenbug, Schizaphis graminum (Rondani). In
addition, some reactions of aphids to wasp presence and attacks were
noted.

Aphelinus asychis was first introduced into the United States in 1968
to help control Schizaphis graminum which was first found in the United
States in 1882 and presently causes millions of dollars damage annually to
grain crops, e.g., Sorghum vulgare (Pers.).

Although Aphelinus asychis is a potentially valuable biocontrol agent
against Schizaphis graminum, there has never been a detailed study of its
Oviposition and host feeding behavior. The only descriptions of host
feeding (Esmaili and Wilde, 1972; Cate et al., 1973) are brief, lack detail

as to the frequency of various behaviors and are inconsistent as to the

442 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

duration of ovipositor insertion. Oviposition by A. asychis has been observed
(Rogers, 1976) but never described in detail.

Materials and Methods

Sorghum vulgare was grown in 355 cm?* plastic pots in a greenhouse.
When about 10 cm tall, plants were transferred to a separate area and
infested with Schizaphis graminum.

Pots with about 4 plants and 10 aphids per plant were transferred to the
laboratory and placed in 42,000 cm* terraria. Each terrarium held 26-32
pots. Terraria were fitted with clear plexiglass tops, each with 2 ventilation
holes covered with nylon mesh and 1 access hole with a 60 cm long nylon
mesh sleeve which was tied closed when not in use. Several Aphelinus
asychis were placed in some terraria and allowed to feed on and parasitize
aphids. All terraria were kept under plant-growth fluorescent bulbs for
12 h per day. Laboratory temperature was from 20.0-26.5°C.

Gelatin capsules (no. 1) were used to capture and carry individual adult
wasps and to isolate and hold individual mummies. This reduced or
eliminated possible injury to wasps which frequently occurs when using
a camel's hair brush.

A total of 54 wasps was observed for 1 h each at about 24°C. To
observe oviposition and host feeding of A. asychis on S. graminum, a
blade of Sorghum, with a minimum of three feeding aphids, was placed in
a plastic petri dish; one A. asychis female was also placed in the dish
which was then covered. Observations were made through a dissecting
microscope. All aphids used for observations were taken from unpara-
sitized cultures. This method is similar to that of Goff and Nault (1974).

Behavior that was observed included oviposition host feeding and re-
actions of aphids to the presence of and attacks by wasps. Attacks were
timed with a stopwatch from the moment the wasp turned and everted her
ovipositor to the moment she retracted it.

In addition, 16 aphids were dissected to determine whether oviposition
actually took place when a wasp’s ovipositor pierced the aphid. Five aphids
were dissected after having been fed on to determine whether oviposition
occurred.

In reporting results, the symbol x is used to denote a mean; M, median;
TEP test for the equality of two percentages (Sokal and Rohlf, 1969);
MWUT, Mann-Whitney U-test; and WMPT, Wilcoxon’s matched pair
test. All numerical data are based on randomly-selected animals and
observations.

Results and Discussion

Ovipositional attacks —In “typical” ovipositional attacks, Aphelinus asy-
chis walked on substrates and tapped surfaces with extended antennae.

VOLUME 80, NUMBER 3 443

On encountering Schizaphis graminum by touching or coming within
about 1 cm of them, wasps either retreated or stopped and withdrew
antennae, leaving only flagellae slightly extended and quivering. If aphids
were not within ovipositor reach, wasps slowly approached and antennated
them with only flagellae extended.

When they were sufficiently close to aphids, wasps began to “sway,”
rocking their bodies from side to side. Occasionally, swaying was inter-
rupted several times for a few seconds each time while wasps lowered
antennae and remained motionless. Swaying was occasionally accompanied
by wasps’ grooming their metasomas with hind legs, grooming hind legs
with middle legs or both. Wasps also either turned their heads slightly
from side to side or kept them still.

After approaching and facing aphids, wasps rapidly turned about 180°
to the left or right, everted ovipositors and stabbed at aphids. If ovipositors
did not make immediate contact with aphids, wasps usually made one or
more stabbing attempts.

When ovipositors pierced aphids, wasps stood motionless and kept an-
tennae lowered for the duration of ovipositor insertion (Fig. 1). Occasionally,
wasps made slight thrusts with ovipositors.

Oviposition occurred when everted ovipositors pierced and were in-
serted in aphids for at least 60 s; this was determined by dissecting 16 aphids
immediately after ovipositional attacks in which everted ovipositors were
in aphids from 4-542 s. Immediately before retracting ovipositors, wasps
raised heads and extended antennae slightly. After wasps oviposited
and retracted ovipositors, they retreated.

Although there are no other published descriptions of oviposition for
Aphelinus asychis, several aspects of this behavior are similar to those
reported for other parasitic hymenopterans. Observations suggest that
A. asychis usually touches or is within 1 em of aphids before attacks are
initiated. Messenger (1967), reported that Praon exsoletum (Nees) detected
hosts only after actually contacting them with antennae, and he concluded
that the wasp’s distance perception was limited. Similar observations have
been made with Lysephlebus testaceipes (Cresson) (Webster and Phillips,
1912), Tetrastichus flavigaster Brothers and Moran (Moran et al., 1969)
and Aphelinus flavus Thompson (Hamilton, 1973). As in most A. asychis
observed in this study, females of each of the above species appeared to
encounter hosts during apparently “nondirected” wanderings and react to
hosts after contact.

Wasps probably gain sensory information as to host suitability and
appropriateness of ovipositor insertion sites when they antennate aphids
and sway prior to ovipositor eversion. Webster and Phillips (1912) ob-
served A. nigritus Howard approach, antennate, and sway in front of
Schizaphis graminum in a manner similar to that of Aphelinus asychis. An-
tennation of hosts prior to ovipositor eversion has also been reported for

444. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

wel

Fig. 1. Everted ovipositor of Aphelinus asychis female piercing Schizaphis graminum.

A. mytilaspidis (Marchal, 1909), Praon aguti Smith, Lysephlebus testaceipes
(Sekhar, 1957), Tetrastichus flavigaster (Moran et al., 1969), and Aphelinus
flavus (Hamilton, 1973). Moran et al., (1969) presumed that antennation
functions in olfactory identification of the host by the wasp. Chalcidoid
antennae have sense organs necessary for host finding, according to
Askew (1971). After their ovipositors are inserted into hosts, sense organs
located on ovipositors probably supply additional information to wasps
concerning host suitability (Moran et al., 1969; Askew, 1971).

The following results, except where otherwise indicated, are based on
the first observed ovipositor eversions, either single or multiple, which each
of 54 wasps made with a different, unparasitized aphid.

When attacking aphids, wasps exhibited either single ovipositor ever-
sions (SE’s) or multiple ovipositor eversions (ME’s); ME’s were comprised
of from 2-14 eversions. Everted ovipositors did not always pierce or
even contact aphids, and piercing was not always followed by oviposition.
In SE’s which they made after approaching aphids, wasps turned about
180° away from aphids, everted and retracted ovipositors once, and re-
treated. In ME’s, wasps turned 180° away from aphids, everted and re-

VOLUME 80, NUMBER 3 445

tracted ovipositors, turned 180° to face aphids, turned 180° again, everted
and retracted ovipositors a second time; this sequence of 3 turns was
repeated from 3-14 times before wasps retreated. Figure 2 shows sequences
of behaviors displayed during first encounters of 54 A. asychis with 54
Schizaphis graminum during | h observation periods.

Single ovipositor eversions, as opposed to ME’s, occurred 64.8% of the
time, and SE’s preceded oviposition in 75.7% of 33 observations. This
suggests that Aphelinus asychis budgeted time and energy efficiently in
that they usually pierced Schizaphis graminum only once before oviposit-
ing. Wasps that pierced aphids several times before ovipositing may have
been attempting to gain additional information as to whether or not aphids
were already parasitized, to place their ovipositors in more favorable sites
within aphids, or both. Wasps that pierced aphids but failed to oviposit
may have received stimuli indicating that aphids were unsuitable for
parasitization. If this is the case, evaluation and rejection of unsuitable hosts
could be accomplished in a relatively brief time of 2 to about 59 s.

Of the total time spent in everting ovipositors, 35 wasps spent 94.0% of
this time in SE’s preceding oviposition; 19 wasps spent 51.6% of this time
in ME’s preceding oviposition. There is a significant difference between
these groups (p = 0.0003, TEP). In SE’s, wasps pierced, presumably evalu-
ated the suitability of aphids and oviposited in significantly less total time
than in ME’s.

Praon exsoletum exhibits similar behavior. In studying its daily oviposi-
tional pattern, Messenger (1967) found that P. exsoletum oviposited from
18-24 times in 20-30 min, making one ovipositor insertion per aphid. After
a period of rapid ovipositing, however, wasps repeatedly struck at aphids be-
fore ovipositing. As in Aphelinus asychis, these ME’s consumed more time
and preceded fewer ovipositions than SE’s. A study to determine the daily
ovipositional pattern of A. asychis could be helpful.

Durations of SE’s which preceded oviposition were not significantly
different from durations of the last eversion of ME’s which preceded
oviposition (U = 94, p > 0.05, N, = 20, No = 8, MWUT). Nor were SE's
which did not precede oviposition significantly different from durations of
ME’s which did not precede oviposition (U = 58, p > 0.05, N; = 10, Ns =
11, MWUT).

Durations of SE’s which preceded oviposition were significantly longer
than durations of SE’s which did not precede oviposition (U = 198, p <
0.002, Ni = 20, N2 = 10, MWUT). Durations of ME’s which preceded
oviposition were also significantly longer than durations of ME’s which did
not precede oviposition (U = 88, p < 0.001, N; = 11, No = 8, MWUT).

Wasps made initial approaches to aphids from the front, rear or either
side. In ME’s, wasps made each ovipositor eversion from the same position
relative to the aphid. Rear approaches seemed to precede oviposition more

446 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

wasp encoun
aphid

wasp antennates
aphid

19 wasp oOvipositor
misses aphid

v2

wasp retracts
ovipositor, turns
180~-, sways, turns,
everts ovipositor,

[ ces aphid,
Pali ES: cee ae

2 3

wasp

retracts ovipositor,

turns, sways, turns,

everts ovipositor,
pierces aphid,

a oviposits

oviposits

$24

wasp retracts
ovi

positor 2

VASD
Ryan retracts ovipositor,
retracts vipositor, turns, sways, turns,
turns, sways, turns, 1 everts ee
byte eis Gneew misses aphid
everts ovipositor,
pierces aphid, retracts ovipositor,
fails t oviposit , turns, sways, turns,
retracts ovipositor, everts ovipositor ,
turns , ys’ tEurns Bie ees aphid ,
everts ovipositor Oviposits 6
misses. aphid,
retracts ovipositor,
turns, sways, turns,
everts ovipositor, wasp

pierces aphid,
fails

retracts ovipositor,

turns 5 Sway ae turns 5
everts ovipositor,
misses aphid,

(from 1 to 6
more times)

to oviposit

Fig. 2. Behaviors exhibited during first encounters of 54 different Aphelinus
asychis with 54 different Schizaphis graminum. Numbers by arrows indicate frequencies
of changes from one behavior (or sequence of behaviors) to another. Question mark
indicates that frequency was not noted. See text for further explanation.

fond

VOLUME 80, NUMBER 3 447

often than front or lateral approaches, but sample sizes were too small
to allow statistical analysis.

Twenty-six wasps were observed for the direction of turn immediately
prior to everting ovipositors in SE’s and ME’s. In ME’s, as noted above,
wasps turned about 180° to face aphids after retracting ovipositors, and
then they turned again immediately prior to everting ovipositors. They did
this two or more times. Turns made immediately prior to ovipositor eversion
were in the alternate direction (e.g., right, left or left, right) to the previous
turn prior to ovipositor eversion in 53.8% of the cases; 46.1% of such turns
were in the same direction (e.g., right, right or left, left) as the previous
turn observed. There was no significant difference in turning direction
compared to a hypothetical turning of either right or left 50.0% of the time
(0:9 > 0:5, TEP).

Stinging and host feeding.—Host feeding is common among chalcidoids
and represents an example of economy in time and effort, in that individual
hosts provide food for both adults and progeny (Askew, 1971). Esmaili and
Wilde (1972) and Cate et al. (1973) found that host feeding by newly
emerged A. asychis almost always preceded oviposition.

Nutrients obtained from host feeding by Aphelinus and other genera are
probably necessary for oogenesis (DeBach, 1943; Moran et al., 1969; Askew,
1971). Oogenesis occurs throughout the adult life of A. semiflavus (Howard)
(Schlinger and Hall, 1959) as it probably does in A. asychis. Therefore,
periodic host feeding is probably necessary to obtain nutrients for con-
tinuous oogenesis.

Twenty A. asychis were observed feeding on 20 different, unparasitized
Schizaphis graminum. Wasps stung and paralyzed aphids, then they fed
from wounds made by ovipositors.

Stinging attacks were initially indistinguishable from ovipositional at-
tacks. Wasps approached and antennated aphids, swayed, turned, everted
ovipositors and pierced aphids in apparently the same manner as in oviposi-
tional attacks. Wasps presumably injected, through their ovipositors, a
paralyzing agent into aphids.

The first noticeable difference in wasp behavior during stinging attacks, as
compared to ovipositional attacks, occurred after aphids were effectively
paralyzed and their movements merely uncoordinated leg waving. Wasps
(with their ovipositors still inserted in aphids) usually backed up slightly and
stood with their hind legs on aphids. Wasps remained in this position,
motionless, except for occasional thrusts of ovipositors and with antennae
lowered, for several more minutes.

As in ovipositional attacks, immediately prior to retracting ovipositors,
wasps raised their heads and slightly extended their antennae. Wasps
then immediately made another stinging attack on the same aphids or an-

448 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tennated aphids to locate wounds made by ovipositors. If they did not
locate wounds, wasps made additional stinging attacks, usually of shorter
duration than the first ones.

When wasps found wounds, they antennated them for several seconds,
then positioned their heads over wounds, lowered their bodies and fed.
Feeding continued for several minutes and usually took longer than stinging.
Wasps either fed once and retreated or alternately stung and fed from the
same aphid before retreating.

Figure 3 shows the behavioral sequence of stinging attacks and feeding
by 20 Aphelinus asychis on 20 Schizaphis graminum. This sequence begins
with wasp ovipositor insertion into aphids; behavioral sequences occurring
before this are similar to those for ovipositional attacks.

Initial approach to and attack of hosts is similar for both oviposition and
stinging in Aphelinus asychis, A. flavus (Hamilton, 1973), A. semiflavus
(Schlinger and Hall, 1959), A. mytilaspidis (Marchal, 1909) and Tetrastichus
flavigaster (Moran et al., 1969). In our study, after initiation of stinging
attacks by Aphelinus asychis, the first noticeable difference in behavior
compared to ovipositional attacks was their placing hind legs on aphids. This
has also been observed by Esmaili and Wilde (1972). The function of this
behavior has not been studied, but it might allow wasps to thrust ovipositors
deeper into aphids.

Paralyzing aphids by A. asychis prior to their feeding probably facili-
tates this activity. Unparalyzed aphids would not be likely to remain still
while wasps fed at their wounds.

Wasps were never observed paralyzing aphids in which they had ovi-
posited, nor has this been reported in the literature for A. asychis. Para-
sitized aphids remain alive and mobile for several days before mummifi-
cation and thus provide one means for wasp dispersal.

Of the 20 wasps observed host feeding, 11 approached aphids from the
front, 7 approached from the rear and 2 approached from the sides of
aphids. All subsequent stinging attacks and feeding were made from the
approach position. In their first stinging attacks on aphids, 10 wasps turned
to the right and 10 turned to the left prior to ovipositor eversion.

In attacks by 18 wasps, durations from the beginning of the initial
paralyzing sting to when wasps placed hind legs on aphids were from
210-570 s (x = 330, M = 390). Esmaili and Wilde (1972) reported durations
of from 120-180 s for this activity. Wasp hind leg placement on aphids
always occurred after their paralysis.

Durations of initial paralyzing stings, before wasps either began feeding
or attempted to find sites of ovipositor insertion, were from 277-1,399 s
(x = 598, M = 838, N = 20). These durations were significantly longer
than durations of SE’s which preceded oviposition (U = 395, ny = 20,
nz = 20, p < 0.001, MWUT). Thus, wasps took significantly longer to para-

VOLUME 80, NUMBER 3 449

wasp ovipositor
erces and stings aphid

wasp places hind a a was

asp
legs on paralyzed remains still
aphid y 9
wasp
4 feeds on aphid
wasp _ wasp
stings aphid |D stings aphid
ee ihe roe 2 wasp
feeds on aphid
wasp ¥v oy
wasp ——— SSIS wasp
stings ——— stings aphid
wasp
wasp feeds on apl
feeds on aphid 1 Lt
wasp was
stings aphid Lateealiae stings
and feeds on acid
times

Fig. 3. Behaviors exhibited during first encounters of 20 different Aphelinus asychis
with 20 different Schizaphis graminum during wasp stinging and feeding attacks on
aphids. Numbers by arrows indicate frequencies of changes from one behavior
(or sequences of behaviors) to another. See text for further explanation.

lyze aphids than to oviposit. Cate et al. (1973) reported that the initial sting
of A. asychis lasted from 300-7,200 s, a much wider range than observed in
our study. Hamilton (1973) reported that the average duration for sting-
ing by A. flavus is 380 s.

After the initial paralyzing sting, most wasps fed and then retreated.
Eight wasps, however, stung aphids one or more times after either feeding
or being unsuccessful in finding sites of ovipositor insertion; these wasps
alternated stinging attacks with feeding. These subsequent stinging attacks

450 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

on the same aphids lasted from 39-821 s (¥ = 399, M = 330). Statistical anal-
ysis suggests that initial stings were longer than subsequent stings (T = 6,
N = 8, p = 0.055, WMPT). The average duration of all stinging attacks,
both initial and subsequent, was 757 s.

Repeated and alternating stinging and feeding has been reported for
A. asychis (Cate et al., 1973), A. mytilaspidis and Tetrastichus xantho-
melaenea (Rond.) (Marchal, 1905). Initial stings made by A. asychis on
aphids seemed to be sufficient to paralyze them and were significantly
longer than subsequent stings on the same aphids. Marchal (1905) and
Cate et al. (1973) also noted that repeated stings were often quite brief.
Subsequent stings might serve to open other wounds in aphids or to en-
large existing wounds rather than to inject paralyzing agent into aphids.

Wasp initial feedings lasted from 141-1,815 s (x = 680, M = 978, N =
20). Durations of subsequent feedings by 5 wasps were from 12-2,916 s
(t = 1,083, M = 1,464). The average duration of all feedings was 952 s.
There was no statistical difference between durations of all stinging attacks
and all feedings (T = 83, N = 20, p > 0.05, WMPT).

The paralyzing agent injected by wasps seemed to cause a number of
changes in the appearance and behavior of stung aphids. These changes
began to occur several minutes after ovipositor insertion. Usually, aphid
abdomens swelled and raised, causing the aphid hind legs and occasionally
the middle legs to be lifted off substrates. Hind legs waved in a weak, un-
coordinated manner in the air. Waving became progressively slower and
weaker until it eventually stopped.

At the same time, aphid antennae, which were usually held against dor-
sums, raised slightly and extended at various angles to aphid bodies. Aphids
were usually prevented from falling by keeping their stylets embedded in
Sorghum blades. In general, paralyzed aphids appeared completely to
lose control of their movements, and they became immobilized and defense-
less. Esmaili and Wilde (1972) reported similar changes in the appearance
and activity of paralyzed Schizaphis graminum.

While wasps were feeding, aphid color changed from the usual green
to yellow-green to yellow and finally to brown, a process which took
from 20-45 min. The paralyzing agent, or substances introduced during
wasps feeding or both, might bring about this gradual color change,
which was also noted by Cate et al. (1973) in the same host. When wasps
finished feeding, aphids were shriveled and shrunken.

Wasps often had dark, flattened abdomens prior to host feeding. Dur-
ing feeding, however, abdomens swelled and became rounded and honey-
colored. This process roughly corresponded to the gradual shriveling of
aphids.

Dissection of 5 aphids that were fed upon Aphelinus asychis did not

VOLUME 80, NUMBER 3 451

reveal any eggs deposited by wasps. Marchal (1909) and Esmaili and
Wilde (1972) reported similar findings and stated that host feeding
renders hosts unsuitable for oviposition. Full larval development of A.
asychis in the dried, shrunken remains of fed-on aphids would seem dif-
ficult.

Aphid reaction to wasps.—Aphids reacted in a variety of ways and
seemed to have some effective defense mechanisms against wasps. The
most conspicuous defense reaction by aphids was “jerking.” Aphids, with
stylets embedded in Sorghum blades, quickly and forcefully flipped up their
abdomens and lifted their hind and middle legs off the substrate. This was
often done continuously for several minutes. Almost always, after one
aphid began jerking, nearby aphids would also begin. A Sorghum blade often
had several small groups of aphids in constant movement of this kind.

Other reactions of aphids included quickly extending antennae forward
to touch wasps, crawling away from wasps and striking with legs at wasps.

Jerking, antennating, crawling away and kicking have been reported
for several aphid species, including Schizaphis graminum, by Webster
and Phillips (1912) and Hamilton (1973). These reactions seemed to
have varying degrees of effectiveness as defense mechanisms depending
on how soon they were initiated.

Jerking by S. graminum seemed to prevent attacks when it was initiated
before or soon after wasps approached. Other aphids jerked after one
started and this probably helped these other aphids from being attacked.
This type of behavior was also noted by Hight et al. (1972).

Wasps apparently required or preferred stationary hosts, as they were
never observed initiating attacks on aphids that were not still when they
were approached. Jerking by aphids before wasps inserted ovipositors was
usually effective in preventing contact because wasps seldom hit moving
aphids with their ovipositors.

Aphid defense behaviors seemed to be stimulated by various kinds of
wasp activity. On 5 occasions, groups of several aphids antennated a wasp
and began jerking when the wasp walked past the aphids, they did not
touch or give any noticeable indication of attacking them. On 29 oc-
casions, wasps, which approached aphids and appeared about to attack
them, seemed to provoke reactions from aphids which seemed to cause
wasps to retreat without attacking. These reactions were all of the types
mentioned above.

Although many aphids showed no noticeable reaction to ovipositional
attacks, some aphids jerked strenuously after being pierced. Once jerking
was initiated, it usually continued for several minutes, even after wasps
retreated. Jerking after wasps inserted ovipositors seemed to have little
or no effect on the outcome of the attacks, either ovipositional or stinging

452 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

attacks. Aphids often completely lifted wasps off the substrate, but wasp
ovipositors remained in the aphids and wasps remained as still as possible,
never raising antennae until just prior to retracting ovipositors.

The most violent and prolonged reactions by aphids were caused by
wasp stinging attacks. In every stinging attack observed, aphids began
jerking violently within from 2-5 min after ovipositor insertion. Aphids
continued violent, strenuous movement for several minutes until they
gradually began to show signs of paralysis. Frequently, aphids twisted
around, using their stylets embedded in Sorghum as pivots. In 8 stinging
attacks, aphids removed stylets from Sorghum and rapidly walked away,
dragging wasps with them. Wasps were never dislodged by these activities,
although they were often dragged several centimeters or flipped over.

Alarm behavior was observed during 4 stinging attacks; immediately
after a stung aphid began jerking, all nearby aphids withdrew their stylets
from Sorghum and quickly scattered. On 3 occasions, stung aphids jerked
violently and produced yellowish droplets from the tips of both cornicles,
after which all nearby aphids began jerking and then scattered. An
alarm pheromone in the cornicle secretions of Schizaphis graminum and
other aphids was demonstrated by Bowers et al. (1972) who identified the
active substance as trans-B-farnesene. Kislow and Edwards (1972) noted
interspecific reactions to the pheromone which is detected by sensoria
located on aphid antennae (Nault et al., 1973). Aphid cornicle secretion is
considered a self-serving and altruistic defensive system by Nault et al.
(1976).

One wasp placed her right foreleg in a cornicle droplet while she was
feeding, and the droplet apparently hardened around her leg. The wasp
struggled for about 45 min before freeing herself. This has also been
noted by Edwards (1966) who found cornicle secretions to be lipid droplets
in water that rapidly crystallizes when in contact with a solid surface. Ed-
wards (1966) found only 2 wasps entrapped in secretions produced by
several thousand aphids; Goff and Nault (1974) never observed wasps
entrapped by cornicle secretions. Our study also suggests that cornicle
secretions rarely entrap wasps.

Schizaphis graminum probably detect approaching Aphelinus asychis
partly through vision. Dixon (1958; 1973) stated that aphids would be at
an advantage when predators approached from the front because they could
be seen more easily. This study supports Dixon’s supposition because al-
though more wasps approached aphids from the front, frontally approached
aphids often reacted defensively, and a larger percentage of rear ap-
proaches preceded oviposition.

Schizaphis graminum probably also use tactile perception to detect
wasps. Aphids which did not show any other reaction to wasp often
jerked only after wasps pierced them. This did not seem to affect the out-
come of the attacks, but it often caused nearby aphids to begin jerking.

VOLUME 80, NUMBER 3 453

Additional observations — Two variations of behavior associated with
ovipositional attacks were occasionally observed. Five wasps approached
single aphids, stopped and briefly antennated them. Then, instead of
swaying, they lowered their antennae and quickly butted the fronts of
their heads against the aphids. Two wasps butted aphids once, and 3
wasps made 2 such charges in rapid succession. All wasps then continued
with the usually observed sequence of turning, everting ovipositors and stab-
bing at aphids. Wasps exhibiting this type of behavior appeared to move
more quickly and to take less time antennating aphids than other wasps,
and in general, appeared more agitated than usual.

Five other wasps “attacked” either cast skins of aphids (2 wasps), aphid
eggs (2 wasps) or a spot on a Sorghum blade where an aphid had recently
been feeding (1 wasp). In all cases, the wasps showed the usual behavioral
sequence prior to Ovipositor eversion. One wasp made a SE and then
retreated. All others made from 3-14 eversions before retreating.

Factors that initiate attacks probably includes wasps’ motivation to
either oviposit or feed and stimuli from aphids, e.g., odors, sizes, shapes.
Wasps that are highly motivated to attack probably do so at lower stimulus
thresholds than wasps that are less highly motivated (Sekhar, 1957). This
may explain wasp attacks on aphid skins, eggs and a recently vacated
feeding site. Each of these objects may have carried sufficient aphid odor,
had other characteristics or both, to stimulate highly motivated wasps to
attack. Similar wasp behavior has been reported for Aphelinus nigritus
(Webster and Phillips, 1912) and Tetrastichus flavigaster (Moran et al.,
1969).

Wasps butting their heads against aphids rather than swaying prior to
turning and everting ovipositors might also be a reflection of the motiva-
tional state of the wasps. Swaying often consumed several seconds and,
as noted, was often interrupted several times while wasps remained motion-
less. Wasps that are highly motivated to attack may have eliminated this
relatively time-consuming behavior and replaced it with a far more rapid
action.

Acknowledgments

The authors thank Roger D. Eikenbary (Oklahoma State Univ.) for pro-
viding Sorghum seeds and live specimens of Aphelinus asychis; Robert
Schroeder (SEA, USDA) for providing Schizaphis graminum; Donald M.
Spoon (Georgetown Univ.) for making available photographic supplies and
facilities; and Gordon Gordh (formerly SEA, USDA) for encouraging this
study, reading a draft of this paper and making numerous helpful sug-
gestions. Lawrence S. Oliver (Georgetown Univ.) also made important
suggestions regarding a preliminary draft of this paper. The first author
gives special thanks to Irving Gray (Georgetown Univ.), Roger Cressey
(Smithsonian Institution), and the late Beulah Howatt McGovern (Dun-

454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

barton College) for their help and encouragement. Janice B. Browne.
helped with the preparation of the manuscript.

Literature Cited

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Bowers, W. S., L. R. Nault, R. E. Webb, and S. R. Dutky. 1972. Aphid alarm
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Cate, R. H., T. L. Archer, R. D. Eikenbary, K. J. Starks, and R. D. Morrison. 1973.
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the parasitoid on aphid mortality. Environ. Entomol. 2:549-553.

DeBach, P. 1943. The importance of host feeding by adult parasites in the reduction
of host populations. J. Econ. Entomol. 36:647-658.

Dixon, A. F. G. 1958. The escape responses shown by certain aphids to the presence

of the coccinillid Adalia decempunctata (L.). Trans. R. Entomol. Soc. London

110:319-334.

1973. Biology of aphids. London: Edward Arnold, Ltd. 55 pp.

Edwards, J. S. 1966. Defense by smear: Supercooling in the cornicle wax of aphids.
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Esmaili, M., and G. Wilde. 1972. Behavior of the parasite Aphelinus asychis in re-
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Goff, A. M., and L. R. Nault. 1974. Aphid cornicle secretions ineffective against
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Hamilton, P. A. 1973. The biology of Aphelinus flavus (Hym. Aphelinidae) a parasite
of the sycamore aphid Drepanosiphum platanoides (Hemipt. Aphididae). En-
tomophaga 18:449-462.

Hight, S. C., R. D. Eikenbary, R. J. Miller, and K. J. Starks. 1972. The greenbug and
Lysephlebus testaceipes (Cresson). Environ. Entomol. 1:205—209.

Kislow, C. J., and L. J. Edwards. 1972. Repellent odor in aphids. Nature 235:
108-109.

Marchal, P. 1905. Observations biologiques sur un parasite de la galerque de l’orme, le
Tetrastichus xanthomelaenae (Rond.). Soc. Ent. de France Bull. (Translation
in Howard, 1908).

—. 1909. La ponte des Aphelinus de Vinteret individuel dans les actes lies
a la conservation de ?Academie des Sciences de Paris 148:1223-1225. (Transla-
tion in Howard, 1910.)

Messenger, P. S. 1967. Bioclimatic studies of the aphid parasites Praon exsoletum:
1. Effects of temperature on the functional response of females to varying host
densities. Can. Entomol. 100:728—741.

Moran, V. C., D. J. Brothers, and J. J. Case. 1969. Observations on the biology of
Tetrastichus flavigaster Brothers and Moran (Hymenoptera: Eulophidae), para-
sitic on psyllid nymphs (Homoptera). Trans. R. Entomol. Soc. London 121:
41-58.

Nault, L. R., J. L. Edwards, and W. E. Styer. 1973. Aphid alarm pheromones: Se-
cretion and reception. Environ. Entomol. 2:101—105.

Nault, L. R., M. E. Montgomery, and W. S. Bowers. 1976. Ant-aphid association:
Role of aphid alarm pheromone. Science 192:1349-1351.

Rogers, C. E. 1976. The greenbug. Insect World Digest 3:12—15.

Schlinger, E. I., and J. C. Hall. 1959. <A synopsis of the biologies of three im-
ported parasites of the spotted alfalfa aphid. J. Econ. Entomol. 52:154—157.

VOLUME 80, NUMBER 3 455

Sekhar, P. S. 1957. Mating, oviposition and descrimination of hosts by Aphidius
testaceipes (Cresson) and Praon aguti Smith primary parasites of aphids. Ann.
Entomol. Soc. Amer. 50:370-375.

Sokal, R. R., and F. J. Rohlf. 1969. Biometry. San Francisco: W. H. Freeman
and Co. 776 pp.

Webster, F. M., and W. J. Phillips. 1912. The spring grain-aphis or greenbug.
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Department of Biology, Georgetown University, Washington, D.C. 20057
(HB mail address: 313-A Broad Run Drive, Sterling, Virginia 22170).

456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
BOOK REVIEW

Taxonomic Study of the Cicadellinae (Homoptera: Cicadellidae). Part 2.
New World Cicadellini and the genus Cicadella. David A. Young. 1977.
1135 pp., 905 figs. North Carolina Agricultural Experiment Station Tech-
nical Bulletin 239. Available for $15. (post paid) from Publications, 318
Ricks Hall, North Carolina State University, Raleigh, North Carolina
27607.

This is the 2nd of 3 volumes of the most ambitious and comprehensive
taxonomic revision ever attempted on a world-wide basis for a large por-
tion of the family Cicadellidae, itself one of the largest families of the
Insecta. Part 1, Proconiini (1968. U.S. Nat. Mus. Bull. 261, 287 pp.) to-
gether with Part 2 complete the subfamily for all of the Americas. Part
3, now in preparation, will treat all of the Old World species.

The Cicadellini, especially in the Neotropics, comprises many species
with bright and striking color patterns. Early describers, often in scattered
and usually unrelated works, had field days with descriptions of hundreds
of new species. No one before Dr. Young was really successful in bringing
together and integrating all of the taxonomic data concerning the New
World Cicadellini within a lifetime; although Signoret (1816-1889) and
Melichar (1856-1924) made partial efforts toward this goal.

Within the present work, Dr. Young provides keys and descriptions for
155 genera, of which 91 are new. He recognizes over 1,100 valid species,
of which 390 are new. The keys to genera and species use characters
which are well illustrated in the text by over 900 figures or line drawings.
Treatment of the previously described species is frugal but highly adequate.
The 1,135 pages of this volume are entirely free of extraneous matter. In
fact, the format is a model of excellence, one in which the data are con-
centrated and concise.

In order to truly appreciate the author’s magnificent effort, one must be
familiar with the confusion and uncertainty in almost all of the past litera-
ture dealing with hundreds of species from Central and South America.
Dr. Young has established both a solid classification upon which others
can build in the future and a system for identification which can be used
today by those concerned with plant-feeding insects. Simply stated, this
is the most outstanding and comprehensive single volume ever produced
on a large segment of the Cicadellidae.

James P. Kramer, Systematic Entomology Laboratory, ITBIII, Fed. Res.,
Sci. Educ. Admin., USDA, c/o U.S. National Museum, Washington, D.C.
20560.

VOLUME 80, NUMBER 3 457
SOCIETY MEETINGS

844th Regular Meeting—6 October 1977

The 844th Regular Meeting of the Entomological Society of Washington
was called to order by President-Elect Sutherland at 8:05 PM on 6 October
1977 in the Ecology Theater of the National Museum of Natural History.
Thirty-six members and 9 guests were present. The minutes of the April
meeting were read and approved.

Membership Chairman Utmar read for the first time the names of the
following new applicants for membership:

W. Nelson Beyer, Patuxent Wildlife Research Center, Laurel, Mary-
land.

Jack L. Boese, 900 Madison Ave., Baltimore, Maryland.

LeMar M. Chilson, 2016 Seattle Ave., Silver Spring, Maryland.

T. P. Copeland, East Tennessee St. Univ., Johnson City, Tennessee.

James H. Kennedy, 105 Tee St., Blacksburg, Virginia.

Brian R. Pitkin, Dept. of Entomology, British Museum (Natural History),
Cromwell Road, London.

President Ramsay was excused due to illness.

Editor Stoetzel announced that Former President George Steyskal’s
“Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae)”
was recently published by the Society and is on sale for $1.50/copy. She
also announced that the October issue of the Proceedings will be about
166 pages and asked the membership for any comments concerning the new
format. Based on the first issue using the new format, the Society saved
about 6%.

President-Elect Sutherland reported that the audit and nominating
committees will be appointed soon. He also announced that Mr. Muesebeck,
a Past President and long-time supporter of the Society had made a con-
siderable donation to the special publications fund.

Don Davis called attention to the new Naturalists Center of the
Museum. He explained the function of the Center, which is to help
serious amateurs, and suggested that the Society might want to hold a meet-
ing there and help with the entomology section.

Joyce Utmar discussed briefly the Maryland Center for Systematic En-
tomology. Don Messersmith further commented on the new program, which
is a cooperative effort by scientists of the USDA, Smithsonian and the
University of Maryland to provide training in systematics.

The main speaker for the evening was Dr. David Inouye, Assistant Pro-
fessor of Zoology, University of Maryland. Dr. Inouye spoke on coevolu-
tion between ants and plants. After reviewing examples from tropical
climates, Dr. Inouye described his research on three species of temperate

458 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Helianthella (Asteraceae) and Formica ants. The presentation was illus-
trated by many excellent Kodachrome slides and was followed by a lively
question and answer period.

Notes and Exhibitions

Manya Stoetzel reviewed a new book entitled “Survey of the World
Aphids” by V. F. Eastop and D. Hille Ris Lambers.

Chris Thompson showed slides of a recent trip to Europe, particularly —

of the Paris and Vienna Museums.

Helen Sollers-Riedel announced that the new Mosquito Catalogue by
Ken Knight and Alan Stone was recently published and Doug Sutherland
showed a copy of Richard Foote’s new “Thesaurus of Entomology.”

Several visitors were introduced.

The meeting was adjourned at 9:17 PM, after which cider and cookies
were served.

Wayne N. Mathis, Recording Secretary

845th Regular Meeting—3 November 1977

The 845th Regular Meeting of the Entomological Society of Washington
was called to order by President Ramsay at 8:00 PM on 3 November
1977 in the Ecology Theater of the National Museum of Natural History.
Twenty-three members and 7 guests were present. The minutes of the
June and October meetings were read and approved.

Acting Membership Chairman Spilman read for the first time the names
of the following new applicants for membership:

Daniel F. Austin, Dept. Biol. Sci., Florida Atlantic University, Boca
Raton, Florida.

Joaquin Bueno-Soria, Instituto de Biologia, U.N.A.M., Apartado Postal
70-153, Mexico 20, D.F. Mexico.

Michael E. Faran, MEP/NHB 165, Smithsonian Institution, Washing-
ton. De:

President Ramsay thanked President-Elect Sutherland for presiding and
conducting while he was ill.

Ted Spilman reminded the membership that dues envelopes were in-
cluded with the October issue of the Proceedings, and he encouraged the
membership to use them. Mignon Davis suggested that a “double” member-
ship category be created for spouses of members. Ted Spilman suggested
that the Executive Committee consider such memberships.

Editor Stoetzel brought copies of George Steyskal’s pictorial key to the
species of Anastrepha and copies of the October issue of the Proceedings
for anyone interested.

VOLUME 80, NUMBER 3 459

Nominating Committee Chairman Steyskal announced the new slate of
Society officers as follows:

President-Elect Donald R. Davis
Recording Secretary Wayne N. Mathis
Corresponding Secretary Donald R. Whitehead
Treasurer F. Christian Thompson
Editor Manya B. Stoetzel
Custodian Sueo Nakahara
Program Chairman Victor E. Adler
Membership Chairman Joyce A. Utmar

The nominating committee included Chairman George C. Steyskal,
Richard Newkirk and Donald Messersmith.

With the approval of Helen Sollers-Riedel, it was moved, seconded, and
unanimously approved that she continue as Hospitality Chairwoman.

Lee Hellman introduced the main speaker for the evening, Dr. F. E.
Wood, Associate Professor, Department of Entomology, University of Mary-
land. Dr. Wood spoke on “New Views on Human Lice.” Dr. Wood ex-
plained how he first became interested in the problem of human head
lice on school children and discussed the results of his research. He noted
that head lice are not found on black children, probably due to the flat
structure of their hair which makes attachment of the nit to the hair strand
less effective. He also presented several sources of evidence which suggest
that the head and body louse are two species. Dr. Wood illustrated his
talk with several slides and entertained an interesting question and answer
session afterwards.

President Ramsay noted that the forthcoming Congress of Plant Pro-
tection would be held in Washington, 5-12 August 1979.

At the suggestion of Ashley Gurney, it was moved and seconded that the
Society further acknowledge the generous contribution of Mr. Muesebeck
with a standing vote of thanks, which followed.

Notes and Exhibitions

Curt Sabrosky announced that the large and valuable collection of
Muscidae and Anthomyiidae of Dr. H. C. Huckett was recently donated
to the Smithsonian. Don Davis announced that the Society could use
the facilities of the Museum’s Naturalists Center for the first meeting of
1978 (January 5th).

Ashley Gurney projected and discussed about 35 Kodachrome pictures
of local insects and spiders, also of scenic outdoor areas in Chile and
southern Argentina which he visited in December 1976. The latter in-
cluded chiefly scenes in national parks in that part of Patagonia near
Bariloche, Argentina.

460 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

John H. Fales reported the collecting of Staphylus hayhurstii (Edwards) —
(Hesperiidae) at two locations in Rock Creek Park, Washington, D.C. on
31 August and 8 and 12 September 1977. These are probably the first oc-
currences of this species in the District of Columbia.

The meeting was adjourned at 9:34 PM, after which punch and cookies
were served.

Wayne N. Mathis, Recording Secretary

846th Regular Meeting—l December 1977

The 846th Regular Meeting of the Entomological Society of Washington
was called to order by President Ramsay at 8:00 PM on 1 December 1977
in the Ecology Theater of the National Museum of Natural History.
Thirty-five members and 31 guests were present. The minutes of the
November meeting were read and approved.

Because of the large number of guests, President Ramsay asked all at-
tending to briefly introduce themselves.

Membership Chairman Utmar reported that 34 new memberships were
accepted in the Society during 1977. She also read for the first time the
name of the following new applicant for membership:

Norman Lin, 1487 E 37 St., Brooklyn, New York.

Treasurer Thompson and Editor Stoetzel gave their annual report to the
Society. Both reports will appear in the April issue of the Proceedings.

President Ramsay introduced the main speaker for the evening, Dr. Larry
Rockwood, Department of Biology, George Mason University, who spoke
on “Howling Monkeys and Leaf-Cutting Ants: Comparative Foraging in
a Tropical Deciduous Forest.” Dr. Rockwood reviewed the biology of the
leaf-cutting ant (Atta columbica) and of the Howling Monkey (Alouatta
palliata) and noted that both are general herbivores but that feeding pref-
erences change during the dry and wet seasons. Dr. Rockwood illustrated
his talk with several Kodachrome slides and led an interesting question
and answer period afterwards.

Notes and Exhibitions

President Ramsay noted that a former student from Japan, Dr. Akira
Kamito, commemorated his 77th birthday with a special publication, en-
titled: “Studies on the broad bean weevil, Bruchus rufimanus Boheman
with special reference to its bionomics and control.”

Nominating Committee Chairman Steyskal read the slate of proposed new
officers for 1978. President Ramsay asked if there were any nominations
from the floor. It was then moved and seconded that the slate be adopted as
announced, which was unanimously accepted by the vote of the Society.

President Ramsay turned the gavel over to President-Elect Sutherland.

VOLUME 8&0, NUMBER 3 461

President Sutherland thanked the Society for their support and adjourned
the meeting at 9:35 PM after which punch and cookies were served.

Wayne N. Mathis, Recording Secretary

847th Regular Meeting—5 January 1978

The 847th Regular Meeting of the Entomological Society of Washington
was called to order by President Sutherland at 8:00 PM on 5 January
1978 in the Naturalists Center of the National Museum of Natural His-
tory. Twenty-seven members and 12 guests were present. The minutes of
the December meeting were read and approved.

Membership Chairman Utmar read for the first time the names of the
following new applicants for membership:

David W. Brassard, 8304 Tobin Rd., Annandale, Virginia.

David G. Casdorph, P.O. Box 1458, Monrovia, California.

William H. Gotwald, Jr., Dept. of Biol., Utica College of Syracuse Univer-
sity, Utica, New York.

George F. Ludvik, 9113 Rockefeller Lane, Springfield, Virginia.

Past President Ramsay noted that the Society has no certificate of mem-
bership or officership. After a brief discussion, it was suggested that any
proposal for certification be floored after consideration by the executive
committee.

President Sutherland announced that Maynard J. Ramsay will retire this
month after 34 years of federal service. A retirement luncheon will be
given 13 January at 11:30 AM at the Royal Arms Restaurant, Hyattsville,
Maryland.

The traditional Past President’s address was given by Maynard J. Ramsay
of the National Program Planning Staff, APHIS, USDA. His topic was
“Exotic Pest Detection and Evaluation—A Regulatory Agency's Dilemma.”
He showed a training film for new recruits to APHIS entitled: New Pest
Detection and Information Program,” explaining the national pest survey
teams work in discovering new pests, especially in corn, soybeans, small
grains, forage and citrus crops. The Agency has two functions:

1. The survey and detection of pests.
2. The action/no action decision making after a new pest has been
discovered.

Ramsay illustrated his remarks after the film with several slides and
charts. A question and answer session followed.
Notes and Exhibitions

Don Davis explained the function of the Naturalists Center, the room

462 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

in which our meeting was held. This new facility at the museum is a place
for serious amateurs to browse through collections of natural history speci-
mens to enable them to identify their own collections or to do research on
hobbies, etc. Collections of specimens of mostly local species were assembled
by members of the museum staff and many other specimens, books, and
equipment were donated by others.

T. J. Spilman showed a colored lithograph of passalid beetles anthropo-
morphically playing a game of pool. The origin of the picture had been
traced, with the help of R. D. Pope, to Theodor Compton, illustrator of
Kaup’s 1871 monograph of the passalids in Berliner Entomologische Zeit-
schrift.

C. W. Sabrosky announced the death on 21 November of Boris B. Rohden-
dorf, a worker on fossil insects and sarcophagid flies.

Helen Sollers-Riedel showed a book on insect flight, Borne on the Wing,
by Steven Dalton.

John H. Fales showed special first day cover stamps featuring four
Lepidoptera prepared by the Maryland Entomological Society and gave
some information on that Society.

Finally, C. W. Sabrosky suggested that any books or other exhibitions
be passed around before the main speaker's talk so everyone gets a chance
to see them.

The meeting adjourned at 9:33 PM, afterwhich punch, cookies, and
cake were served.

Joyce Utmar, Acting Recording Secretary

848th Regular Meeting—2 February 1978

The §48th Regular Meeting of the Entomological Society of Washington
was called to order by President Sutherland at 8:00 PM on 2 February
1978 in the Ecology Theater of the National Museum of Natural History.
Twenty-nine members and 5 guests were present. Minutes of the January
meeting were read and approved.

Membership Chairman Utmar read for the first time the names of the
following new applicants for membership:

Antonio Paulo Assis de Moraes, Acacia Ct., Mission Viejo, California
92675.

Richard Duffield, Department of Biology, Howard University, Wash-
ington, D.C. 20059.

Michael Kosztarab, Department of Entomology, Virginia Polytechnic
Institute and State University, Blacksburg, Virginia 24061.

Roy A. Norton, Department of Environmental and Forest Biology, College
of Environmental Science and Forestry, State University of New York,
Syracuse Campus, Syracuse, New York 13210.

VOLUME 80, NUMBER 3 463

Durward D. Skiles, Institute of Geophysics and Planetary Physics,
University of California, Los Angeles, California 90024.
Greg Spicer, 405 Cedar Creek Ct., Fort Worth, Texas 76103.

Announcement was made of our Annual Banquet, to be held at the Ft.
McNair Officers Club, 4th and P Sts., Washington, D.C., members attending
as guests of Dr. Dale W. Parrish.

The Washington, D.C. Science Fair requested members to assist in judg-
ing exhibits at the 18 March Science Fair.

Exhibitions

John Horne displayed a mulberry stake from his tomato garden, show-
ing vespid wasps which had bored 1'2 feet in the twig.

The main speaker of the evening was Dr. David Stiller, Animal Para-
sitology Institute, ARS, Beltsville, MD, who spoke on “Entomological studies
in Malaysia.” In a travelogue section, he noted the diversity of cultures of
the people inhabiting Malaysia, then spoke on his research. Included were
studies of the humidity requirements of the ticks Dermacentor atrosignatus
and Hemaphysalis semiermis, ticks of a monitor lizard, lung mites of
snakes, and lung mites of the silvered leaf monkey. Attempts were made
to find a suitable alternate host for laboratory study of treatment of
these lung mites. The mites also affect Rhesus monkeys, widely used in
research. There is presently no treatment for this infestation. Dr. Stiller's
talk was illustrated with many Kodachrome slides. Question and answer
session followed.

Notes

Ashley B. Gurney reported on his research on the history of stereoscopic
microscopes of the 1890's and early 1900's, especially those made by Zeiss
in Jena, Germany.

The meeting was adjourned at 9:35 PM, after which punch and cookies
were served.

Joyce A. Utmar, Acting Recording Secretary

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PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

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CONTENTS

(Continued from front cover)

The allotype of Arenasella maldonadoi and change of depository for two of
Fennah’s holotypes (Homoptera: Tropiduchidae and Issidae)
J. MALDONADO CAPRILES and A. BERRIOS

Biology and immature stages of Antichaeta borealis (Diptera: Sciomyzidae), a
predator of snail eggs W. H ROBINSON and B. A. FOOTE

Neotype designation of Culex quinquefasciatus Say (Diptera: Culicidae)
S. SIRIVANAKARN and G. B. WHITE

Taxonomy and description of two prionine Cerambycidae from southern Arizona:
A new species of Stenodontes and new status for Neomallodon arizonicus
(Coleoptera) D. D. SKILES

An infestation of miltogrammine Sarcophagidae (Diptera: Sarcophagidae) in a
population of Hybomitra lasiophthalma (Macquart) (Diptera: Tabanidae)
P. H. THOMPSON

NOTE:
A new name for Colpocephalum abbotti Price (Mallophaga: Menoponidae)
R. D. PRICE

BOOK REVIEW:
Taxonomic Study of the Cicadellinae (Homoptera: Cicadellidae). Part 2. New
World Cicadellini and the Genus Cicadella (D. A. Young, 1977)
J. P. KRAMER

SOCIETY MEETINGS

380

388

360

407

373

343

456

VOL. 80 OCTOBER 1978 NO. 4
s: 70673
Ent,

‘

PROCEEDINGS

of the

ENTOMOLOGICAL SOCIETY
of WASHINGTON

DEPARTMENT OF ENTOMOLOGY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY

CONTENTS

The genus Tychius Germar (Coleoptera: Curculionidae): Larvae and pupae of
some species, with evaluation of their characters in phylogenetic analysis
W. E. CLARK, H. R. BURKE and D. M. ANDERSON 626

A new genus and two new species of Eosentomoidea (Protura: Eosentomidae)
T. P. COPELAND 473

An new species of Eosentomon (Protura: Eosentomidae) from North Carolina and
Tennessee T. P. COPELAND and C. WHITE 485

Sandfly distribution in the United States, with a first record for Colorado (Dip-
| tera: Psychodidae) Ro BY EADS) 38

A new species of Asphondylia (Diptera: Cecidomyiidae) from Costa Rica with
taxonomic notes on related species R. J. GAGNE ‘514

New synonymy and a review of Haplusia (Diptera: Cecidomyiidae)
R. J. GAGNE

Description of the male of Neurobezzia granulosa (Wirth) (Diptera: Cera-
topogonidae) W. L. GROGAN, JR. and W. W. WIRTH 548

Description of a new Polymerus, with notes on two other little known mirids
from the New Jersey Pine-Barrens (Hemiptera: Miridae) T. J: HENRY

A new genus and species of Cardiastethini from Peru (Hemiptera: Anthocoridae)
T. J. HENRY and J. L. HERRING

(Continued on back cover)

ENTOMOLOGICAL SOCIETY

OF WASHINGTON
OrcanrzEp Marcu 12, 1884

OFFICERS FOR 1978

Doucias W. S. SUTHERLAND, President HELEN SOLLERS-RIEDEL, Hospitality Chairwoman
DonaLp R. Davis, President-Elect Vicror E. ADLER, Program Chairman
Wayne N. Martuis, Recording Secretary Joyce A. Urmar, Membership Chairwoman
DonaLp R. WHITEHEAD, Corresponding Secretary SuEO NAKAHARA, Custodian
F. Curistian THOMPSON, Treasurer D. W. S. SuTHERLAND, Delegate, Wash. Acad. Sci.

Manya B. STrorerzeEL, Editor
Publications Committee
EARLENE ARMSTRONG ASHLEY B. GURNEY
WayneE E,, CLark GEorRGE C, STEYSKAL

Honorary President
C. F. W. MurEsEBECK

Honorary Members
FREDERICK W. Poos Ernest N. Cory RayMonp A. ST. GEORGE

All correspondence concerning Society business should be mailed to the appropriate officer at the following
address: Entomological Society of Washington, c/o Department of Entomology, Smithsonian Institution,
Washington, D.C. 20560.

MEETINGS.—Regular meetings of the Society are held in the Ecology Theater, Natural History Building,
Smithsonian Institution, on the first Thursday of each month from October to June, inclusive, at 8 P.N
Minutes of meetings are published regularly in the Proceedings. 7

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All remittances should be made payable to The Entomological Society of Washington.

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STATEMENT OF OWNERSHIP
Title of Publication: Proceedings of the Entomological Society of Washington.
Frequency of Issue: Quarterly (January, April, July, October).

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use

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 465-472
A SYNOPSIS OF NEOTROPICAL ELELEIDES CRESSON
(DIPTERA: EPHYDRIDAE )

Wayne N. Mathis

Abstract—A synopsis of Neotropical Eleleides is presented and includes
a key to all species, appropriate figures, description of a new species (E.
penai), distributional notes on E. liroceras and a general discussion of the
genus. Eleleides liroceras and E. penai are shown to be sister-species. The
former is newly described from Chile (type-locality: Anticura, Osorno
Province) and fulfills an earlier prediction that additional members of
Eleleides would be found in western South America. The known distribu-
tion of E. liroceras is increased to southern Brazil (Nova Teutonia) and
southern Chile (Lampa and Anticura ).

Since the revision of Eleleides Cresson by Mathis (1977), additional
specimens of both E. liroceras Mathis and a new species have become
available. The new species is the third to be described in Eleleides and
the second from South America. The purpose of this paper is to report
these discoveries and to comment on the distribution and relationships of
the species. The methods used are those of my earlier revision, particularly
the descriptive portions which follow the same format as used previously.

The new specimens of E. liroceras were collected in the state of Santa
Catarina, Brazil, and in the provinces of Osorno and Santiago, Chile, which
substantially increases the known distribution of that species. Likewise,
the occurrence of a second Neotropical species on the west slope of the
Andes Mountains is an important discovery. Not only does the distribution
of the genus now appear more plausible, but the new species partially
fulfills an earlier prediction that other members of the genus are likely to
be found in western South America ( Mathis, 1977).

Key to species of Eleleides Cresson

1. Third antennal segment and maxillary palp black; eye-to-cheek ratio
less than 1:0.25; 1 or more pairs of larger acrostichal setae in addi-
tion to prescutellar acrostichal pair; supra-alar seta well developed,
at least 42 length of postalar seta (Australia) E. chloris Cresson
— Third antennal segment and apex of maxillary palp orange to yel-
lowish orange; eye-to-cheek ratio more than 1:0.40; all acrostichal
setae except for prescutellar acrostichal pair uniform in size, small;

bo

supra-alar seta lacking or greatly reduced

466 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2. Tibiae yellowish orange, concolorous with basitarsi; mesopleuron
completely silvery gray, pollinose, contrasting distinctly with bronze-
colored mesonotum; abdominal terga usually with pollinose grayish
wedges extending dorsally from ventral margins (Argentina, Brazil
and Chile ) E. liroceras Mathis

— Tibiae mostly black, concolorous with femora; mesopleuron mostly
brown to bronze colored, concolorous with mesonotum; abdominal
terga unicolorous or at most slightly grayish along ventral margins
( Chile ) E. penai, new species

Eleleides liroceras Mathis
Fig. 4

Eleleides liroceras Mathis, 1977:560 (figures of head, thorax, wing, and
male genitalia ).

Remarks.—When this species was described, specimens were available
only from the type-locality, Hurlingham, Buenos Aires, Argentina. I have
now examined five additional specimens as follows: Brazil: Santa Catarina,
Nova Teutonia (27°11’S, 52°23’W), Sept. 1970, Fritz Plaumann (2°,
MZUSP). Chile: Osorno Province, Anticura (1 km W.), 1-3 Feb. 1978,
W.N. Mathis (1°, USNM). Santiago Province, Lampa (22 km N. Santi-
ago), 21 Jan. 1978, W. N. Mathis (26, USNM). Although I am confident
that these specimens are conspecific with those from the type-series, males
were available only from the Lampa locality for confirmation.

Based on the locality data now at hand, I would expect to find specimens
of this species throughout the Araucaria zone of southern South America
where sedge-meadow habitats exist.

Eleleides penai Mathis, new species
Figs. 1-4

Diagnosis —Specimens of E. penai are distinguished from those of both
congeners by the following combination of characters: Third antennal
segment pale, yellowish orange to orange; frons mostly pollinose, grayish-
bronze colored, except for subshiny vitta extending from median ocellus
to ptilinal suture; maxillary palp pale, yellowish orange on apical % to 7%;
eye-to-cheek ratio more than 1:0.40; mesonotum pollinose to subshiny,
mostly bronze colored, becoming grayer toward lateral margins, scutellum
and posterior portion of mesonotum not distinctly darker in color; bristles
in general more strongly developed; no larger pairs of acrostichal bristles
except for prescutellar pair; supra-alar seta reduced greatly or lacking;
mesopleuron mostly brown to bronze colored, concolorous with mesonotum
and contrasting distinctly with silvery-gray color of notopleuron; tibiae

VOLUME 80, NUMBER 4 467

Figs. 1-3. Eleleides penai. 1, Epandrium, cerci, surstyli and hypandrium, posterior
aspect. 2, Anterior surstylus, lateral aspect. 3, Epandrium, cerci, surstyli and hypan-
drium, lateral aspect.

dark, grayish black to black, concolorous with femora; abdominal terga
lacking distinct grayish wedges extending dorsally from ventral margins,
at most slightly grayish along ventral margin.

Description.—Length 2.36 to 2.97 mm (averaging 2.66 mm).

Head: Head width-to-height ratio averaging 1:0.79; frons width-to-length
ratio averaging 1:0.61; frons subshiny, black with dense bronzish-brown
pollinose vestiture, pollinosity becoming weaker anteriorly; frontal vitta
between median ocellus and ptilinal suture, subshiny, becoming narrower
anteriorly, bronze colored; fronto-orbital plate slightly raised in relief from
remainder of frons. First and 2nd antennal segments black, mostly dull;
3rd segment slightly longer than wide, mostly pale, yellowish orange to
orange, becoming darker along dorsal edge in some specimens, macro-
pubescent along rounded apical edge. Face and gena concolorous, silvery
gray, densely pollinose to tomentose, face height-to-width ratio averaging
1:0.67; clypeus black, with pollinose vestiture less dense than face; maxillary
palp pale on apical ' to %, yellowish orange, concolorous with 3rd antennal
segment, becoming darker, brownish black basally. Eye height-to-width
ratio averaging 1:0.90; eye-to-cheek ratio averaging 1:0.43; ventral portion
of gena with a shallowly impressed groove running parallel with edge.

Thorax: Black, covered with pollinose vestiture; mesonotum mostly dull

468 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

anteriorly, becoming subshiny posteriorly, brown to bronze colored, be-
coming grayer anteriorly and laterally; humerus and notopleuron con-
colorous, gray, contrasting distinctly with mesonotum; mesopleuron and
pteropleuron concolorous mostly grayish bronze, posterior margins gradu-
ally becoming grayer; propleuron, front coxa, sternopleuron and_hypo-
pleuron nearly concolorous, mostly gray to silvery gray, becoming slightly
darker posteriorly. Femora and tibiae of all legs concolorous, subshiny to
pollinose, bronzish black, base and apex of tibiae paler, yellowish; basitarsus
of each leg pale, yellowish; other tarsomeres becoming darker apically, apical
ones mostly black. Wing entirely hyaline; wing length-to-width ratio aver-
aging 1:0.44; costal vein ratio averaging 1:0.50; M,,. vein ratio averaging
1:0.80; posterior crossvein perpendicular with vein M,,».. Halter pale,
capitellum white, pedicel yellowish.

Abdomen: Subshiny to shiny dark metallic bronzish-black coloration;
some specimens with ventral margin of each tergum slightly more pollinose,
grayish. Male terminalia as follows (Figs. 1-3): Posterior surstylus tri-
angular in posterior aspect, medium edge crenulate to serrate, with medio-
posterior enlargement in lateral aspect, wider dorsally; anterior surstylus
with anterior margin setose, posterior margin angulate; epandrium 2X as
wide as high, narrowing ventrally. Female ventral receptacle similar to
that of E. liroceras (Figs. 6-7, Mathis, 1977).

Type-material—Holotype ¢, labelled: “CHILE: Osorno Pr. Anticura
(1 km W) 432 m elev. 1-3 Feb. 1978 WNMathis.” Allotype ° and 2 para-
types (16, 12; USNM): with same locality data as the holotype; dates
from 3-6 February 1978. Other paratypes as follows: CHILE: Santiago
Province: El Alfalfal, 22 Jan 1978, W. N. Mathis (14, 22; USNM);
Coquimbo Province: El] Naranjo, Tilama, October 1967, L. E. Pefia (1¢,
42: MZUSP). Chiloe Province: Chepu, ‘April 1968, L> Ee Pena
MZUSP). The holotype is in the National Museum of Natural History
Smithsonian Institution, Washington, D. C., (USNM type-number 75764).

Geographic distribution (Fig. 4)—Specimens were examined from four
localities in Chile between 32° and 42° south latitude. This distribution
substantiates an earlier prediction (Mathis, 1977) that other members of
the genus would be discovered in western South America.

Natural history—All specimens taken by me were collected in sedge-
meadow habitats. The surrounding environs of each sedge-meadow habitat
varied considerably from scrub-covered foothills southeast of Santiago (El
Alfalfal) to Nothofagus forests (Anticura) in the Lake district of southern
Chile.

Etymology.—The species epithet, E. penai, is a genitive patronym honor-
ing Luis E. Pena G., J. I. Molina Institute, who collected part of the type-

VOLUME 80, NUMBER 4 469

%& Eleleides liroceras

@ Eleleides penai

© both species

series and who graciously hosted me in Chile while I collected the re-
mainder.

Relationships —This species and E. liroceras are sister-species. This
relationship is corroborated by numerous synapotypies as indicated in Fig.
5 and Table 1.

General Discussion

Although the distribution of Eleleides remains disjunct, the discovery
of an additional species in western South America and of the extended
range of E. liroceras substantiates their south temperate distribution. Still,
I am of the opinion that more members of the genus will yet be found
and suggest that South Africa or New Guinea will be productive in this
regard.

The distribution of Eleleides species in South America is not related to

470 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Argumentation scheme for the hypothetical phylogeny of the genus Eleleides.
Filled squares = apotypic character states; open squares = plesiotypic character states.

471

VOLUME 80, NUMBER 4

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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

conventional biogeographic provinces used to partition that continent
(Cabrera and Willink, 1973). This is evident from the distribution of either
species. The distribution of E. liroceras, for example, includes four of
Cabrera and Willinks biogeographic provinces: Paranense, Pampeana,
Chilena and Subantartica. The Paranense and Pampeana provinces are
adjacent, along the east coast near the Parana and La Plata Rivers, but the
Subantartica and Chilena provinces are far removed, being situated along
the western slope of the Andes Mountains. Likewise, E. penai is known
to occur in more than one biogeographic province in western South
America. What seems to be more important to the distribution of Eleleides
is the occurrence of aquatic systems, almost without regard to the surround-
ing habitat. Where sedge-meadow habitats occur in temperate South
America, it is likely that specimens of Eleleides will also be found there.

The widespread distribution of Eleleides species also indicates that the
species of Eleleides are probably not closely associated with a particular
plant species. The biogeographic provinces, as defined by Cabrera and
Willink (1973), are based primarily on the distribution of plants; and,
because the distribution of Eleleides species broadly overlaps these, the
likelihood of a specific plant association seems remote.

The relationships among species of Eleleides is summarized in Fig. 5
and the accompanying list of character evidence (Table 1). The numbered
squares refer to the list of character states on the table.

Acknowledgments

I am grateful to Mr. Luis E. Pena G. for his assistance while collecting
in Chile and to Dr. Nelson Papavero (Museu de Zoologia da Universidade
de Sao Paulo, MZUSP) for hosting me in Brazil and for the loan of
Eleleides specimens. For financial assistance through a fluid research grant,
I thank Mr. S. Dillon Ripley, Secretary of the Smithsonian Institution. For
critical review and constructive commentary of this paper, I thank Dr. Paul
J. Spangler.

Literature Cited

Cabrera, A. L. and A. Willink. 1973. Biogeografia de America Latina. Monografia
no. 13, serie de biologia, Organizacion de los Estados Americanos. 120 pp.
Mathis, W. 1977. Key to the Neotropical genera of Parydrinae with a revision of
the genus Eleleides Cresson (Diptera: Ephydridae). Proc. Biol. Soc. Wash.

90( 3) :553-565.

Department of Entomology, NHB 169, Smithsonian Institution, Wash-
ington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 473-484
A NEW GENUS AND TWO NEW SPECIES OF
EOSENTOMOIDEA (PROTURA: EOSENTOMIDAE)

T. P. Copeland

Abstract—The genus Styletoentomon is erected and two new species are
described, Styletoentomon styletum and Eosentomon erwini. Eosentomon
rostratum Ewing is moved to Styletoentomon.

Until 1974 the suborder Eosentomoidea contained only the family
Eosentomidae with a single genus, the Eosentomon. Tuxen (1964) in his
monumental work made provisional groupings within the genus on the
basis of the female squama genitalis but without setting them up as genera.
In 1974, Yen described an eosentomid without spiracles from China and
established a second genus, Antelientomon, to contain it. Tuxen (1975)
erected Isoentomon, a third genus, to contain several species with spiniform
foretarsal sensilla e and g. In 1977, Yen erected a fourth genus within the
Eosentomoidea, Anisentomon, from China. There still remains well over
a hundred species within the original genus, Eosentomon. All species in
the four genera have chewing-type mandibles each possessing three to
several small apical teeth except for Eosentomon rostratum Ewing (1940)
and the one described herein both of which have stylet-shaped mandibles
for piercing. The genus Styletoentomon is erected to contain these two
with Eosentomon rostratum as the type-species.

Styletoentomon Copeland, new genus

Diagnosis —Eosentomids with spiracles and 3 pairs of 2-segmented ab-
dominal appendages. Mandibles very long, extremely slender and sharp
pointed. Labrum longer than mandibles and very narrow. Foretarsal
sensilla e and g clavate. The only characters not shared by 1 or more
species in the other genera are the stylet-shaped mandibles and extremely
narrow labrum.

Styletoentomon rostratum (Ewing), NEW COMBINATION

Eosentomon rostratum Ewing, 1940:520.

Styletoentomon styletum Copeland, new species

This form is closely related to Styletoentomon rostratum (Ewing, 1940)
in that they both possess long, slender, stylete-shaped mandibles and in this

A474 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ee
0.05 mm

Figs. 1-2. Styletoentomon styletum. 1, Labrum and mouthparts; RS I, rostral seta;
GL, gland pore; Cl. Apd., clypeal apodeme. 2, Abdominal terga I-II.

VOLUME 80, NUMBER 4 475

respect differ from all other Eosentomidae. It differs from S. rostratum in
the absence of foretarsal sensillum b’-1 and in smaller body size. Body
length fully extended averages 1300 » and foretarsal length 115 ». In S.
rostratum b’-1 is present, body length averages 1700 » and tarsal length
125 p.

Holotype.—? , 1125 » long but not fully distended.

Description —Head: Almost round excluding labrum; capsule length
139 », with labrum 175 ». Labrum (Fig. 1) extremely narrow and without
labral setae. Mandibles rapier like and not flattened at base; rostral setae
III % as long as RS-I, RS-III:I = 0.53. Lacinia I poorly sclerotized, blunt
tipped and not hooked; lacinia II well sclerotized and strongly hooked.
Maxillary palpi resembling those of Eosentomon vermiforme Ewing and
S. rostratum. Clypeal apodemes distinct, connected anteriorly.

Thorax: Three wedge-like sensilla in each pleural membrane between
thorax I-II. Filamento de sostegno in prothorax. Mesothoracia seta P 1
shorter than distance to its homolog (35:43) and longer than P 1’. Setae-
sensilla P 3’ setiform.

Tarsi: Foretarsus (Figs. 3-4) broader relative to length than in E.
veriforme, length excluding claw 99 ,. Sensillum t-1 inserted on level of
a 3; t-2 setiform, long and inserted on level of a 4. Sensillum a’ setiform,
shorter than distance to a 4, 15:19; bl absent; b’—2 setiform, broad and
long; c’ present but indistinct except for base; sensillum a extending 4 dis-
tance to y 2; b strong, tip at base of B 6; c long, not setiform; g clavate
with small club, long shank, inserted near level of 2 8. Tarsal pit 1 distinct
and nearer a’ than a 3’; pit 2 nearer d than y. Ratios: BS 1.44; TR 5.7;
EU 0.90.

Tarsus III bearing the usual strong, dorsal, subapical spine; empodium
very short. Claws II-III with very fine tooth on upper surface.

Abdomen: Posterior row of setae on tergum I (Fig. 2) with 2 primary,
2 accessory and 1 microchaeta on each side. Primary setae longer than
corresponding accessories, 34:20. Seta P 1’ on t VII (Fig. 5) very short,
peg-like, without terminal “brush” and inserted on posterior margin of
tergum. Sternum VIII with 2 anterior and 7 posterior setae, the P 2 almost
in line with P 1-P 3; sterna IX-X with 4 setae each. Abdominal chaetotaxy
shown in Table 1.

Genitalia: Female squama genitalis (Fig. 6) similar to that of S.
rostratum and E. veriforme. The processus sternales sharp-pointed and
evenly bent giving a stoop-shouldered appearance. No unusual features
noted in male apparatus.

Discussion.—The most characteristic features of this species are the very
long narrow labrum, stylete-shaped mandibles, prominent clypeal apodemes,
the 2:2:1 seta ratio on tergum I and the absence of tarsal sensillum b’-1.
The positions, shapes and sizes of the tarsal sensilla conformed closely to

476 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Z,

—
0.01 mm

Figs. 3-4. Styletoentomon styletum. 3, Foretarsus, dorsal view. 4, Foretarsus,

ventral view.

VOLUME 80, NUMBER 4

perlite

alee | |

on

oo EE
“Vp l Aan
of: r

t
EE

5 0.05 mm

Figs. 5-6. Styletoentomon styletum. 5, Abdominal terga WVII-VIII.

genital apparatus.

6,

Female

478 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Abdominal chaetotaxy for Styletoentomon styletum.

Abdomen I II-III IV vV-VI VIL VIIL IX-xX XI XII
Tergum 4 10 10 8 6 6 8 8 9
10° 16 16 16 16 9 sc _ 7
Sternum 4 6 6 6 6 2, 4 8 1
4 4 10 10 10 “ - a -_

* Two primary, two accessory, and one microchaeta on each side.

those on the type as did the numbers, positions and shapes of the abdominal
setae, specifically the P 1’ of t VII, the P 1” and P 2 on @ Viiiihe
P 1-2-3 on that sternum and the 4-4 seta number on s IX-X.

The species is known from approximately 175 specimens taken at altitudes
of less than 775 m in Tennessee, Arkansas, North and South Carolina and
Virginia. Samples from elevations over 775 m yielded over 400 examples
of the closely related form S. rostratum but only 10 of S. styletum.

Holotype and type-locality.—The holotype 2°, on slide TN 283-1 taken by
T. P. Copeland from deciduous leaf litter 8 miles east of Speedwell (Clai-
borne Co.), TN., 21 July 1953.

Material examined.—The species is known from the holotype and para-
types from the following localities; collector, T. P. Copeland unless in-
dicated otherwise: Speedwell, Claiborne Co., TN., 21 July 1953. Sex and
slide numbers: 2, 280-1: 4, 281-2; 292-1: Imm., 293-1. Anderson Co.,
TN., 21 July 53. ?, 287-3: Imm., 286-3. Campbell Co., TN., 21 July 53.
?, 292-1; ¢, 293-3: Imm., 293-1. Blount Co., TN., 1 July 53. 2 9, 98X-1;
99-1; 99-3; 99-4; 99-5; 99-6; 99-8: ¢ 99X-1. Smoky Mt. Nat. Park, Cades
Cove, Blount Co., TN., 1 July 53. 22, 101-15; 101-17: Imm., 101-8. Sevier
Co., Dupont Mt. TN., 20 Apr. 52. 2.9, 91-95: 91-96) C252 ieseoleas
Sequatchie Co., TN., 30 July 53. 2, 395-1. Hamblen Co., TN., 26 July 53.
Imm., 296-2 (2). Jefferson Co., TN., 20 June 59. Imm., 270-2. Lenoir
City, TN., 8 Oct. 59. 4, 520-1. Cedars of Lebanon State Park, TN., 12
Oct. 62. ?%, 723E-3; 723EH-5; 723E-12; 724G-5, 724-4: 724-1. 724@ 212:
36, 723-3; 723E-7; 723E-11; 723E-15: Imm., 724-13. Shelby Forest State
Park, TN., Coll. J. S. Henderson, summer, 1969. 2, 110-1: 66, 111-4:
111-5; 124-13. Chickasaw State Park, TN., Coll. G. Hunnicutt, 26 June 64.
? 2, 25-7; 27-10; 27-13; 28-3, 30-1; 30-2, 30-4; 96-16; 109-8: ¢ 6, 16-2; 27-11;
27-18; 28-6; 30-9; 30-11. Natches Trace State Park, TN., Coll. T. D. Dia-
mond, 26 June 64. 22, 1-1; 1-6; 3-6; 30-4; 40-2; 41-2; 42-1; 42-6; 42-7; 44-3;
44-4; 46-1; 48-2; 50-12; 74-1; 74-3; 74-4; 74-7; 75-1; 75-2; 75-3; 76-2:
76-3; 76-9; 76-11; 76-15; 79-3; 88-3; 98-1; 98-2; 98-6; 98-7; 98-9: 34,
1-7; 3-9; 48-6; 50-14; 74-5; 75-3; 98-8; 76-5; 76-10; 98-3. Reelfoot Lake, TN..,

VOLUME 80, NUMBER 4 479

Coll. R. Davis. Aug. 63. 2°, 4-3; 4-9; 4-11; 4-12; 4-13; 4-20; 4-22; 4-24:
8 6, 4-5; 4-7; 4-8; 4-18; 4-21; 4-23; 9-7. Highlands, No. Car., Coll. T. Cope-
land, 25 June 77. 2°, 880-1; 880-4; 881-1: ¢ 880-2: Imm., 880-7; 880-9;
880-10; 880-12; 880-14.

Deposition of type-material—The holotype and a male paratype will be
deposited in the National Museum of Natural History, Smithsonian In-
stitution. Male and female paratypes will be given to Dr. S. L. Tuxen,
Zoological Museum, Copenhagen, Denmark and to Dr. Gentaro Imadate,
Tokyo Medical and Dental University, Tokyo, Japan. The remainder will
be retained by the author.

Eosentomon erwini Copeland, new species

This species belongs to the wheeleri (2/7) group of Bonet and Tuxen
(1960) and to the 3:1:1 sub-group. Its closest relatives are most likely
members of the latter group and in the United States only Eosentomon
wheeleri Silvestri, Eosentomon pseudowheeleri Copeland, Eosentomon
tennesseense Copeland, Eosentomon pusillum Ewing, Eosentomon quad-
ridentatum Copeland, Eosentomon yosemitense Ewing and_ possibly
Eosentomon christianseni Bonet possess the requisite features.

This form can be easily differentiated from all other U.S. Eosentomon
species by the massive size of the mouthparts, especially the mandibles.
In having a long empodium on tarsus III, it resembles Eosentomon pallidum
Ewing and E. pusillum but differs from those species in that E. pallidum
has no anterior setae on sternum VIII, six setae each on sterna IX-X and
t-1 inserted nearer a 3 than to a 3’. In E. erwini sternum VIII has two
anterior setae, four setae each on sterna IX-X and sensillum t-1 inserted
near level of a 3’. In E. pusillum the clypeal apodemes are very con-
spicuous, mouthparts including labrum very short, t-1 placed nearer seta
a 3, body length 570 microns and foretarsus without claw 55 microns. In
E. erwini clypeal apodemes indistinct or absent, with huge mouthparts,
t-l nearer a 3’ than to a 3, body length 1500 » and tarsal length 100 1.

Holotype. —?, on slide TN. 727-23, 1515 » in length and narrow body
giving a long slender appearance.

Description —Head: Egg shaped, capsule excluding rostrum 115 p. La-
brum (Fig. 7) very wide at base and long, length 32 », terminating in a
broad V-shaped apical notch with a wide, shallow, median cleft; LR 3.65.
Pseudoculus divided and much broader than long, 15:9. Labral setae
absent. Rostral setae I extremely short, 7 «, not flattened and only % as
long as RS III. Rostral setae IT longer than labrum and flattened (winged )
in basal %. Mandibles (Figs. 7-8) very large and massive, terminating in
5 prominent teeth. Clypeal apodemes either absent or obscured.

Thorax: Three prominent wedge-shaped sensilla (Fig. 9) in each pleural

480 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

10

Figs. 7-11. Eosentomon erwini. 7, Labrum and mouthparts; RS I, rostral seta. 8,
Mandible. 9, Mesothoracic tergum. 10, Tarsus II claw and empodium. 11, Tarsus
III claw and empodium.

membrane between thorax I-II. Glands of filamento de sostegno in meso-
thorax: Seta P 1 on that tergum far shorter than distance to its homolog
(27:35) but longer than. P 1’ (27:14). Tergal setae P 3’ on thorax II-III
long for these setae; another sensillum immediately adjacent to seta A 4 on
metathorax plus 1 further removed from that seta.

Tarsi: Foretarsus (Figs. 12-13) relatively broad, length 103 ». The t-1
inserted slightly proximal to level of a 3’; t-2 narrow lanceolate and in-
serted on level of a 4; t-3 extending to base of a 7; a’ setiform, tip reaching
base of a 4; b’-1 absent; b—2 lanceolate, equal length to t-2 but slightly

VOLUME 80, NUMBER 4 481

q!

12

Figs. 12-13. Eosentomon erwini. 12, Foretarsus, dorsal view. 13, Foretarsus,
ventral view.

broader; ¢’ very short; sensillum a indistinct except for base but in paratypes
length varied from '% to % distance to y 2; b setiform and extending to
midpoint between £ 6 and £ 7; c indistinct except for socket but in para-
types varied from ‘2 to % distance to y 3; e and g clavate with long shanks
and small clubs; f-1 clavate, club approximately equal in size to those of
e and g. Tarsal pits 1 and 2 prominent. Shapes, sizes and locations of all
setae and sensilla as represented in the figures. Tarsus II empodium (Fig.
10) % length of claw; tarsus III empodium (Fig. 11) longer, % length of
claw; claws not toothed.

Abdomen: Abdominal tergum I with 3 primary, 1 accessory and 1
microchaeta on each side in posterior row. Abdominal accessory setae

longer than corresponding primaries, t IV P 1-P 1’ 23:28. On t VII P 1’

482 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

15

Figs. 14-16. Eosentomon erwini. 14, Abdominal terga VII-VIII. 15, Abdominal
sternum VIII. 16, Female genital apparatus.

(Fig. 14) very short setiform, not at all filamentous, inserted along the
posterior margin of the tergum and not “brush” tipped. On t VIII the
P 1” peg-like with long terminal filament. Sternum VIII (Fig. 15) with 2
anterior and 7 posterior setae, P 2 almost in line with P 1-P 3. Sterna
IX-X with 4 setae each. Abdominal chaetotaxy represented in Table 2.
Unfortunately, the type has a setal abnormality in anterior row of tergum
VI, 3 on 1 side instead of 4. Paratypes have 4 on this segment.

Genitalia: The female apparatus (Fig. 16) characterized by prominent
basal apodeme possessing a long plate at base; processus sternales with
very sharp pointed, dense tips; posterior valves and filaments long.

Discussion.—The species is known from 61 females, 13 males and 20

VOLUME 80, NUMBER 4 483

Table 2. Schematic representation of abdominal chaetotaxy for Eosentomon erwini.

Abdomen I II-III IV vV-vI_ VII VIII IxX-xX XI XII

Tergum a 4 10 10 8 6 6 8 8 9
p 108 16 16 16 16 9 a - %
Stemum a 4 6 6 6 6 2 4 8 12
Dp 4 4 10 10 10 % 7 2 _

Abbreviations: a = anterior row; p = posterior row.
* Three primary, one accessory and one microchaeta on each side.

immatures, an unusual ratio of females to males. There is little variation
among the individuals for all characters studied. The most characteristic
features are: Size and shape of labrum, massive appearance of mandibles,
each with five prominent teeth; and extremely short rostral setae I giving
RS III:I ratio of 2.0 and winged condition of rostral setae II. The arrange-
ment of setae P 1-2-3 on sternum VIII is uniform in all specimens. The
pseudoculi are never prominent and often difficult to find. Clypeal
apodemes are either absent or obscured. Sizes, shapes and locations of
all sensilla conformed closely to those on the type. Three sensilla are
present in pleural membranes between thorax 1-2 on all specimens. On
the foretarsus, sensillum c’, when it could be clearly seen, is very short,
little more than nipple-like in appearance; b’-1 is always absent; sensillum
s has a very small or no club; the f-1 is always clavate, and tarsal pits 1
and 2 are prominent and uniform in location. There was no variation in
the 3:1:1 seta ratio on abdomen I and in the relative lengths of primary
to accessory setae, the latter being longer. On sternum VIII the P 2 is
always nearly in line with P 1-P 3. In other species this is not always so
but it is consistent for each species. Four adult individuals had variations
in abdominal setae number but these involved only the loss of a single
seta on one side. The female genital apparatus is consistently uniform in
appearance except for minor distortions apparently caused by pressure
from the cover glasses.

Type-locality—Near Tennessee, State Fish Hatchery, Erwin, Tennessee.
All specimens taken in leaf litter from black cherry trees and honeysuckle
vines.

Types and deposition—The holotype, 2, on slide TN 727-23 and 93
paratypes all with numbers TN 727, collected by T. P. Copeland, 23 May 77.
The type will be temporarily retained in East Tennessee State University
museum but eventually it and a male will be deposited in the National
Museum of Natural History, Smithsonian Institution. A paratype male and
female will be deposited with Dr. S. L. Tuxen, Zoological Museum, Copen-

484 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

hagen, Denmark and Dr. Gentaro Imadaté, Tokyo Medical and Dental
University. All others will be retained by the author.

Literature Cited

Bonet, F. and S. L. Tuxen. 1960. Reexamination of species of Protura described by
H. E. Ewing. Proc. U.S. Nat. Mus. No. 3437:265-305.

Ewing, H. E. 1940. The Protura of North America. Ann. Entomol. Soc. Am. 33:
495-551.

Tuxen, S. L. 1964. The Protura. Herman, Paris. 360 pp.

1975. Isoentomon, A new genus within the Eosentomoidea (Protura:
Eosentomidae). Entomol. Scand. 6:89-101.

Yen, Wen-Ying. 1974. Studies on Chinese Protura III. A new genus of Protentomidae

and its phylogenetic significance. Acta Entomol. Sin. 17:49-54.

1977. Two new genera of Protura. Acta Entomol. Sin. 20:85—-94.

Department of Biology, East Tennessee State University, Johnson City,
Tennessee 37601.

PROC. ENTOMOL. SOC. WASH.

80(4), 1978, pp. 485-492
A NEW SPECIES OF EOSENTOMON
(PROTURA: EOSENTOMIDAE) FROM
NORTH CAROLINA AND TENNESSEE

T. P. Copeland and Charlie White

Abstract.—Eosentomon pseudoyosemitense White, new species, is de-
scribed. Its distribution in Tennessee-North Carolina and a table sum-
marizing differences among 10 species of U.S. Eosentomon are given.

Eosentomon pseudoyosemitense White, new species

White using material that had been collected by Copeland in 1959-1961
originally described this species in his M.S. thesis (1966) but this has not
been published. In 1977 Copeland made additional collections in the
vicinity of Highlands, North Carolina and the following description and
drawings were made by him from a specimen taken near Bridal Veil Falls.

In most characters but especially in the shape of the female genital
apparatus E. pseudoyosemitense is closely related to Eosentomon yosemi-
tense Ewing (1940), both have the processus sternales in the form of two
semicircles. However, E. yosemitense has no anterior setae on sternum
VIII while E. pseudoyosemitense possesses two. These and other differ-
ences in related species are summarized in Table 1, some of the informa-
tion taken from Bonet and Tuxen (1960).

Holotype.—& , 725 » long but not fully distended and poorly sclerotized.

Description —Head: Egg shaped, length excluding labrum 89 ». Labrum
length 6 ,; terminating in a flat V-shaped notch with relatively broad,
deep, median cleft (Fig. 1); LR 14.8. Labral setae present. Rostral setae
I much broadened in basal % and equal length to III. Pseudoculi large,
11 »; PR 8.0, PR derived by dividing the pseudoculus length into head
capsule length. Mandibles partially obscured but in paratypes terminate
in 3 teeth. Lacinia II hooked but not as strongly as in Eosentomon
vermiforme Ewing. Clypeal apodemes connected anteriorly.

Thorax: Glands of filamento de sostegno in mesothorax (Fig. 4). Spi-
racular setae P 3’ setiform and relatively long for these setae. Two sensilla
located in each membranous pleural area between thorax I-II and 1 im-
mediately adjacent to setae A 4 on metathoracic tergum. Mesothoracic
seta P 1 length % as long as distance to its homolog and slightly longer
than p I’.

Tarsi: Foretarsus (Figs. 2-3) short and broad, length 65 »; BS 0.77.
Sensilla t-1, e, g and s with huge clubs. The t-1 inserted very near to a 3;
e between @ 6 and y 4; g closer to £ 8 than y 4. Sensillum b narrow

486 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Summary of similarities and differences among selected species of U.S.
Eosentomon.

A B G D E
Labrum length 6.8 31 13 19 14
Labral setae present + 0 + + +
Ros. setae III:I 1.0 0.3 1.0 0.5 1.0
Foretarsus length 69 125 89 103 113
BS ratio 0.80 1.45 1.40 135 iepl
Sensillum b’ 1 pres. + 0 0 0 0
t-1 closer to a 3’ than a3 0 + + + +
Seta no. ant. row t IV 8 10 10 10 8
Seta no. ant. row t V-VI 8 8 8 8 8
Abd. acc. setae longer + 0 + 0 +
Setae no. s VII 2/7 2/7 2/7 2/7 2/7
Setae no. s IX-X 6 6 6 4 4
F G H I J

Labrum length > ir 15 15 32
Labral setae present + Ie + + 0
Ros. setae III:I Ir P 0.8 0.7 2.0
Foretarsus length rg 80 ie 108 103
BS ratio 0.81 0.90 0.98 0.84 1.42
Sensillum b’ 1 pres. 0 P + ae 0
t-1 closer to a 3’ than a 3 0 0 0) 0 tL
Seta no. ant. row t IV 10 8 10 10 10
Seta no. ant. row t V—-VI 10 6 8 8 8
Abd. acc. setae longer =f + 0 0) a
Setae no. s VIII ayia 0/7 0/7 0/7 2/7
Setae no. s IX-X 6 4 6 6 4

A. E. pseudoyosemitense D. E. pseudowheeleri G. E. yosemitense

B. E. quadridentatum E. E. tennesseense H. E. dureyi

C. E. wheeleri F. E. pusillum I. E. pallidum

J. E. erwini

/

lanceolate, tip barely reaching base of 6 6; f-1 setaceous; a’ narrow
lanceolate, inserted near the level of t-1; pits 1 & 2 extremely small, rep-
resented only as tiny white spots, the former near a 3’ and the latter very
close to and on level of y. Sensillum b’-1 present; c’ absent. Shapes, sizes
and locations of all sensilla as figured.

Tarsus III with the usual strong dorsal spine; claw not toothed; empodium
very short.

Abdomen: Posterior row on tergum I with 3 primary, 1 accessory and
1 microchaeta (3:1:1) on each side. Abdominal accessory setae longer
than corresponding primaries, t IV P 1:P l’ = 15:18. On t VII (Fig. 5)

VOLUME 80, NUMBER 4 487

Sue
cae

Figs. 1-3. Eosentomon pseudoyosemitense, holotype. 1, Labrum; Cl. Apd., clypeal
apodeme. 2, Foretarsus, dorsal view. 3, Foretarsus, ventral view.

488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 4 489

seta P 1 not displaced anteriorly; P 1’ filamentous, relatively long for this
seta, terminating in a “brush” tip and inserted near level of P 1-P 2.
Arrangements of setae on t VIII as in Fig. 6. Sternum VIII (Fig. 7) with
2 anterior and 7 posterior setae, the P 2 not greatly displaced anteriorly.
Sterna IX-X with 6 setae each. Abdominal chaetotaxy shown in Table 2.

Genitalia: The processus sternales of the female apparatus (Fig. 8)
roughly in shape of 2 semicircles. Laterally just posterior to the basal
apodeme arms a thin dark line present and internally from each, 2 more
or less oval structures present.

Discussion—tThe species is known from Highlands and Mount Mitchell,
North Carolina and Roan Mountain, Smoky Mountain National Park,
Chickasaw State Park and Newport, Cocke County, Tennessee. All speci-
mens except those from Newport, were collected at or above 1100 m
elevation. Repeated collecting at the Newport site failed to produce ad-
ditional specimens. Total number of adults collected were 52, plus im-
matures which were not examined.

There are no variations in the possession of labral setae, only two
sensilla in each pleural area between thorax I-II, the 3:1:1 tergum I seta
ratio, two anterior and seven posterior setae on sternum VIII and _ six
setae on each sterna IX-X. One individual had an extra seta on one
abdominal tergum, and one specimen had one less. On tergum VII the
positions of setae P 1 were constant as were the shapes and positions of the
P 1’. On tergum VIII the positions of the P 2 and P 1” were relatively
constant as were the positions of the setae on that sternum, specifically
the setae P 2.

The labrum always terminated in a flat V-shaped notch with a broad,
deep, median cleft; LR ranged from 11.4 to 16.6 with a median of 14.6.
The PR ranged from 7.4 to 10.6, median 9.3.

The foretarsal sensilla shapes, sizes and locations were as on the holo-
type with most characteristic features being the huge clubs on sensilla t-1,
e, g and s, the t-1 always inserted very close to seta a 3 and c’ absent on
all specimens. The BS ranged from 0.69 to 0.86, median 0.76.

Location of the glands of the filamento de sostegno varied from most
posterior portion of prothorax to anterior half of the mesothorax, more
often found in the latter. It is felt that they lie within the mesothorax near
its anterior border unless displaced by pressure from the cover glass.

Some variation existed in the appearance of the female apparatus but

<

Figs. 4-6. Eosentomon pseudoyosemitense, holotype. 4, Mesothorax; G. Fil., glands
filamento de sostegno, P 1, P 1’, P 3’ setae. 5, Abdominal tergum VIL. 6, Abdominal
tergum VIII.

490 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 80, NUMBER 4 491

Table 2. Schematic representation of abdominal chaetotaxy for Eosentomon pseu-
doyosemitense.

Abdomen I I-III IV-VII VIII IDK XI XII

Terga a 4 8 8 6 8 8 9
Dp 10° 16 ‘16 9

Sterna a 4 6 6 2 6 8 12
Dp 4 4 10 7

Abbreviations: a = anterior row; p = posterior row.
* Three primary, one accessory and one microchaeta on each side.

it is felt this is more apparent than real, probably distortions caused by
pressure of the cover glasses. The four oval spots shown in the genital
apparatus appear to be of no taxanomic significance as they are often
not present in female paratypes. No unusual features in the male ap-
paratus observed. A North Carolina specimen was selected for the holo-
type because White’s original specimen is in poor condition.

Type-locality —Bridal Veil Falls on Highway 64, two miles west of High-
lands, North Carolina.

Types and deposition.—The holotype, a female on slide NC 881-13 col-
lected 25 June 1977 by T. P. Copeland from deciduous leaf litter, 3600 ft.,
Bridal Veil Falls, Natahala National Forest, approximately 2 miles west
of Highlands, North Carolina.

Paratypes: The species is also known from 51 paratypes, 32 2 and
19 é. Paratypes from the following localities: Highlands, North Carolina,
Coll. T. Copeland, 25 June 1977, deciduous leaf litter, 3600-3800 ft., slide
nos.; °°, X-5; 886-1; 887-10; 888-21; 888-45; 888-49; 888-58; 889-8;
889-32; 889-39; 890-F-Z; 890-I-1; 890-S-2; 891-F-1; 44, 879-8; 881-22;
881-31; 885-3; 887-12; 888-67; 889-1A; 890-0-3; 891B-23. Mt. Mitchell,
North Carolina, Summer 1962, Coll. R. A. Durey, 4500-6500 ft., slide nos.;
22 45-12/1; 48-8/5; 60-4/7; 65-8/2; 65-15/1; 65-15/2; 65-15/3; 65-15/7;
65-15/8. Smoky Mt. Nat. Park, Sevier Co., Tenn., 4000 ft., Hwy 441, Coll.
T. Copeland, 27 Oct. 59, slide nos: ? 2, 621-8; 622-8; 622-9: Trout Branch,
S.M.N. Park, 27 Sept. 61, 712-X8-2; 712-X8-5; 712-X20-2; $4, 712-X8-2;
712-X8-5; 712-X20-2; 712-X8-3. Newport, Tenn. Coll. T. Copeland, 23 Nov.
61; 646, 717-3; 718-11. Roan Mountain, Tenn., Coll. T. Copeland, 11 Oct.
59; 8 558-3. Chickasaw State Park, Tenn., Coll. G. Hunnicutt, 26 June
1964, slides GSH; ? 2, 55-5; 97-5; 132-17; 6 6, 117-30; 120-3; 122-28.

<

Figs. 7-8. Eosentomon pseudoyosemitense, holotype. 7, Sterna VIII-XII. 8, Female
genital apparatus.

492, PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The holotype and a male paratype will be deposited in the National
Museum of Natural History, Smithsonian Institution. Male and female
paratypes will be given to Dr. S. L. Tuxen, Zoological Museum, Copen-
hagen, Denmark and to Dr. Gentaro Imadate, Tokyo Medical and Dental
University, Tokyo, Japan. All others will be retained in the East Tennessee
State University Museum.

Literature Cited

Bonet, F. and S. L. Tuxen. 1960. Re-examination of species of Protura described
by H. E. Ewing. Proc. U.S. Nat. Mus. 112:265-305.

Ewing, H. E. 1940. The Protura of North America. Ann. Entomol. Soc. Am. 33:
495-551.

White, Charlie. 1966. A new species of Protura from Tennessee. Unpublished M.S.
Thesis. East Tennessee State University.

(TPC) Department of Biology, East Tennessee State University, Johnson
City, Tennessee 37601; and (CW) Department of Biology, New River
Community College, Dublin, Virginia 24084.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 493-507
STUDIES ON THE GENUS FORCIPOMYIA. V. KEY TO
SUBGENERA AND DESCRIPTION OF A NEW
SUBGENUS RELATED TO EUPROJOANNISIA
BRETHES (DIPTERA: CERATOPOGONIDAE )

Willis W. Wirth and Niphan Chanthawanich Ratanaworabhan

Abstract.—Saliohelea, new subgenus of the genus Forcipomyia Meigen,
is proposed for F. leei, new species (type-species), from the Nearctic and
Neotropical regions, F. brevicosta (Clastrier) from West Africa, F.
deminuta Tokunaga and Murachi from the western Pacific, and F. stami,
new species, from Zaire. A provisional key to the subgenera of Forcipomyia
is presented. Saliohelea is most closely related to Euprojoannisia Brethes
and differs mainly in the complete fusion of the fourth and fifth palpal
segments, the absence of mandibular teeth in the female, the character-
istically sclerotized margins of the male aedeagus and the shape and
arrangement of the spiracular openings on the pupal respiratory horn. It
is also related to Warmkea Saunders, from which it differs in its shorter
costa and distal antennal segments, the shape of the fourth palpal segment,
the absence of mandibular teeth in the female and the arrangement of the
spiracular openings on the pupal respiratory horn.

We have become especially interested in the classification of the small,
inconspicuously brownish, hairy, biting midges of the genus Forcipomyia
Meigen because of their importance in the pollination of cacao (Theobroma
cacao L.) and other tropical crop plants (Saunders, 1956 and 1959; Dessart,
1961 and 1963; Barroga, 1964; Soria and Wirth, 1974; Kaufmann, 1975;
Winder, 1977). There is a small group of species for which we have found
characters of the female, male and pupa that in our opinion, justify the
description of a new subgenus.

The modern classification of the genus Forcipomyia is well grounded
in the recent works of Saunders (1956 and 1959), Tokunaga and Murachi
(1959), Dessart (1963), and Chan and LeRoux (1965 and 1971). Readers
are referred to these authors for explanation of our terminology and a fuller
discussion of subgeneric classification.

We are especially indebted to Paul G. Bystrak, Maryland Department
of Agriculture, Annapolis, for his excellent research on the taxonomy of all
stages of the North American species of the subgenus Euprojoannisia
Brethes. His work has given us a much deeper insight into the taxonomy
of this subgenus as well as those others most closely related to the new
subgenus. Mr. Bystrak provisionally included our new species F. leei in

494 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

his unpublished thesis (Bystrak, 1974) and has permitted us to draw
freely upon his description and discussion.

Tokunaga and Murachi (1959) gave a short diagnosis of what they
termed “Subgenus C” for a provisional group containing only the species
Forcipomyia deminuta Tokunaga and Murachi from Micronesia. They
compared “Subgenus C” with the subgenera Proforcipomyia Saunders and
Synthyridomyia Saunders. Dessart (1963) commented on Tokunaga and
Murachi’s “Subgenus C” and included it in his keys to the African sub-
genera of Forcipomyia; he assigned the African species Lasiohelea brevi-
costa Clastrier to this subgenus. We have found two additional undescribed
species that fit in this group and with sufficiently distinct characters of the
adult and pupa to prompt us to described a new subgenus to contain them.

Forcipomyia, subgenus Saliohelea Wirth and Ratanaworabhan,
new subgenus

Fig. 1

Forcipomyia, subgenus C; Tokunaga and Murachi, 1959:219, Dessart,
1963:24.

Type-species—Forcipomyjia (Saliohelea) leei Wirth and Ratanaworabhan,
new species.

Description—Small, unmarked, yellowish to brownish species. Female
antenna (Fig. 1b) with five distal segments not much longer than preced-
ing segments, AR 0.78-1.20. Palpus (Fig. la) 4-segmented, primitive 4th
and 5th segments fused without trace of articulation, characteristically
tapering to slender tip; 3rd segment with or without sensory pit. Mandible
teeth absent.

Body, wings and legs usually more or less clothed with elongate, slender,
scalelike hairs with 1-3 stripes or striations. Costa (Fig. 1d) short, costal
ratio 0.41 to 0.52 in female. Hind tibial comb (Fig. lh) with 4-5 slender
spines in distal series. Hind tarsal ratio varying from 1.7 to 2.7; hind
basitarsus of male sometimes expanded dorsally. Empodium (Figs. 1g and
1j) well developed in both sexes; claws rather slender and curved.

Female with a single well-developed spermatheca (Fig. li), this usually
tapering to slender neck, a minute rudimentary 2nd spermatheca usually
present. Male genitalia (Figs. lm and In) similar to those of Euprojoannisia
or Warmkea Saunders. Ninth sternum more or less transverse distally, with
long, slender, anterolateral projections; 9th tergum short and tapering to
moderately separated, setose, apicolateral lobes. Basistyle moderately
stout; dististyle long and slender, nearly straight, tapering to tip. Aedeagus
somewhat variable in form, usually with triangular basal portion without
well-developed basal arch or lateral arms, this portion usually with distinct

VOLUME 80, NUMBER 4 495

marginal sclerotization; slender distal median process usually present,
rounded or peglike, in 1 species with a submedian pair of slender posterior
processes. Parameres with basistylar apodemes forming a more or less
V-shaped basal arch, with a distinct mesal connective which is usually
platelike but may be narrow; a pair of submedian posterior sclerotized
processes may be present (completely absent in 2 species, short and pointed
in 1 species, long and curved in 1 species ).

Pupa with slender respiratory horn (Fig. le) bearing 6 spiracular open-
ings spaced loosely around the apical margin giving it a crinite appearance.

Etymology.—The name Saliohelea is an anagram of Lasiohelea Kieffer,
another subgenus of Forcipomyia.

Discussion—Species of Saliohelea are most closely related to the sub-
genus Euprojoannisia (synonyms Proforcipomyia and Euforcipomyia
Malloch), which they resemble in their small, unmarked, yellowish-brown
appearance; tarsal ratio greater than 1.0; curved, slender tarsal claws,
rather short distal antennal segments; and reduction of the last two palpal
segments. Species of Euprojoannisia, however, differ in having palpal seg-
ments four and five both present, although their articulation is lost and
they are partially fused; mandibular teeth are usually present in the fe-
male; striated scales are lacking on the body; two spermathecae are
usually present; the mesal connective of the male basistylar apodemes is
usually slender and rarely are indistinct posterior processes present; and
the pupal respiratory horn is usually short and swollen with the spiracles
arranged in a tight circle or partial circle at the apex.

The subgenus Warmkea is also close to Saliohelea. The resemblance is
especially close in the complete loss of the fifth palpal segment, the
presence of only one spermatheca, the general features of the male para-
meres, and the shape of the pupal respiratory horn. However, in Warmkea
the distal antennal segments are usually much elongated, female mandibular
teeth are present, the costa is much longer, the male aedeagus has a distinct
shieldlike shape with a well-developed basal arch and no lateral thickenings,
and the spiracles are arranged in a tight row on the pupal respiratory horn.

Biological information is available only for Forcipomyia (Saliohelea)
leei, new species. Saunders reared the species from a pupa taken under
bark of a fallen tree in the forest in Brazil, and Williams reared it from
rotting pods of cacao in Trinidad. Numerous collections of F. leei were
made in rain forest locations.

The following provisional key to the subgenera of Forcipomyia is
presented for the convenience of workers trying to identify adult
Forcipomyia material. The table to the subgenera presented by Saunders
(1956) and the key in Dessart (1963) are now incomplete because of the
recent description of additional subgenera. There admittedly will be many

496

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

difficulties in using this key because some subgenera are most distinct in
the male sex, others in the female. The subgeneric classification of
Forcipomyia is based primarily on the relatively good characters of the
immature stages, and identification of the adults probably will always be
difficult.

Key to the Subgenera of Forcipomyia
(for adults, primarily females )

Female antenna with distal 6 segments elongated; empodium
large and broad, adapted for clinging to wings of insects; TR 3.0
or more Pterobosca Mactie
Female antenna with distal 5 segments, if any, elongated,
rarely distal 6; empodium not greatly enlarged or modified;

TR 0.4-3.2 2
Palpus 4-segmented, only 1 segment distal to the 3rd (which
bears the sensory organ) 3

Palpus 5-segmented (segments 4 and 5 incompletely fused and

non-articulated in Euprojoannisia ) 8
Female with 2 large functional spermathecae 4
Female with one large functional spermatheca 6
Body with deep greenish or bluish subcutaneous pigmentation,

especially in abdomen Caloforcipomyia Saunders (part)

Body without greenish or bluish subcutaneous pigmentation 5
Female antenna with 5 distal segments greatly elongated, the
proximal ones short and globular; TR 2.66-3.00
Blantonia Wirth and Dow
Female antenna with distal segments not much longer than
those in proximal series, all segments elongate tapering; TR
about 2.0 Metaforcipomyia Saunders
Female antenna with 5 distal segments much longer than the
short proximal segments; costa long, CR usually about 0.67;
TR 1.3-1.75; female mandibular teeth well developed
Warmkea Saunders
Female antenna with distal segments not much longer than
those in proximal series, all segments short ovoid to moderately
long tapering; costa variable; TR variable; mandibular teeth not
developed 7
Basitarsi short, TR about 1.0; costa short, CR less than 0.5
genus A near Lepidohelea Kiefter
Basitarsi elongate, TR about 2.0; costa short or moderately
long Saliohelea, new subgenus

VOLUME 80, NUMBER 4 497

8(2).

10.

i

13.

14.

Female antennal segments of proximal series much shorter than

distal 5, compressed, usually transverse 9
Female antennal segments of proximal series not much shorter
than distal segments, usually not compressed 12

Costa extending to well beyond middle of wing; 2nd _ radial
cell much longer than Ist, very narrow; 1 subspherical sperma-
theca present, this without neck; female sucking vertebrate
blood Lasiohelea Kieffer
Costa usually ending at or near middle of wing; 2nd radial cell
not unusually long and narrow; 2 spermathecae with distinct
necks present; not known to suck vertebrate blood 10
Second and 3rd palpal segments stout, 3rd with scattered
sensilla or shallow pit; mandible with proximal teeth very
strong; male empodium present; male parameres U-shaped
Rhynchoforcipomyia Wirth and Dow
Third palpal segment with definite, deep sensory pit; mandib-
ular teeth small and more uniform; male empodium and _para-
meres various 11
Females suck blood from insects; male empodium present; male
parameres U-shaped; larvae breed in moss and wet wood
Trichohelea Goetghebuer
Female feeding habits unknown; male empodium usually ab-
sent; male parameres H-shaped; larvae breed in plant leaf axils
Phytohelea Remm

. Fourth and 5th palpal segments incompletely fused, immovable;

smalled, unmarked, brownish midges; TR greater than 1.0
Euprojoannisia Brethes
Fourth and 5th palpal segments distinctly separated, articulated;
size, color, and TR various 13
Palpus with 3rd segment broadly swollen to past middle,
usually nearly to tip, with sensory pit deep, extending nearly
to base of segment; slender, hyaline, peglike, sensory spines
present on surface near sensory pore; TR usually less than 0.5;
large species, females suck insect blood Microhelea Kieffer
Palpus various, 3rd segment rarely swollen past midlength and
not bearing peglike sensory spines on surface near sensory
pore; TR usually more than 0.5; size and habits various 14
Body usually with metallic jade green or deep blue subcutane-
ous pigmentation; antenna unusually elongate and _ slender;
palpus slender; TR 1.36-2.32; wing and body often with ornate
color pattern Caloforcipomyia Saunders (part)

498 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

- Body without greenish or bluish subcutaneous pigmentation;
antenna usually much shorter; palpus, TR and coloration
various 15

15. Small grayish or brownish unmarked species; TR usually
greater than 2.0; antenna short, proximal segments subspheri-
cal, gradually more elongated distally; 1 spermatheca present 16

- Larger species, often with conspicuous markings on body, legs
or wings; TR usually about 1.0 (0.5-1.5); 2 well-developed

spermathecae present ily
16. Male genitalia with club-shaped parameres extending caudad
from basistylar apodemes Synthyridomyia Saunders

- Male genitalia without club-shaped parameres
Thyridomyia Saunders
ie Body with conspicuous, numerous, flattened scales in addition
to normal setae and hairs; male genitalia with parameres not
joined or fused at bases, dististyle elongate, sinuate with
distinct distal expansion Lepidohelea Kieffer
- Body usually without scales, or if present they are usually not
broad; male genitalia with parameres joined or fused at bases,
dististyle not expanded distally 18
18. Male genitalia with aedeagus V-shaped, bearing a pair of small,
sharp processes at tip Schizoforcipomyia Chan and LeRoux
- Male genitalia with aedeagus shield-shaped, with low basal
arch Forcipomyia, s. str.

Forcipomyia (Saliohelea) leei Wirth and Ratanaworabhan, new species
Fig. 1

Female.—Wing length 0.62 mm; breadth 0.26 mm.

Head: Brownish, palpus paler. Proboscis (Fig. 1f) short. Antenna
(Fig. lb) with lengths of flagellar segments in proportion of 17-15-15-16-
17-17-17-19-25-23-22-20-28; antennal ratio 0.89; segment 11 unusually long
and segments 12-14 becoming progressively shorter; segments 4-10 sub-
spherical to slightly elongate, 5 distal segments distinctly tapering to slender
distal portions; 15 with distinct terminal papilla. Palpus (Fig. la) 4-seg-
mented; lengths of segments in proportion of 10-12-38-23; PR 2.2; 3rd
segment ovoid with shallow, round, sensory pit; 4th segment markedly
tapered to slender tip. Mandible without visible teeth.

Thorax: Dark brown, pleuron paler. Legs (Fig. 1k) variably brownish
to pale yellowish; with sparse setae and 1-striped slender scales; no broad
scales; hind tibial comb as in Fig. lh. Tarsi (Fig. 11) unmodified; hind
TR 2.7. Empodium (Fig. 1g) well developed; claws curved, moderately

VOLUME 80, NUMBER 4 499

= = (Lok EERE A
ee ee aeeee =
LS RI at =e

4

~S

Wigs
(| iiny

Z < Sears
paras ee
RESET

:

‘cae
g h
\

)

Fig. 1. Forcipomyia (Saliohelea) leei. a, female palpus; b, female antenna; c,
male wing; d, female wing; e, pupal respiratory horn; f, female head; g, female fifth
tarsomeres and claws of (left to right) front, middle and hind legs; h, hind tibial comb;
i, spermatheca; j, male claws of (left to right) front, middle and hind legs; k, femora
and tibia, of (left to right) hind, middle and front legs of female; 1, tarsi of (left to right)

hind, middle and front legs of female; m, male parameres; n, male genitalia, parameres
removed.

500 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

stout. Wing (Fig. ld) pale brownish due to moderately large micro-
trichia; macrotrichia especially long and coarse, decumbent, resembling
slender, 1-striped scales; Ist radial cell obsolete, 2nd long and narrow;
CR 0.52. Halter brownish.

Abdomen: Brownish, terga dark brown; last segment yellowish. Sperma-
theca as in Fig. lh; one large ovoid functional spermatheca with long
slender neck, usually with a minute irregularly tubular rudimentary
spermatheca present; functional spermatheca measuring 0.052 by 0.030 mm
including neck.

Male.—Wing length 0.64 mm; breadth 0.25 mm. Similar to female with
usual sexual differences. Antenna with lengths of flagellar segments in
proportion of 20-16-17-19-16-16-18-20-20-58-35-27-39; plume moderately
long and dense, brownish; segments with 7-10 fused. Wing (Fig. Ic)
with macrotrichia sparser than in female; CR 0.60. Hind TR 2.5; hind
basitarsus slightly enlarged and darkened. Empodium moderately de-
veloped; claws (Fig. 1j) more slender than in female, without bifid tips.

Genitalia (Fig. In): Ninth sternum short with long anterolateral pro-
jections; 9th tergum short and tapering to moderately separated, setose,
apicolateral lobes. Basistyle simple, about 2x as long as broad; dististyle
long and slender, nearly straight, tapering to slightly bent, pointed tip.
Aedeagus triangular in ventral view, basal margin transverse or nearly so,
with short anterolateral processes; proximal and lateral margins all with
similar internal sclerotized thickening; apex with distinct slender terminal
papilla. Parameres (Fig. lm) with diagonal basal apodemes connected by
a well-developed transverse connective, slightly arcuate, the concave side
anteriad; a short pair of slender, pointed processes projecting caudad from
junction of apodeme and transverse connective, the length of these pro-
cesses slightly variable, but usually less than length of aedeagus.

Pupa—tLength 1.7 mm. Color pale ochreous throughout. Head with a
prominent tubercle at each anterolateral angle of antennal cases; a single
low papilla and 2 pairs of minute spines on medium triangle; a small in-
conspicuous papilla on each lateral triangle. Thorax with 3 pairs of bulbous
papillae on dorsum, and 1 minute pair at base of posterior prolongation,
the latter extending across Ist abdominal segment. Prothoracic respiratory
horn (Fig. le) very distinctive; with posterior basal shoulder similar to that
of Euprojoannisia; distal portion enlarged, with 6 double or triple spiracular
papillae arranged at intervals around margin giving a crinate appearance;
tracheae of each papilla radiating from central felt chamber. Abdomen with
many microchaetae on all surfaces of each segment; no large spines. Ter-
minal abdominal processes long and slender, 1.5 length of basal portion
of 9th segment; male appendages dorsal, very short, each with a small
lateral spine.

|

VOLUME 80, NUMBER 4 501

Distribution Eastern U.S.A., Neotropical Region to southern Brazil.

Types.—Holotype, °, Rio Raposo, Valle, Colombia, June-July 1963,
V. H. Lee (Type no. 70440, USNM). Allotype, ¢, same data but July
1965, in light trap.

Paratypes, 43 ¢é and 352 as follows: COLOMBIA: Rio Raposo, Valle,
Jan.—Dec. 1963-1965, V. H. Lee, light trap, 22 ¢, 18°. Rio Anori, Antioquia,
Sept. 1970, D. G. Young, light trap in rain forest, 2 4, 2°. BRAZIL: Belem,
Para, April 1970, T. H. G. Aitken, light trap, 1¢. Nova Teutonia, Santa
Catarina, Sept. 1961, F. Plaumann, 12. Rio de Janeiro, 31 July 1923, L. G.
Saunders, reared from beneath bark of fallen tree in forest, 1¢ and
associated pupal exuviae. Rio Preto, Amazonas, 7 June 1962, E. J. Fittkau,
at light, 1¢. TRINIDAD: No locality, Aug. 1963, R. W. Williams, reared
from rotting cacao pods, 1 ¢, 42. DOMINICA: Cabrit Swamp, 23 Feb.
1965, W. W. Wirth, 1 2. Central Forest Reserve, 11 May 1968, P. C. Drum-
mond, black light, 12. d’Leau Gommier, 17 March 1956, J. F. G. Clarke,
1é. Fond Figues River, 11 Feb. 1965, W. W. Wirth, rain forest, 1é. Pont
Casse, April 1964, O. S. Flint, at light, 1 ¢, 1 2; same, 12 Feb. 1965, W. W.
Wirth, rain forest, 2°. PUERTO RICO: El Verde, Barrio Rio Grande,
G. E. Drewery, sticky trap, 1 2. Mayaguez, Univ. Puerto Rico Campus,
13 Aug. 1969, T. Walker and P. Drummond, 12. JAMAICA: Hardwar
Gap, 10 May 1970, W. W. Wirth and T. Farr, malaise trap, 36, 22. U.S.A.:
FLORIDA: Alachua Co., Gainesville, June 1967, F. S. Blanton, light trap,
26,12. Jefferson Co., Monticello, Oct. 1969, W. H. Whitcomb, light trap,
1é. Leon Co., 3 mi N Tallahassee, May 1970, F. S. Blanton, light trap, 1 ¢.
Marion Co., Juniper Springs, 28 Apr. 1970, W. W. Wirth, light trap, 1 ¢.
Orange Co., Lake Magnolia Park, 6 Aug. 1970, E. Irons, light trap, 1 2°.
SOUTH CAROLINA: Georgetown Co., Hobcaw House, 20 Apr. 1972,
Mrs. L. Henry, light trap, 2 ¢. VIRGINIA: Fairfax Co., Falls Church,
Holmes Run, 7 Sept. 1961, W. W. Wirth, light trap, 1 ¢. NEW YORK:
Newcomb, Lake Harris, 19 Aug. 1972, L. Knutson, malaise trap, 1 ¢.

Etymology.—This species is named for Vernon H. Lee in recognition of
his extensive contributions to our knowledge of Colombian biting midges.
Dr. Lee collected ceratopogonids extensively for us in Colombia as a mem-
ber of a Rockefeller Foundation arbovirus research team.

Discussion.—F orcipomyia leei is closely related to the African species F.
stami, new species, with differences as discussed under that species.
Forcipomyia leei is a very widespread species, occurring from the north-
eastern United States to southern Brazil and, as might be expected, varies
somewhat in leg color and in the length of the posterior processes of the
male parameres. We have been unable to correlate this variation with geo-
graphical distribution or other factors and conclude that our material is
conspecific.

502 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Forcipomyia (Saliohelea) stami Wirth and Ratanaworabhan, new species
Fig. 2

Female—Wing length 0.66 mm; breadth 0.34 mm.

Head: Brown, including palpi and antennae. Antenna (Fig. 2a) with
lengths of flagellar segments in proportion of 20-14-15-15-15-16-17-20-26-27-
27-24-34; antennal ratio 1.00; segments 4-10 progressively globular to short
tapering, 11-14 moderately elongate, tapering. Palpus (Fig. 2d) with
lengths of segments in proportion of 14-13-36-34; 3rd segment ovoid with
large, round, shallow sensory pit; PR 1.9; 4th segment tapering to moder-
ately slender tip. Mandible without visible teeth.

Thorax: Dark brown, pleuron slightly paler. Legs yellowish, distal
tarsomeres slightly infuscated; hind tibial comb (Fig. 2c) with 5 subequal
setae; hind TR 1.86; empodium well developed; claws slender and curved,
pointed distally (Fig. 2f). Wing (Fig. 2b) pale brownish due to moder-
ately large microtrichia; macrotrichia long and coarse, 1-striated, moderately
dense, without color pattern; Ist radial cell obsolete, 2nd slitlike; costa
exceptionally short, CR 0.41. Halter brownish.

Abdomen: Brownish including terminal segments and cerci. Sperma-
thecae (Fig. 2e): 1 large ovoid functional spermatheca with long slender
neck, measuring 0.058 by 0.032 mm; a tiny, ovoid, well-sclerotized, rudi-
mentary spermatheca present.

Male——Wing length 0.74 mm; breadth 0.25 mm; CR 0.44. Similar to
female, with usual sexual differences. Hind TR 1.90.

Genitalia as in Fig. 2h: Ninth sternum moderately short, with distinct
caudomedian excavation; 9th tergum tapering to rather closely approxi-
mated apicolateral lobes. Basistyle rather slender; dististyle straight to
slender, slightly pointed tip. Aedeagus slightly longer than basal breadth;
basal arch low; outline triangular in ventral view, the sides moderately
sclerotized with a thinner, slightly concave, marginal rim; apex produced
slightly in a slender terminal papilla. Parameres (Fig. 2g) with slender,
diagonal basal apodemes joined mesally in a short, quadrate, transverse
connective; no trace of submedian caudal processes from this platelike
connective.

Distribution.—Zaire.

Types.—Holotype, °, allotype, ¢, Zaire, Coquilhatville, March-April
1972, A. B. Stam (Type no. 17305, USNM ).

Paratypes, 2 ¢,1 2, same data.

Etymology.—This species is named for its collector, Professor A. B.
Stam, Director of the Institute of Entomology and Parasitology of Africa,
Kumasi, Ghana, in recognition of his keen interest in the taxonomic study
of African biting midges.

VOLUME 80, NUMBER 4 503

Fig. 2. Forcipomyia (Saliohelea) stami. a, female antenna; b, female wing; c, hind
tibial comb; d, female palpus; e, spermatheca; f, fifth tarsomere and claws of (left to
right) front, middle and hind legs of female; g, male parameres; h, male genitalia,
parameres removed.

Discussion.—Forcipomyia stami is closely related to the American F. leei,
appearing almost identical to some variants of leei, but differing as follows:
In stami the costa is shorter, the distal antennal segments are longer, the
hind basitarsus is shorter and the posterior processes are lacking in the
male parameres. The latter character causes a problem with the subgeneric
diagnosis, but occasional variation of the same character in the related
subgenus Euprojoannisia shows the same instability and is not considered
critical to subgeneric recognition.

504 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Forcipomyia (Saliohelea) brevicosta ( Clastrier )
Fig. 3

Lasiohelea brevicosta Clastrier, 1960:520 (2; Congo; fig. wing, antenna,
palpus).

Forcipomyia (subgenus C) brevicosta (Clastrier); Dessart, 1963:47 (re-
described; compared with deminuta T. & M.; combination).

Female.—Wing length 0.83 mm; breadth 0.33 mm.

Head: Brown. Antenna (Fig. 3a) with lengths of flagellar segments in
proportion of 25-20-20-20-20-21-23-25-37-37-40-40-53; AR 1.20; distal 5
segments distinctly more elongate than in F. leei, without the peculiar
shortening of the subapical segments. Palpus (Fig. 3b) with lengths of
segments in proportion of 10-15-36-34; 3rd segment without definite sensory
pit; palpal ratio 2.1 (Clastrier’s figure shows the 3rd palpal segment stouter,
with palpal ratio 1.8).

Thorax: Dark brown. Legs uniformly yellowish brown; hind TR 1.83
(2.0 in holotype according to Clastrier), much lower than the TR of 2.7
found in F. leei; hind basitarsus not swollen. Tibial comb as in Fig. 3d,
5th tarsomeres and claws as in Fig. 3f. Wing (Fig. 3c) with CR 0.48;
macrotrichia broader than in F. leei, more scalelike with up to 3 or 4
striations, compared with a maximum of | stripe in F. leei. Halter brown.

Abdomen: Uniformly brown. Spermatheca (Fig. 3e) ovoid to pyriform,
tapering to duct; without the slender neck found in F. leei; measuring
0.058 by 0.033 mm; a minute, oval, rudimentary 2nd spermatheca present.

Male.—As in the female with the usual sexual differences. Genitalia
(Fig. 3h) with 9th segment similar to those of F. leei. Basistyle moderately
slender, slightly curved; dististyle slightly more slender and more curved
than in leei. Aedeagus about as broad as long, basal arch low, basal arms
stout and scarcely differentiated; distal portion cleft a short distance in a
contiguous pair of short, curved, strongly sclerotized processes. Parameres
(Fig. 3g) with long, slender, oblique basal apodemes, anterior connective
short, bearing a posterior pair of long, curved, moderately stout, strongly
sclerotized posterior processes, their apices slightly crossed on midline just
distad of tip of aedeagus.

Distribution—West Africa (Rep. Congo, Nigeria).

Type.—Holotype, ?, Nuku N’Situ, Rep. Congo, May 1956, R. Taufflieb,
(in Inst. Pasteur d’Algerie, Algiers).

Material examined—NIGERIA: Redescribed from 5 ¢, 2 ?, Ibadan,
Aug. 1962, D. C. Eidt, malaise trap (through Canadian National Collection,
Ottawa).

Discussion —The present material agrees well with Clastrier’s original
description, except as noted above. The most important differences from

VOLUME 80, NUMBER 4 505

Fig. 3. Forcipomyia (Saliohelea) brevicosta. a, female antenna; b, female palpus;
c, female wing; d, hind tibial comb; e, spermatheca; f, fifth tarsomere and claws of
(left to right) front, middle and hind legs of female; g, male parameres; h, male
genitalia, parameres removed.

F. leei, which while still permitting assignment to the subgenus Saliohelea
require a slight adjustment in the subgeneric concept, are the more elongate
distal antennal segments, the lack of a sensory pit on the third palpal seg-
ment, the more elongate, curved parameres and the distinctly cleft tip of
the male aedeagus.

Forcipomyia (Saliohelea) deminuta Tokunaga and Murachi

Forcipomyia (subgenus C) deminuta Tokunaga and Murachi, 1959:219
(4, 2; Palau and Caroline Islands; figs.).

Female.—Wing length 0.62 mm; breadth 0.25 mm. A tiny, uniformly
pale yellowish-brown species without distinctive markings. Antenna with
all flagellar segments short tapering, lengths in proportion of 10-8-8-8-8-9-9-

506 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

9-9-9-9-10-15; AR 0.78. Palpal segments with lengths in proportion of 4-10-
24-19; PR 2.6; 3rd segment slightly swollen at base, slender distally, with
an indistinct, small, shallow, round sensory pit at midlength; 4th segment
quite slender, slightly narrowed on distal portion. Mandible without visible
teeth. Legs with TR 2.2 on front leg, TR 1.71 on middle leg, and TR 1.67
on hind leg; tarsi with numerous 1-3 striped, slender, scalelike hairs. Wing
with CR 0.48. Halter pale yellowish. Abdomen yellowish; spermatheca
single, ovoid, tapering to slender neck (Tokunaga and Murachi’s figure
showing two spermathecae is apparently in error; it was stated to be single
in their description ).

Male.—Wing length 0.80 mm; breadth 0.24 mm. As in female with usual
sexual differences; CR 0.47; TR 2.38 on front leg, 1.81 on middle leg, and
1.81 on hind leg; hind basitarsus distinctly swollen. Genitalia with aedeagus
small and triangular without well-developed basal arch or lateral arms, the
anterior and lateral margins with distinct narrow internal sclerotization
(not shown in Tokunaga and Murachi’s figure); median distal process
slender, peglike. Parameres with diagonal basistylar apodemes, the sclero-
tization of the median connective indistinct and poorly visible (shown as
quite extensive in Tokunaga and Murachi’s figure); posterior processes not
visible.

Distribution.—Micronesia, Philippines.

Types.—Holotype, ¢, allotype, 2°, Mt. Temwetemwensekir, Ponape,
Caroline Islands, 180 m, 19 Jan. 1953, J. L. Gressitt (in Bishop Museum,
Honolulu ).

Material examined—CAROLINE ISLANDS: 2 ¢, same data as types
(in USNM). PHILIPPINE ISLANDS: 7 2, Mindanao, Mt. Apo, Davao
Prov., 15 Nov. 1947, H. Hoogstraal and F. Werner (Chicago Nat. Hist.
Mus. and USNM).

Discussion—This species was well illustrated by Tokunaga and Murachi
(1959). The Philippine material conforms closely to the original descrip-
tion of the female from Micronesia, and we add this new distribution
without hesitation.

Acknowledgment

Acknowledgment is gratefully made by the junior author to the Southeast
Asia Treaty Organization of Bangkok for financial assistance for study at
the U.S. National Museum.

Literature Cited

Barroga, S. F. 1964. Progress report on the study of insects, particularly midges
associated with pollination of Theobroma cacao, April, 1963. Philipp. J. Plant
Ind. 29:123-133.

VOLUME 80, NUMBER 4 507

Bystrak, P. G. 1974. A revision of the Nearctic species of Euforcipomyia Malloch, a
subgenus of Forcipomyia Meigen (Diptera: Ceratopogonidae). Unpublished
M.S. Thesis, University of Maryland, College Park. August 1974. 121 pp.

Chan, K. L. and E. J. LeRoux. 1965. Description of Forcipomyia (Neoforcipomyia )

sp. n. and redescription of Forcipomyia (Neoforcipomyia) eques (Johannsen )

(Diptera: Ceratopogonidae), with an account of the digestive and reproductive

systems. Phytoprotection 46:74—104.

1971. Phyogenetic relationships in the Forcipomyiinae (Diptera: Cerato-

pogonidae ). Can. Entomol. 103:1323-1335.

Clastrier, J. 1960. Notes sur les cératopogonidés XI.—Cératopogonidés de la Ré-
publique du Congo (3). Arch. Institut Pasteur d’Algérie 38:510-526.

Dessart, P. 1961. Contribution a l'étude des Ceratopogonidae (Diptera). Les

Forcipomyia pollinisateurs du cacaoyer. Bull. Agric. Congo Belge 52:525-540.

1963. Contribution a l’étude des Ceratopogonidae (Diptera) (VII). Tab-
leaux dichotomiques illustrés pour la détermination des Forcipomyia Africains.
Mem. Inst. R. Sci. Nat. Belge (2 Ser.) 72:1-151.

Kaufmann, T. 1975. Ecology and behavior of cocoa pollinating Ceratopogonidae in
Ghana, W. Africa. Environ. Entomol. 4:347-351.

Saunders, L. G. 1956. Revision of the genus Forcipomyia based on characters of all
stages (Diptera, Ceratopogonidae). Can. J. Zool. 34:657—705.

1959. Methods for studying Forcipomyia midges, with special reference to
cacao-pollinating species (Diptera, Ceratopogonidae). Can. J. Zool. 37:33-51.
Soria, S. de J. and W. W. Wirth. 1974. Identidade e caracterizacao taxondmica

preliminar das mosquinhas Forcipomyia (Diptera, Certopogonidae) associadas
com a polinizacao do cacaueiro na Bahia. Rev. Theobroma 4(1):3-12.
Tokunaga, M. and E. K. Murachi. 1959. Insects of Micronesia. Diptera: Cerato-
pogonidae. B. P. Bishop Museum. Insects Micronesia 12:103—434.
Winder, J. 1977. Field observations on Ceratopogonidae and other Diptera: Nemato-
cera associated with cocoa flowers in Brazil. Bull. Entomol. Res. 67:57-63.

(WWW) Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. and
Educ. Admin., USDA, c/o U.S. National Museum, Washington, D.C.
20560; and (NCR) Applied Scientific Research Corporation, Bangkok, Thai-
land.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 508-513
APHIDS OF SUNFLOWER: DISTRIBUTION AND HOSTS IN
NORTH AMERICA (HOMOPTERA: APHIDIDAE)'

C. E. Rogers, T. E. Thompson and M. B. Stoetzel

Abstract.—Distribution and host data are given for aphids on 15 species
of Helianthus collected in the United States in 1976 and 1977. Aphids of
the genus Dactynotus were found primarily on eastern perennial species
of Helianthus. Aphis helianthi Monell and Masonaphis masoni (Knowlton)
were found mostly on western annual species of Helianthus. A summary
of previous records for aphids on Helianthus is also given.

Little is known about the bionomics or effects of aphids on sunflower.
Rogers et al. (1972) reported that Aphis helianthi Monell on Helianthus
annus L. served as a good alternate host for parasitoids released to control
the greenbug, Schizaphis graminum (Rondani), on sorghum. Most reports
of aphids on sunflower have been included in regional surveys (Williams,
1910; Hottes and Frison, 1931; McGillivray, 1958; Leonard, 1959; Palmer,
1952; Leonard and Tissot, 1965; Leonard, 1968; and Olive, 1963) and
taxonomic studies (Olive, 1965). Some of the aphids that are known from
sunflower are well-known vectors of virsuses (Kennedy et al., 1962).

Sunflower has become an important oilseed crop in the United States.
Because of the importance of aphids as pests on other crops (Gibson and
Plumb, 1977), this study was designed to determine the species and
abundance of aphids on native Helianthus species.

Materials and Methods

Two of us (CER and TET) drove about 24,000 km in the southern half
of the United States in 1976 and 1977 and collected native species of
Helianthus and associated aphids. We paid particular attention to the
effect of the aphids on their host plants. Aphids were collected in 70%
ethyl alcohol for subsequent sorting and identification. Plants were identi-
fied in situ according to Heiser (1969). Specimen mounts of plants were
also prepared and later confirmed by Dr. C. B. Heiser, Jr. MBS identified
the aphids, and voucher specimens have been placed in the U.S. National
Museum Collection at Beltsville, Maryland.

Results and Discussion

Some species of aphids reported from sunflower normally use other
genera as primary hosts (Table 1). For example, Aphis gossypii Glover

VOLUME 80, NUMBER 4 509

Table 1.—Aphids known from Helianthus species in North America.

Aphid species Helianthus host* Location Authority
Aphis armoraciae Cowen — pumilis Nuttall (P) Rocky Mtns. Palmer (1952)
A. debilicornis annuus (A) Rocky Mtns. Palmer (1952)
(Gillette & Palmer) grossesserratus MO Leonard (1963 )
Martens (P)
tuberosus L. (P) Rocky Mtns.; Palmer (1952)
MO
Leonard (1959)
A. gossypii Glover cultivar (A) TX Leonard & Tissot
(1965 )
species (?) DEX Leonard & Tissot
(1965 )
A. helianthi Monell annuus Rocky Mtns.; KS Palmer (1952);

Walker (1936)
petiolaris Nuttall (A) Rocky Mtns. Palmer (1952)

tuberosus MO Leonard (1963 )

species (?) KS; MO; NE; Walker (1936);
OK; Rocky Leonard (1963);
Mtns.; TX Williams (1910);

Rogers et al.
(1972); Palmer
(1952); Leonard

and Tissot
(1965)
Bipersona ochrocentri annuus MO Leonard (1962)
(Cockerell )
Dactynotus ambrosiae annuus Rocky Mtns. Palmer (1952)
(Thomas ) grosseserratus MO Leonard (1963)
species (?) TX; (?) Leonard & Tissot
(1965 ); Olive
(1965 )
tuberosus MO Leonard (1963)
D. helianthicola Olive atrorubens L. (P) NC Olive (1963)
microcephalus Torrey NC Olive (1963)
& Gray (P)
strumosus L. (P) NC Olive (1963 )
tuberosus (P) NC Olive (1963 )
species (?) TX Leonard & Tissot
(1965 )
D. illini species (?) (?) Olive (1965)

(Hottes & Frison )

@ A = annual; P = perennial

510 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Continued.

Aphid species Helianthus host* Location Authority
D. obscuricaudatus Olive — strumosus PA Olive (1965)
D. parvtotubercultus Olive atrorubens NC Olive (1965)
D. pseudambrosiae Olive — microcephalus NC Olive (1965)
D. rudbeckiae ( Fitch ) species (?) ex Leonard & Tissot
(1965 )
D. ruralis species (?) Pp Olive (1965)
(Hottes & Frison )
Dactynotus sp. grosseserratus MO Leonard (1959)
tuberosus MO Leonard (1963 )
Macrosiphum euphorbiae — annuus Rocky Mtns. Palmer (1952)
( Thomas ) species (?) Rocky Mtns. Palmer (1952)
Macrosiphum sp. tuberosus MO Leonard (1959)
Masonaphis masoni annuus (?) MacGillivray
(Knowlton ) (1958 )
species (?) (?) MacGillivray
(1958 )
species (?) CO Palmer (1952)
Prociphilus erigeronensis annuus Rocky Mtns. Palmer (1952)
( Thomas ) species (?) CO Palmer (1952)

(cotton aphid) and Macrosiphum euphorbia (Thomas) (potato aphid) are
common pests on the crops indicated by their common names. Also, the
species names for Aphis amoraciae Cowen, Dactynotus ambrosiae
(Thomas), D. rudbeckiae (Fitch), and Prociphilus erigeronensis (Thomas)
implicate genera other than Helianthus as the primary host plants for these
aphids. The published records suggest that Aphis species commonly colon-
ize annual species of Helianthus, whereas Dactynotus species colonize
perennial species of Helianthus.

We collected 47 species of Helianthus, 15 of which harbored aphids
(Table 2). Aphids were also collected from “Hybrid 896° and from a
cultivar of unknown parentage. Our data verified that Dactynotus species
colonize primarily perennial Helianthus species and that other genera occur
mostly on annual sunflowers. We have shown that several species of
perennial Helianthus are resistant to Masonaphis masoni (Knowlton), a
species that is common on annuus types and cultivated sunflower (Rogers
and Thompson, 1978). Also, Dactynotus helianthicola Olive failed to
survive in the laboratory when transferred from the perennial H. occiden-

VOLUME 80, NUMBER 4 511

Table 2.—Aphids collected from Helianthus species in the United States during
1976 and 1977.

Aphid species Helianthus host" Location
Aphis deblicornis nuttallii Torrey & Gray (P) CO
(Gillette & Palmer )
A. helianthi Monell annuus (A) AR; CA; CO;
KS: NM; NV;
AD Ke WAL
neglectus Heiser (A) NM
petiolaris (A) CO
Hybrid 896 (A) TX
Dactynotus ambrosiae petiolaris CO
(Thomas )
D. helianthicola heterophyllus Nuttall (P) MS
Olive tuberosus (P) OK; SC
microcephalus (P) AL
occidentalis (P ) MO
longifolius Pursh (P) AL
atrorubens (P) NC
silphioides Nuttall (P) OK
Dactynotus sp. argophyllus Torrey & Gray (A) TX
grosseserratus (P) TX
maximiliani Schrader (P) TX
petiolaris NM
tuberosus xX
Macrosiphum euphorbiae __ petiolaris CO
(Thomas ) cultivar CA
Hybrid 896 ADS
Masonaphis masoni paradoxus Heiser (A) TX
( Knowlton ) Hybrid 896 TX
nuttallii (P) CO
Myzus persicae (Sulzer) cultivar CA
Hybrid 896 TX
Rhopalosiphum sp. annuus NM
Hybrid 896 TX

* A = annual; P = perennial

talis Riddell to Hybrid 896. These narrow host preferences among sun-
flower aphids become very important in the development of aphid-resistant
sunflower hybrids.

The host and distribution data shown in Table 2 (but not in Table 1)
probably represent new records. For the most part, Dactynotus species

512 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

were found primarily on eastern perennial species of Helianthus, and A.
helianthi mostly on western annual species of Helianthus. Dactynotus spe-
cies feed mainly on the upper stems and leaf petioles of sunflower. Aphis
helianthi and M. masoni feed mainly on lower leaf surfaces and on the
underside of receptacles. Macrosiphum euphorbia feeds on the terminal of
plants and on ray flowers around the receptacle. One alate Hysteronuera
setariae (Thomas) adult was found on H. microcephallus Torrey and Gray
in North Carolina, probably as a result of an accidental landing.

Acknowledgments

We are grateful to Dr. C. B. Heiser, Jr., Department of Biology, Indiana
University, Bloomington, Indiana, for verifying the identification of
Helianthus species.

Travel for collecting Helianthus species was made possible by a grant
from the Deputy Administrator's reserve funds, Fed. Res., Sci. and Educ.
Admin., USDA, New Orleans, Louisiana.

Literature Cited

Gibson, R. W. and R. T. Plumb. 1977. Breeding plants for resistance to aphid
infestation. Chapter 20:473-500. In Aphids as Virus Vectors. K. F. Harris
and K. Maramorosch [eds.]. Academic Press. 559 pp.

Heiser, C. B., Jr. 1969. The North American Sunflowers (Helianthus). Mem. Torrey
Bot: Club, 22. 218) pp:

Hottes, F. C. and T. H. Frison. 1931. The plant lice, or Aphididae, or Illinois. MII.
Nat. Hist. Surv. Bull. 19:121—447.

Kennedy, J. S., M. F. Day and V. F. Eastop. 1962. A conspectus of aphids as vectors
of plant viruses. Commonwealth Institute of Entomology. London. 114 pp.

Leonard, M. D. 1959. A preliminary list of the aphids of Missouri. J. Kans. Entomol.
Soc. 32:9-18.

Leonard, M. D. 1968. Additional records of Missouri aphids. Ibid. 32:65-84.

Leonard, M. D. and A. N. Tissot. 1965. A preliminary list of Texas aphid. FI.
Entomol. 48:255—264.

MacGillivray, M. E. 1958. <A study of the genus Mansonaphis Hille Ris Lambers,
1939 (Homoptera: Aphididae). Temminckia 10:1—131.

Olive, A. T. 1963. The genus Dactynotus Rafinesque in North Carolina (Homoptera:
Aphididae). Misc. Publ. Entomol. Soc. Am. 4. 66 pp.

Olive, A. T. 1965. Two new species of Dactynotus from Pennsylvania (Homoptera:
Aphididae) Ann. Entomol. Soc. Am. 59:786-791.

Palmer, M. A. 1952. Aphids of the Rocky Mountains Region. Thomas Say Founda-
tion. Vol. 5. Hirschfield Press. Denver, Colorado. 452 pp.

Rogers, C. E., H. B. Jackson, R. D. Eikenbary and K. J. Starks. 1972. Host-parasitoid
interaction of Aphis helianthi on sunflowers with introduced Aphelinus asychis,
Ephedrus plagiator, and Praon gallicum, and native Aphelinus nigritus and
Lysiphlebus testaceipes. Ann. Entomol. Soc. Am. 65:38—41.

Rogers, C. E. and T. E. Thompson. 1978. Resistance of wild Helianthus species
to an aphid, Masonaphis masoni. J. Econ. Entomol. 71:221—222.

VOLUME 80, NUMBER 4 513

Walker, F. H., Jr. 1936. Observations on sunflower insects in Kansas. J. Kans.
Entomol. Soc. 9: 16-25.

Williams, T. A. 1910. The Aphididae of Nebraska. In Univ. Studies 10. Univ.
Nebraska, Lincoln. 91 pp.

(CER and TET) Southwestern Great Plains Research Center, Fed. Res.,
Sci. and Educ. Admin., USDA, Bushland, Texas 79102; and (MBS) Sys-
tematic Entomology Laboratory, Fed. Res., Sci. and Educ. Admin., USDA,
Beltsville, Maryland 20705.

Footnote

‘In cooperation with The Texas Agricultural Experiment Station, Texas A&M Uni-
versity, College Station, Texas. Approved for publication as TA No. 14292 by the
Director, The Texas Agricultural Experiment Station.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 514-516
A NEW SPECIES OF ASPHONDYLIA (DIPTERA:
CECIDOMYIIDAE) FROM COSTA RICA WITH
TAXONOMIC NOTES ON RELATED SPECIES

Raymond J. Gagné

Abstract—A new species, Asphondylia enterolobii, a gall former on
Enterolobium cyclocarpum in Costa Rica, is described with illustrations.
It and its closest congeners form bud and pod galls on various Mimosaceae.
Hemiasphondylia Mohn is synonymized under Asphondylia and H. mimo-
sae, preoccupied in Asphondylia, is renamed Asphondylia mimosicola
Gagne.

This paper was intended to be simply a description of a new species,
the subject of a biological study now in progress in Costa Rica. But search-
ing a large genus for close relatives of a species often uncovers taxonomic
complications as well as leads to interesting realizations of systematic
relationships as related below.

The new species, Asphondylia enterolobii, was reared from flower galls
on Enterolobium cyclocarpum (Jacq.) Griseb. (Mimosaceae) in Costa
Rica. Specimens were submitted for identification by Dr. D. H. Janzen of
the University of Pennsylvania, Philadelphia, who is interested in the
biology of the gall midge in view of its apparently devastating effect on
the host’s seed crop production.

Interestingly, the three species most closely related to A. enterolobii have
been reared from Mimosaceae also: Asphondylia mimosae Felt from bud
and pod galls on Mimosa sp. (undet.) in Texas; Hemiasphondylia mimosae
Mohn on bud and pod galls of Mimosa albida H. & B. in El Salvador; and
Asphondylia prosopidis Cockerell from buds of Prosopis glandulosa Torr.
in New Mexico and Texas. The four gall midge species share the derived
character states of a reduced shaft of the larval spatula, the development
of a pair of corniform setae on the terminal segment of the larval abdomen
and anisomorphic tarsal claws. The pupae of all are similar in that they
have a simple upper frontal crest and a trifid lower one, this with the
medial point shortest.

Mohn (1960) erected Hemiasphondylia for his new species mimosae on
the basis of the characters listed above and the fact that the sternal spatula
of mimosae is bifid, a condition that does not obtain in prosopidis. I see
no practical reason for segregating H. mimosae and the other species on
Mimosaceae from Asphondylia and so consider Hemiasphondylia a synonym
of Asphondylia. Asphondylia mimosae (Mohn), new combination, is con-
sequently a secondary homonym and is renamed here A. mimosicola
Gagné.

VOLUME 80, NUMBER 4 515

ee Tene De cial

5 7
Figs. 1-2. Pupal head of Asphendylia enterolobii (ventral and lateral views, re-
spectively ). Fig. 3. Pupal head of A. prosopidis (ventral view). Figs. 4-7. Asphondylia
enterolobii. 4, Larval spatula. 5, Larval terminal segments (dorsal). 6, Front, middle
and hind claws of female. 7, Same, of male.

Asphondylia enterolobii Gagné, new species

Adult.—Habitus and terminalia as in other Asphondylia. Palpus 3-seg-
mented. Legs covered with brownish scales, unbanded. Tarsal claws
(Figs. 6-7) anisomorphic, middle claws largest, front claws smallest, at
least middle and hind claws enlarged near middle.

516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pupa—Head (Figs. 1-2): Antennal horns conic, tapering to point with-
out crenulations on anteromedial surface; upper frontal crest single, lower
frontal crest trifid, lateral points longer than medial point.

Larva.—Sternal spatula (Fig. 4) bifid, shaft not developed. Terminal
abdominal segment (Fig. 5) with 3 papillae per side, 2 with short setae,
1 with corniform seta.

Holotype.—Pupa, ex Enterolobium cyclocarpum gall, 10 March 1977,
Santa Rosa National Park, Guanacaste Province, Costa Rica, D. H. Janzen,
USNM Type No. 75229. Paratypes: 4 ¢, 2 2, 21 pupae and 6 larvae, all
with same data as holotype (USNM).

Discussion.—Asphondylia enterolobii is the only one of the 4 related taxa
with conical pupal horns. The other species have wider, less tapered horns
shaped as in A. prosopidis (Fig. 3) with crenulations on the anteromedial
edge. The larval spatula of enterolobii lacks a shaft; those of mimosicola
and prosopidis are quadridentate. The larva of mimosae is unknown.

Literature Cited
Mohn, E. 1960. Gallmiicken (Diptera, Itonididae) aus E] Salvador 3. Teil. Sencken-
bergiana Biol. 41:333-358.

Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. and Educ.
Admin., USDA, c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 517-519
NEW SYNONYMY AND A REVIEW OF HAPLUSIA
(DIPTERA: CECIDOMYIIDAE)

Raymond J. Gagne

Abstract—New synonymy of Haplusia is made to validate use of the
name in a forthcoming key to the genera of Cecidomyiidae. Chastomera,
Palaeocolpodia and Johnsonomyia are treated as junior synonyms, and a
list is given of the species referable to Haplusia.

This paper is written mainly to report new synonyms and to validate
the use of the name Haplusia Karsch that will be used in a forthcoming
key to the genera of Nearctic Cecidomyiidae, but also to review the
scattered writings concerning the genus. Haplusia contains 14 described
and many undescribed species from all over the world and from Baltic
amber. The venation is characteristic and distinct from the other genera
of Porricondylinae, a subfamily of mycophagous cecidomyiids containing
many genera of worldwide distribution. The rs crossvein is far distant
from the wing base, R; bends abruptly at rm and joins C considerably
caudad of the wing apex, and Cu is simple. All species of Haplusia lack
antennal circumfila and most have very long palpi and well-marked wing
spots and leg bands.

The proposed synonymy is as follows:

Haplusia Karsch, 1877:15, 16. Type-species, plumipes Karsch, by original
designation.

Chastomera Skuse, 1888:112. Type-species, bella Skuse, by monotypy.

Palaeocolpodia Meunier, 1904:18. Type-species, eocenica Meunier, by
monotypy. NEW SYNONYM.

Johnsonomyia Felt, 1908:417. Type-species rubra Felt, by original designa-
tion. NEW SYNONYM.

Riibsaamen (1892) was first to point out the similarity between the wing
of Haplusia plumipes and that drawn for Chastomera bella by Skuse (1888)
and to treat the 2 genera as synonyms; but his observation was ignored
until recently when Panelius (1965) cited it but, not having seen specimens
of Chastomera, declined to follow Riibsaamen. Dr. D. H. Colless, CSIRO,
Canberra, Australia kindly sent me an Australian specimen he compared
with the type of Chastomera bella and considered to be the same in
obvious respects. The specimen has the same venation and general habitus
as North American specimens of what has been known as Johnsonomyia.
Chastomera was most recently used by Mamaey (1964, 1966), first as a

518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

senior synonym of Johnsonomyia, then as a genus distinct from the latter
and distinguished on the basis of the eye bridge length, a character that I
do not consider will necessarily distinguish natural subdivisions of the
genus.

Palaeocolpodia eocenica Meunier is known from Baltic amber fossils
with wings typical of the modern species. That an Eocene-Oligocene fossil
can be congeneric with an extant genus is not surprising considering that
such genera as Lestodiplosis and Contarinia were well established in the
Oligocene-Miocene (Gagné, 1973). Mamaev (1964) also pointed out the
resemblance of Palaeocolpodia eocenica to Chastomera and considered the
2 genera to be synonyms.

Haplusia may be separated into 2 or more genera someday, but that
decision should best follow a study of the fauna on a world basis and not
arbitrary splitting resulting from superficial study of limited material and
narrow geographic scope.

Following is a list of species referable to Haplusia. Haplusia bella is a
restored combination; all others except plumipes are new combinations.

alexanderi (Felt), 1921:96 (Johnsonomyia). “Cameroun.”
bella (Skuse), 1888:112 (Chastomera). Australia.

braziliensis (Felt), 1915:153 (Johnsonomyia). Brazil.
brevipalpis (Mamaev), 1964:903 (Chastomera). Russia.

cincta (Felt), 1912:103 (Johnsonomyia). Guatemala.

eocenica ( Meunier ), 1904:18 ( Palaeocolpodia). Baltic amber.
fusca (Felt), 1908:417 (Johnsonomyia). Eastern United States.
hondrui (Mamaev), 1964:902 (Chastomera). Rumania.
longipalpis (Mamaev), 1964:902 (Chastomera). Russia.
pallida (Mamaev), 1966:220 (Johnsonomyia). Eastern USSR.
palpata (Mamaev), 1966:219 (Johnsonomyjia). Russia.
plumipes Karsch, 1877:16. Brazil.

rubra (Felt), 1908:417 (Johnsonomyia). Eastern United States.
spiculosa (Barnes), 1927:271 (Chastomera). Malaya.

Literature Cited

Barnes, H. F. 1927. Some Cecidomyiidae from the Federated Malay States. J. Fed.
Malay States Mus. 13:269-274.

Felt, E.P. 1908. Appendix D. N.Y. State Mus. Bull. 124:286—422.

1912. New Itonididae (Dipt.). J.N.Y. Entomol. Soc. 20:102—107.

—. 1915. New South American gall midges. Psyche 22:152—157.

——. 1921. Observations on Johnsonomyia Felt with a description of a new
species. Can. Entomol. 53:96.

Gagné, R. J. 1973. Cecidomyiidae from Mexican tertiary amber (Diptera). Proc.
Entomol. Soc. Wash. 75:169-171.

VOLUME 80, NUMBER 4 519

Karsch, F. 1877. Revision der Gallmiicken. Minster i. W., E. C. Brunn. 57 pp.
Mamaev, B. M. 1964. Gall midges of the USSR. 6. New species of the tribe Porri-
condylini (Diptera, Cecidomyiidae). Entomol. Obozr. 43:894-913; Entomol.
Rev. 43:456—465.
1966. New and little known palaearctic gall midges of the tribe Porri-
condylini (Diptera, Cecidomyiidae). Acta. Entomol. Bohemoslov. 63:213-239.
Meunier, F. 1904. Monographie des Cecidomyidae, des Sciaridae, des Mycetophilidae
et des Chironomidae. Ann. Soc. Sci. Bruxelles 28;12—275.

Panelius, S. 1965. A revision of the European gall midges of the subfamily Porri-
condylinae (Diptera: Itonididae). Acta Zool. Fenn. 113:1—157.

Ribsaamen, E. H. 1892. Die Gallmiicken des K6nigl. Museums for Naturkunde
zu Berlin. Berlin Entomol. 37:321—411.

Skuse, F. A. A. 1888. Diptera of Australia. Proc. Linn. Soc. NSW (2) 3:17-145.

Systematic Entomology Laboratory, I[BII, Fed. Res., Sci. and Educ.
Admin., USDA, c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 520-523

A NEW GENUS AND SPECIES OF CARDIASTETHINI FROM
PERU (HEMIPTERA: ANTHOCORIDAE )

Thomas J. Henry and Jon L. Herring

Abstract.—A new genus and species, Dolostethus pubescens, is described
from the Pichis River region of Pasco, Peru. Its placement in the tribe
Cardiastethini is discussed, and figures of the habitus of the adult male
and the metapleural scent gland are included.

Recently, while searching for undetermined Isometopinae (Miridae) in
the Cornell University collections, A. G. Wheeler, Jr. discovered an interest-
ing and peculiar anthocorid. At first, this bug appeared to belong to the
isometopines because of its large eyes, type of pronotum and other features.
Upon closer examination, the single male specimen proved to be a new
Neotropical genus and species of the tribe Cardiastethini in the anthocorid
subfamily Lyctocorinae.

Dolostethus Henry and Herring, new genus

Description—Characterized by the large eyes, narrow vertex, deeply
emarginate posterior margin of the pronotum, distinctive fracture between
the cuneus and membrane, and the unique character of scent gland canal.

Head wider than long, eyes large, granulate, not touching anterior margin
of pronotum, dorsal width of eye much greater than width of vertex; ocelli
set near posterior margin of head; tylus broad, truncate; rostrum short,
not reaching beyond middle of sternum; antennae simple, thickly pubescent,
segment I short, visible from dorsal aspect; segment II longest, thickest,
gradually enlarged to apex; segments III and IV each greater than %
length of II, their combined lengths 1.5x length of segment II, clothed
with pilose setae, length of some more than 2X diameter of segments.
Pronotum, width 3x length, subquadrate, collar weakly formed, anterior
margin straight, lateral margins weakly rounded, distinctly carinate,
posterior margin deeply sinuate, humeral angles bearing blunt nodes, calli
narrow, shiny, smooth, rugose between; scutellum large, set into pronotal
sinuation, basal % level. with pronotum, transversely impressed through
middle, without circular depressions, apical % level with clavus. Hemelytra
opaque, punctate, thickly pubescent, radial vein clearly delimited to cuneal
fracture, cuneus much broader than long, posterior fracture before membrane
deeply incised, membrane very finely pubescent, margins ciliate, veins linear,
indistinct. Venter dark, shining, abdomen strongly pubescent; scent gland

VOLUME 80, NUMBER 4 521

1

Figs. 1-2. Dolostethus pubescens. 1, Male habitus. 2, Metasternal scent gland.

canal curved forward along rim of metapleura (Fig. 2) to anterior margin;
legs slender, pale, weakly spined, hind tibiae (of male, at least) distinctly
bowed. Genital paramere sickle-shaped, curved to the left and dorsad
around genital tergite.

Type-species.—Dolostethus pubescens, new species.

Discussion.—Dolostethus keys to Cardiastethus Fieber in Herring (1976)
but may be separated by its broader form, proportionately larger eyes and
narrow vertex (which is much narrower than diameter of an eye), the
abruptly emarginate posterior margin of the pronotum and the distinctive
scent gland canal. It resembles and could easily be mistaken for a member
of the Isometopinae. However, the presence of a 3-segmented rostrum, a
single genital paramere and the absence of closed cells on the membrane will
distinguish it from that mirid subfamily.

5

bo
bo

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Dolostethus pubescens Henry and Herring, new species
Fig. 1-2

Male.—Length 1.76 mm, width 1.00 mm, suboval, generally shiny, dark
brown, densely clothed with erect and suberect, pale pubescence. Head:
Length 0.28 mm, width across eyes 0.52 mm, ocelli 0.14 mm apart (mea-
sured from middle of each) width of vertex 0.12 mm, dorsal width of eye
0.20 mm; eyes dark brown, granulate, finely pubescent; ocelli red, set near
posterior margin of head; basal carina rounded, distinct. Rostrum: Length
(0.42 mm, testaceous, basal segment reddish, apex not surpassing mesocoxae
or middle of sternum. Antennae: Length of segments I-IV, 0.10: 0.28:
0.18: 0.18 mm; testaceous, in part infuscated, thickly clothed with long,
erect setae, III and IV only slightly thinner than I and II; IV fusiform.
Pronotum: Length 0.26 mm, width at base 0.80 mm, shiny dark brown,
posterior angles more testaceous; subquadrate, strongly punctate, lateral
margins weakly rounded, distinctly carinate, base deeply emarginate,
humeral angles bearing blunt indistinct nodes, calli smooth, slightly raised,
transversely rugose between, mesoscutum not visible. Scutellum: Length
0.40 mm, width at base 0.52 mm; dark brown, shiny, basal % finely punctate,
anterior % level with pronotum, transversely impressed through middle,
impression without circular depressions often found in cardiastethines.
Hemelytra: Dark brown, punctate, densely clothed with short, erect, pale
setae, radial vein clearly delimited to cuneal fracture, cuneus much broader
than long, with pronounced incision at junction with membrane; membrane
brownish, paler along margins, only outer and inner veins evident. Venter:
Dark brown, pleura shining, sternum and abdomen more fuscous; meta-
pleural scent gland curved forward along outer margin to apex of metapleura
(Fig. 2); abdomen clothed with pale, short to long hairs. Legs: Testaceous,
sparsely pubescent, coxae distinctly reddish, hind tibiae distinctly curved on
basal 4, tibial spines weakly formed, tarsi 3-segmented. Paramere: Slender,
sickle shaped.

Holotype.—4, Azupiza [River] to Miriatiriani [River], Cam. del Pichis
[Dept. of Pasco], Peru, July 9, 1920, W. T. Forbes and J. C. Bradley colls.,
Cornell University Expedition, Lot 607, Sublot 132 (Cornell University
Type Collection ).

Acknowledgments

We kindly thank Dr. L. L. Pechuman, Cornell University, for the loan
of the unique specimen of Dolostethus pubescens. We also give thanks to
Dr. A. G. Wheeler, Jr., Pennsylvania Department of Agriculture, for dis-
covering this new anthocorid and for his helpful review of the manuscript.

VOLUME 80, NUMBER 4 523

Literature Cited

Herring, J. L. 1976. Keys to genera of Anthocoridae of America north of Mexico
with description of a new genus (Hemiptera: Heteroptera). Fla. Entomol. 59:
143-150.

(TJH) Bureau of Plant Industry, Pennsylvania Department of Agriculture,
Harrisburg, Pennsylvania 17120; and (JLH) Systematic Entomology Labo-
ratory, Fed. Res., Sci. and Educ. Admin., USDA, c/o U.S. National Museum,
Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 524-537
A NEW GENUS NEAR CANACEOIDES CRESSON, THREE NEW
SPECIES AND NOTES ON THEIR CLASSIFICATION
(DIPTERA: CANACIDAE)

Wayne N. Mathis and Willis W. Wirth

Abstract—A diagnosis is given of Paracanace, new genus, and a key is
presented for the separation of the seven known species, including three
new species which are described and illustrated: hoguei, a new species from
Cocos Island, Costa Rica; lebam, a new species from Jamaica; and aicen,
a new species from Dominica. A cladogram is diagrammed for the genus
and gives apotypic character states for the phylogeny of the species of
Paracanace and of its sister-group, Canaceoides Cresson.

Surf or beach flies were last reviewed as a family by Wirth (1951). In
that publication, Wirth described Canace maritima as a new species from
the Galapagos Islands. Five years later, Wirth (1956) described two addi-
tional species, C. blantoni (Panama) and C. oliveirai (Brazil); both are
closely related to C. maritima. In 1969(b), Wirth described a fourth related
species, C. cavagnaroi, which like C. maritima is also from the Galapagos
Islands. The present paper is a continuation of this series. The species
noted above plus three new ones are distinguished as a new genus,
Paracanace. A key to the species of Paracanace and illustrations of the
new species are provided for their identification. [Illustrations of the
epandrium and surstylus of the male terminalia accompanied the descrip-
tions of each previously described species; neither the descriptions nor the
illustrations are republished here.

Paracanace Mathis and Wirth, new genus

T ype-species.—Paracanace hoguei Mathis and Wirth, new species, by
present designation.

Species included.—Paracanace aicen, new species; P. blantoni (Wirth),
new combination; P. cavagnaroi (Wirth), new combination; P. hoguei,
new species; P. lebam, new species; P. maritima (Wirth), new combina-
tion; P. oliveirai (Wirth); new combination.

Diagnosis —Paracanace is distinguished from other genera of the family
by the following combination of characters (an asterisk indicates apotypic
character states): General coloration light to dark gray; head in lateral
aspect higher than wide; 3 pairs of eclinate fronto-orbital setae; *2 pairs of
proclinate interfrontal setae, posterior pair aligned with, or posterior of

VOLUME 80, NUMBER 4 525

median ocellus; mesofrons mostly dull, pollinose; eye in lateral aspect as
high as wide, with oblique orientation to general plane of body; 4 pairs of
genal setae, anterior pair cruciate anteriorly, 3 posterior pairs dorsally-
curved; *anterior notopleural seta reduced, inconspicuous; front femur
lacking row of anteroventral, spinelike setae; *middle femur of male with
posteroventral row of closely set setae, proximal ones frequently pale;
*scutellum bare except for 2 pairs of lateral setae; female cerci with 2
pairs of stout apical setae.

Description—Small to moderately small beach flies, length 1.43 to
2.57 mm; coloration mostly gray; vestiture mostly pollinose, dull in ap-
pearance.

Head: Lateral aspect higher than wide; mesofrons mostly full, pollinose,
but specimens of some species with some lustrous, semimetallic reflections
as viewed from some angles; mesofrons triangular, nearly equilateral,
anterior angle napiform; ocelli arranged to form isosceles triangle, distance
between posterior ocelli larger than between either posterior ocellus and
median 1; parafrons somewhat membranous and distinguished in color and
conformation from mesofrons; fronto-orbits evenly wide along dorsal and
anterodorsal margin of eye, continuing ventrally as parafacies. Chaetotaxy
of frons as follows: Ocellar bristles large, divergent, inserted between
posterior and median ocelli; 2 pairs of proclinate interfrontal bristles, pos-
terior pair more or less aligned with ocellar bristles; 1 pair of proclinate to
divergent postocellar bristles inserted posteromedially of posterior ocelli;
mesofrons with several other smaller setae, especially anterolaterally; 3
pairs of larger eclinate fronto-orbital bristles and several smaller setae;
parafrons with 3-4 smaller setae; inner and outer vertical bristles well
developed, inserted posteriad of posterior ocelli. Antenna dark colored,
brownish black to black; 3rd segment subcircular with pubescent vestiture;
arista slightly longer than combined length of first 3 segments, micro-
pectinate on dorsal and ventral surfaces. Face silvery gray to charcoal
gray, bare, broadly carinate medially, ventral margin concave. Gena nearly
concolorous with face, slightly darker, about % as high as eye anteriorly,
becoming higher posteriorly; 4 pairs of genal bristles, anterior pair con-
vergent anteriorly, next 3 pairs dorsally curved; postgena with several pale
setae toward ventral margin. Eye in lateral aspect as high as wide, broadly
oval to subrectangular, oriented at oblique angle to remainder of head.
Clypeus broad, protruding through ventral facial concavity. Maxillary
palpus somewhat pale, sparsely setose with fine pale setae.

Thorax: Mesonotum darker than pleural areas, dark brownish gray to
charcoal gray, pleural areas uniformly gray. Chaetotaxy of thorax as fol-
lows: Acrostichal setae mostly small, hairlike except for 1 pair of larger

526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

prescutellar bristles; 4 pairs of dorsocentral bristles, posterior pair dis-
placed laterally; 1 pair of humeral bristles; 1 pair of presutural bristles;
often with 1 or 2 pairs of moderately large posthumeral setae; 1 pair of
supra-alar bristles; 1 pair of postalar bristles; scutellum bare except for 2
pairs of lateral bristles; mesonotum with several scattered smaller setae in
addition to larger bristles; 2 pairs of notopleural bristles, seta at anterior
angle much smaller than posterior setae, 1-2 other smaller setae; meso-
pleuron with 3-4 bristles and several smaller setae, larger bristles inserted
on posterior %; 1 larger ventrally-curved lower mesopleural seta; sterno-
pleuron with 1 larger bristle toward posterodorsal angle and _ several
moderately larger setae anteroventrally; mesopleuron and hypopleuron
bare; front and middle coxae setose, several setae pale. Legs generally
concolorous with pleural areas, basitarsi of middle and hind leg frequently
pale; legs setose, development of setae varying with species. Halter pale.
Wing slightly infuscated to mostly hyaline; vein R, long; see figures for
details.

Abdomen: Mostly concolorous with thoracic pleural areas; symmetrically
setose bilaterally; males with 6 terga and epandrium exposed; females with
8 terga and genital cerci exposed. Surstylus of male terminalia simple,
digitiform to lobelike except for P. maritima, in P. maritima with subbasal
anterior setulose lobe and constriction. Eighth tergum of female with 1-2
pairs of larger setae; female genital cerci with 2 pairs of apical stout
spines; female atrium oval.

Geographic distribution—The composite distribution of the genus is
Neotropical, between 19° north latitude and 23° south latitude. Five of
the seven species are known only from islands in the Caribbean Sea and
Pacific Ocean.

Key to Species of Paracanace

1. Costal vein between humeral crossvein and subcostal break with a
row of long, spinelike setae, setal length equal to, or greater than
width of Ist costal cell; middle femur of male with posteroventral
row of closely set setae along entire length, those along proximal
% pale; front femur with 3-4 long setae along posteroventral mar-
gin, setal length subequal to 2* femoral width 9)

— Setae along anterior margin of wing much shorter, not more than %
width of Ist costal cell; middle femur of male with posteroventral
row of closely set setae along distal % only; front femur lacking 3-4
setae as described above 4

2. Three dorsally-curved genal setae subequal in length; body strongly
setose (Cocos Island, Costa Rica ) P. hoguei, new species

VOLUME 80, NUMBER 4 527

- Middle dorsally-curved genal seta about ”% length of setae on either
side; body moderately setose 3

3. Surstylus broader on distal %, especially evident in lateral view;

ventral surstylar margin broadly truncate in lateral and posterior

views; posterior margin of surstylus bearing distinct row of longer
setae (Jamaica) P. lebam, new species

— Surstylus in lateral view swollen along anterior margin near middle,

tapering ventrally to broadly rounded ventral margin; posterior mar-

gin of surstylus lacking distinct row of longer setae; posteroventral

angle of surstylus noticeably produced apically (Dominica )

P. aicen, new species

4, Front and middle femora of male with row of about 20 long white

setae along proximal % of posteroventral margin; surstylus with sub-

basal anterior setose lobe and constriction before apical enlargement
(Galapagos Islands ) P. maritima (Wirth)

— Front and middle femora of male with not more than 10 long white

setae along posteroventral margin at base; surstylus simple, lacking
anterior setose lobe or subbasal constriction 5

5. Tarsi mostly dark, concolorous with tibiae (Galapagos Islands)
P. cavagnaroi (Wirth)
— Tarsi mostly pale, yellowish, especially basitarsus of hind leg 6

6. Surstylus slender, angulate, length about 3x width (Brazil)

P. oliveirai (Wirth)

— Surstylus broad, truncate ventrally, length not more than 2 width,
posteroventral angle slightly produced (Panama) _ P. blantoni (Wirth)

Paracanace hoguei Mathis and Wirth, new species
Figs. 1-5

Diagnosis.—Specimens of this species closely resemble those of P. aicen
and P. lebam but are distinguished by the following combination of char-
acters: Middle dorsally-curved genal seta subequal in length to setae on
either side; spinelike setae along costal margin larger, those between
humeral crossvein and subcostal break longer than width of Ist costal cell;
body more setose; surstylus of male terminalia long and slender, digitiform,
ventral margin rounded in lateral view, pointed medially in caudal view.

Description.—As in generic description but with the following additional
details. Length 1.68 to 2.57 mm.

Head: Mesofrons yellowish gold to faintly lavender, appearing semi-
lustrous from some angles; parafrons yellowish gold to rusty orange; middle
dorsally-curved genal bristle subequal in length to setae on either side.

Thorax: Mesonotum charcoal gray. Front femur with 3-4 long setae

528 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-2. Canace hoguei. 1, Left front leg, posterior view. 2, Femur of middle
leg, posterior view.

along posteroventral margin, length more than 2X greatest width of femur
(Fig. 1), front femur also with 5-7 slender pale setae along proximal % of
ventral surface; middle femur of male with row of closely set setae along
posteroventral margin extending to base, setae on proximal % or less pale
(Fig. 2); middle and hind basitarsi yellowish brown to grayish brown,
front basitarsus paler than tibia but not distinctly yellowish. Wing length
averaging 1.91 mm; setae along costal margin spinelike, those between
humeral crossvein and subcostal break longer than width of Ist costal cell
(Hig. 5).

Abdomen: Surstylus of male terminalia (Figs. 3-4) long and slender,
slightly angulate in lateral view; ventral margin rounded in lateral view;

VOLUME 80, NUMBER 4 529

Figs. 3-4. Canace hoguei. 3, Epandrium and surstylus, lateral view. 4, Same,
caudal view.

ventromedial angle pointed in caudal view; posterior margin lacking an
isolated row of distinctive setae.

Types.—Holotype ¢, labelled: “COCOS ISLAND (Costa Rica) Wafer
Bay 17-22 Apr. 75 C. L. Hogue.” Allotype ° and 170 paratypes (34 4,
136 2) with same label data as the holotype. The holotype, allotype, and
most of the paratypes will be deposited in the Los Angeles County Museum
of Natural History. Paratypes will also be deposited in the National
Museum of Natural History, Washington, D.C., the British Museum
(Natural History), London, and the California Academy of Sciences, San
Francisco.

530 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Meng SQ
SENECA NY

Figs. 5-6. Wing. 5, Canace hoguei. 6, Canace aicen.

Geographic distribution—This species is known only from the type-
locality, Isla del Coco (Cocos Island). This island is situated approximately
550 km from the Costa Rican coast in the Pacific Ocean 5°33’ north
latitude; 87° west longitude).

Etymology.—tThe species epithet hoguei is a genitive patronym honoring
the distinguished entomologist Dr. C. L. Hogue, collector of the type-series.

Paracanace lebam Mathis and Wirth, new species
Figs. 7-9

Diagnosis—Specimens of P. lebam are similar and closely related to
those of P. aicen and P. hoguei but are distinguished by the following
combination of characters: Middle dorsally-curved genal seta about %
length of setae on either side; spinelike setae along costal margin, particu-
larly those between humeral crossvein and subcostal break, about as long
as width of Ist costal cell; general appearance setose, like specimens of P.

VOLUME 80, NUMBER 4 531

Fig. 7. Canace lebam, head, lateral view.

aicen; surstylus of male terminalia about 1% as long as wide in lateral
view, broadly truncate along ventral margin in lateral and caudal views;
ventromedial angle pointed in caudal view; posterior margin with distinctive
row of longer setae.

Description.—As in generic description but with the following additional
details. Length 1.76 to 1.94 mm.

Head (Fig. 7): Mesofrons dark, grayish brown to black, dull, only
slightly darker than general coloration of mesonotum; parafrons grayish
olivaceous to brownish green, contrasting distinctly with darker mesofrons;
middle dorsally-curved genal seta about % length of setae on either side.

Thorax: Mesonotum grayish brown to charcoal gray. Front femur with
34 long setae along posteroventral margin, length more than 2X greatest
width of femur, front femur also with 2-3 slender pale setae along proximal

532 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 8-9. Canace lebam. 8, Epandrium and surstylus, lateral view. 9, Same, caudal
view.

% of ventral surface; middle femur of male with row of closely set setae
along posteroventral margin extending nearly to base, setae on proximal “
or less pale; basitarsi of all legs nearly concolorous, grayish brown, only
slightly paler than tibia. Wing length averaging 1.65 mm; setae along
costal margin spinelike, those between humeral crossvein and subcostal
break equal in length to width of Ist costal cell.

Abdomen: Surstylus of male terminalia (Figs. 8-9) about 1%~x as long
as broad, in lateral view; ventral margin broadly truncate in lateral and
caudal views; ventromedial angle of surstylus in caudal view somewhat
pointed; posterior margin with distinctive row of setae.

Types.—Holotype 4, labelled: “JAMAICA Runaway Bay Feb. 1969 W. W.
Wirth bay shore.” Allotype ? and 1 é paratype with same label data as
the holotype. The type-series is in the National Museum of Natural
History, Washington, D.C., type number 75303.

Geographic distribution—Specimens of P. lebam have been collected
only at the type-locality, Runaway Bay, Jamaica.

VOLUME 80, NUMBER 4 533

Etymology.—The species epithet lebam is an anagram of the proper name
Mabel (Wirth), after whom this species is named. The epithet is a noun
in apposition to the generic name.

Paracanace aicen Mathis and Wirth, new species
Figs. 6, 10-11

Diagnosis —Paracanace aicen is closely related and similar to P. lebam
and P. hoguei but is distinguished from either of the latter by the following
combination of characters: Middle dorsally-curved genal seta about %
length of setae on either side; spinelike setae along costal margin about as
long as width of Ist costal cell; general appearance setose but less so than
in P. hoguei; surstylus of male terminalia with anterior swelling near
middle in lateral view and broadly rounded ventrally; in caudal view with
developed ventromedial angle but not pointed; posterior margin of sur-
stylus not notably setose.

Description—As in generic description but with the following additional
details. Length 1.72-2.24 mm.

Head: Mesofrons dark, grayish brown to black, dull, nearly concolorous
with mesonotum; parafrons yellowish to olivaceous brown, contrasting dis-
tinctly with darker mesofrons; middle dorsally-curved genal seta about %
length of setae on either side.

Thorax: Mesonotum dark grayish brown. Front femur with 3-4 long
setae along posteroventral margin, length more than 2 greatest width of
femur, front femur also with 5-7 slender pale setae along proximal % of
ventral surface; middle femur of male with row of closely set setae along
posteroventral margin extending nearly to base, setae on proximal % or
less pale; basitarsi of all legs nearly concolorous with tibiae. Wing length
averaging 1.76 mm; setae along costal margin spinelike, those between
humeral crossvein and subcostal break equal in length to width of Ist
costal cell (Fig. 6).

Abdomen: Surstylus of male terminalia (Figs. 10-11) about 2x as
long as broad; anterior margin swollen near middle in lateral view; ventral
margin broadly rounded in lateral view; ventromedial angle of surstylus
produced but not pointed in caudal view; posterior margin lacking obvious
row Of setae.

Types—Holotype 2, labelled: “DOMINICA, W. I. Calibishie 27 Feb.
1965 W. W. Wirth sea shore/Bredin-Archbold Smithsonian Bio. Surv.
Dominica.” Allotype ° and 3 paratypes (1¢, 22) with same label data
as the holotype. The type-series is in the National Museum of Natural
History, Washington, D.C., type number 75304.

Geographic distribution—Specimens of P. aicen have been collected
only from the type-locality, Calibishie, Dominica.

534 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 10-11. Canace aicen. 10, Epandrium and surstylus, lateral view. 11, Same,
caudal view.

Natural history——Wirth collected the type-series on an algae covered,
intertidal rock shelf on the beach to the west of the village.

Etymology.—tThe species epithet aicen is an anagram of the proper name
Necia (Dianne Mathis), after whom this species is named. The epithet is
a noun in apposition to the generic name.

General Discussion

Paracanace belongs to a group of related taxa of the family Canacidae
that is recognized as follows: Body coloration generally gray; head in
lateral aspect narrow, higher than wide; 3 pairs of larger eclinate fronto-
orbital setae; frons short; eye as high as wide, suboval to subrectangular,
and oriented at an oblique to nearly vertical angle to general plane of head;
1 distinct, ventrally-curved lower mesopleural seta. Although the relation-
ships among the included taxa of this assemblage have not been adequately
clarified, several taxa can be grouped into closely related, monophyletic
groups as evidenced by apotypic character states. Among these taxa are
the genera Canaceoides Cresson (Wirth, 1969) and Paracanace. We believe

'
|

VOLUME 80, NUMBER 4 535

&
AY
S S S
ea & = s S
é = g g & § g =
& a © 5 v EN = =
> v © ” ay ey ¥ Si
a 13 (] in|
12 []
a 10
9 [J
8 [J
[] 7
L) 6
5 CL]
B 4
3 (]
2 L)

Fig. 12. Argumentation scheme for the hypothetical phylogeny of the genus Para-
canace. Filled squares indicate apotypic character states; open squares indicate plesio-
typic character states.

that these two genera are sister-groups. This relationship is corroborated
by the reduced anterior notopleural seta, a character state we interpret to
be apotypic.

We suggest that the phylogeny of the species of Paracanace is as
diagrammed in Fig. 12. Numbers in the cladogram correspond with charac-
ters listed in Table 1. The relative plesiotypic-apotypic states were deter-
mined primarily from out-group comparisons.

536

Table 1.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of Paracanace.

CHARACTER
1. Anterior notopleural seta
2. Disc of scutellum
3. Number of large
interfrontal setae
4. Setae on 8th
tergum of female
5. Setae along posteroventral
margin of middle femur of
male
6. Shape of female atrium
7. 3-4 long slender black setae
along posteroventral margin
of front femur
8. Setae along costal
margin of wing
9. Row of setae along
posteroventral margin of
middle femur of male
10. Male terminalia, shape
of surstylus
11. Color of basitarsus
12. Length of middle dorsally-
curved genal seta
13. Coloration of mesofrons

Characters and character states used in cladistic analysis of the species

CHARACTER STATE

Apotypic

reduced, less than 14 length
of posterior seta

bare

2. pairs
setae subequal

partial to complete row
of closely set setae

U-shaped, incomplete
posterior end

absent

large, spinelike

extending along entire
length

complicated with subbasal
constriction and anterior
subbasal setose lobe

dark

short, about % length
of setae on either side
semilustrous, faintly
lavender

Plesiotypic

subequal to posterior
seta

with seta

1 pair

with 1 or 2 pairs of
larger setae

lacking row of closely set
setae

oval

present

normally developed

evident only along
distal half

simple, fingerlike

pale

long, subequal to length
of setae on either side

dull, dark greenish to
grayish brown or black

The monophyly of Paracanace is clearly evident, being confirmed by

three apotypic character states as noted in the diagnosis and Table 1. We
suggest that the genus differentiated into two lineages. The first lineage
includes the three new species described above. These three species are
closely related as evidenced by the large spinelike setae along the costal
wing margin and the nearly complete row of closely set setae along the
posteroventral margin of the middle femur. The two West Indian species,
P. aicen and P. lebam, of this lineage are distinguished from P. hoguei by

VOLUME 80, NUMBER 4 537

the short middle dorsally-curved genal seta and the dull coloration of the
mesofrons.

The second lineage of the genus comprises the remaining four species,
P. blantoni, P. cavagnaroi, P. maritima and P. oliveirai. These species are
characterized by the loss of the three or four long slender black setae along
the posteroventral margin of the front femur. Of this group, P. maritima
is characterized by the unique conformation of the male terminalia and
the 20 or so long white hairs on the front femur. This distinctive species
probably arose from a lineage apart from the other three species as indicated
on the cladogram. The remaining three species are all quite similar in
general appearance; and, except for specimens of P. cavagnaroi, reference
to structures of the male terminalia will usually be necessary to accurately
identify them. Unlike the other two species, the tarsi of specimens of P.
cavagnaroi are dark.

Acknowledgments

We are grateful for the helpful comments of Oliver S. Flint, Jr., Raymond
J. Gagné, Ronald W. Hodges and Theodore J. Spilman; all of whom were
kind enough to read through the first draft of the manuscript. For technical
assistance, we thank Mary Jaque Mann (scanning electron micrographs),
Victor E. Krantz (wing photographs), L. Michael Druckenbrod (Fig. 7),
George L. Venable (photograph retouching) and Hollis B. Williams
(general assistance). For allowing us to study the specimens of P. hoguei,
we are indebted to C. L. Hogue, Los Angles County Museum of Natural
History.

Literature Cited

Wirth, W. W. 1951. A revision of the dipterous family Canaceidae. Occas. Pap.
Bernice Pauahi Bishop Mus. 20:245-275.

1956. Two new Neotropical species of surf flies of the genus Canace (Dip-
tera, Canaceidae). Rev. Bras. Entomol. 5:161—166.
1969a. The shore flies of the genus Canaceoides Cresson (Diptera: Cana-
ceidae ). Proc. Calif. Acad. Sci. 36:551-570.

1969b. New species and records of Galapagos Diptera. Proc. Calif. Acad.
Sci. 36:571-594.

(WNM) Department of Entomology, NHB 169, Smithsonian Institution,
Washington, D.C. 20560; and (WWW) Systematic Entomology Labora-
tory, Fed. Res., Sci. and Educ. Admin., USDA, c/o U.S. National Museum,
Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 538-542
SANDFLY DISTRIBUTION IN THE UNITED STATES,
WITH A FIRST RECORD FOR COLORADO
(DIPTERA: PSYCHODIDAE)

R. B. Eads

Abstract.—The 11 species and subspecies of Lutzomyia known to occur
in the United States are listed, with their known distribution and host
preferences. Lutzomyia oppidana (Dampf) is recorded in Colorado for the
first time.

The isolation of three strains of a new Phlebotomus fever group virus
from the grey woodrat, Neotoma micropus, in Cameron County, Texas, in
December 1973 and March 1974, (Calisher et al., 1977) has focused atten-
tion upon the species and distribution of sandflies in the United States and
their possible role in virus transmission. The International Catalogue of
Arboviruses (Berge, 1975) lists 20 serologically related viruses in the
Phlebotomus fever group, 13 of which have been recovered from phle-
botomine flies. The new virus, called Rio Grande, was not recovered from
hematophagous insects (principally mosquitoes ), including several hundred
Lutzomyia spp., tested by this laboratory during the 1971 outbreak of
Venezuelan equine encephalitis (VEE) in south Texas and during the
next two years (Vector-Borne Diseases Division (VBDD) unpublished
data).

VBDD personnel collected three species of Lutzomyia—texana (Dampf),
cruciata (Coquillett) (=diabolica) and anthophora (Addis)—in CDC
light traps on the same ranch in Cameron County, Texas, during 1971-73,
from which the Rio Grande virus isolations were made. The most likely
candidate for Neotoma to Neotoma transmission of Rio Grande virus is L.
anthophora, since it is known to feed on rabitts and rodents (Addis, 1945),
and it uses woodrat nests as diurnal resting sites (Young, 1972). Lutzomyia
cruciata feeds readily on rodents in the laboratory. The only thing known
of its feeding habits in south Texas is that in localized areas it is annoying
to human beings. Addis (1945) reported that this species entered homes
in Uvalde, Texas, and bit people during the summer of 1944. We en-
countered a similar situation at Del Rio, Texas, in 1964. Lutzomyia texana
feeding habits are not known, but it is considered likely that reptiles are
the usual hosts.

The fact that New World sandflies reach greatest specific diversity and
population density in the tropics has, perhaps, kept to a minimum the
interest displayed in these insects in temperate to cold zones. However,

539

VOLUME 80, NUMBER 4

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VOLUME 80, NUMBER 4 541

the growing number of reports of sandfly recoveries in the United States
and southern Canada suggests that the comparatively small number of
species present are widely distributed (Fairchild and Harwood, 1961; Har-
wood, 1965; Eads et al., 1965; Easton et al., 1967; Easton et al., 1968;
Downes, 1972; Chaniotis, 1974; and Aitken et al., 1977).

The 11 species and subspecies of Lutzomyia recorded from the United
States are listed in Table 1, with their known distribution and host prefer-
ences. Selected authorities for these data are included. A key to the
Lutzomyia species known to inhabit the United States has been provided
by Rosabal and Miller (1970).

Only two of the tropical, man-biting species of Lutzomyia seem suf-
ficiently adaptable to diverse climatic conditions to extend their distribution
into the United States. Lutzomyia cruciata has been reported from Central
America, Mexico and Texas. Lutzomyia shannoni (Dyar) is even more
widely distributed, having been taken from Argentina to Louisiana, North
Carolina and Florida. Lutzomyia anthrophora is the only other species
recorded from the United States known to feed on mammals.

The species collected in the more northern latitudes of the United States
are known or suspected to feed on coldblooded vertebrates. Ground squirrel
and other rodent burrows are used for larval and pupal development
(Chaniotis, 1967).

We have recently collected a female (7/19/77) and a male and female
(7/23/77) L. oppidana in Larimer County, Colorado, in CDC light traps.
To our knowledge, those were the first Lutzomyia recovered in the state.
The CDC light traps were operated some 200 trap nights in Larimer
County during June-September 1977, in conjunction with extensive western
encephalitis investigations. Our small catch indicates low population
densities of this sandfly—or that the CDC traps are not attractive to these
insects. Thirteen-lined ground squirrel (Spermophilus tridecemlineatus )
burrows were common around both trapping sites from which L. oppidana
were taken.

Literature Cited

Addis, C. J. 1945. Phlebotomus (Dampfomyia) anthophorus, n. sp. and Phlebotomus
diabolicus Hall from Texas. J. Parasitol. 31(1):119-127.

Aitken, T. H. G., A. J. Main and D. G. Young. 1977. Lutzomyia vexator (Coquillett )
in Connecticut. Proc. Entomol. Soc. Wash. 79( 4) :582.

Berge, T. O. (Ed.). 1975. International Catalogue of Arboviruses. U.S. Dept. of
Health, Education, and Welfare Publ. No. 27-8301 (CDC). 898 pp. (Second
Edition. )

Calisher, C. H., R. G. McLean, G. C. Smith, D. M. Szmyd, D. J. Muth and J. L.
Lazuick. 1977. Rio Grande—a new Phlebotomus fever group virus from south
Texas. Am. J. Trop. Med. Hyg. 26(5) :997-1002.

542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Chaniotis, B. N. 1967. The biology of California Phlebotomus (Diptera: Psychodidae)
under laboratory conditions. J. Med. Entomol. 4(2):221-233.

1974. Phlebotomine sandflies in Montana: First Report. Mosq. News 34(3):
334-335.

Christensen, H. A. and A. Herrer. 1973. Attractiveness of sentinel animals to vectors
of Leishmaniasis in Panama. Am. J. Trop. Med. Hyg. 22(5):578-584.

Downes, J. A. 1972. Canadian records of Phlebotomus vexator, Trichomyia nudata
and Maruina lanceolata (Diptera: Psychodidae). Can. Entomol. 104:1135-1136.

Eads, R. B., H. A. Trevino and E. G. Campos. 1965. Additional records of Phle-
botomus texanus. Proc. Entomol. Soc. Wash. 67 (4) :251—252.

Easton, E. R., M. A. Price and O. H. Graham. 1967. The occurrence of Phlebotomus

californicus Fairchild and Hertig and Phlebotomus oppidanus Dampf in Texas.

Mosq. News 27(3):429.

1968. The collection of biting flies in west Texas with malaise and animal-
baited traps. Mosq. News 28(3) :465—469.

Fairchild, G. B. and R. F. Harwood. 1961. Phlebotomus sandflies from animal
burrows in eastern Washington. Proc. Entomol. Soc. Wash. 63:239-245.
Fairchild, G. B. and M. Hertig. 1957. Notes on the Phlebotomus of Panama XIII.
The vexator group, with descriptions of new species from Panama and California.

Ann. Entomol. Soc. Am. 50(4):325-334.

—. 1959. Geographic distribution of the Phlebotomus sandflies of Central Amer-
ica (Diptera: Psychodidae). Ann. Entomol. Soc. Am. 52(2):121-124.

Fairchild, G. B. and H. Trapido. 1950. The West Indian species of Phlebotomus
(Diptera, Psychodidae). Ann. Entomol. Soc. Am. 43(3):405-417.

Hall, D. G. 1936. Phlebotomus (Brumptomyia) diabolicus, a new species of biting
gnat from Texas (Diptera: Psychodidae). Proc. Entomol. Soc. Wash. 38:27-29.

Harwood, R. F. 1965. Observations on distribution and biology of Phlebotomus
sandflies from northwestern North America. Pan-Pac. Entomol. 41:1-4.

Mangabeira, O. and P. Galindo. 1944. The genus Phlebotomus in California. Am.
J. Hyg. 40(2):182-194.

Rosabal, R. and A. Miller. 1970. Phlebotomine sandflies in Louisiana (Diptera:
Psychodidae ). Mosq. News 30(2):180-187.

Shannon, R. C. 1913. Feeding habits of Phlebotomus vexator Coq. Proc. Entomol.
Soc. Wash. 15( 4): 165-167.

Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson. 1965. A
catalog of the Diptera of America north of Mexico. U.S. Dept. Agric. Handbook
No. 276. Washington, D.C. 1696 pp.

Vargas, L. and A. Diaz Najera. 1953. Lista de Flebotomus Mexicanos y su distri-
bucion geografia (Diptera: Psychodidae). Rev. Inst. Salubr. Enferm. Trop.
Mexico City 13(4):309-314.

Young, D. G. 1972. Phlebotomine sand flies from Texas and Florida (Diptera:
Psychodidae ). Fla. Entomol. 55(1):61-64.

Vector-Borne Disease Division, Center for Disease Control, Public Health
Service, Department of Health, Education and Welfare, Fort Collins, Colo-
rado 80522.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 543-547

DESCRIPTION OF A NEW POLYMERUS, WITH NOTES ON
TWO OTHER LITTLE KNOWN MIRIDS FROM THE
NEW JERSEY PINE-BARRENS (HEMIPTERA: MIRIDAE)!'

Thomas J. Henry

Abstract.—The new species, Polymerus rostratus, is described from the
New Jersey Pine-Barrens. Hudsonia ericoides L. is shown to be the true
host of Parthenicus vaccini Van Duzee and Polymerus rostratus, and
Arenaria caroliniana Walt. the host of Polymerus nigropallidus Knight.

Several interesting species of Miridae have been described from pine-
barren habitats along the eastern coast of the United States. Knight
(1923), and in other papers, described from the New Jersey Pine-Barrens
several mirids, most of which have not been recorded since. The hosts of
these bugs have been confused or, in most cases, have remained unknown.

On a recent collecting trip to the New Jersey Barrens, my colleague,
A. G. Wheeler, Jr., and I discovered the hosts of several species known
only from one or a few localities and one new species. The description of
a new Polymerus and notes on two other mirids are presented here.

Polymerus rostratus Henry, new species

Figs. 1-3

Male holotype.—Length 3.60 mm, width 1.36 mm, generally brown, with
the head and pronotum largely black; clothed with silvery, silky or tomen-
tose pubescence, intermixed with simple, semierect setae. Head: Length
0.40 mm, width 0.82 mm, black, basal carina, part of lorum and median
line to base of tylus brownish yellow; vertex 0.30 mm, dorsal width of eye
0.26 mm; vertex and part of front set with a few silvery, silky setae.
Rostrum: Length 1.98 mm, reaching well beyond hind coxae to 6th
abdominal segment, segments I and II pale, III dark brown, IV _ black.
Antennae: I, length 0.36 mm, black, extreme base pale; II, length 1.40 mm,
dark brown or reddish brown, base black, thickly clothed with fine, brown,
recumbent setae; III, length 0.76 mm, reddish brown; IV, length 0.56 mm,
reddish brown. Pronotum: Length 0.72 mm, width at base 1.24 mm,
largely black, with the basal %, collar and median line lighter brown (some
specimens are entirely black, except for the basal margin and narrow
median line); surface weakly rugose, calli slightly raised; calli and area
immediately behind thickly clothed with silvery, silky pubescence;
scutellum pale brown, basal angles black (some specimens are more nearly

544 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

black with only the apical 3rd brown), thickly set with silky pubescence.
Hemelytra: Largely brown, with the clavus darker brown, its margins and
inside 2 often approaching black, the costal and radial veins fuscous, often
tinged with red, apical % of embolium and apex of corium red, cuneus
bright red with the apex and outer margin pale; clothed with silvery, silky
pubescence. Membrane: Translucent grayish brown, anal area more black,
veins pale brown. Venter: Pale yellow, dorsal ’ of pleura black, abdomen
greenish yellow with a black line across pleural region, genital segments
more testaceous; thoracic segments thickly set with silvery, silky pubescence,
abdomen clothed with longer, simple setae. Legs: Testaceous to yellowish
orange, front and middle femora with 2 subapical red bands, the basal band
often fading into broad reddish area, hind femora with 3 reddish bands,
these often becoming infuscated, the basal 1 often broadly faded; tibiae
testaceous, tinged with red, spines black, these sometimes with indistinct
black spots at base; tarsi testaceous, 3rd segment and claws_ black.
Genitalia: Parameres typical of genus, not distinguishable from other
Polymerus; spiculum of aedeagus slender, apex acutely produced, covered
with microspines.

Allotype female.—Length 3.72 mm, width 1.64 mm; very similar to male
in coloration and markings, differing largely in the broader form and more
brown pronotum. Head: length 0.44 mm, width 0.90 mm, black, median
line, basal carina and spot on either side of vertex pale; vertex 0.42 mm,
dorsal width of eye 0.24 mm. Rostrum: Length 2.20 mm, reaching beyond
base of ovipositor to 7th abdominal segment. Antennae: I, length 0.34 mm,
black, extreme base and apex pale; I, length 1.28 mm, brown, base black;
III, length 0.78 mm, reddish brown; IV, length 0.50 mm, reddish brown.
Pronotum: Length 0.84 mm, width at base 1.36 mm, largely brown, calli
and anterior angles black; scutellum pale yellowish, black across basal
angles. Hemelytra: more brown, less fuscous and red than males.

Type-data.—Holotype: ¢, Burlington Co., N.J., near Tabernacle, June
13, 1977, taken on Hudsonia ericoides, T. J. Henry and A. G. Wheeler, Jr.
coll. (USNM type no. 75743). Allotype: ¢, same data as holotype
(USNM).° Paratypes: 1 ¢, 9 ?, same data as holotype; 1a ycyyuigee
Ocean Co., N.J., Rt. 37 near Lakehurst, 14 June 1977, taken on H. ericoides,
T.J.H. and A.G.W. coll. (Am. Mus. Nat. Hist., Pa. Dept. Agric., Pa. State
Univ., USNM); 1 2, Ocean Co., N.J., near Tom’s River along Rt. 37,
14 June 1977, taken on H. ericoides, T.J.H. and A.G.W. coll. (Pa. Dept.
NETIC).

Remarks.—Polymerus rostratus is very similar in coloration and markings
to Polymerus basalis Reuter, 1876. When first collected, this species was
thought to be basalis; but after some field observation, its smaller size and
quicker, more erratic flight habit strongly suggested a new form.

Polymerus rostratus keys to basalis in Blatchley (1926) and Knight (1923

VOLUME 80, NUMBER 4

hone A 4,
Pye P| bp
SE ghd

Fig. 1. Typical New Jersey Pine-Barrens habitat for Arenaria caroliniana and
Hudsonia ericoides. Fig. 2. Hudsonia ericoides. Fig. 3. Arenaria caroliniana.

and 1941). It is easily separated from basalis by its smaller size (¢, N = 10,
x = 3.75, 3.48-4.04; 2, N = 10, x = 3.91, 3.68-4.32), mostly black head,
longer rostrum that reaches well beyond the hind coxae to the 6th or 7th
abdominal segment, the relative lengths of the antennal segments (rostratus,

546 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

segment II 2X length of segment III vs. basalis, segment II nearly 4x
length of segment IIL) and the male genitalia.

Extensive collecting throughout much of the Pine-Barrens region revealed
that golden heather, Hudsonia ericoides L. (Cistaceae) (Figs. 1 and 2) is
the only host of rostratus. Only several miles away, but outside the un-
disturbed Barrens, 13 examples of basalis were found breeding on ox-eye
daisy, Chrysanthemum leucanthemum L., in an old field along Rt. 37 near
Tom’s River, June 14, 1977.

Polymerus nigropallidus Knight

Polymerus nigropallidus (Knight, 1923), one of our prettiest Polymerus
species, is known only from Brown’s Mills Junction, New Jersey. Con-
centrated collecting in the Pine-Barrens disclosed that this species is
actually very common and breeds on the low growing, pine-barren sand-
wort, Arenaria caroliniana Walt. (Caryophyllaceae) (Figs. 1 and 3).
Numerous adults and a few nymphs were taken June 13 and 14, 1977
at Lakehurst, Ocean Co. and Tabernacle, Burlington Co.

Parthenicus vaccini Van Duzee

Parthenicus vaccini (Van Duzee, 1915) is known only from Long Island,
N.Y., Massachusetts and Florida (Knight, 1923). The name vaccini would
suggest that this species breeds on Vaccinium spp. and, in fact, numerous
authors have carried this misnomer through the literature. A special effort
was made to collect vaccini on cranberry and blueberry, but only after
carefully beating plants of Hudsonia ericoides did we discover both adults
and nymphs. Parthenicus vaccini was common wherever Hudsonia ericoides
was growing. Eight males, 1 macropterous female and 21 brachypterous
females were taken near Tabernacle, June 13; 1 male and 11 brachypterous
females were taken along Rt. 37 near Tom’s River, June 13; and 1 male
and 4 brachypterous females were taken at Lakehurst, June 14.

Acknowledgments

I wish to give special thanks to Dr. K. C. Kim (Pa. State Univ.) for
supporting this project and to Drs. Kim and A. G. Wheeler, Jr. (Pa. Dept.
Agric.) for reading the manuscript and making useful comments.

Literature Cited

Blatchley, W. S. 1926. Heteroptera or true bugs of eastern Northern America. The
Nature Publishing Co., Indianapolis. 1,116 pp.
Knight, H. H. 1923. Family Miridae (Capsidae). pp. 422-658. In Britton, W. E.

VOLUME 80, NUMBER 4 547

[Ed.], The Hemiptera or sucking insects of Connecticut. Bull. Conn. State Geol.
Nat. Hist. Surv. No. 34. 807 pp.
1941. The plant bugs, or Miridae, of Illinois. Ill. Nat. Hist. Surv. Bull.
No. 22. 234 pp.
Van Duzee, E. P. 1915. New genera and species of North American Hemiptera.
Pomona J. Entomol. Zool. 7:109-121.

The Frost Entomological Museum, Department of Entomology, The
Pennsylvania State University, University Park, Pennsylvania 16802 (mail-
ing address: Bureau of Plant Industry, Pennsylvania Department of
Agriculture, Harrisburg, Pennsylvania 17120).

Footnote

* Authorized for publication Mar. 27, 1978 as paper no. 5480 in the Pennsylvania
Agricultural Experiment Station, University Park, Pennsylvania 16802.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 548-552
DESCRIPTION OF THE MALE OF NEUROBEZZIA GRANULOSA
(WIRTH) (DIPTERA: CERATOPOGONIDAE)

William L. Grogan, Jr. and Willis W. Wirth

Abstract—The male of Neurobezzia granulosa (Wirth) is described and
illustrated for the first time. Locality records are presented. The genus is
compared with its nearest relative, Neurohelea Kieffer.

Wirth and Ratanaworabhan (1972) proposed the genus Neurobezzia for
Bezzia granulosa Wirth (1952) from California and indicated its relation-
ship to Neurohelea Kieffer in the predaceous midge tribe Heteromyiini.
The genus Neurobezzia has been known only from the female sex of
granulosa; in this paper we provide the first description and illustrations of
the male. For an explanation of special terminology of Ceratopogonidae
see Wirth (1952); terms dealing with male genitalia are those of Snodgrass
(1957).

Neurobezzia probably evolved from an ancestor most similar to
Neurohelea. We believe that these two genera are plesiotypic in comparison
with other genera in the tribe Heteromyiini and that they are probably
annectant types similar to ancestral heteromyiines. This is plausible in
view of the fact that they lack elongated hind 4th tarsomeres, elongated
hind claws, bifid 4th tarsomeres, or spinose fore femora, apotypic characters
present in other genera in this tribe. Wirth et al. (1974) included these two
genera in their key to the Ceratopogonidae. They may be further dif-
ferentiated by the following characters:

Neurobezzia Neurohelea
Wing with | radial cell Wing with 2 radial cells
Hind claws without basal Hind claws with basal
inner teeth inner teeth
Antennal ratio 0.89-0.94 Antennal ratio 1.28—1.33
Claspettes narrowly separated Claspettes broadly separated
basally basally

Neurobezzia granulosa ( Wirth)
Fig. 1

Bezzia granulosa Wirth, 1952:240 (female; California).

Neurobezzia granulosa (Wirth); Wirth and Ratanaworabhan, 1972:244
(combination; fig. female flagellum, wing, mandible, palpus, legs, claws,
genitalia ).

VOLUME 80, NUMBER 4 549

Fig. 1. Neurobezzia granulosa, male. A, Wing. B, Genitalia.

550 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Female.—For a detailed description of the female see Wirth (1952) and
Wirth and Ratanaworabhan (1972). To these descriptions may be added
the following: Wing length 1.85 mm; breadth 0.68-0.74 mm; costal ratio
0.91-0.93; vein M2 complete, forking at r-m crossvein as described by
Wirth (1952), not obsolescent at base as figured by Wirth and Ratana-
worabhan (1972). Antennal flagellomeres with lengths in proportion of
93-15-15-15-15-16-18-19-23-23-24-23-28; palpal ratio 2.59-2.67. Claws of fore
and mid legs with basal inner teeth; hind claws without basal inner teeth.

Male.—Wing length 0.94-1.00 mm; breadth 0.35 mm.

Head: Dark brown. Eyes narrowly separated. Antennal pedicel darker
brown; flagellum brown; proximal 10 flagellomeres ovoid, distal 3
flagellomeres elongate; plume moderately well developed. Palpus light
brown, relatively short, extending just beyond tip of proboscis.

Thorax: Brown. Legs lighter brown; 4th tarsomeres cordiform; 5th
tarsomeres slightly swollen; claws small, equal, without basal inner claws,
tips bifid. Wing (Fig. 1A) hyaline, more slender than in female; anterior
veins pale gray, just slightly darker than posterior veins; radial cell extend-
ing to 0.66 of wing length, costa extending beyond to 0.95 of wing length
and bearing a sparse fringe; media forking just proximad of r-m crossvein,
mediocubital fork slightly beyond r-m crossvein. Halter brown.

Abdomen: Brown; intersegmental areas pale; pleuron granular purple.
Genitalia as in Fig. 1B. Ninth sternum 3.3 broader than long, base
curved with a broad deep caudomedian excavation, ventral membrane
spiculate; 9th tergum tapering slightly distally to a broad rounded tip with
a subapical row of 7 large setae, cerci short, each bearing a single large
stout seta. Basimere curved, 2.25 longer than broad; telomere nearly
straight, covered with dense fine pubescence, tip curved and _ pointed.
Aedeagus about as long as broad, membrane and ventral surface spiculate,
basal arch broad, reaching to 0.35 of total length; basal arm long, slightly
recurved, very heavily sclerotized; distal portion more lightly sclerotized,
tapering slightly distally on proximal % then abruptly on distal % to a
rounded tip; underlying membrane broadly crescent-shaped and extending
beyond tip of aedeagus. Claspettes narrowly separated; distal portion
heavily sclerotized, expanded distally into broad paddle shaped structures;
basal arm more heavily sclerotized, recurved nearly 180°, tip slightly
bulbous.

Distribution —California and Oregon; locality records plotted in Fig. 2.

New records—CALIFORNIA: Tulare Co., Elderwood, 14 May 1976,
W. D. Murray, 2 6. OREGON: Deschutes Co., Sparks Lake, 24 July 1969,
K. Goeden, light trap, 1 °.

Discussion-—Our presumption that the Tulare County males are

VOLUME 80, NUMBER 4 551

Fig. 2. Locality records for Neurobezzia granulosa.

Neurobezzia granulosa is based on a careful comparison with the male
characters of other genera of Heteromyiini, and the elimination of those
whose males differ markedly, or which are not known to occur in the
western United States. We are troubled by the fact that the males have
legs without distinct paler markings, while the females of N. granulsoa
have bicolored legs. However the leg color varies somewhat in our females,
and sexual dimorphism in leg color is common in known species of Hetero-
myiini. The position of the base of vein M2, arising at the r-m crossvein
in the female, and arising slightly proximad in the male also forms a
discrepancy which possibly can be explained by the usual sexual di-
morphism in wing venation common in this tribe.

Acknowledgments

We are grateful to Ethel L. Grogan for preparing the illustrations. Thanks
are due W. D. Murray of the Delta Mosquito Abatement District in Visalia,
California, for submission of the lot of ceratopogonid specimens in which
the males were discovered.

Literature Cited

Snodgrass, R. E. 1957. A revised interpretation of the external reproductive organs
of male insects. Smithson. Misc. Collect. 135:1-60.

Wirth, W. W. 1952. The Heleidae of California. Univ. Calif. Publ. Entomol. 9:
95-266.

5o2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
A new genus of biting midge re-

1972.
Pan-Pac. Entomol.

Wirth, W. W. and N. C. Ratanaworabhan.
Ceratopogonidae).

lated to Neurohelea Kieffer (Diptera:

48 :244-945,
Wirth, W. W., N. C. Ratanaworabhan and F. S. Blanton.
genera of Ceratopogonidae (Diptera). Ann. Parasitol. Hum. Comp. 49:595-613.

1974. Synopsis of the

(WLG) Department of Entomology, University of Maryland, College
Park, Maryland 20742 (now at: Department of Biological Sciences, Salis-
bury State College, Salisbury, Maryland 21801); and (WWW) Systematic
Entomology Laboratory, IIBIII, Fed. Res., Sci. and Educ. Admin., USDA,
c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 553-555

A NEW SPECIES OF HYLEMYA FROM WYOMING
(DIPTERA: ANTHOMYIIDAE )

George C. Steyskal

Abstract.—The new species, Hylemya (Delia) evansi, is described from
Teton County, Wyoming.

Specimens submitted for determination were collected by Howard E.
Evans during behavioral studies on solitary wasps. An undescribed species
was included and is described at this time to make its name available.

Hylemya (Delia) evansi Steyskal, new species
Figs. 1-4

Male.—Very similar to H. angustitarsis Malloch, to which it will run in
the keys of Huckett (1965, 1971) and Hennig (1966-1976). It is also
similar to H. antiqua (Meigen), the alternate to H. angustitarsis in Huckett’s
(1971:191) key; but the male terminalia differ in more ways from H.
evansi than do those of H. angustitarsis. Hylemya tenuiventris (Zetter-
stedt), the alternate to H. angustitarsis in Hennig’s key (1966-1976:703),
is also similar to H. evansi. Both H. angustiventris and H. tenuiventris have
the 5th sternum of the male abdomen with narrower and more elongate
lobes than those of H. evansi, and both also lack the 3 contiguous blunt
apical setae on each lobe. Hylemya gracilipes Malloch and H. curvipes
Malloch also bear considerable resemblance to H. evansi; but both of these
species, among other differences, have only one, much longer, blunt apical
seta on each 5th sternal lobe. The most diagnostic characters seem to be
those of the 5th sternum, the curved hindfemur and the short anterodorsal
setae on hindtibia.

The following characters will distinguish H. evansi from its nearest
relatives:

Antenna as in Fig. 4; 3rd segment with short pubescence, 1.8 as long
as wide; aristal hairs 2 as long as basal diameter of arista.

Thorax dull, ochreous gray; notopleural area without fine setae; pre-alar
seta present, shorter than posterior notopleural seta.

Wing 4.4 to 4.5 mm long; tinged with yellowish; setulae lacking on under-
side of costa beyond humeral crossvein.

Legs black, base of tibiae sometimes a little piceous; hindfemur bowed
upward, lower margin concave in profile; tarsi simple. Forefemur with
only minute setulae posteriorly besides macrochaetae; foretibia with blunt
posteroventral apical seta hardly longer than apical tibial diameter. Mid-

554 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-4. Hylemya (Delia) evansi, male. 1, Abdominal sterna 3 to 5, right half.
2, Postabdomen, left profile. 3, Terminalia, rear view. 4, Antenna.

femur with 3 or 4 posteroventral setae in basal %; midtibia without distinct
non-apical erect setae; midtarsus with neither elongate dorsal basitarsal
setae nor elongate nor enlarged apical segmental setae. Hindfemur with
2 or 3 conspicuous ventral setae near apex and about 4 or 5 moderately
long ventral setae basad of midlength; hindtibia without erect ventral
setae, but with 4 or 5 anterodorsal setae not longer than tibial diameter
and 3 posterodorsal setae, the subbasal one little more than % as long as
the other 2.

VOLUME 80, NUMBER 4 555

Abdominal sterna 3 to 5 as in Fig. 1, sternum 3 slightly longer than
sternum 4. Fifth sternum with short stout processes (lobes) bearing 3
closely adjacent, blunt-tipped, apical spinules; 3 long and a few short
lateral setae; and no setae on mesal margin.

Postabdomen as in Figs. 2 and 3; mesolobus (fused cerci) from posterior
view (Fig. 2) slightly concave laterally and nearly evenly tapering; surstyli
well separated, bowed outwards, their depressed apices bearing backwardly
extending fringe; distiphallus with strong posterior (ventral) spicules, most
evident in profile in medial %, but present to apex, where a few are also
lateral.

Female unknown.

Holotype and 5 paratypes, males, Wyoming: 14 mi E Moran Postoffice,
Teton County (just E of Grand Teton National Park), 29 July 1977, note-
book no. 2549 (H. E. Evans), USNM Type #75716 in United States Na-
tional Museum of Natural History.

I am happy to name the species for my friend, the well-known hyme-
nopterist Howard E. Evans; the epithet is a noun in the genitive case.

Literature Cited

Hennig, W. 1966-1976. 63a. Anthomyiidae. In Lindner, E., Die Fliegen der pale-
arktischen Region VII, (Lfg. 315, 262, 268, 271, 272, 276, 278, 280, 282, 283,
294, 295, 296, 297, 306, 307, 308, 314). I-LXXVIII, 1-974 pp., pls. I-XIX.
Huckett, H. C. 1965. The Muscidae of Northern Canada, Alaska, and Greenland
(Diptera). Mem. Entomol. Soc. Can. 42. 369 pp.
1971. The Anthomyiidae of California exclusive of the subfamily Scato-
phaginae (Diptera). Bull. Calif. Ins. Surv. 12. (vi), 121 pp.

Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. and Educ.
Admin., USDA, c/o U.S. National Museum, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.

80(4), 1978, pp. 556-574
NEW SPECIES, RECORDS AND KEY TO
TEXAS LIPOSCELIDAE (PSOCOPTERA)

Edward L. Mockford

Abstract.—Three new species of Liposcelis and one of Belaphotroctes are
described from Texas: Liposcelis hirsutoides, L. ornatus, L. pallidus and
Belaphotroctes alleni. New state records and additional distribution records
of eight other species are included as well as a key to the 13 species of
Liposcelidae now recorded from Texas.

The family Liposcelidae contains small psocids from 1-2 mm in body
length, usually of somewhat flattened form with broad hind femora. Many
of the species commonly invade houses and stored foods; these are called
booklice. They have been incriminated along with dust mites as probable
causative agents of asthmatic reactions (Spieksma and Smits, 1975). Some
of the Liposcelids that occur in rangeland grasses are thought to be inter-
mediate hosts of the fringed tapeworm of sheep (Allen, 1973).

Mockford and Gurney (1956) reviewed the psocids of Texas but did not
include the Liposcelidae. Sommerman (1957) described three new species
of Liposcelis from Texas, and Mockford (1963) described a new species
of Belaphotroctes from that state. To date, these are the only published
records of Liposcelidae from Texas. The present paper includes descriptions
of three new species of Liposcelis, L. hirsutoides, L. ornatus and L. pallidus,
and one new species of Belaphotroctes, B. alleni. Three species of
Liposcelis, one of Belaphotroctes and one of Embidopsocus are recorded
from Texas for the first time. These are Liposcelis bostrychophilus Badon-
nel, L. knullei Broadhead, L. liparus Broadhead, Belaphotroctes badonneli
Mockford and Embidopsocus sp. Thus, the Texas liposcelid fauna is raised
from 4 to 13 species. Records constituting important range extensions are
included for Belaphotroctes hermosus Mockford, Liposcelis deltachi Som-
merman and L. nasus Sommerman. A key to the species is included. Col-
lecting for liposcelids in Texas has as yet been very scanty, and it is likely
that additional collecting will produce more species.

Measurements were made on _ slide-mounted material with a_ filar
micrometer. The micrometer unit = 0.462 ». Abbreviations used in the
measurements and descriptions are explained as follows: Mx4 = distal
segment of maxillary palpus and its length (Table 1); H = greatest head
width; f,, f., f; = first, second and third flagellomeres and their lengths
(Table 1); F + Tr = length of posterior femur + trochanter; FW = greatest

VOLUME 80, NUMBER 4 557

Table 1. Measurements (in w) of Texas species of Liposcelidae. Abbreviations
explained in text.

Name and
Catalog Number Mx4 H f, fo im Ie to tee Iai Tt

Belaphotroctes alleni 100 312 66 55 52 294 141 262
9, ELM #2118

B.alleni 2, ELM 99 298 63 57 53 284 137 244
#2552

Liposcelis hirsutoides 84 282 53 53 52 284 160 222
9, ELM #1673

L. hirsutoides 2°, 80 276 54 53 oil 288 164 21)
ELM #1674

L. hirsutoides @ , 89 292, 48 54 60 288 179 216
ELM #1674

L. hirsutoides @ , 84 296 60 68 al 284 179 218
ELM #4617

L. ornatus 2, 78 264 63 — — 274 170 200
ELM #4617

L. ornatus 2 78 256 59 86 — 264 168 198
ELM #4617

L. pallidus ° , — 326 108 161 187 386 208 280
ELM #2117

Name and
Catalog Number th te ts SI SII MdIx Se

Belaphotroctes alleni 55 38 49 60 — 139 114
9, ELM #2118

B. alleni 2, ELM 54 35 48 58 — 134 138
#2552

Liposcelis hirsutoides 88 34 48 48 52 78 62
9, ELM #1673

L. hirsutoides @ , 88 37 49 43 45 80 73
ELM #1674

L. hirsutoides @ , 89 36 50 44 46 80 76
ELM #1674

L. hirsutoides @ , 87 33 43 44 46 82 =
ELM #4617

L. ornatus @ , 89 33 46 46 46 80 67
ELM #4617

L. ornatus @ , 84 36 50 40 42, lat, 69
ELM #4617

L. pallidus 2, 141 41 53 65 — 120 97
ELM #2117

558 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

width of posterior femur; T = length of posterior tibia; t,, to, t; = lengths
of first, second and third posterior tarsomeres; SI = longest lateral seta of
pronotum and its length (Table 1); SII = longest anterolateral marginal
seta of mesonotum and its length (Table 1); MdIX = a specific long ab-
dominal seta (Fig. 21) (nomenclature of Badonnel, 1962) and its length
(Table 1); Se = longest seta of epiproct and its length (Table 1);
Tg,-Tg,, = abdominal terga 1-11.

The taxonomic categories of section, group and subgroup used in
Liposcelis follow Badonnel (1962, 1963).

In the distribution records, the author was collector unless otherwise
indicated. Catalogue numbers referred to in Table 1 are included with
the records.

Subfamily Embidopsocinae
Embidopsocus sp.

The single specimen is a nymph and cannot be determined to species.
Record.—Texas: Cameron County: Southmost Palm Grove near Browns-
ville, 27 January 1958, beating branches of thorny trees, 1 nymph.

Belaphotroctes alleni Mockford, new species

Diagnosis.—Similar to B. simberloffi Mockford and B. hermosus, differ-
ing from both in having no closely-set group of sense clubs on ventral
surface of Mx4 but having instead diffusely-arranged sense clubs in the
same area (Fig. 45).

Female.—Measurements as given in Table 1. Morphology: Flagellum
with short terminal segment partially fused with subterminal 1. Ocelli
absent (only apterous forms known). Two units in each eye. Lacinial tip
(Fig. 1) with outer and inner prongs bifid distally; denticle between
these relatively broad. Mx4 (Fig. 2) with 3 blunt and 4 pointed sensilla
dorsally; ventrally (Fig. 45) with scattered blunt sensilla in distal %. An-
tennal sensilla: f, with 2 distal rods, 1 stouter than other; f. with 1
slender distal rod, f; and f; each with a stout distal rod; fg with a slender,
curved distal rod, f;5 with a stout distal rod, f;, with a slender curved distal
rod. Pretarsal claw with denticle near tip and short, acuminate appendage
near base. Spermapore sclerite (Fig. 3, compare to Fig. 7 of B. hermosus)
slender, tapering toward opening. Sculpture of integument: Vertex with
faint curved depressed lines enclosing (or partially enclosing) areoles; a
series of fine vermiculate marks between antennal socket and eye on each
side. Abdominal terga without sculpture except for faint transverse lines

VOLUME 80, NUMBER 4 559

bordering posterior ends of segments. Chaetotaxy: Vertex with scattered
sparse setae showing much variation in length, laterals longer than centrals
(a representative lateral = 35 p, a representative central = 17 »). Antennal
orbit with a short posteromedian (< scape), long median (> scape +
pedicel) and short anteromedian (< scape) seta. Pronotum (Fig. 4):
Median lobes each with 2 setae near anterior margin; each lateral lobe
with long SI and short scattered setae including 1 slightly longer than
others near base of median lobe. Meso-metanotum with few, scattered
setae. Prosternum (Fig. 5), with 4-5 setae on each lateral margin including
an anterior minute 1, followed by a somewhat longer 1, followed just in
front of middle by a much longer 1, followed near posterior margin by 1
somewhat shorter than previous 1. Meso-metasternal setae as in Fig. 5.
Setae of Tgs., arranged as in Fig. 6; 1 pair of epiproctal setae much
longer than others and curved. Color (in alcohol 14 years): Eye patches
black. Well-sclerotized body areas yellowish brown, slightly darker in
bands across abdominal terga 1-8.

Holotype 2 and 3 & paratypes, Texas, Jeff Davis County, Davis
Mountains State Park, 25-26 July 1962, sifting ground litter of oak leaves
and juniper debris, ELM #2118. The types are in my collection.

Additional record—New Mexico: Chaves County: Roswell, Diamond A
Ranch, 24 November 1959, R. W. Allen collector, 18 2, ELM #2552.

Note.—The material on which the description of B. hermosus was based
contained a mixture of true hermosus and this species. Figure 14b in
Mockford (1963) refers to this species; all other figures, all measurements
and the description except for reference to this figure are based on true
hermosus.

Etymology.—The species is named for Dr. Rex W. Allen, whose studies
have strongly suggested that psocids may be vectors of the fringed tape-
worm of sheep, Thysanosoma actinioides Diesing. This is one of the
psocid species that Allen has used in his studies.

Belaphotroctes badonneli Mocktord
Belaphotroctes badonneli Mockford, 1972:115.

This species was described from Alachua County, Florida and was pre-
viously known only from that area. The present record marks an extension
of the range westward by approximately 750 miles and suggests a range
around the northern Gulf Coast.

Record.—Texas: Matagorda County: State Highway 35, 8.3 miles along
highway southwest of Old Ocean, 29 November 1975, beating broad-leaved
trees, 1 ¢.

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560

VOLUME 80, NUMBER 4 561

Belaphotroctes hermosus Mockford
Belaphotroctes hermosus Mockford, 1963:27.

The published records of this species as restricted by the description,
above, of B. alleni are entirely from the Lower Rio Grande Valley. The
following record extends the known range somewhat northward.

Record.—Texas: Brooks County: Laguna Salada, 5 miles southwest of
Falfurrias, 5 September 1974, collector not indicated, 1 °&.

Genus Liposcelis
Section I, Group A, Subgroup Aa
Liposcelis liparus Broadhead

Liposcelis liparus Broadhead, 1947:42.

This species has not been recorded previously from the Western
Hemisphere but is known from England and South Africa. In addition to
the Texas record cited below, I have several records from Arizona and New
Mexico, and one from Nebraska.

Record.—Texas: Pecos or Terrell County (about at county line), 42.4
miles along U.S. Highway 90 east of Marathon, 25 July 1962, beating
juniper, 1 °.

Section I, Group A, Subgroup Ab
Liposcelis deltachi Sommerman
Liposcelis delta-chi, Sommerman, 1957:127.

Published records of this species are only from the type-locality, Garner
State Park, Uvalde County, Texas. The following records extend the
known range east and west.

Records.—Texas: Jeff Davis County, Davis Mountains State Park, 25-26
July 1962, sifting ground litter, 1 2; Kleberg County, Kingsville, 7 October
1972, collector not indicated, 1 2; Pecos or Terrell County, 42.4 miles east
of Marathon along U.S. Highway 90, 25 July 1962, beating junipers, 1 °?;
same locality and date, beating yucca, F. Hill collector, 1 &.

<

Figs. 1-6. Belaphotroctes alleni. 1, Lacinial tip, scale of Fig. 3. 2, Distal segment
of maxillary palpus (Mx4), dorsal view. 3, Spermapore sclerite. 4, Thoracic terga
showing chaetotaxy. 5, Thoracic sterna showing chaetotaxy, scale of Fig. 4. 6, Ab-
dominal segments 8-11 showing dorsal chaetotaxy. 7, Belaphotroctes hermosus, sperma-
pore sclerite, scale of Fig. 3. Fig. 8, Liposcelis hirsutoides, lacinial tip, scale of Fig. 3.
Scales in mm.

562 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Liposcelis hirsutoides Mockford, new species

Diagnosis.—Similar to L. hirsutus Badonnel, L. distinctus Badonnel and
L. puber Badonnel. Differing from L. hirsutus and L. puber in paler
coloration, from L. distinctus in having short, truncated setae of the
hirsutus type on all abdominal terga. Also differing from L. hirsutus in
smaller size and in having SII approximately parallel-sided.

Female.—Measurements as given in Table 1. Morphology: Median
suture of vertex recognizable as a break in sculpture, edges of areoles
abutting on it appearing scalloped. Eight units in each eye. Lacinial tip
with prongs strongly diverging; outer prong slightly indented apically,
inner acute apically. Mesothoracic parapsidal sutures visible as thin lines.
Tg, with 3 transverse sclerotized bands, the anterior 2 interrupted medially.
Intersegmental lines 2-3, 3-4, 4-5, 5-6, 6-7 and 7-8 of abdominal terga each
marked in middle by a narrow dark band (Fig. 9). Common trunk of
gonapophyses (Fig. 19) short and broad. T-shaped sclerite as in Fig. 18.
Sculpture: Vertex (Fig. 12) with impressed lines enclosing transverse
areoles, most of them bearing tubercles, usually arranged in rows. Ab-
dominal terga (Fig. 13) with rather regularly spaced tubercles, some
slightly darker than others, arranged in arcs, vaguely setting out areoles, but
tubercles not in areolate pattern in some areas. Weakly sclerotized portions
of Tg;-; with sculpture in same pattern but less pronounced. Chaetotaxy:
Setae of vertex long (a typical central seta = 20 ».) and tapering distally
but not pointed at tips (Fig. 12), from about as long to about 2X as long
as distances between their bases. Pronotum (Fig. 34) with SI decidedly
longer than other setae of lateral lobe, slightly widened in its middle; 2
other setae forming anterior row, both slightly widened in their middles;
3 shorter setae posterior to these. Meso-metathorax with SII about same
length as SI and almost imperceptibly widened in its middle. Six prosternal
setae in an arc; mesosternal row of 9 setae (Fig. 20). Short (a typical seta
of Tg; = 10.6 «) truncated setae abundant on all abdominal terga. Setae
of Tgs4, as in Fig. 21. Color (in alcohol 3 months; specimens in alcohol
16 years generally paler): Eye patches black. Head, body and appendages
generally medium grayish brown; abdominal terga slightly darker on sides
than in middles; head with a slight orange hue. Tg,,., except anterior 2
sclerotized strips, paler than rest of body. A narrow dark line between each
pair of adjacent abdominal terga from 2-8. Each of Tg;-, pale along its
entire posterior border.

Male.—The single male on hand is not in suitable condition to allow
preparation of a description. The sexes were associated by similarity in
color, sculpture of integument and chaetotaxy. There are five units in the
eye.
Holotype female.—Texas, Cameron County, State Hwy. 4, 11 miles along

VOLUME 80, NUMBER 4 563

Fig. 9. Liposcelis hirsutoides 9, dorsal view; appendages, except antennal bases,
not shown; scale of Fig. 10. Fig. 10, Liposcelis ornatus @, dorsal view; appendages,
except antennal bases, not shown. Fig. 11, Liposcelis deltachi 2, dorsal view; ap-
pendages, except antennal bases and maxillary palpi, not shown. Fig. 12, Liposcelis
hirsutoides 2, sculpture of vertex bordering median ecdysial line. Fig. 13, Liposcelis
hirsutoides 2, sculpture of 4th abdominal tergum near middle. Fig. 14, Liposcelis
ornatus ¢, sculpture of vertex near median ecdysial line. Fig. 15, Liposcelis ornatus
@, sculpture of 4th abdominal tergum near middle. Fig. 16, Liposcelis pallidus °&,
sculpture of vertex in parietal region. Fig. 17, Liposcelis pallidus 2, sculpture of 6th
abdominal tergum near middle. Figs. 13-17 to scale of Fig. 12.

564 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

18

4
ty

Figs. 18-21. Liposcelis hirsutoides 9. 18, T-shaped sclerite, scale of Fig. 19.
19, Common trunk of gonapophyses. 20, Thoracic sterna showing chaetotaxy. 21, Ab-
dominal terga 8-11 showing chaetotaxy.

highway east of junction with Farm Rd. 511, 30 January 1958, beating
branches of thorny shrubs on old dune ridge, ELM #1674. Ten para-
types, 1 nymph, and 14 (not designated a paratype due to its poor
condition), same data as holotype; 1 ° paratype, same data except one
mile farther east on same highway. The types are in my collection.

Additional records—Texas: Jim Wells County, 18 miles north of Alice
along U.S. Highway 281, 10 June 1962, beating vegetation, F. Hill and
E. L. Mockford collectors, 1 2; Matagorda County, State Hwy. 35, 2 miles
along highway northeast of Van Vleck, 29 November 1975, beating broad-
leaved trees and on tree trunks, 2 °, ELM #4617.

Liposcelis nasus Sommerman
Liposcelis nasus Sommerman, 1957:128.

The following record extends the range of this species north of the
Lower Rio Grande Valley, the area to which other published records are
restricted.

VOLUME 80, NUMBER 4 565

Record.—Texas: Bexar County, San Antonio, Northwest Preserve Park,
29 June 1973, beating dried cut grass and oak litter, 1 °.

Liposcelis ornatus Mockford, new species

Diagnosis—Marked with a striking pattern (Fig. 10) somewhat similar
to those of L. marginepunctatus Badonnel, L. nigrofasciatus Badonnel and
L. fasciatus Enderlein. Differing from L. marginepunctatus in lacking
series of lateral spots on each side of abdomen. Differing from L. nigro-
fasciatus and L. fasciatus in lacking transverse dark bands of terga in
posterior half of abdomen.

Female.—Measurements as given in Table 1. Morphology: Median
suture of vertex indicated by slight break in sculpture. Eight units in each
eye. Lacinial tip (Fig. 22) normal for the genus. Mesothoracic parapsidal
sutures developed as distinct arched lines. Tg, apparently uniformly
sclerotized. Intersegmental lines 2-3, 3-4, 4-5, 5-6, 6-7 and 7-8 of abdominal
terga each marked in middle by a narrow dark band, those of 3-4 and 4-5
more obvious than others (Fig. 10). Common trunk of gonapophyses (Fig.
23) relatively short and broad. T-shaped sclerite as in Fig. 24. Sculpture:
Vertex (Fig. 14) with impressed lines partially enclosing transverse
areoles, most of them bearing vague tubercles, the tubercles, instead of
lines, forming margins of areoles in places. Abdominal terga (Fig. 15)
with regularly spaced minute tubercles. Weakly sclerotized portions of
Tg;-; with same pattern, somewhat less pronounced. Chaetotaxy: Setae
of vertex as described for L. hirsutoides. Pronotum (Fig. 32) with SI
decidedly longer than other setae of lateral lobe, but other (3) setae of
anterior row each at least % length of SI; 3 short setae posterior to these.
Meso-metathorax with SII about same length as SI. Six prosternal setae
in row of 4 and 2 more posterior; mesosternal row of 9 setae (Fig. 25).
Setae of Tgs1, as in Fig. 26. Color (in alcohol 3 months): Eye patches
black. Ground color of head dull ivory, of thorax and abdomen dull ivory
to white. Postclypeus grayish brown. A medium brown band along mid-
line of vertex to frontal area, there dividing into 2 arms, each running to
antennal base. Antennae grayish brown. Propleura medium brown. A
medium brown area covering each side of meso-metanotum and upper
parts of corresponding pleura. A brown spot on each side of midline in
Tg:, a broad brown band, somewhat irregular on its lateral edges, covering
most of Tg; and Tg, and extending on each side of midline onto Tg;. A
brown spot surrounding the spiracle on each side of Tg; and another
pair of such spots on Tgs. Tgs.: with a slight brownish wash. Antennae
pale brown. Legs each with a diffuse brown spot dorsally on the femur;
tibiae each with a diffuse brown band near distal end and scattered
brown pigment granules more basally. Tarsomeres pale brown, somewhat
darker in first than in others.

566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

0.1

Figs. 22-26. Liposcelis ornatus 9. 22, Lacinial tip, scale of Fig. 23. 23, Com-
mon trunk of gonapophyses. 24, T-shaped sclerite, scale of Fig. 23. 25, Thoracic
sterna showing chaetotaxy. 26, Abdominal terga 8-11 showing chaetotaxy.

Male.—Unknown.

Holotype female.—Texas, Matagorda County, State Highway 35, 2 miles
along highway northeast of Van Vleck, 29 November 1975, beating broad-
leaved trees and on tree trunks. Five 2 paratypes, same data as for holo-
type. The types are in my collection.

Additional records: UNITED STATES: Florida: Alachua County,
Cross Creek, 15 November 1952, beating red maples along creek, 1 ?;
Gainesville, 16 November 1952, beating palm leaves, 1 2°; Newnan’s Lake,
28 March-11 June 1952, trunks and branches of broad-leaved trees and
vines in hammock, 4 2, 3 nymphs; Glades County, 8.6 miles south of
Brighton on State Highway 781, 18 April 1954, beating live oak in palmetto-
dominated hammock, 1 2; Hendry County, State Highway 833, locality

VOLUME 80, NUMBER 4 567

not recorded, 16 April 1954, beating broad-leaved trees and shrubs, and
on trunks and foliage of Sabal palmetto, 2 2°, 1 nymph; Highlands County,
Highlands Hammock State Park, 3 March 1956, beating dead cabbage palm
leaves in hammock, 4 2°; Indian River County, Vero Beach, 18 April 1954,
beating broad-leaved shrubs and trees and Sabal palmetto leaves in dune
hammock, 5 2; Levy County, Seahorse Key, 28 June 1953, beating broad-
leaved shrubs, 1 °; Inglis, collected 9 June 1965 from laboratory culture
from this locality, 5 2; Polk County, Avon Air Force Base, 4 March 1956,
beating saw palmetto leaves in cypress dome, 1 nymph; Sarasota County,
Myakka River State Park, 30 August 1951, ex cabbage palm trunk, 1 2°.
Louisiana: Orleans Parish, New Orleans, 1 December 1965, beating
vegetation in Audubon Park, 1 2°. MEXICO: San Luis Potosi, Tama-
zunchale, 18 June 1966, beating ferns, bromeliads, and cabbage palms,
6 2, E. L. Mockford, R. Sloan and A. Manzano collectors. Tabasco,
4 miles southwest of Frontera, 3 July 1966, beating cacao foliage, 1 °.

Note.——The species was reared in culture at this laboratory through
several generations in 1966. It proved to be obligatorily thelytokous, lacking
males entirely.

Liposcelis pallidus Mockford, new species

Diagnosis.—Similar to L. villosus Mockford in color and chaetotaxy but
differing in sculpture, resembling L. reticulatus Badonnel, L. laparvensis
Badonnel, and L. discalis Badonnel in having finely reticulate sculpture
on membranous zones of abdomen; but this sculpture irregular in size of
reticulations and in relative fineness (Fig. 17). Similar in color to L.
nasus and L. pallens Badonnel, differing from both in details of sculpture
and in possessing only one relatively long seta anteriorly on each lateral
pronotal lobe in addition to SI.

Female.—Measurements as given in Table 1. Morphology: Region of
median suture of vertex not showing a break in sculpture. Eight units in
each eye. Lacinial tip as in Fig. 27. Mesothoracic parapsidal sutures not
visible. Tg, partly obscured by material in digestive tract; visible part
appearing uniform in sclerotization. Intersegmental lines 4-5 and 5-6
each marked dorsally in middle by a narrow, dark, double band; a nar-
rower single band marking intersegmental line 6-7. Common trunk of
gonapophyses elongate (Fig. 28). T-shaped sclerite as in Fig. 29. Sculpture:
Vertex (Fig. 16) with impressed lines enclosing transverse, smooth areoles;
occasional short segments of the lines faint or absent. Abdominal terga
beset with closely-spaced, small polygonal tubercles; on Tg, and Tg; these
changing abruptly in weakly sclerotized posterior ’% of tergum to faint
lines in reticulate pattern becoming bolder more posteriorly (Fig. 17).

568 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

29

Figs. 27-31. Liposcelis pallidus 2. 27, Lacinial tip, scale of Fig. 29. 28, Common
trunk of gonapophyses, scale of Fig. 29. 29, T-shaped sclerite. 30, Thoracic sterna
showing chaetotaxy. 31, Abdominal terga 8-11 showing chaetotaxy.

Chaetotaxy: Setae of vertex (Fig. 16) long and tapering to fine points.
Pronotum (Fig. 33) with SI approximately parallel-sided and decidedly
longer than single other seta of lateral lobe. Other seta of lateral lobe
anterior, slightly swollen medially. Meso-metatergum with SII somewhat
shorter than SI. Prosternal setae apparently 6 in an anterior are with 2
lateral setae longest (Fig. 30). Mesosternal row of 9 or 10 setae. Tgi-«
bearing very few, scattered, short, pointed setae. Setae of Tgs+; as in
Fig. 31. Color (in alcohol 14 years): Eye patches black. Head, body,
and appendages generally dull yellow; reddish-brown subcuticular pigment
granules scattered along sides of head, thorax and abdomen.

Holotype 2 and one ? paratype, Texas, Jeff Davis County, 8.2 miles
southeast of Fort Davis along State Highway 118, 25 July 1962, beating
yucca, ELM #2117. The types are in my collection.

VOLUME 80, NUMBER 4 569

Fig. 32. Liposcelis ornatus @, right halves of pro- and mesonota showing chaetotaxy,
scale of Fig. 34. Fig. 33, Liposcelis pallidus 2, left halves of pro- and mesonota showing
chaetotaxy. Fig. 34, Liposcelis hirsutoides 2, right halves of pro- and mesonota show-
ing chaetotaxy.

Section I, Group B, Subgroup Bb
Liposcelis knullei Broadhead
Liposcelis knullei Broadhead, 1971:264.

The species was previously known from Wooster, Ohio and Ottawa,
Ontario. With the present record, a wide distribution both north-south
and east-west in eastern North America is indicated.

Record.—Texas: Panola County, 5.6 miles south along U.S. Highway 59
from Carthage, 28 November 1975, beating oaks and pines, 2 °.

570 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Section II, Group D
Liposcelis bostrychophilus Badonnel
Liposcelis bostrychophilus Badonnel, 1931:251.

The species is widely distributed. In North America, it has been
recorded from the boundary area of Georgia and Florida, and from
Ottawa, Ontario.

Records.—Texas: Cameron County, State Highway 4, 11 miles east of
junction with County Road 511, 30 January 1958, beating dead branches
of thorny shrubs on old dune ridges, 1 ?; Hidalgo County, Bentsen Rio
Grande Valley State Park, 28 January 1958, beating palms and branches
of thorny trees and shrubs, and sifting ground litter, 24 2, 8 nymphs;
Kleberg County, Kingsville, 28 August 1977, in stored grain, numerous 2
and nymphs, R. Schmidt collector; Lee County, 2.5 miles north of Giddings
on Highway 77, 10 June 1962, 1 2°, E. L. Mockford, F. Hill and J. M.
Campbell collectors.

Key to the Texas Species of Liposcelidae

1. Hind femur bearing a lateral protuberance. Female subgenital
plate with a T-shaped sclerite. Apterous forms only (Figs. 35
and 36) Subfamily Liposcelinae, genus Liposcelis 5

— Hind femur lacking a lateral protuberance. Female subgenital
plate without a T-shaped sclerite. Males apterous, females generally
in macropterous and apterous forms (Figs. 37-41)

Subfamily Embidopsocinae 2

2. Mx4 less than 1.5x as wide in middle as next segment. Tg;-, each
with a slender, heavily sclerotized transverse strip (Fig. 37)

Genus Embidopsocus (E. sp.).

- Mx4 at least 1.5 as wide in middle as next segment. Tg;-s lacking
sclerotized transverse strips (Fig. 42). Genus Belaphotroctes 3

3. Female with Mx4 fully 2 as wide in middle as next segment and
bearing on its lower surface 2 groups of closely-set rod-like sensilla
(Fig. 43). Corticolous species Belaphotroctes badonneli Mockford

— Female with Mx4 about 1.5 as wide in middle as next segment
and bearing on its lower surface a single group of closely-set
rod-like sensilla, or scattered, short sensilla (Figs. 44, 45). Ground

litter species +
4. Female with 1 group of closely-set rod-like sensilla on lower sur-
face of Mx4 (Fig. 44) Belaphotroctes hermosus Mockford

— Female with scattered rod-like sensilla on lower surface of Mx4
(Fig. 45) Belaphotroctes alleni, new species

VOLUME 80, NUMBER 4 571

Figs. 35-36. Liposcelis liparus 9. 35, Dorsal view with left legs. 36, Subgenital
plate with T-shaped sclerite. Figs. 37-41, Embidopsocus laticeps 2°. 37, Dorsal view
of apterous female with right legs, scale of Fig. 35. 38, Subgenital plate, scale of Fig.
36. 39, Head and thorax of macropterous female, dorsal view. 40, Forewing of macrop-
terous female, scale of Fig. 35. 41, Hindwing of macropterous female, scale of
Fig. 35. Fig. 42, Belaphotioctes sp., abdominal terga.

Ot

Tg;-, uniform in color, not presenting a pale posterior mem-
branous band (Fig. 46) Section I 6
— Tg,-; annulate, i.e., each presenting a pale posterior membranous
band with sculpture different from that of anterior portion of ter-
gum (Fig. 47) Section II —Liposcelus bostrychophilus Badonnel
. SI of pronotum long and strong, about equal in length to distance
between its base and anteromedial margin of lateral pronotal lobe;
lateral pronotal lobe with either a transverse row of long, strong
setae or a single 1 in addition to SI (Figs. 32-34) Group A 7
— SI of pronotum not so long and strong, in length decidedly less

[op)

572 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

45

43

Fig. 43. Belaphotroctes badonneli 92, distal 2 segments of maxillary palpus, ventral
view. Fig. 44, Belaphotroctes hermosus @, distal 2 segments of maxillary palpus,
ventral view. Fig. 45, Belaphotroctes alleni ?, distal 2 segments of maxillary palpus,
ventral view. All to same scale.

than distance between its base and anteromedial margin of lateral
pronotal lobe; all other setae of lateral pronotal lobe much shorter
than SI (Fig. 48) Group B12
7. Two very long, curved, fine setae on epiproct (Fig. 49)
Liposcelis liparus Broadhead

— Epiproct lacking pair of long, curved, fine setae 8
8. A single long, strong seta on lateral lobe of pronotum in addition

to SI (Fig. 33) Liposcelis pallidus, new species
— A transverse row of long, strong setae on each lateral lobe of

pronotum in addition to SI (Figs. 32 and 34) )
9. Body color essentially uniform pale brown or buffy yellow. 10
— Body color a contrasting pattern of brown or reddish-brown marks

on a creamy white background I:

10. Body color pale brown except white on Tg,,. (Fig. 9); short,
truncate setae abundant on all abdominal terga (Fig. 50)
Liposcelis hirsutoides, new species
- Body color buffy yellow except anterior % of head, gradually
darkening to rusty brown on clypeus and labrum; short setae of
abdominal terga slender and sparse (Fig. 51)
Liposcelis nasus Sommerman
11. Body marked with a series of reddish-brown spots along each side
of abdomen (Fig. 11) Liposcelis deltachi Sommerman

VOLUME 80, NUMBER 4 573

Fig. 46. Abdominal terga of Liposcelis species of Section I (L. simulans Broad-
head shown here), scale of Fig. 47. Fig. 47, Abdominal terga of Liposcelis species of
Section II (L. bostrychophilus shown here). Fig. 48, Pronotal chaetotaxy of Liposcelis
species of Group B (L. knullei shown here). Fig. 49. Liposcelis liparus @ epiproct.
Fig. 50, Liposcelis hirsutoides 2, setae of 5th abdominal tergum in middle, scale of Fig.
51. Fig. 51, Liposcelis nasus @ , setae of 5th abdominal tergum in middle.

— Body marked with variegated brown pattern on head and thorax,

a broad, brown crossband on Tg;-;, lateral brown spots on Tgo-11
(Fig. 10) Liposcelis ornatus, new species

12. Body color medium brown; lacinial tip with inner prong grooved,

a short denticle at base of inner prong (Sommerman, 1957, Fig. 1)
Liposcelis lacinia Sommerman

— Body color dark brown; lacinial tip with inner prong not grooved
and lacking a basal denticle Liposcelis knullei Broadhead

574 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Acknowledgments

Field work on my 1962 trip was supported by a National Science
Foundation grant, NSF G-19263, to Illinois State University. My collecting
trip in 1975 was supported in part by the College of Arts and Sciences,
Illinois State University. Material of several species was lent by Dr. Joel
Hallan and Dr. James Gillaspy of Texas A & I University, Kingsville.

Literature Cited

Allen, R. W. 1973. The biology of Thysanosoma actinioides (Cestoda: Anoplo-
cephalidae ) a parasite of domestic and wild ruminants. N.M. State Univ. Agric.
Expt. Stn. Bull. 604: 1-69.

Badonnel, A. 1931. Contribution a l’étude de la faunne du Mozambique. 4° note.—

Copeognathes. Ann. Sci. Nat. Zool. 14:229-260.

1962. Psocoptéres. Biologie de Amérique Australe I: 185-229.

——. 1963. Psocoptéres terricoles, lapidicoles, et corticoles du Chili. Biologie
de Amérique Australe I1:291-338.

Broadhead, E. 1947. New species of Liposcelis Motschoulsky (Corrodentia, Lipo-
scelidae) in England. Trans. R. Entomol. Soc. London 98:41-58.

1971. A new species of Liposcelis (Psocoptera, Liposcelidae) from North

America with records of another species. J. Nat. Hist. 5:263-270.

Mockford, E. L. 1963. The species of Embidopsocinae of the United States
(Psocoptera: Liposcelidae ). Ann. Entomol. Soc. Am. 56:25-37.

—. 1972. New species, records, and synonymy of Florida Belaphotroctes
(Psocoptera: Liposcelidae). Fla. Entomol. 55:153-163.

Mockford, E. L. and A. B. Gurney. 1956. <A review of the psocids, or book-lice and
bark-lice, of Texas. J. Wash. Acad. Sci. 46:353-368.

Sommerman, K. M. 1957. Three new species of Liposcelis (= Troctes) (Psocoptera)
from Texas. Proc. Entomol. Soc. Wash. 59:125-129.

Spieksma, F. T. M. and C. Smits. 1975. Some ecological and biological aspects of
the booklouse Liposcelis bostrychophilus Badonnel 1931 (Psocoptera). Neth.
J. Zool. 25:219-230.

Department of Biological Sciences, Illinois State University, Normal,
Illinois 61761.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 575-605
ADDITIONAL APHID-HOST RELATIONSHIPS AT THE
LOS ANGELES STATE AND COUNTY ARBORETUM
(HOMOPTERA: APHIDIDAE)

Harry G. Walker, Manya B. Stoetzel and Leonid Enari

Abstract.—This is the eighth report on collections of aphids from plants
in the Los Angeles State and County Arboretum at Arcadia, California.
About 140 species of aphids are listed with information on their host
plants, date of collection and abundance.

Aphids are plant feeders and many are efficient vectors of plant viruses.
Information on the host plant relationships and distribution of aphids is
becoming increasingly important. This is the eighth in a series of papers
which contribute to the knowledge of the aphids and their host plants
in the Los Angeles State and County Arboretum, Arcadia, California. As
in three previous papers (Leonard et al., 1972; Leonard and Walker, 1973
and 1974), this one deals with a variety of aphids and their host plants.
Four papers (Leonard et al., 1970, 197la, 1971b and 1971c) dealt with six
specific aphids and their host plants.

Walker has been responsible for the collection of the aphids and Enari
for the identification of the host plants. In this report, Stoetzel is respon-
sible for the identification of some of the aphids and for help in the
publication. In addition the authors would like to express their deep
appreciation to the following for the identification of many of the aphids
included in this report: R. L. Blackman (RLB), T. L. Bissell (TLB),
C. S. Wood-Baker (W-B), V. F. Eastop (VFE), A. K. Ghosh (AKG),
T. Kono (Kono), M. D. Leonard (MDL) deceased, R. J. Nielson (RJN),
F. W. Quednau (FWQ), A. G. Robinson (AGR), C. F. Smith (CFS) and
D. J. Voegtlin (DJV).

Aphid, host plant relationships at the Los Angeles State and County
Arboretum are given and include a list of the aphid, the host, the date of
collection, the relative abundance of the aphid and the initials of the person
making the aphid identification.

Acyrthosiphon (Metopolophium) dirhodus Stereospermum kunthianum Cham.
( Walker ) 15/1X/69 Abundant (W-B)
Bromus carinatus Hook. & Arn. Acyrthosiphon pisum (Harris )
3/IV/71 Abundant (CFS) Acacia gillii Maiden & Blakely
Acyrthosiphon pelargonii (Kaltenbach ) 2/XI1/67 Moderate (MDL)
Pelargonium quercifolium Baum. Sophora tetraptera F. Mill.

24/XI/69 Moderate (AKG) 13/V/69 Scarce (AKG)

576 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Teucrium scorodonia L.

23/11/70 Scarce (RJN)
Acyrthosiphon (Rhodobium) porosus

(Sanderson )

Rosa hugonis Hemsl.

17/I11/71 Abundant (MBS)

Rosa sp.

4/XII/69 Abundant (MBS)

Tecoma fulva G. Don
1/V/69 Abundant (W-B)

Acyrthosiphon sp.

Rhamnus californica Eschsch.
31/XII/69 Scarce (AKG)

Amphorophora rubitoxica Knowlton

Rubus palmatus Thunb.
18/V/76 Abundant (RLB)
Amphorophora sp.
Callitris propinqua R. Br.
20/1/71 Abundant (Kono )
Hymenanthera dentata R. Br.
13/V/70 Moderate (RJN)
Salvia microphylla HBK
20/III/70 Moderate (RJN )
Aphis armoraciae Cowen
Conyza bonariensis (L.) Cronq.
11/XI/73 Moderate (CFS)
Aphis sp. near armoraciae Cowen
Conyza bonariensis (L.) Cronq.
30/1IX/75 Abundant (RLB)
Aphis cephalanthi Thomas
Clerodendrum trichotomum Thunb.
27/V1/69 Moderate (RJN )
Salvia greggii A. Gray
23/1/69 Abundant (RJN)
Aphis citricola van der Goot
Acacia albida Lindl.
30/X/70 Scarce (AKG)
Actinostrobus pyramidalis Miq.
25/1V/68 Moderate Vagrant (VFE)
Aesculus pavia L.
11/III/70 Scarce (AKG)
Agathis palmerstoni F. Muell.
3/V1/68 Moderate (VFE)

Aloe wickensii Pole-Evans
27/XII/69 Moderate (AKG)

Aloysia triphylla (L’Hér.) Britt.

28/1V/70 Moderate (RJN)

Araucaria heterophylla (Salisb. )
Franco

27/TII/70 Scarce (VFE)

Bauhinia blakeana Dunn

29/X/70 Moderate (AKG)

Britoa guazumifolia Diego Legrand

19/VI/70 Moderate (AKG)

Broussonetia papyrifera Vent.

24/VI/70 Moderate (AKG)

Buddleia saligna Willd.

16/VI/70 Scarce (AKG)

Buxus microphylla Sieb & Zuce. var.
japonica (Muell. Arg.) Rehd. &
E. H. Wils.

25/V1I/70 Abundant (AKG)

Callicarpa dichotoma (Lour.) C. Koch

30/XII/69 Scarce (RJN)

Callitris propinqua R. Br.

30/V/68 Vagrant (VFE)

26/V/69 Vagrant (VFE)

C. rhomboidea R. Br.

5/VI/70 Vagrant (VFE)

Calocedrus decurrens (Torr. )

Florin

24/1V/71 Vagrant (VFE)

Calodendrum capense (L.F.) Thunb.

5/V/70 Moderate (AKG)

Camptotheca acuminata Decne.

14/VII/70 Scarce (AKG)

Carissa grandiflora DC.

28/V1/68 Moderate Vagrant (VFE)

Cassia abbreviata D. Oliver subsp.
beareana (Holmes) Brenan

7/V1/68 Scarce (VFE)

C. bicapsularis L.

17/11I1/70 Abundant (AKG)

C. excelsa Schrad

11/III/70 Scarce (AKG)

Catalpa bignonioides Walt.

15/V/69 Scarce Vagrant (VFE)

Cedrus deodara (D. Don) G. Don

29/1V/69 Moderate Vagrant (VFE)

Celastrus loeseneri Rehder & Wilson
2/VII/70 Abundant (AKG)

VOLUME 80, NUMBER 4

Chamaecyparis lawsoniana (A. Murr. )
Parl.

4/VI1/69 Moderate Vagrant (VFE)

C. nootkatensis (D. Don) Spach

1/VII/69 Moderate Vagrant (VFE)

Chamelaucium ciliatum Desf.

5/V/72 Abundant (AKG)

Chomelia obtusa Cham. & Schlect.

16/III/70 Abundant (AKG)

Chorisia insignis HBK

19/II/70 Scarce (AKG)

Cistus incanus L.

20/1V/70 Scarce Vagrant (VFE)

Citharexylum myrianthum Cham.

19/VII/69 Scarce (RJN)

Citrus australis Planch.

27/111/70 Abundant (AKG)

C. “Sweet Tangor’

8/1V/70 Abundant (VFE)

Clerodendrum japonicum (Thunb.) Sweet

18/VI/69 Moderate (RJN)

C. nutans Wallich

10/XI/70 Scarce (RJN)

C. trichotomum Thunb.
27/V1/69 Moderate (RJN )
11/VII/70 Scarce (RJN)

Cleyera japonica “Tricolor’
19/V1/69 Scarce (AKG)
Cocculus laurifolius DC.
26/V1/70 Moderate (AKG)
Combretum microphyllum Klotzsch.
13/VI1/70 Moderate (AKG)
Cotoneaster cooperi Marquand
29/V1/70 Abundant (AKG)

C. frigidus Wall.

24/V1/69 Abundant (AKG)

C. pannosus French.

11/XII/69 Abundant (AKG)

C. wardii W. W. Smith

6/1I1/75 Abundant (RLB)
Crataegus laevigata ‘Pauls’ Scarlet’
23/V/75 Abundant (RLB)
Cryptocarya rubra Skeels
16/I11/70 Abundant (AKG)
Cryptomeria japonica ‘Araucarioides’
22/1V/71 Scarce Vagrant (VFE)

C. japonica ‘“Viminalis’

22/1V/71 Moderate (VFE)
Cryptostegia grandiflora R. Br.
4/1/71 Moderate (Kono)

Cupressus cashmeriane Royle
17/1IV/71 Abundant Vagrant (VFE)
C. forbesii Jeps.

13/X/69 Scarce Vagrant (VFE)

C. lusitanica Mill.

17/IV/71 Abundant Vagrant (VFE)
C. pygmaea (Lemm.) Sarg.
1/1V/70 Abundant Vagrant (VFE)

C. sempervirens L.
20/V1/69 Abundant Vagrant (VFE)

Cydonia sinensis Thovin
17/V/76 Abundant (RLB)

Duranta repens L.
26/VI/70 Scarce (RJN )

D. repens ‘Alba’

18/XII/68 Moderate (RJN )
19/VII/69 Scarce (RJN )
5/11I/70 Abundant (RJN )

D. stenostachya Tod.
30/XII/69 Abundant (RJN )

Echium wildpretti H. Pearson
19/V/69 Scarce Vagrant (VFE)
Eucalyptus amygdalina Labill.
14/V/70 Moderate (AKG)

E. cladocalyx F. Muell.

21/1/71 Moderate (Kono)
Euonymus europaea L.

7/X/76 Scarce (RLB)

Ficus sakalavarum Baker
19/III/70 Scarce (AKG)

Gardenia globosa Hochst.

14/1/71 Abundant (Kono)
Hymenosporum flavum F. Muell.
20/1/75 Abundant (RLB)
4/1I1/75 Abundant (RLB)
Jasminum subhumile W. W. Smith
26/V1/69 Scarce (AKG)
Juniperus chinensis ‘Armstrongii’
19/V/69 Scarce Vagrant (VFE)
J. chinensis “Blaauw’

2/V/70 Scarce Vagrant (VFE)

578 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

J. chinensis ‘Fruitlandii’

19/V/69 Scarce Vagrant ( VFE)
J. scopulorum ‘Lakewood Globe’
19/V/69 Scarce Vagrant (VFE)
Malus floribunda Siebold
30/VI/69 Moderate (AKG)
16/1X/70 Abundant (AKG)
5/III/77 Moderate (RLB )

M. ionensis (A. Wood) Britt.
23/V/70 Abundant (AKG)
Ophiopogon japonicus Ker.-Gawl.
16/IV/69 Scarce (AKG)
Oreopanax peltatus Linden
8/1/69 Abundant (AKG)
Osmaronia cerasiformis Greene
24/V1/69 Moderate (AKG)
Owenia acidula F. Muell.
29/V/69 Scarce (AKG)
Paulownia fortunei (Seem. ) Hance
20/VI1/69 Scarce (AKG)
Peltophorum africanum Sond.
18/VI/69 Abundant (AKG)
Phaedranthus buccinatorium Miers
19/V1/69 Scarce (AKG)
Phellodendron lavallei Dode
26/V1/69 Abundant (AKG)
Physalis floridana Rydb.
17/XII/69 Scarce (AKG)
Pittosporum daphniphylloides Hayata
14/1/71 Moderate (Kono )

P. mannii Hook.

5/11/75 Abundant (RLB)
Poncirus trifoliatus Rafin
6/VI1/69 Scarce (AKG)

Prunus caroliniana ( Mill.) Ait.
19/V/69 Abundant (RLB)

P. mahaleb L.

9/VI/69 Abundant (RLB)

P. yedoensis Matsum.

23/V1/69 Scarce (AKG)
Pseudocydonia sinensis C. K. Schneider
15/1/69 Abundant (AKG)
Pseudopanax ferox T. Kirk.
24/V1I/69 Abundant (AKG)
Psidium sp.

18/VII/69 Moderate (AKG)

Pyracantha coccinea M. J. Roem.
5/1/71 Scarce (Kono)

P. koidzumii (Hayata) Rehder
8/VII/69 Abundant (AKG)
Pyracomeles vilmorini Rehder
13/III/69 Abundant (AKG)
Pyrus kawakamii Hayata
7/X/76 Abundant (RLB)

P. sikkimensis (Wenz. ) Kochne
14/V1/69 Abundant (AKG)
Raphiolepis indica Lindl.
6/11/75 Abundant (RLB)
Rhus chinensis Mill.

8/VII/69 Scarce (AKG)

Rosa pendulina L.

9/III/76 Scarce (MBS)

R. pisocarpa A. Gray

23/11/71 Abundant (MBS)

R. spinosissima L.

10/VII/68 Scarce (MBS)

Rosa sp.
29/V1/69 Abundant (MBS)

Salvia greggii A. Gray
10/VII/68 Scarce (RJN)
28/1X/70 Abundant (RJN)
30/XII/70 Scarce (RJN)

S. mellifera Greene
21/11/70 Abundant (RJN)
S. microphylla HBK
21/1V/69 Scarce (RJN)
26/II1/70 Abundant (RJN)
S. officinalis L.

16/VII/69 Scarce (RJN)

S. purpurea Cav.
8/1/70 Moderate (RJN )

Schefflera arboricola Hayata
9/X/76 Abundant (RLB)
Sorbus aria (L.) Crantz.
19/V/76 Abundant (RLB)
Spiraea chamedryfolia L.
3/1/69 Abundant (AKG)

S. trilobata L.

20/V1/69 Moderate (W-B)

Stranvaesia nussia Decne.
4/1V/69 Moderate (AKG)

VOLUME 80, NUMBER 4

Tabebuia impetiginosa “Roxo’

19/VII/69 Scarce (AKG)
Teucrium chamaedrys L.
22/II1/69 Moderate (RJN )
Trema guinensis Priemer
5/1/71 Scarce (Kono)
Ulmus laevis Pall.

8/VII/69 Scarce (AKG)
Viburnum judii Rehder.
9/VI/69 Abundant (AKG)
V. macrocephalum Fort.
3/1V/69 Scarce (AKG)

V. odoratissimum Ker.-Gawl.
6/III/75 Abundant (RLB)
7/X/76 Abundant (RLB)
5/11/77 Abundant (RLB)
V. rufidulum Raf.
24/V1/69 Abundant (AKG)
V. tinus L.

6/11/75 Abundant (RLB)
24/V1/69 Moderate (AKG)
Vitex negundo L.

21/V1/69 Abundant (AKG)
Watsonia sp.

17/V/69 Scarce (AKG)
Weigela floribunda “Variegata’
8/VII/69 Moderate (AKG)

Aphis craccivora Koch

Abutilon mauritianum Sweet.
23/1X/70 Abundant (W-B)
Acacia farnesiana Willd.
7/V/70 Scarce (W-B)

A. pinnata Willd.

11/IV/66 Moderate (MDL)

Callistemon acuminatus Cheel.

28/V/70 Scarce (AKG)

Caragana densa Kom.
13/VII/70 Abundant (AKG)

Colutea istria Mill.
7/VII/70 Abundant (AKG)

Cupressus lindleyi Klotzsch.

17/IV/71 Abundant Vagrant (VFE)

Cytisus sp.

23/V/75 Abundant (RLB)
Indigofera australis Willd.
17/VI/70 Moderate (W-B)

I. gerardiana R. Grah.
10/X/71 Abundant (Kono)
Juniperus chinensis “V iridis’
27/V/71 Abundant (AKG)
Phaseolus caracalla L.

2/X/75 Abundant (RLB)
Pittosporum phillyraeoides DC.
7/X/76 Moderate (RLB )
Salsola iberica Sinnen & Pall.
9/V/69 Moderate (W-B)

Aphis fabae Scopoli

Achillea tomentosa L.
20/V/76 Abundant (RLB)
Agave sp.

23/V/75 Abundant (RLB )
Althaea setosa Boiss.
28/XII/70 Moderate (W-B )
Bulbinella robusta Kunth.
5/11/75 Abundant (RLB)

579

Cordia cylindristachya Roem. & Schult.

4/1/71 Scarce (Kono)

Cynara cardunculus L.
19/V/76 Abundant (RLB)

Eucalyptus maidenii F. J. Muell.

8/III/77 Abundant (RLB)
Spathodea campanulata Beauv.
31/I11/69 Abundant (W-B)
Tecoma fulva G. Don.

1/V/69 Abundant (W-B)
Tulipa gesneriana L.

1/1V/69 Abundant (W-B)
6/11/75 Abundant (RLB)

Aphis gossypii Glover

Abutilon indicum Sweet.
16/V1/70 Abundant (W-B)
A. mauritianum Sweet.
27/1X/68 Moderate (AKG)
23/1X/70 Abundant (W-B)
Achillea tomentosa L.
13/V/69 Scarce (AKG)
Agave sp.

7/11/70 Abundant (W-B)
Aloe ciliaris Haw.

19/11/70 Abundant (AKG)

Amelanchier sanguinea DC.
12/XI1/70 Abundant (W-B)

580 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Arctostaphylos oppositifolia Parry C. stans Sieb. & Zucc.

30/XII/70 Abundant (W-B) 18/VI/70 Scarce (AKG)

Aspidosperma australe Muell. Clerodendrum trichotomum Thunb.

13/V/70 Scarce (AKG) 27/V1/69 Moderate (RJN )

Bidens vulgatus Greene 11/VII/70 Scarce (RJN )

7/11/70 Abundant (W-B) Cordia cylindristacha Roem. & Schult.

Bombax sp. 4/1/71 Scarce (Kono)

5/X/70 Moderate (W-B ) Cordyline rubra Hueg.

Bowkeria gerardiana Harv. 28/V/69 Moderate (AKG)

10/V1/68 Scarce Vagrant (VFE) Coreopsis tinctoria Nutt.

Buddleia saligna Willd. 24/VII/70 Abundant (W-B)

16/V1I/70 Scarce (AKG) Cotoneaster dammeri ‘Skogsholmen’

Callistemon acuminatus Cheel. 13/VII/70 Abundant (W-B)

28/V/70 Scarce (AKG) C. henryanus (C. K. Schneid.) Rehd. &

C. phoeniceus Lindl. Wils.

2/X/75 Abundant (RLB) 13/VIII/68 Scarce (AKG)

Calotropis procera Ait. C. salicifolius Franch

15/X1I/70 Moderate (W-B) 5/VII/70 Abundant (W-B)

Camellia sinensis Kuntze Crassula argentea Thunb.

22/1X/70 Abundant (W-B) 11/XI/70 Abundant (W-B)

Cassia abbreviata D. Oliver subsp. Crataegus lavallei Herincq
beareana (Holmes) Brenan 25/1V/70 Abundant (W-B)

7/V1/68 Scarce (VFE) Crotalaria retusa L.

C. nemophila coriacea Symon. 28/V/70 Abundant (W-B)

15/1/71 Moderate (Kono) Crowea exalata F. Muell.

Catalpa bignonioides Walt. 29/XII/70 Scarce (W-B)

15/V/69 Scarce Vagrant (VFE ) Cucurbita foetidissima HBK

C. bungei C. A. Mey. 28/X/70 Abundant (AKG)

16/X/70 Abundant (AKG) Echium fastuosum Jacq.

C. speciosa Warder 5/1/71 Moderate (Kono)

29/1X/75 Moderate (RLB) E. wildpretti H. Pearson

Ceiba pentandra Gaertn. 19/V/69 Scarce Vagrant (VFE)

12/X1/70 Abundant (AKG) Eucalyptus leucoxylon F. J. Muell.

Celtis willdenowiana Roem. & Schult. 20/1/71 Abundant (Kono)

23/VII/70 Abundant (W-B) E. sideroxylon A. Cunn.

Celtis sp. 20/1/71 Moderate (Kono )

6/VII/70 Abundant (W-B ) E. stowardii Maiden

Centranthus ruber DC. 30/IX/75 Abundant (RLB)

23/11/70 Abundant (W-B) Euonymus japonica Thunb.

19/VII/70 Abundant (AKG) 22/1V/70 Scarce (AKG)

Chiranthodendron pentadactylon Larreat. — Forsythia ‘Lynwood Gold’

29/1/70 Scarce (W-B) 6/1/71 Moderate (Kono)

Cissus antartica Venten. Hibiscus hamabo Sieb. & Zuce.

3/XI1/70 Scarce (AKG) 4/1/71 Abundant (Kono)

Clematis armandii Franch H. syriacus “Paeonaeaflorus’

20/V1I/70 Abundant (W-B ) 6/1/71 Moderate (Kono )

VOLUME 80, NUMBER 4

Hypericum chinense L.
3/11/70 Scarce (AKG)
Leptospermum patersonii F. M. Bailey
14/1/71 Scarce (Kono)

Liquidambar orientalis Mill.
14/1/71 Abundant (Kono)

Manfreda maculosa (Hook. ) Rose
7/V/70 Abundant ( W-B )
Parmentiera edulis DC.
18/VII/69 Scarce (AKG)
Phlomis taurica Hartwiss
15/III/69 Scarce (AKG)
Pittosporum sp.
14/1/71 Abundant (Kono)
Plumeria rubra L.
21/III/69 Scarce (AKG)
Rosa banksiae Ait. f.
24/1V/70 Scarce (MBS)
R. “Margo Koster’
22/V/68 Abundant (MBS)
R. “Wind Chimes’
9/11/76 Abundant (MBS)
Rudbeckia hirta L.
29/1X/69 Moderate (W-B)
Ruttya fruticosa Lindau.
16/V/69 Scarce (AKG)
Solanum robustum H. Wendl.
15/III/69 Moderate (AKG)
Stigmaphyllon affine A. Juss.
1/X/75 Abundant (RLB)
Tecoma fulva G. Don
1/V/69 Abundant (W-B)
T. mollis HBK
19/VII/69 Scarce (AKG)
Xylosma congestum (Lour.) Merrill
8/1V/69 Scarce (AKG)
Aphis helianthi Monell

Ribes aureum Pursh var. gracillimum
(Cov. & Britt.) Jeps.
21/X1/69 Abundant (W-B)

Schefflera arboricola Hayata
27/111/69 Abundant (W-B)
Schinus terebinthifolius Raddi
18/VI/68 Scarce (W-B)

581

Aphis sp. probably helianthi Monell
Raphiolepis indica “Tashiroi’
31/1/69 Abundant ( W-B)
Sanguisorba minor Scop.
21/VII/70 Moderate ( W-B)

Aphis ilicis Kaltenbach

Ilex cassine L.
24/X1/71 Abundant (W-B)

Aphis maidiradicis Forbes
Portulaca oleracea L.
8/IX/72 Abundant (CFS)
Aphis sp. near maidiradicis Forbes
Sonchus oleraceus L.
30/1X/75 Abundant (RLB)
Portulaca oleracea L.
29/1X/75 Abundant (RLB)
Aphis nasturtii Kaltenbach
Acacia arabica Willd.
6/VI/69 Moderate (W-B)
Catalpa japonica Dode
10/VII/70 Scarce (AKG)
Aphis sambuci L.
Aloysia triphylla (L’Hér) Britt.
8/XII/69 Scarce (RJN )
Duranta repens ‘Alba’
5/11/70 Abundant (RJN )
Sambucus kamtschatica E. Wolf.
24/X/69 Abundant (W-B)
Aphis sedi Kaltenbach
Sedum spectabile Bor.
4/11/69 Abundant (W-B)
Aphis sp.
Albizia procera Benth.
13/VI1/70 Scarce (W-B)
Amaranthus graecizans L.
2/X/72 Abundant (CFS)
30/1X/75 Abundant (RLB )
Angianthus tomentosus Wend.
4/11/75 Abundant (RLB)
Basancantha spinosa K. Schum.
14/1/71 Scarce (Kono)
Carya ovata ( Mill.) C. Koch
1/VII/68 Scarce (TLB)
Celtis africana Burm.
5/1/75 Abundant (RLB)

582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Cerastium tomentosum L.

13/11/70 Moderate (RJN )

Clarkia amoena A. Nelson & MacBride
2/X/75 Abundant (RLB)
Cupressus lusitanica Mill.

23/V/70 Scarce Vagrant (VFE )
17/1V/71 Abundant Vagrant (VFE)
Cussonia paniculata Eckl. & Zeyh.
20/1/75 Moderate (RLB)

Dioscorea floribunda Mart. & Gal.
6/1/71 Moderate (Kono )

Echeveria gibbiflora DC

20/1/75 Abundant (RLB)

Erigeron linifolius Willd.

7/11/75 Abundant (CFS)

Eriobotrya deflexa (Hemsl.) Nakai var.

koshunensis Kaneh. & Sasaki
23/V/75 Abundant (RLB)

Eucalyptus niphophila Maiden & Blakely

6/1/71 Moderate (Kono )

Rothmannia globosa ( Hochst.) Keay
23/V/75 Scarce (RLB)

Hovenia dulcis Thunb.
5/III/77 Scarce (RLB)

Juglans californica Wats.
26/1II/68 Abundant (TLB)
16/V1/69 Abundant (TLB )

Juniperus scopulorum “Moffeti’
2/V/70 Abundant (DJV) (VFE)

Pinus halepensis Mill.
10/X/68 Moderate (VFE)

Pittosporum bicolor Hook.
14/1/71 Moderate (Kono)

Podocarpus faleatus (Thunb.) R. Br.
16/VI/70 Scarce (VFE)

P. macrophyllus (Thunb. ) D. Don
22/VI/70 Moderate (VFE)

Portulaca oleracea L.
19/1X/74 Moderate (CFS)

Pterocarya fraxinifolia (Lam.) Spach
5/1V/68 Scarce (TLB)

3/1X/70 Scarce (TLB)

P. rehderiana C. K. Schneider
30/11I/68 Moderate (TLB)
27/V1/70 Scarce (TLB)

P. rhoifolia Sieb. & Zucc.
16/VII/70 Scarce (TLB)
22/1I1/70 Moderate (TLB)

Pyracomeles vilmorini Rehder
5/1/71 Abundant (Kono)
Salix matsudana “‘Umbraculifera’
6/1/71 Moderate (Kono)
Senecio alpinus L.
5/1/71 Abundant (Kono)
Stranvaesia nussia Decne.
11/VI/69 Abundant (W-B)
Teucrium scorodonia L.
23/I1II/70 Scarce (RJN)
Aphis (Cerosipha) sp.
Eucalyptus delegatus R. T. Baker
29/1V/75 Abundant (RLB)
Viburnum odoratissimum Ker.-Gawl.
17/1/75 Abundant (RLB)
Aphis (Zyxaphis) sp.
Euphorbia supina Rafin
30/1X/75 Scarce (RLB)
Appendiseta robinea (Gillette )
Robinia pseudoacacia L.
25/VII/69 Scarce (AKG)
Aulacorthum solani (Kaltenbach)
Aesculus pavia L.
11/III/70 Scarce (AKG)
Aloysia triphylla (L.’Hér.) Britt.
8/XII/69 Scarce (RJN)
28/1V/70 Moderate (RJN)
Argania spinosa (L.) Skeels
6/VI/69 Scarce (AKG)
Asparagus setaceus (Kunth) Jessop
2/1V/69 Scarce (AKG)
Cassia atomaria L.
17/11/70 Abundant (W-B)
C. bicapsularis L.
17/11/70 Abundant (AKG)
Cerastium tomentosum L.
13/V/69 Scarce (RJN)
Clerodendrum nutans Wall.
10/XI/70 Scarce (RJN)

Coreopsis lanceolata L.
28/1V/69 Moderate (AKG!

VOLUME 80, NUMBER 4 583

Dianthus ‘China Doll’ V. wittrockiana Gams.

12/11/70 Moderate (RJN ) 13/II/70 Scarce (RJN)

D. deltoides L. Aulacorthum (Neomyzus) circumflexum
8/1IV/69 Abundant (RJN) (Buckton )

Dieffenbachia picta Schott. Artemisia dracunculus L.

3/VI/69 Moderate (W-B) 25/11/70 Moderate (AKG)

Duranta repens L.

Aulacorthum sp.
5/III/70 Moderate (RJN)

Cedrus deodara (D. Don) G. Don

D. repens ‘Alba’ 29/1V/69 Moderate Vagrant (VFE)
ae ee ae Brachycaudus (Acaudus) cardui (L.)
loistenostachyal Tod! Chrysanthemum maximum Ramond
5/I1I/70 Abundant (RJN) Ta) abun car)
Hagenia abyssinica J. F. Gmel. Brachycaudus helichrysi (Kaltenbach )
6/IV/69 Moderate (RJN) Achilla taygetea Boiss & Heldr.
16/VI/70 Abundant (RJN) 12/11/70 Abundant (W-B)
Hymenanthera dentata R. Br. Athanasia parviflora L.

13/V/70 Moderate (RJN) 5/V/70 Abundant (AKG)
Nephrolepsis exaltata Schott Brassaia actinophylla Endl.
23/I1I/75 Scarce (AGR) 1/VIII/69 Abundant (W-B)
Phlomis caucasica K. H. Rechinger Chrysanthemum balsamita L.
8/1V/69 Abundant (AKG) 21/11/70 Scarce (AKG)

Pieris taiwanensis Hayata Cordia cylindristachya Roem. & Schult.

4/1V/69 Moderate (AKG) 4/1/71 Scarce (Kono)
Polygonum chinense L.
16/IV/69 Moderate (AKG)
Rhamnus frangula L.
15/1/69 Moderate (W-B)
Rudbeckia hirta L.

a Brevicoryne brassicae (L.)
IX/69 N W-
ee) Actinidia chinensis Planch.

26/III/70 Scarce (AKG)

Brassica campestris Oed.

10/11/70 Abundant (AKG)

B. rapa L.
23/V/70 Abundant (W-B)

Raphanus sativus L.

Rhus sylvestris Sieb. & Zucc.
14/VIII/68 Moderate (W-B)

Senecio mikanioides Otto
21/1V/69 Abundant (W-B)

Rumex crispus L.

12/1II/69 Moderate (W-B)
Salvia microphylla HBK
26/111/70 Abundant (RJN)
Santolina virens Mill.
17/1V/69 Scarce (W-B)

Solanum dulcamara L.

24/XI/Abundant (W-B) SUSE es puoi (2)
Spathodea campanulata Beauv. Calaphis betulella Walsh
31/111/69 Abundant (W-B) Betula papyrifera Marsh.
iy Leen 2/V/67 Moderate (FWQ)
5/V/69 Moderate (AKG) Calaphis flava Mordvilko
Teucrium scorodonia L. Achillea tomentosa L.
23/11/70 Scarce (RJN) 18/IV/71 Abundant (VFE)
Viola cornuta L. Betula andrewsii A. Nelson

3/V1/69 Scarce (RJN) 27/V1/70 Moderate (AKG)

584 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

B. davurica Pall.

18/V/76 Abundant (RLB)
4/X/76 Scarce (RLB)
5/III/77 Scarce (RLB)

B. forrestii Hand-Mazz.
17/V/76 Scarce (RLB)

B. lutea Michx.

8/VIII/67 Scarce (FWQ)

B. maximowicziana Regel.
18/V/76 Scarce (RLB)

B. nigra L.

19/V/76 Scarce (RLB)

B. papyrifera Marsh.
2/V/67 Moderate (FWQ)
5/VI1/69 Moderate (VFE)
24/VI/70 Abundant (VFE)
19/V/76 Moderate (RLB )
B. pendula Roth

15/VII/69 Moderate (VFE)
B. pendula ‘Dalecarlica’
3/VI/69 Moderate (VFE)
B. pendula ‘Fastigata’
5/VI1/70 Abundant (VFE)
B. pendula ‘Tristis’
10/VI/69 Abundant (VFE)
2/VIII/70 Abundant (VFE)
19/V/76 Scarce (RLB)

B. platyphylla Sukachev
17/V/76 Scarce (RLB)
5/III/77 Scarce (RLB)
B. turkestanica Litw.
19/V/76 Scarce (RLB)
B. pubescens J. F. Ehrh.
11/V1I/69 Abundant (VFE)
4/VII/70 Abundant (VFE)
21/1V/71 Moderate (VFE)
Calaphis sp.
Betula jaquemontii Spach.
28/VI/70 Moderate (AKG)
Callipterinella callipterus Hartig
Betula pendula Roth
17/V/76 Moderate (RLB)
7/X/76 Abundant (RLB)
B. pendula ‘Dalecarlica’
29/X/70 Abundant (VFE)
Capitophorus elaeagni (del Guercio )
Dianthus ‘China Doll’
12/11/70 Moderate (RJN)

Capitophorus sp.

Salix lasiolepis Benth.
11/XII/69 Abundant (AKG)
Teucrium flavum L.
2/1V/69 Scarce (RJN)

Cavariella aegopodii (Scopoli)

Brassaia actinophylla Endl.
1/VIII/69 Abundant (W-B)
Carum carvi L.

25/V1/70 Abundant (W-B)
Euonymus japonica Thunb.
2/1V/70 Scarce (AKG)

Cavariella pustula Essig

Salix matsudana ‘Umbraculifera’
6/1/71 Moderate (Kono)

Chaetosiphon fragaefolii (Cockerell)

Baeckea camphorata R. Br.
16/V/70 Moderate (W-B)
Cerastium tomentosum L.
23/X/69 Abundant (RJN)
Rosa banksiae ‘Lutea’
5/V/69 Moderate (MBS )
R. centifolia L.

20/V/68 Scarce (MBS)

R. ‘Manon Cochet’
9/III/76 Abundant (MBS)
R. ‘Newport Fairy’
9/III/76 Moderate (MBS)
R. picoscarpa A. Gray
23/11/71 Abundant (MBS)
9/III/76 Abundant (MBS)

R. ‘Wind Chimes’
9/III/76 Abundant (MBS)

Chromaphis juglandicola (Kaltenbach )

Pterocarya rhoifolia Sieb. & Zucc.
20/X/69 Scarce (TLB)

Cinara arizonica ( Wilson)

Pinus coulteri D. Don
24/VII/69 Abundant (VFE)
P. glabra Walt.

24/1V/71 Scarce (VFE)

P. wallichiana A. B. Jack.
22/1V/71 Abundant (DJV)

Cinara californica Hottes and Essig

Pinus douglasiana Martinez
4/XI/69 Abundant (VFE)

VOLUME 80, NUMBER 4

P. duragnensis Roezl.

22/1V/71 Moderate (VFE)

P. glabra Walt

11/XI/69 Abundant (DJV) (VFE)
P. lumholzii Robinson & Fernald
7/X1/69 Abundant (DJV) (VFE)
P. nigra Arnold

23/VII/69 Scarce (DJV) (VFE)
P. occidentalis Sw.

24/1V/71 Abundant (DJV)

P. oocarpa Schiede

29/VII/68 Abundant (DJV)
31/X/69 Abundant (DJV)

P. patula Schlechtend. & Cham.
5/XI/69 Abundant (VFE)

P. roxburghii Sarg.

26/X1I/69 Abundant (VFE)
5/11/70 Abundant (VFE) (DJV)

Cinara fresai E. E. Blanchard

Cupressus macnabiana A. Murr.
19/V1/68 Scarce (VFE)

C. pygmaea (Lemm.) Sarg.
1/IV/70 Abundant (VFE)

C. sempervirens ‘Stricta’
5/VI/71 Scarce (VFE)
Juniperus chinensis “Armstrongi’
19/V/69 Scarce (VFE)

J. chinensis ‘Blaauw’

19/V/69 Scarce (VFE)
2/V/70 Scarce (VFE)
27/V/71 Abundant (VFE )

J. chinensis ‘Columnaris Glauca’
4/V/70 Scarce (VFE)

J. chinensis “Hetzii’
1/V/70 Abundant (VFE)
26/V/71 Abundant (VFE)
J. chinensis “Keteleeri’
27/V/71 Abundant (VFE )
J. chinensis ‘Pfitzeriana’
6/V/69 Abundant (VFE )
24/11/70 Abundant (VFE )

J. chinensis ‘Pfitzeriana Aurea’
19/V/69 Moderate (VFE )

J. chinensis “Robusta Green’
26/III/70 Moderate (VFE )

J. chinensis “Sea Green’
26/V/71 Abundant (VFE)

585

J. chinensis “Winter Green’
14/III/70 Moderate (VFE)
J. chinensis procumbens “Aureo-variegata’
26/V/71 Scarce (VFE)

J. communis “Prostrata’
24/V/71 Moderate (VFE)
J. ‘Gold Coast’

24/1II/69 Abundant (VFE)
J. horizontalis ‘Bar Harbor
19/XI/68 Scarce (VFE)

J. horizontalis ‘Hughes’
1/V/70 Abundant (VFE)
27/V/71 Moderate (VFE)
J. horizontalis ‘Plumosa Compacta’
1/V/70 Abundant (VFE)

J. monosperma Sargent
27/V/71 Abundant (VFE)
J. occidentalis ‘Sierra Silver
13/11I/70 Moderate (VFE)
10/IV/71 Abundant (VFE)
J. sabina ‘Arcadia’
13/11/70 Abundant (VFE)
31/V/71 Moderate (VFE)
J. sabina ‘Scandia’
24/11/69 Moderate (VFE)
14/11/70 Abundant (VFE)
2/V/70 Moderate (VFE)
7/1V/71 Scarce (VFE)
27/V/71 Scarce (VFE)

J. scopulorum ‘Erecta Glauca’
31/11/70 Abundant (VFE)
10/IV/71 Abundant (VFE)
J. scopulorum “Gray Gleam’
2/V/70 Abundant (VFE)
31/V/71 Moderate (VFE)

J. scopulorum “Lakewood Globe’
19/V/69 Scarce (VFE)

J. scopulorum ‘Rependens’
26/V/71 Abundant (VFE)

J. scopulorum ‘Staver’
13/11/70 Abundant (VFE)

J. scopulorum “Table Top Blue’
19/XI/68 Moderate (VFE)
19/V/69 Moderate (VFE)
1/V/70 Scarce (VFE)
27/V/71 Abundant (VFE)

J. squamata “Parsoni’
26/V/71 Scarce (VFE)

586 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

J. virginiana ‘Cupressifolia’
13/11/70 Abundant (VFE)
31/11/70 Scarce (VFE)
10/IV/71 Abundant (VFE)
J. virginiana “Manhattan Blue’
19/V/69 Moderate (VFE)
2/V/70 Abundant (VFE)
31/V/71 Abundant (VFE )
J. virginiana ‘Silver Spreader’
14/III/70 Abundant (VFE)
10/1V/71 Moderate (VFE)
Pinus coulteri G. Don
1/IV/70 Abundant (VFE)

Cinara juniperi (de Geer)

Calocedrus decurrens ( Torr.) Florin
24/1V/71 Abundant (VFE)
Cupressus pygmaea Sarg.

1/IV/70 Scarce (VFE)

V. virginiana Burki

26/1IV/71 Scarce (VFE)

Cinara pilicornis (Hartig )
Cryptomeria japonica “Araucarioides’
22/1V/71 Scarce (VFE)

Picea pungens Engelm.

24/1V/71 Abundant (VFE)
Cinara ponderosae (Williams )

Pinus canariensis Sweet

31/X/69 Abundant (VFE)

27/11I/70 Moderate ( VFE )

P. douglasiana Martinez

4/X1/69 Abundant (VFE)

17/1/75 Abundant (RLB)

P. greggii Engelm.

20/1/75 Abundant (RLB)

P. michoacana Martinez

20/VIII/68 Moderate (VFE) (DJV)

3/11/69 Moderate (VFE) (DJV)

P. nigra Arnold

23/X1/69 Scarce (VFE) (DJV)

26/1V/71 Scarce (VFE) (DJV)

P. occidentalis Sw.

24/1V/71 Abundant (DJV)

P. quadrifolia Parl.

18/V/71 Moderate (VFE) (DJV)

25/V/71 Abundant (VFE) (DJV)
Cinara sibiricae (Gillette and Palmer )

Juniperus scopulorum ‘Moffeti’
2/V/70 Abundant (VFE) (DJV)

Cinara tujafilina (del Guercio )

Callitris columellaris F. J. Muell.
13/1/70 Scarce (VFE)

C. rhomboidea R. Br.
29/1V/68 Scarce (VFE)
31/1/71 Moderate (VFE)
4/11/69 Abundant (VFE)
13/1/70 Abundant (VFE)
1/11/71 Abundant (VFE)

C. drummondii Benth.

11/III/69 Moderate (VFE)

27/VIII/68 & 13/1/70 Abundant (VFE)
C. gracilis R. T. Baker

11/III/69 Scarce (VFE)

15/1/70 Abundant (VFE)

1/II/71 Scarce (VFE)

C. morrisonii R. T. Baker

30/IV/68 Scarce (VFE)

28/VIII/68 Abundant (VFE) (RLB)
11/111/69 Abundant (VFE) (RLB)
13/1/70 Moderate (VFE)

1/II/71 Abundant (VFE) (RLB)
1/X/75 Abundant (VFE) (RLB)

C. murrayensis Miq.

13/1/70 Abundant (VFE)
1/II/71 Abundant (VFE)
17/1/75 Moderate (RLB)
C. propinqua R. Br.
31/XII/69 Abundant (VFE)

Chamaecyparis lawsoniana Parl.
4/VII/69 Moderate (VFE) (DJV)
1/IV/70 Scarce (VFE)

31/11/71 Abundant (VFE)
10/III/70 Abundant (VFE)

Cupressus pygmaea Sarg.
1/IV/70 Abundant (VFE)

Juniperus “Gold Coast’
24/III/69 Abundant (VFE)

Pinus coulteri G. Don
1/IV/70 Abundant (VFE)

Platycladus orientalis (L.) Franco
4/11/69 Abundant (VFE)
14/VI/69 Moderate (VFE)
30/1/70 Abundant (VFE)
1/VII/70 Abundant (VFE)
23/11/71 Abundant (VFE)

Widdringtonia juniperoides Endl
20/1/75 Abundant (RLB)

VOLUME 80, NUMBER 4

Cinara wahluca Hottes group

Cupressus glabra Sudw.
5/X/70 Abundant (VFE)

Cinara sp.

Actinostrobus pyramidalis Miq.
25/I1V/68 Moderate (VFE)
Cedrus deodara Loud.
29/IV/69 Moderate (VFE)
Cupressus guadalupensis S. Wats.
31/III/70 Moderate (VFE)
Juniperus scopulorum “Lakewood Globe’
2/V/70 Abundant (VFE)
3/V/71 Abundant (VFE )
Pinus canariensis Sweet
26/1V/71 Abundant (VFE)

P. caribaea Morelet

2/1V/70 Abundant (VFE)

P. cembroides Zucc.

22/V/71 Moderate (VFE)

P. douglasiana Roezl.
16/XII/73 Moderate (VFE)

P. durangensis Roezl.
22/1V/71 Moderate (VFE)

P. greggii Engelm.

18/11/69 Scarce (VFE)

P. muricata D. Don

2/1V/70 Scarce (VFE) (DJV)

Cinara sp. (Delta)

Pinus douglasiana Martinez
4/X1/69 Abundant (DJV)

P. glabra Walt.

11/XI/69 Abundant (DJV)

P. greggii Engelm.

12/1I/71 Abundant (DJV)

P. lomholzii Robinson & Fernald
7/X1/69 Abundant (VFE) (DJV)
P. nigra Arnold

23/VII/69 Scarce (VFE) (DJV)
P. occidentalis Sw.

21/1V/71 Abundant (DJV)

P. oocarpa Schiede

29/VII/68 Abundant (DJV)
31/X/69 Abundant (DJV)

P. patula Schlechtend. & Cham.
5/XI/69 Abundant (VFE)

P. roxburghii Sarg.

26/X1/69 Abundant ( VFE)

Dactynotus ambrosiae (Thomas )
Baeckea camphorata R. Br.
16/V/70 Moderate (W-B)
Brickellia guatemalensis
B. L. Robinson
25/1X/70 Abundant (W-B)
Doronicum pardalianches L.
20/V/76 Abundant (RLB)
Santolina virens Mill.
17/1IV/69 Scarce (W-B)

Dactynotus sp.

587

Maclura pomifera ( Raf.) C. K. Schneid.

19/VII/70 Moderate (AKG)

Olearia pimeleoides Benth.
17/11/69 Abundant (AKG)

Tanacetum vulgare L.
24/XI/69 Moderate (AKG)

Drepanaphis sp.

Acer saccharinum ‘Laciniatum’
16/X/70 Abundant (W-B)

Dysaphis tulipae (Boyer de Fonscolombe )

Buphthalum salicifolium L.
5/1/71 Scarce (Kono)

Dysaphis sp.

Belamcanda chinensis DC.
23/1/69 Abundant (AKG)

Osteospermum fruticosum (L.) Norlindh.

19/III/69 Moderate (AKG)

Sisyrinchium bellum S. Wats.
21/X1/69 Abundant (AKG)
Eriosoma lanigerum (Hausmann )

Malus domestca Baumg.
29/XI/73 Moderate (CFS)
Malus sp.
2/X/75 Abundant (RLB)
Essigella californica (Essig )
Cedrus deodara (D. Don) G. Don
29/1V/69 Moderate (VFE)
Pinus canariensis Sweet
21/V1I/68 Moderate (VFE)
P. cooperi C. E. Blanco
4/XI/69 Moderate ( VFE)
P. coulteri G. Don
1/VIII/68 Moderate (VFE)
7/X1/69 Abundant (VFE)
7/X/70 Abundant (VFE)

588 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

P. durangensis Roezl.
1/IV/70 Moderate (VFE )
P. douglasiana Martinez
16/XI/73 Moderate (VFE)
17/1/75 Abundant (RLB)
P. halepensis Mill.
5/X1/69 Moderate (VFE)
P. michoacana Martinez
16/X1/73 Abundant (VFE)
1/X/75 Moderate (RLB )
P. wallichiana A. B. Jacks.
31/VII/68 Scarce (VFE)
5/X1/69 Abundant (VFE)
Pinus sp.
3/XI/69 Moderate (VFE)
Essigella essigi Hottes
Pinus canariensis Sweet
24/11/69 Abundant (VFE)
27/111/70 Moderate (VFE)
9/XI/70 Moderate (VFE)

Pinus sp.
3/X1/69 Moderate (VFE)

Essigella fusca Gillette and Palmer

Callitrus drummondii Benth.
31/1/71 Moderate (VFE)
Pinus attenuata Lemm.
4/V/68 Abundant (VFE)

P. coulteri G. Don
24/VII/69 Abundant (VFE)
26/IV/71 Abundant (VFE)

Essigella pini Wilson
Pinus cembroides Zucc.
21/XI/69 Moderate (VFE)

P. cooperi C. E. Blanco
1/1V/70 Scarce (VFE)
24/1V/71 Scarce (VFE)

P. wallichiana A. B. Jacks.
2/1V/70 Scarce (VFE)
7/X/70 Moderate (VFE)

Pinus sp.

27/11/69 Abundant ( VFE )
3/XI/69 Moderate (VFE )
31/III/70 Moderate (VFE)
12/1II/71 Abundant (VFE)

Essigella sp.

Cupressus lusitanica Hill.
31/X/69 Scarce (VFE)

Picea gemmata Rehder & Wilson
4/X1/69 Scarce (VFE)

Pinus caribaea Morelet
22/VII/69 Abundant (VFE)
5/XI1/69 Abundant (VFE)
2/1V/70 Abundant (VFE)
9/X/70 Abundant (VFE)
10/IV/71 Scarce (VFE)

P. douglasiana Martinez
4/X1/69 Abundant (VFE)

P. durangensis Roezl.

92/1V/71 Moderate (VFE )

P. greggii Engelm.

12/11/71 Abundant (DJV) (VFE)
P. lumholzii Robinson & Fernald
7/X1/69 Abundant (DJV) (VFE)

P. michoacana Martinez
20/VIII/68 Moderate (VFE) (DJV)
3/11/69 Moderate (VFE) (DJV)

P. muricata D. Don

2/1V/70 Scarce (VFE) (DJV)
P. nigra Arnold

3/X1/69 Scarce (VFE) (DJV)
26/1V/71 Scarce (VFE) (DJV)
P. occidentalis Sw.

24/1V/71 Abundant (DJV)
P. oocarpa Schiede

31/X/69 Abundant (DJV)

P. patula Schlechtend. & Cham.
5/X1I/69 Abundant (VFE)

P. pinea L.

13/X/69 Moderate (VFE)
5/XI/69 Moderate (VFE )
22/1V/71 Moderate (VFE)

P. roxburghii Sarg.

26/X1/69 Abundant (VFE)
5/11/70 Abundant (VFE )

P. sylvestris L.

4/X1/69 Moderate (VFE )
31/III/70 Moderate (VFE)

Eucallipterus tiliae (1. )

Centranthus ruber (L.) DC.
23/11/70 Abundant ( W-B )

Euceraphis betulae (Koch)

Betula pendula Roth
17/V/76 Moderate (RLB )
19/V/76 Scarce (RLB)

VOLUME 80, NUMBER 4

Euceraphis gilletti Davidson
Alnus koehnei Callier
4/VII/70 Scarce (AKG)
A. rhombifolia Nutt.
5/III/76 Moderate (RLB)
17/V/76 Moderate (RLB )
Eulachnus rileyi (Williams )

Pinus canariensis Sweet
21/V1/68 Moderate (VFE)
24/11/69 Abundant (VFE)
9/XI/70 Moderate (VFE )

P. caribaea Morelet

5/XI/69 Abundant ( VFE)
2/1V/70 Abundant (VFE)
9/X/70 Abundant (VFE)
10/IV/71 Scarce (VFE)

P. durangensis Roezl.
22/1V/71 Moderate (VFE)
P. glabra Walt.

11/XI/69 Abundant (DJV) (VFE)
P. greggii Engelm.
31/VII/68 Moderate (VFE)
18/11/69 Scarce (VFE)
11/XI/69 Abundant (VFE )
1/IV/70 Scarce (VFE)
7/X/70 Moderate (VFE)
24/1V/71 Abundant (VFE )
16/XI/73 Abundant (VFE)
20/1/75 Abundant (RLB)

P. halepensis Mill.
2/1V/70 Scarce (VFE)
10/IV/71 Scarce (VFE)

P. halepensis brutia (Ten.) A. Henry
7/11/69 Scarce (VFE)

P. michoacana Martinez
16/XI/73 Abundant (VFE)
17/1/75 Abundant (RLB)
1/X/75 Moderate (RLB)

P. pinea L.

29/VIII/69 Moderate (VFE)
13/X/69 Moderate (VFE)
5/XI/69 Moderate (VFE)
7/X/70 Abundant (VFE)
12/11/71 Abundant (VFE)
22/1V/71 Moderate (VFE)
P. sylvestris L.

3/VIII/68 Moderate (VFE)
22,/V11/69 Moderate (VFE)

4/X1/69 Moderate (VFE)
31/III/70 Moderate (VFE)
6/X/70 Scarce (VFE)

P. wallichiana A. B. Jacks.
5/XI1/69 Abundant (VFE)
2/1V/70 Scarce (VFE)
Pinus sp.

27/11/69 Abundant ( VFE)
12/11/71 Abundant (VFE)
Thuja occidentalis “Cristata’
11/XI/69 Scarce (VFE)

Euthoracaphis umbellulariae Essig
Cinnamomum brevifolium Miq.

6/11/75 Abundant (RLB)
C. daphnoides Seib. & Zucc.
31/1/69 Abundant (VFE)

Forda formicaria von Heyden
Bromus carinatus Hook. & Arn.

21/1/70 Abundant (CFS)
9/X/70 Abundant (CFS)
1/11/73 Moderate (CFS )

Forda marginata Koch

Agrotis alba L.
3/X/72 Abundant (CFS )

Aristida adscensionsis L.
10/IV/73 Abundant (CFS )

Bromus carinatus Hook. & Arn.

29/1/69 Abundant (CFS )
21/1/70 Abundant (CFS )
13/1X/72 Abundant (CFS )
1/11/73 Moderate (CFS )
26/II1/73 Abundant (CFS )
28/I1I/73 Abundant (CFS )
16/1V/73 Abundant (CFS )
8/V/73 Abundant (CFS )
17/V/73 Abundant (CFS )
11/VI/73 Moderate (CFS )
23/V/75 Abundant (RLB)
30/1X/75 Abundant (RLB)
2/X/75 Abundant (RLB)
17/III/77 Scarce (RLB)

B. catharticus Vahl.
8/V/73 Abundant (CFS )
10/V/73 Abundant (CFS )
18/V/73 Abundant (CFS )
5/VI/73 Abundant (CFS )

B. racemosus L.

6/VI/73 Abundant (CFS )

589

590 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

B. rubens Cav.
12/V/73 Abundant (CFS )
18/V/73 Abundant (CFS )
Cynodon dactylon Pers.
12/V/73 Abundant (CFS)
Echinochloa crus-galli Beauv.
2/1X/72 Abundant (CFS)
8/V/73 Scarce (CFS)
5/VI/73 Abundant (CFS )
Festuca megalura Nutt.
28/III/73 Moderate (CFS )
28/1V/73 Scarce (CFS)
Hordeum stebbinsii L.
10/V/73 Abundant (CFS )
Paspalum dilatatum Poir.
12/V/73 Moderate (CFS)
Poa annua L.
26/III/73 Moderate (CFS )
8/V/73 Moderate (CFS )
12/V/73 Abundant (CFS)
2/V/74 Abundant (CFS )
23/V/75 Abundant (RLB)
Schismus barbatus (L.) Thell.
10/IV/73 Abundant (CFS)
Setaria glauca (L.) Beauv.
14/V/73 Scarce (CFS )
Triticum aestivum L.
5/VI/73 Scarce (CFS)
Hyadaphis erysimi ( Kaltenbach )
Capsella bursa-pastoris Medi
17/11/70 Abundant (AKG)
Matthiola incana (L.) R. Br.
8/1II/77 Abundant (RLB)
Hyadaphis foeniculi ( Passerini )
Oncoba routledgei T. Sprague
4/1/71 Scarce (Kono)
Hyalomyzus monardae ( Davis )
Rumex crispus L.
12/III/69 Moderate (W-B)
Hyperomyzus lactucae (L.)
Picris echioides L.
10/V/69 Abundant (AKG)
Hysteroneura setariae (Thomas)
Bougainvillaea spectabilis Willd.
28/X/70 Scarce (AKG)

Macchiatiella rhami ( Boyer de
Fonscolombe )

Aloysia triphylla (L’Hér. ) Britt.
23/1X/70 Moderate (RJN )
15/XII/71 Moderate (RJN )
Clerodendrum bungei Steud.
3/1V/69 Abundant (RJN )
Dianthus deltoides L.
8/1V/69 Abundant (RJN)
Hagenia abyssinica J. F. Gmel.
6/1V/69 Moderate (RJN )
Hymenanthera dentata R. Br.
18/V/69 Scarce (RJN )
Sagina apetala L.

6/VI/69 Scarce (RJN )

Viola cornuta L.

3/V1/69 Scarce (RJN)

V. odorata L.

21/XI1/69 Scarce (RJN)

V. wittrockiana Gams.
13/VI/69 Scarce (RJN )
Viola sp.

9/XI/70 Moderate (RJN)

Macrosiphoniella sanborni ( Gillette )

Salvia microphylla HBK
26/11/70 Abundant (RJN)

Macrosiphoniella sp.

Artemisia californica Less.
21/11/70 Searce (AKG)

Macrosiphum (Sitobion) avenae ( Fabricius )

Ampelodesmos mauritanicus ( Poir. )
T. Durant & Schinz.

18/1V/70 Abundant (AKG)

Bromus carinatus Hook. & Arn.

3/11/70 Moderate (AKG)

3/1V/71 Abundant (CFS )

Oryzopsis miliacea (L.) Benth.

30/1IV/69 Moderate (AKG)

Macrosiphum californicum (Clarke )

Salix taxifolia HBK
6/XII/69 Abundant (W-B)

Macrosiphum euphorbiae (Thomas )

Abutilon indicum Sweet
31/11I/69 Moderate (AKG)
16/VI/70 Abundant (W-B)
Acacia arabica Willd.
6/V1/69 Moderate (W-B )

Acer tartaricum L.
11/III/70 Moderate (AKG)

VOLUME 80, NUMBER 4

Agrimonia eupatoria L.
23/11/70 Scarce (AKG)
Albizia thorelli Pierre
24/III/70 Scarce (W-B)
Alnus tenuifolia Nutt.
18/V/76 Scarce (RLB )
Aloe ciliaris Haw.

18/II/70 Abundant (AKG)
A. immaculata Pillans
23/1/70 Moderate (AKG)

A. wickensii Pole-Evans
27/XIII/69 Moderate (AKG)

Anemopaegma chamberlaynii (Sims )

Bur. & K. Schum.
19/VI/70 Moderate (W-B)
Anigozanthos flavidus Redouté
28/XII/70 Scarce (AKG)
Anoda triangularis DC.
25/I11/70 Abundant (AKG)
Antigonon leptopus Hook. & Arn.
10/VI/70 Abundant (W-B)
Baccharis glutinosa Pers.
16/11/70 Abundant (W-B)
Berberis triacanthophora Fedde.
27/1V/70 Scarce (AKG)
Brassaia actinophylla Endl.
1/VIII/69 Abundant (W-B)

Bromus carinatus Hook. & Arn.
3/1V/71 Abundant (CFS)

Caesalpinia japonica Sieb. & Zucc.

12/I11/70 Abundant (AKG)
Caragana aurantiaca Koehne
12/III/70 Moderate (AKG)
Cassia atomaria L.

17/111/70 Abundant (W-B)
C. bicapsularis L.

17/11/70 Abundant (AKG)

C. mimosoides L.

31/XII/70 Abundant (W-B)
C. nemophila coriacea Symon
15/1/71 Moderate (Kono)
Centranthus ruber DC.
23/11/70 Abundant (W-B)
Chenopodium album Bosc.
17/11/70 Scarce (AKG)

Chorisia insignis HBK
19/II/70 Scarce (AKG)

Cistus incanus L.

6/1/71 Scarce (Kono )
Clematis orientalis L.
19/VI/70 Moderate ( W-B )

Clerodendrum nutans Wall.
4/1V/69 Scarce (RJN )

Combretum microphyllum Klotzsch.

13/VI/70 Moderate (AKG)

Convolvulus mauritanicus Boiss.
5/V/70 Scarce (AKG)

Cordia escalyculatha Vell.
21/11/70 Moderate (AKG)

Coreopsis lanceolata L.
28/1V/69 Moderate (AKG)

Crataegus lavallei Herincg
95/1V/70 Abundant (W-B)

Croton megalocarpus Hutchinson
31/1II/69 Moderate (AKG)

Dianthus ‘China Doll’
12/11/70 Moderate (RJN )

D. deltoides L.
8/1IV/69 Abundant (RJN)

Dombeya buettneri K. Schum.
4/1/71 Moderate (Kono)

Duranta repens L.
5/11/70 Moderate (RJN )

D. repens “Alba’
5/11/70 Abundant (RJN)

D. stenostachya Tod.
18/XII/68 Moderate (RJN )
29/XI1/69 Scarce (RJN)
5/11/70 Abundant (RJN)

Echium wildpretii H. Pearson
19/V/69 Scarce Vagrant (VFE)
Eryngium planum L.

5/1/71 Scarce (Kono )

Eucalyptus cladocalyx F. Muell.
21/1/71 Moderate (Kono)

Euonymus japonica Thunb.
27/1V/70 Scarce (AKG)
Ficus iteophylla Miq.
19/III/70 Moderate (W-B)
Ficus sp.

6/1/71 Scarce (Kono )

Filipendula hexapetala Gilib.
5/1/71 Abundant (Kono)

592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Forsythia ‘Lynwood Gold’ S. officinalis L.

6/1/71 Moderate (Kono) 16/VII/69 Scarce (RJN )

Hagenia abyssinica J. F. Gmel. Schefflera arboricola Hayata
6/1V/69 Moderate (RJN) 27/III/69 Abundant (W-B )
26/1/70 Abundant (RJN ) Senecio adonidifolius Loisel.
16/V1/70 Abundant (RJN) 5/1/71 Moderate (Kono )
Hypericum elegans Steph. Spathodea campanulata Beauv.
5/1/71 Abundant (Kono) 31/11/69 Abundant (W-B )

Iris spuria L. Stereospermum kunthianum Cham.
5/1/71 Abundant (Kono) 15/1X/69 Abundant (W-B)
Lampranthus sp. Teucrium scorodonia L.

14/1/71 Scarce (Kono) 23/11/70 Scarce (RJN)

Linum perenne Lam. Tulipa gesneriana L.

1/1/71 Scarce (Kono) 1/IV/69 Abundant (W-B)
Nandina domestica “‘Compacta Nana’ Viola wittrockiana Gams.
16/IV/69 Moderate (AKG) 13/II/70 Scarce (RJN )
Osteospermum ecklonis (DC.) Norlindh. Watsonia sp.

9/V/69 Scarce (AKG) 17/V/69 Scarce (AKG)

Phlomis caucasica K. H. Rechinger Withania somnifera Dunal.
8/1V/69 Abundant (AKG) 5/1/71 Moderate (Kono)
Potentilla rupestris L. Zinnia elegans “Wild Cherry’
5/1/71 Moderate (Kono) 23/X/69 Scarce (AKG)
Raphanus sativus L. Macrosiphum (Sitobion) fragariae
8/V/69 Abundant (W-B) ( Walker )

Rhus sylvestris Sieb. & Zucc. Bromus carinatus Hook. & Arn.
16/III/68 Abundant (AKG) 3/IV/71 Abundant (CFS)
14/VII1/68 Moderate (W-B ) Chamaecyparis lawsoniana

Rosa banksiae ‘Lutea’ (A. Murr.) Parl.

5/V/69 Moderate (MBS) 10/III/70 Abundant (VFE) (DJV)
R. hugonis Hemsl. Juniperus occidentalis “Sierra Silver’
17/11/71 Abundant (MBS) 10/IV/71 Abundant Vagrant (VFE)
R. ‘Margo Koster’ Macrosiphum rosae (L.)

22/V/68 Abundant (MBS ) Brunfelsia uniflora (Pohl) D. Don
Rosa sp. 17/III/70 Moderate (AKG)
29/V1/69 Abundant (MBS ) Rosa ‘American Beauty’

5/XI11/69 Abundant (MBS) 9/III/76 Moderate (MBS)
7/III/71 Abundant (MBS) R. banksiae Ait.

Rudheclkiahires aie 9/III/76 Moderate (MBS)
29/1X/69 Moderate (W-B ) R. banksiae “Lutea’

Rumex crispus L. 5/V/69 Moderate (MBS)
12/T1I/69 Moderate (W-B) R. ‘Buff Beauty’

ealisieroee ener 9/III/76 Moderate (MBS)
10/VII/68 Scarce (RJN) R. “Glorie Des Mosseux’

23/1/69 Abundant (RJN) 21/X/70 Abundant (MBS )

S. mellifera Greene R. ‘Cecil Brunner’

21/11/70 (RJN) 9/III/76 Abundant (MBS)

VOLUME 80, NUMBER 4

R. hugonis Hemsl.
17/11/71 Abundant (MBS)
R. “Margo Koster’

22/V/68 Abundant (MBS)
R. ‘La France’

9/III/76 Abundant (MBS)
R. ‘Manon Cochet’

9/I1i/76 Abundant (MBS)
R. ‘Mme Pierre Oger’
9/III/76 Moderate (MBS)
R. moschata J. Herrm.
9/III/76 Scarce (MBS)

R. moyesii Hemsl. & E. H. Wils.
9/III/76 Scarce (MBS)

R. ‘Newport Fairy’

9/11/76 Moderate (MBS)
R. pendulina L.

9/III/76 Scarce (MBS)

R. pisocarpa A. Gray
23/11/71 Abundant (MBS )
9/III/76 Abundant (MBS)
R. ‘Sombrevil’

9/TII/76 Scarce (MBS)

R. spinosissima L.
10/VII/68 Scarce (MBS )
7/1V/70 Abundant (MBS)
R. rugosa Thunb.

9/III/76 Moderate (MBS)
R. ‘Wind Chimes’

9/III/76 Abundant (MBS)
Rosa sp.

7/V/69 Scarce (MBS)
4/XII/69 Abundant (MBS)
7/I11/71 Abundant (MBS)
8/III/76 Abundant (MBS)

Macrosiphum sp.

Nephrolepis exaltata (L.) Schott.

27/1I1/75 Scarce (AGR)
Pinus halepensis Mill.
2/1V/70 Scarce (VFE)
10/1V/71 Scarce (VFE)
Polypodium aureum L.
96/I11/69 Moderate (AKG)
Potentilla detommasii Tenore
5/1/71 Moderate (Kono)

Rhamunus crocea Nutt.
20/1/71 Scarce (Kono)

593

Rosa centifolia L.

20/V/68 Scarce (MBS)
Rosa sp.

4/XII/69 Abundant (MBS)
Senecio mikanioides Otto
21/1V/69 Abundant (W-B)
Sonchus oleraceus L.
6/III/77 Abundant (RLB)
Tulipa sp.

6/III/75 Abundant (RLB)

Masonaphis azaleae (Mason)

Rhododendron sp.
3/11/70 Scarce (W-B)

Masonaphis brevitarsis Gillette and Palmer
Salvia aurea L.
2/V1I/69 (RJN)

Masonaphis lambersi MacGillivray

Rhododendron sp.
17/11I/69 Moderate (AKG)

Masonaphis morrisoni (Swain )
Actinostrobus pyramidalis Miq.
25/1V/68 Moderate (VFE)
10/11/71 Abundant ( VFE)
Araucaria heteophylla (Salisb.) Franco
27/1I1/70 Scarce (VFE)
Callitris murrayensis Miq.
30/1V/68 Scarce (VFE)
12/III/69 Abundant (VFE )
3/VI/70 Abundant (VFE)

C. propinqua R. Br.
30/V/68 Scarce (VFE)
26/V/69 Moderate (VFE)
20/1/71 Abundant (VFE)

Calocedrus decurrens ( Torr.) Florin
24/1V/71 Abundant ( VFE)

Calothamnus chrysanthereus F. Muell.

18/VI/70 Moderate (VFE)

Catalpa bignonioides Walt.

15/V/69 Scarce (VFE)

Cedrus deodara (D. Don) G. Don

29/1V/69 Moderate ( VFE )

Chamaecyparis lawsoniana (A. Murr. )
Parl.

4/VI1/69 Moderate (VFE )

C. nootkatensis (D. Don.) Spach

1/VII/69 Moderate (VFE)

594

C. pisifera “Cyanoviridis’
26/V/71 Moderate ( VFE )
Cistus incanus L.
20/1V/70 Scarce (VFE)

Cryptomeria japonica “Araucarioides’

22/1V/71 Scarce (VFE)

C. japonica “Viminalis’
22/TV/71 Moderate (VFE)
Cupressus arizonica Greene
22/VII/69 Scarce (VFE)

C. cashmeriana Royle
21/V1I/69 Moderate (VFE)
17/1IV/71 Abundant ( VFE )
C. forbesii Jeps.

13/III/69 Abundant (VFE )
18/VII/69 Scarce (VFE)
27/11/70 Abundant (VFE)
26/IV/71 Moderate ( VFE)

C. guadalupensis S. Wats.
31/III/70 Moderate (DJV)
C. lusitanica Mill.

13/TII/69 Scarce (VFE)
24/VII/69 Scarce (VFE)
2/1V/70 Moderate (VFE)
17/IV/71 Abundant ( VFE )

C. macnabiana A. Murr.
19/V1/68 Scarce (VFE)

C. pygmaea (Lemm.) Sarg.
3/III/69 Scarce (VFE)
23/XII/69 Scarce (VFE)
1/IV/70 Abundant (VFE)
5/VI/71 Scarce (VFE)

C. sempervirens L.
20/VI/69 Abundant (VFE)

Juniperus bermudiana L.
13/VII/69 Moderate ( VFE )
28/1V/71 Abundant (VFE)

J. chinensis “Blaauw’
19/V/69 Scarce (VFE )
27/V/71 Abundant (VFE)

J. chinensis ‘Columnaris Glauca’
21/V/68 Abundant (VFE)
4/V/70 Scarce (VFE)

J. chinensis ‘Fruitlandii’
19/V/69 Scarce (VFE)
27/V/71 Moderate (VFE)

J. chinensis “Hetzii Columnaris’
19/V/69 Scarce ( VFE )

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

J. chinensis ‘Hetzii’
19/V/69 Scarce (VFE)
26/V/71 Abundant (VFE)
J. chinensis ‘Japonica’
12/VI/71 Moderate ( VFE )
J. chinensis ‘Keteleeri’
19/V/69 Scarce (VFE)
27/V/71 Abundant (VFE)
J. chinensis ‘Pfitzeriana’
21/VI/68 Abundant (VFE)
6/V/69 Abundant (VFE)
19/V/69 Moderate (VFE )
1/V/70 Moderate (VFE)
26/V/71 Scarce (VFE)

J. chinensis “Robusta Green’

13/VII/69 Scarce (VFE)

26/III/70 Moderate (VFE)

28/1IV/71 Abundant ( VFE)

J. chinensis ‘Sea Green’

26/V/71 Abundant (VFE)

J. chinensis “Torulosa’

27/V/71 Moderate (VFE)

J. chinensis procumbens ‘Aureo-variegata’
26/V/71 Scarce (VFE)

J. monosperma (Engelm.) Sarg.
27/V/71 Abundant (VFE)

J. occidentalis “Sierra Silver’
10/IV/71 Abundant (VFE)

J. sabina ‘Arcadia’

31/V/71 Moderate (VFE )
J. sabina ‘Variegata’
27/V/71 Scarce (VFE)

J. scopulorum “Erecta’
22/VII/69 Scarce (VFE)
10/IV/71 Abundant (VFE)
26/V/71 Abundant (VFE)
J. scopulorum “Gray Gleam’
31/V/71 Moderate (VFE)
J. scopulorum “Lakewood Globe’
19/V/69 Scarce (VFE)
31/V/71 Abundant (VFE)

J. scopulorum ‘Reptans’
26/V/71 Abundant (VFE)

J. scopulorum ‘Steel Blue’
27/V/71 Scarce (VFE)

J. scopulorum “Table Top Blue’
19/V/69 Moderate (VFE )

VOLUME 80, NUMBER 4 595

J. squamata ‘Meyeri’ Carya ovata ( Mill.) C. Koch
19/V/69 Moderate (VFE) 28/VIII/70 Scarce (TLB)
J. virginiana “Cupressiformis’ Monellia caryella (Fitch )

19/V/69 Scarce (VFE)
11/VII/69 Moderate (VFE )
10/1V/71 Abundant ( VFE)
26/V/71 Scarce (VFE)

Carya illinoinensis (Wangenh. ) C. Koch
9/X/67 Moderate (TLB)

1/VII/68 Moderate (TLB)

8/VII/69 Moderate (TLB)

J. virginiana ‘Silver Spreader’ 21/VIIL/69 Scarce (TLB)
10/1IV/71 Moderate (VFE) 13/X/69 Scarce (TLB)
Metasequoia glyptostroboides H. 24/X/69 Scarce (TLB)

H. Hu & Cheng 5/XI1/69 Scarce (TLB )
27/1V/70 Scarce (VFE) 5/XII/69 Scarce (TLB)
Taxodium mucronatum Ten. 23/VII/70 Moderate (TLB )
23/1V/70 Scarce (VFE) 3/1X/70 Moderate (TLB)
22/V1/70 Scarce (VFE) 21/X/70 Abundant (TLB)

Thuja occidentalis “Cristata’ 17/VII/71 Scarce (TLB)
22/V1/69 Moderate (VFE) Monellia costalis (Fitch )

Masonaphis sp. Carya illinoinensis (Wangenh. ) C. Koch
Callitris quadrivalvis Vent. 1/VII/68 Moderate (TLB)
13/VI/70 Abundant (AKG) 23/VII/70 Moderate (TLB )
Chamaecyparis lawsoniana (A. Murr. ) 27/VII/70 Moderate (TLB )

Parl. Pinus canariensis Sweet
31/1II/71 Abundant (VFE) 9/XI/70 Moderate Vagrant (VFE)
Cotoneaster frigidus Wall. Monelliopsis bisetosa Richards
24/V1I/69 Abundant (AKG)

Pterocarya fraxinifolia (Lam.) Spach
Cupressus lusitanica “Pendula’ 1/XII/69 Scarce (TLB)
13/III/69 Moderate ( VFE )
31/III/70 Abundant (VFE)
17/I1I/71 Abundant (VFE) Carya illinoinensis (Wangenh.) C. Koch
9/VI/71 Moderate (TLB)

Monelliopsis californica (Essig )

Juniperus occidentalis “Sierra Silver’

Juglans californica Wats.
ma cen) te ey
undan

an /N/ TY Abundant (VEE) 16/VI/69 Abundant (TLB)

Solanum robustum H. Wend]. 21/VIII/69 Scarce (TLB)

15/III/69 Moderate (AKG) 13/X/69 Scarce (TLB)

Vitex pseudo-negundo Hand.-Mazz. 3/XII/69 Moderate (TLB )

7/1V/69 Scarce (AKG) 27/1V/70 Moderate (TLB )
Melanocallis fumipennellus (Fitch) 1/VU/70 Abundant (TLB)

27/VII/70 Moderate (TLB)
28/VIII/70 Scarce (TLB)
21/X/70 Abundant (TLB )
12/V1/71 Scarce (TLB)

Carya illinoinensis (Wangenh.) C. Koch
1/VII/68 Moderate (TLB )
16/V1I/69 Abundant (TLB)
8/VII/69 Moderate (TLB)

24/X/69 Scarce (TLB) Pterocarya fraxinifolia (Lam.) Spach
5/XI1/69 Scarce ( TLB ) 29/V/68 Scarce ( TLB )

23/VII/70 Moderate (TLB) AN 0 ence (UES)

27/VII/70 Moderate (TLB) 3/X11/69 Scarce (TLB )

3/1X/70 Moderate (TLB ) P. rehderana C. K. Schneider

21/X/70 Abundant (TLB) 1/XII/69 Scarce (TLB)

596 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Myzaphis rosarum ( Kaltenbach )

Rosa “Buff Beauty’

9/11/76 Moderate (MBS )

R. ‘La France’

9/1I1/76 Abundant (MBS)

R. “Manon Cochet’

9/1III/76 Abundant (MBS)

R. “Mme. Pierre Oger’

9/III/76 Moderate (MBS )

R. ‘Wind Chimes’

9/11/76 Abundant (MBS)

Rosa sp.

8/I1I/76 Abundant (MBS)
Myzocallis sp. possibly castanicola Baker

Quercus robur L.

9/XI/73 Abundant (VFE)
Myzocallis coryli (Goeze )

Corylus avellana L.

29/1X/75 Scarce (RLB)

C. avellana “Fuscorubra’

10/V1/69 Abundant (VFE)

30/VI/70 Abundant (AKG)

18/1V/71 Moderate (VFE)

16/XI/71 Abundant (VFE)
Myzocallis punctatus (Monell )

Ceratonia siliqua L.

18/I1I/70 Scarce (AKG)

Quercus agrifolia Née

A/NI1/75 Abundant (RLB)

Q. engelmannii Greene

17/1/75 Abundant (RLB)
Myzocallis sp. near punctatus (Monell)

Quercus agrifolia Née

26/V1/75 Moderate (VFE )

Q. leucotrichophora A. Camus

18/VI/73 Scarce (VFE)
Myzocallis walshii (Monell)

Quercus agrifolia Née

26/V1/73 Moderate (VFE)
Myzocallis sp.

Cedrus deodara (D. Don) G. Don

29/1V/69 Moderate Vagrant (VFE )

Citharexylum montevidense (Spreng )

Moldenke
18/XII/68 Scarce (RJN)

Coleonema album E. Mey.

7/V/70 Moderate Vagrant (W-B)
Duranta repens ‘Alba’

18/XII/68 Moderate (RJN )
Rothmannia globosa ( Hochst.) Keay
23/V/75 Scarce Vagrant (RLB)

Myzus ascalonicus Doncaster

Allium ascalonicum L.
26/11/70 Abundant (W-B)

Myzus certus (Walker )

Cerastium tomentosum L.
13/V/69 Scarce (RJN)
23/X/69 Abundant (RJN )
13/11/70 Moderate (RJN )
3/XI/70 Moderate (RJN )
Dianthus “China Doll’
12/11/70 Moderate (RJN )
Hymenanthera dentata R. Br.
28/V/69 Scarce (RJN )
13/V/70 Moderate (RJN )
Viola wittrockiana Gams.
13/II/70 Scarce (RJN)
Viola sp.

9/XI/70 Moderate (RJN )

Myzus ornatus Laing

Agrimonia eupatoria L.
23/11/70 Scarce (AKG)
Amygdalus bucharica Korsh.
20/1II/71 Moderate (AKG)
Bergenia ciliata (Haw. ) Sternb.
3/III/70 Scarce (W-B)
Caragana aurantiaca Koehne
12/III/70 Moderate (AKG)
Centranthus ruber DC.
23/11/70 Abundant (W-B)
Cerastium tomentosum L.
13/V/69 Scarce (RJN )
13/II/70 Moderate (RJN )
Dianthus “China Doll’
12/II/70 Moderate (RJN)
Duranta repens L.

15/V/69 Moderate (RJN )
18/II/70 Abundant (RJN )
5/III/70 Moderate (RJN )
D. repens ‘Alba’

30/XII/69 Moderate (RJN )
31/XII/70 Moderate (RJN )

VOLUME 80, NUMBER 4

D. stenostachya Tod.
18/XII/68 Moderate (RJN)
15/III/69 Moderate (RJN)
30/XII/69 Abundant (RJN)
Echium wildpretii H. Pearson
19/V/69 Scarce Vagrant ( VFE)
Ficus iteophylla Miq.

19/TII/70 Moderate ( W-B )
Nephrolepis exaltata Schott.
27/I11I/75 Scarce (AKG)
Rosmarinus officinalis “Prostratus’
13/III/69 Abundant (AKG)
Salvia aurea L.

2/V1/69 Moderate (RJN)

S. greggii A. Gray

10/VII/68 Scarce (RJN )
23/1/69 Abundant (RJN)

S. mellifera Greene
21/11/70 Abundant (RJN)
S. microphylla HBK
21/IV/69 Scarce (RJN)
Sambucus racemosa L.
20/1II/69 Moderate (AKG)
Sarcococca saligna (D. Don) Muell.
15/1/69 Scarce (AKG)
Sorbaria tomentosa (Lindl.) Rehd.
15/1/69 Scarce (AKG)
Teucrium chamaedrys L.
22/11/69 Moderate (RJN)
T. fruticans L.
20/XI/69 Scarce (RJN)
T. scorodonia L.
23/11/70 Scarce (RJN )
Thymus serphyllum Coss.
12/V1I/68 Moderate (RJN)
Viola odorata L.
21/XI/69 Scarce (RJN)
V. wittrockiana Gams.
13/VI/69 Scarce (RJN )
13/II/70 Scarce (RJN )
25/1V/70 Moderate (RJN )
Myzus persicae (Sulzer)
Abutilon indicum (L.) Sweet
31/III/69 Moderate (AKG)
16/VI/70 Abundant (W-B )

Acer tataricum L.
11/III/70 Moderate (AKG)
Agathis robusta (F. J. Muell. )
F. M. Bailey
24/V/69 Scarce (AKG)
Agonis flexuosa Lindl.
5/VI/70 Scarce (AKG)
Agrimonia eupatoria L.
23/11/70 Scarce (AKG)
Ajuga reptans L.
29/V/68 Scarce (RJN )
17/V1/68 Moderate (RJN)
Akebia quinata (Moutt.) Decne.
21/1/70 Abundant (AKG)
Albizia thorelli Pierre
24/1II/70 Scarce (W-B)
Aloe ciliaris Haw.
19/11/70 Abundant (AKG)
Aloysia triphylla (L’Hér.) Britt.
8/XII/69 Scarce (RJN )
28/1V/70 Moderate (RJN )

A. virgata Juss.

5/11/70 Abundant (RJN )
Alcea rosea L.

21/11/70 Moderate (AKG)
Althaea setosa Boiss.
28/XII/70 Moderate (W-B)
Amelanchier sanguinea (Pursh. ) DC.
19/V/70 Abundant (AKG)
12/XII/70 Abundant (W-B)
Ammi majus L.

22/V1/70 Abundant ( W-B )
Amygdalus bucharica Korsh.
20/III/71 Moderate (AKG)

Anemopaegma chamberlaynii (Sims. )
Bur. & K. Schum.
19/VI/70 Moderate (W-B )

Anoda triangularis (L.) Schlechtend.
25/III/70 Abundant (AKG)

Antigonon leptopus Hook & Arn.
10/VI/70 Abundant (W-B)

Aphelandra squarrosa Nees
11/11/69 Abundant (AKG)

Aralia chinensis L.
2/II/70 Scarce (AKG)

597

598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Aristolochia petersiana Klotzsch.
11/VI/70 Abundant (W-B)

A. tagala Cham.

6/1/71 Abundant (Kono )
Artemisia dracunculus L.
25/11/70 Moderate (AKG)
Asparagus setaceus (Kunth) Jessop
2/V1/69 Scarce (AKG)

A. densiflorus (Kunth) Jessop
8/III/77 Abundant (RLB )
Baccharis vaccinioides Gardn.
11/11/70 Abundant (AKG)

B. glutinosa Pers.

16/11/70 Abundant (W-B)
Baeckea camphorata R. Br.
16/V/70 Moderate (W-B)
Balmea stormae Martinez
31/XII/70 Abundant (AKG)
Barleria obtusa Nees

31/XII/70 Scarce (W-B)
Basanacantha spinosa K. Schum.
14/1/71 Scarce (Kono)
Bocconia frutescens L.
18/1X/70 Abundant (AKG)

Brassaia actinophylla F. Muell.
1/VIII/69 Abundant (W-B)
25/V1/70 Moderate ( W-B )
Brassica campestris Oed.
10/11/70 Abundant (AKG)
Brunfelsia uniflora ( Pohl.) D. Don
17/11/70 Moderate (AKG)
Buddleia nivea Duthie

5/II/70 Scarce (AKG)
Buettneria urticifolia K. Schum.
19/III/70 Moderate (AKG)
Bulbinella robusta Kunth.
5/III/75 Abundant (RLB)

Buxus sempervirens “Arborescens’

20/1V/70 Moderate (AKG)

Caesalpinia japonica Sieb. &
Zuce.

12/I1I/70 Abundant (AKG)

Calliandra ‘Minima’
9/IV/69 Abundant (AKG)

Candollea tetrandra Lindl.
29/XII/70 Scarce (AKG)

Capsella bursa-pastoris Medic
17/11/70 Abundant (AKG)

Caragana aurantiaca Koehne

12/III/70 Moderate (AKG)

Carum carvi L.

25/V1/70 Abundant (W-B)

Casimiroa edulis Llave

21/V1/70 Scarce (W-B)

Cassia abbreviata D. Oliver subsp.
beareana (Holmes) Brenan

7/V1/68 Scarce (VFE)

C. atomaria L.

17/I1I/70 Abundant (W-B)

C. bicapsularis L.

17/I1/70 Abundant (AKG)

C. mimosoides L.
31/XII/70 Abundant (W-B )

C. nemophila coriacea Symon
15/1/71 Moderate (Kono )

Catalpa bignonioides Walt.
15/V/69 Scarce (VFE)

Cecropia peltata L.
14/III/70 Moderate (AKG)

Celtis africana Burm.
5/11/75 Abundant (RLB)

Centranthus ruber DC.
23/11/70 Abundant (W-B)
19/VII/70 Abundant (AKG)

Cerastium tomentosum L.
13/11/70 Moderate (RJN )

Cercidium sonorae Rose & I. M.
Johnst.
7/1V/70 Scarce (AKG)

Chamaecyparis nootkatensis (D. Don)
Spach
1/VII/69 Moderate Vagrant (VFE )

Chenopodium album Bosc.
17/II/70 Scarce (AKG)

Chlorophytum elatum R. Br.
25/V1/70 Abundant (W-B)
Chomelia obtusa Cham. & Schlect.
16/III/70 Abundant (AKG)

Cistus monspeliensis L.
21/IV/70 Scarce (AKG)

C. incanus L.
6/1/71 Scarce (Kono)

VOLUME 80, NUMBER 4

Citharexylum myrianthum Cham.

18/XII/68 Moderate (RJN )

19/VII/69 Scarce (RJN)

Clematis dioica L.

23/1/70 Scarce (W-B)

C. orientalis L.

19/V1/70 Moderate (W-B)

Clerodendrum bungei Steud.

3/1IV/69 Abundant (RJN)

10/XI/70 Scarce (RJN )

C. japonicum (Thunb. ) Sweet

18/VI1/69 Moderate (RJN )

C. myricoides ( Hochst.) R. Br.

17/V/69 Scarce (RJN )

C. nutans Wallich.

4/TV/69 Scarce (RJN)

10/XI/70 Scarce (RJN )

C. trichotomum Thunb.

27/V1/69 Moderate (RJN)

Coccoloba sagittifolia Ortega

26/1/70 Scarce (AKG)

Coffea arabica L.

19/V/70 Scarce (AKG)

Coleonema album (Thunb. ) Bartl. &
H. L. Wendl.

7/V/70 Moderate (W-B )

Cordia cylindristachya Roem. & Schult.

4/1/71 Scarce (Kono )

C. escalyculatha Vell.

21/III/70 Moderate (AKG)

Coronilla varia L.
19/III/69 Scarce (AKG)

Crataegus laevigata “Paul’s Scarlet’
23/V/75 Abundant (RLB)

Croton megalobotrys Muell.
11/VI/70 Abundant (W-B)
Cryptocarya rubra ( Mol.) Skeels
16/11/70 Abundant (AKG)

Cucumis sativus L.
17/XII/69 Scarce (AKG)

Cupaniopsis anacardioides (A. Rich. )
Radlk.
28/1V/69 Scarce (AKG)

Cuphea hyssopifolia HBK
29/XI/69 Scarce (AKG)

Cydonia sinensis Thovin
17/V/76 Abundant (RLB)

599

Dianthus deltoides L.
8/1IV/69 Abundant (RJN )
Duranta erecta L.
18/XII/68 Moderate (RJN )
15/V/69 Moderate (RJN )
18/11/70 Abundant (RJN )
5/I11/70 Moderate (RJN )
D. repens L.

5/11/70 Moderate (RJN )
D. repens ‘Alba’

18/XII/68 Moderate (RJN )
3/1IV/69 Abundant (RJN )
19/VII/69 Scarce (RJN)
29/X1/69 Scarce (RJN)
30/XII/69 Moderate (RJN )
5/11/70 Abundant (RJN)
31/XII/70 Moderate (RJN)
D. stenostachya Tod.
18/XII/68 Moderate (RJN )
15/III/69 Moderate (RJN )
29/XI1/69 Scarce (RJN )
30/XII/69 Abundant (RJN)
5/11/70 Abundant (RJN)
Echium fastuosum Jacq.
5/1/71 Moderate ( Kono)

E. wildpretii H. Pearson
19/V/69 Scarce Vagrant (VFE)
Eucalyptus baeuerlenii F. Muell.
25/1/71 Abundant (Kono)
E. caesia Benth.

7/V/70 Abundant (W-B)
E. calophylla R. Br.

20/1/71 Abundant (Kono)
E. cladocalyx F. Muell.
21/1/71 Moderate (Kono)
E. fasiculata Deane & Maiden
25/1/71 Abundant (Kono)

E. punctata DC.

20/1/71 Moderate (Kono)
Ficus iteophylla Miq.
19/III/70 Moderate (W-B)

Ficus sp.

6/1/71 Scarce (Kono)
Hagenia abyssinica J. F. Gmel.
6/1V/69 Moderate (RJN )
26/1/70 Abundant (RJN )

Halimium umbellatum (L.) Spach
5/1/71 Scarce (Kono)

600 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Hibiscus hamabo Sieb. & Zucc.
4/1/71 Abundant (Kono)
Hymenanthera dentata R. Br.
28/V/69 Scarce (RJN)
Hymenosporum flayum ( Hook.) F. L.
Muell.
4/111/75 Abundant (RLB)
23/V/75 Moderate (RLB)
30/1X/75 Moderate (RLB )
Hypericum elegans Steph.
5/1/71 Abundant (Kono)
H. prolificum L.
5/1/71 Scarce ( Kono )
Tris spuria L.
5/1/71 Abundant (Kono)
Juglans californica Wats.
28/11/68 Abundant (TLB)
26/11I/68 Abundant (TLB)
Kalanchoe densiflora Rolfe
5/11/75 Abundant (RLB)
Lagerstroemia indica L.
25/1V/70 Scarce (VFE)
Lampranthus sp.
14/1/71 Scarce (Kono)
Malus floribunda Sieb.
30/V1/69 Moderate (AKG)
Messerschmidia fruticosa L.
7/11/77 Scarce (RLB)
Nageliella purpurea ( Lindl.) L. O.
Williams
16/V1/69 Abundant (AKG)
Nephrolepis exaltata (L.) Schott.
27/111/75 Scarce (AGR)
Odontoglossum cariniferum Rchb.f.
12/1X/69 Abundant (AKG)

Olearia pimeleoides Benth.
17/111/69 Abundant (AKG)
Oncidium ampliatum Lindl.
16/V1I/69 Abundant (AKG)

O. cabagrae Schlechter
16/V1/69 Scarce (AKG)

O. sphacelatum Lindl.
16/VI/69 Abundant (AKG)
Oroxylum indicum (L.) Venten.
1/V/69 Abundant (AKG)

Owenia acidula F. Muell.
29/V/69 Scarce (AKG)

Pavonia sepium St.-Hil.
12/VII/69 Moderate (AKG)
Phaedranthus buccinatorium Miers.
19/VI1/69 Scarce (AKG)
Philodendron selloum C. Koch
27/111/69 Moderate (AKG)
Phlomis taurica Hartwiss
15/III/69 Scarce (AKG)
Physalis floridana Rydb.
17/XII/69 Scarce (AKG)
Pittosporum illicoides Makino
24/V1/69 Scarce (AKG)

P. mannii Hook.

5/11/75 Abundant (RLB)

Plantago major L.

30/11/71 Abundant (VFE)
Polycarpon tetraphyllum L.
2/V/69 Abundant (RJN)
Portulacaria afra Jacq.
22/1V/69 Abundant (AKG)
Potentilla rupestris L.

5/1/71 Moderate (Kono)
Prostanthera sp.

8/IV/69 Scarce (AKG)
Prunus yedoensis Matsum.
23/V1/69 Scarce (AKG)
Pseudobombax ellipticum (HBK) Dug.
10/X1I/70 Moderate (W-B)
Pseudocydonia sinensis C. K. Schneider
5/III/77 Scarce (RLB)
Pterocarya fraxinifolia (Lam.) Spach
5/1V/68 Scarce (TLB )
22/VI/70 Moderate (TLB)
3/1X/70 Scarce (TLB)

P. rehderiana C. K. Schneider
30/11/68 Moderate (TLB )
Puya alpestris C. Gay
9/V/69 Scarce (AKG)
Raphanus sativus L.

8/V/69 Abundant (W-B)
Rhagodia mutans R. Br.
8/IV/69 Scarce (AKG)

Rhus sylvestris Sieb. & Zuce.
14/VIII/68 Moderate (W-B)
19/VI1/69 Scarce (AKG)

Ricinus communis L.

20/V/69 Moderate (AKG)

VOLUME 80, NUMBER 4

Rosa banksiae Ait.
24/1V/70 Scarce (MBS)
Rumex crispus L.

12/11/69 Moderate (W-B)
Ruttya fruticosa Lindau.
16/V/69 Scarce (AKG)
Sagina apetala L.

6/VI/69 Scarce (RJN)
Salsola iberica Sennen & Pau.
9/V/69 Moderate (W-B)
Salvia aurea L.

2/V1/69 Moderate (RJN )
S. greggii A. Gray
10/VII/68 Scarce (RJN )
23/1/69 Abundant (RJN )
5/III/77 Scarce (RLB )

S. mellifera Greene
21/11/70 Abundant (RJN )
S. microphylla HBK
21/IV/69 Scarce (RJN )
26/II1/70 Abundant (RJN)
S. nutans L.

21/XI/69 Moderate (RJN)
S. pratensis L.

16/1X/69 Moderate (RJN )
Sambucus mexicana Presl.
21/11/70 Abundant (AKG)
S. nigra L.

8/VII/69 Scarce (AKG)
Santolina virens Mill.
17/IV/69 Scarce (W-B)
Sarcococca saligna (D. Don) Muell
15/1/69 Scarce (AKG)
21/V1/69 Moderate ( W-B )
Schinus polygamus ‘Cabrera’
18/VII/69 Scarce (AKG)
Senecio adonidifolius Loisel.
5/1/71 Moderate (Kono)
Sida rhombifolia L.
1/VIII/69 Scarce (W-B)
Solanum hispidum Pers.
4/1V/69 Moderate (AKG)

S. seaforthianum Andr.

31/III/69 Abundant (AKG)

Sterculia caudata Heward
1/VIII/69 Moderate (W-B)

Stereospermum kunthianum Cham.
15/1X/69 Abundant ( W-B )

Strongylodon macrobotrys A. Gray
16/1X/69 Abundant (AKG)
Tagetes patula L.

5/V/69 Moderate (AKG )

Tecoma fulva G. Don
1/V/69 Abundant (W-B)
Teucrium chamaedrys L.
22/TII/69 Moderate ( RJN )
T. flavum L.

23/XII/69 Scarce (RJN)
T. scorodonia L.

23/11/70 Scarce (RJN)

Tibouchina multiflora (G. Gardn.)
Gogn.
31/III/69 Moderate (AKG)

Tilia amurensis Komarov.
3/1V/69 Scarce (AKG)

601

Tithonia rotundifolia ( Mill.) S. F. Blake

1/VIII/69 Scarce (W-B)
Tubocapsicum anomalum Makino

1/V/69 Abundant (AKG)

Verbena carolina L.

16/IX/69 Abundant (AKG)
Veronica perfoliata R. Br.
4/III/75 Abundant (RLB)
Vinca minor L.

22/11/69 Scarce (AKG)

Viola wittrockiana Gams.
13/VI1/69 Scarce (RJN )
13/11/70 Scarce (RJN)
25/1V/70 Moderate (RJN )

V. wittrockiana ‘Majestic Giant’
4/X1/70 Abundant (RJN )
Vitex negundo L.

21/VI/69 Abundant (AKG)

V. pseudo-negundo Hand.-Mazz.
7/1V/69 Scarce (AKG)

Viola sp.

9/XI/70 Moderate (RJN )
Withania riebeckii Schweinf.
16/1X/69 Scarce (AKG)

Zexmenia frutescens Blake
16/1X/69 Moderate (AKG)

602 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Myzus sp. Pterocallis alni (de Geer )
Hymenosporum flavum F. Muell Alnus arguta Benth.
20/1/75 Abundant (RLB) 18/V/76 Scarce (RLB )

Neophyllaphis podocarpi Takahashi A. rhombifolia Nutt.
Podocarpus macrophyllus (Thunb. ) 17/V/76 Scarce (RLB)

D. Don A. tenuifolia Nutt.
1/V/69 Abundant (VFE) 18/V/76 Scarce (RLB )
2/X/69 Abundant (VFE) Pterocallis sp.

14/1V/70 Abundant (VFE) Alnus formosana Makino
22/V1/70 Moderate (VFE)

4/11/77 Scarce (RLB)
13/11/71 Moderate (VFE ; ela a
BINT Alandont a Rhopalosiphum fitchii (Sanderson )

Bromus carinatus Hook. & Arn.
All | L 5/11/70 Moderate (AKG)

um schoenoprasum L. i
24/V1/70 Abundant (AKG) Rhopalosiphum insertum (Walker) group

Poa annua L.
7/Il1/77 Scarce (RLB )

Rhopalosiphum maidis ( Fitch )

Neotoxoptera oliveri (Essig )

A. tuberosum Rottl.
25/11/70 Abundant (AKG)

Neotoxoptera violae (Pergande )

; ron rinatus Hook. & Arn.
Cerastium tomentosum L. Bromus carina

13/V/69 Scarce (RJN) 5/11/70 Moderate (AKG)

23/X/69 Abundant (RJN) Echinochloa crus-gallis (L.) Beauv.
13/11/70 Moderate (RJN) 14/1/71 Abundant (CFS )

5/XI/70 Moderate (RJN ) Oryzopsis miliaceae Benth. & Hook.
Dianthus deltoides L. 30/1V/69 Moderate (AKG )
8/1V/69 Abundant (RJN) Sida rhombifolia L.

Hymenanthera dentata R. Br. 1/VIII/69 Scarce (W-B)

18/V/69 Scarce (RJN) Sorghum halepense (L.) Pers.

5/V1/69 Abundant (W-B)
8/VII/69 Abundant (AKG)
S. bicolor (L.) Moench
16/X/69 Abundant (AKG)
5/III/77 Abundant (RLB)

Rhopalosiphum padi (L.)

Teucrium chamaedrys L.

22/111/69 Moderate (RJN )

Viola odorata L.

21/XI/69 Scarce (RJN)
Ovatus crataegarius (Walker)

Teucrium fruticans L.

é Bromus carinatus Hook. & Arn.
22/VII/68 Scarce (RJN ) 8/1V/70 Moderate (CFS)
Ovatus phloxae (Sampson) 9/V/70 Abundant (CFS)
Cerastium tomentosum L. 3/1V/71 Moderate (CFS)
13/V/69'Scarce (RJN ) Chamaecyparis lawsoniana (A. Murr.)
Periphyllus negundinis (Thomas ) Parl.
Acer negundo L. 10/III/70 Abundant (VFE)
27/1V/70 Abundant (W-B ) Cupressus lusitanica Mill.
Pleotrichophorus glandulosus (Kaltenbach ) 17/1V/71 Abundant Vagrant (VFE)
Artemisia nutans Willd. Oryzopsis miliacea Benth. & Hook.
22/V/67 Abundant (MDL) 30/1V/69 Moderate (AKG)
Prociphilus fraxinifolii (Riley ) Rhopalosiphum rufiabdominalis (Sasaki)
Fraxinus uhdei “Majestic Beauty’ Conyza bonariensis (L.) Cronq.

23/V/75 Moderate (RLB) 17/X/73 Moderate (CFS)

VOLUME 80, NUMBER 4

Echinochloa crus-galli (L.) Beauv.

5/VI/73 Abundant (CFS )
Rhopalosiphum sp.

Rosa spinosissima L.

10/VII/68 Scarce (MBS )

Tecoma fulva G. Don

1/V/69 Abundant (W-B)
Schizolachnus curvispinosus Hottes, Essig

and Knowlton

Pinus durangensis Roezl.

22/1V/71 Moderate (VFE)

P. glabra Walt.

11/XI/69 Abundant (VFE) (DJV)

P. roxburghii Sarg.

5/TiI/70 Abundant (VFE)

Siphonatrophia cupressi (Swain)

=.

Cupressus lusitanica Mill.
7/X1/69 Scarce (VFE)
2/1V/70 Moderate (VFE)
26/1V/71 Abundant (VFE)
C. pygmea Sarg.
1/1V/70 Abundant (VFE )
C. sempervirens ‘Stricta’
25/1V/70 Scarce (VFE)
Ervatamia coronaria Stapf.
12/X1I/70 Abundant (VFE)
Smynthurodes betae Westwood
Portulaca oleracea L.
2/X/72 Abundant (CFS)
Stegophylla essigi Hille Ris Lambers
Quercus agrifolia Née
29/V1/73 Abundant (VFE)
4/111/75 Abundant (RLB)
Takecallis arundicolens (Clarke )
Arundinaria simonii (Carriére) A. & C.
Riviere
6/XII/69 Abundant (MDL)
Pseudosasa japonica Makino
23/1V/70 Abundant (MDL)
Sasa pygmaea Mitf.
28/1/70 Abundant (MDL)
Takecallis arundinariae (Essig )

Arundinaria amabilis McClure
7/11/68 Moderate (MDL)
13/1V/70 Moderate (MDL)

603

A. simonii (Carriére) A. & C. Riviere

15/1V/70 Scarce (MDL)

Bambusa glaucescens ‘Stripestem
Fernleaf’

15/IV/70 Scarce (MDL)

B. oldhamii Munro

6/XII/69 Abundant (MDL)

Bambusa sp.

23/11/68 Abundant (MDL)

Chimonobambusa hookerianum Nakai

13/IV/70 Scarce (MDL)

C. marmorea ( Mitt.) Nakino

13/IV/70 Scarce (MDL)

Phyllostachys aurea Carriere

6/XII/69 Abundant (MDL)

27/1/70 Abundant (MDL)

20/1/75 Abundant (RLB)

P. bambusoides Sieb. & Zucc.

13/1V/70 Moderate (MDL)

P. bambusoides ‘Castillon’

7/11/68 Abundant (MDL)

P. flexuosa A. & C. Riviére

7/11/68 Abundant (MDL)

P. meyeri McClure
20/1/68 Abundant (MDL)
13/1V/70 Moderate (MDL)

P. nigra (Lodd.) Munro
6/XII/69 Moderate (MDL)

P. nigra “‘Henon’

6/XII/69 Scarce (MDL)

P. viridis (R. A. Young ) McClure
7/11/68 Abundant (MDL)
13/IV/70 Scarce (MDL)

Phyllostachys sp.

6/XII/69 Abundant (MDL)
10/X/70 Abundant (MDL)
6/XII/69 Moderate (MDL)
15/1V/70 Moderate (MDL)

Pleioblastus simoni Nakai
16/IV/70 Scarce (MDL)

Semiarundinaria fastuosa ( Marliac)
Makino
15/IV/70 Scarce (MDL)

Tinocallis platani ( Kaltenbach )

Metrosideros kermadecensis W. Oliver
15/1/71 Moderate (Kono)

604

Tinocallis ulmifolii (Monell )

Carya illinoinensis (Wangenh.) C. Koch.
27/VII/70 Moderate (TLB)

Toxoptera aurantii (Boyer de Fonscolombe )
Acer cappadocicum Gled.
17/VII/70 Scarce (AKG)
Agonis flexuosa (K. Spreng.) Schauer.
5/V1I/70 Scarce (AKG)
Camellia fraterna Hance
22/1X/70 Scarce (AKG)

C. granthamiana Sealy
21/1X/70 Scarce (W-B )
Camellia sp.
1/X/75 Scarce (RLB)
Clausena lansium (Lour.) Skeels
27/III/70 Scarce (AKG)
Clematis armandii Franch
2/XI/70 Scarce (AKG)
Cordia superba Cham.
30/1X/70 Abundant (AKG)
Murraya paniculata (L. )
Jack
7/X/76 Moderate (RLB)
Olearia argophylla F. Muell
20/1/71 Moderate (Kono)
Quillaja saponaria Molina
29/X/69 Scarce (AKG)
Rhamnus californica Eschsch.
10/11/70 Abundant (W-B )
Xylosma congestum (Lour.) Merrill
23/V1/69 Scarce (AKG)
18/XI/68 Abundant (W-B)

Tuberculatus (Pacificallis )
columbiae Richards
Quercus bebbiana C. K.

Schneid.

18/IV/73 Scarce (VFE)

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Tuberculatus (Pacificallis ) sp.
near columbiae Richards
Quercus alba L.

12/XI/73 Moderate (VFE)
Q. agricolia Née

26/V1I/73 Moderate (VFE)
Q. emoryi Torr.

13/VI/73 Scarce (VFE)

Q. ganderi C. Wolf

18/V1/73 Moderate (VFE)
Q. leucotrichophora A. Camus
18/VI/73 Scarce (VFE)

Tuberculatus (Pacificallis )
maureri (Swain )

Quercus agrifolia Née
26/V1/73 Moderate (VFE)
29/V1/73 Abundant (VFE)

Tuberculatus (Tuberculoides ) annulatus
( Hartig )

Quercus robur L.

9/VII/73 Abundant (VFE)
1/XI/73 Abundant (VFE)
8/VI/75 Abundant (RLB)

Tuberolachnus salignus (Gmelin)
Carpenteria californica Torr.
30/1/70 Moderate (AKG)
Salix taxifolia HBK
6/XII/69 Abundant (W-B)

Wahlgreniella nervata (Gillette )
Rosa hugonis Hemsl.
17/11I/71 Abundant (MBS)
R. ‘Manon Cochet’

9/III/76 Abundant (MBS)
R. pisocarpa A. Gray
23/11/71 Abundant (MBS)
Rosa sp.

29/V1I/69 Abundant (MBS)

Literature Cited

Leonard, M. D. and H. G. Walker. 1973.
and County Arboretum (Homoptera
75(2):209-212.

1974.

Aphids collected in the Los Angeles State
: Aphididae). Proc. Entomol. Soc. Wash.

Additional aphids collected in the Los Angeles State and County
Arboretum (Homoptera: Aphididae).
Leonard, M. D., H. G. Walker and L. Enari.

Coop. Econ. Ins. Rpt. 24(39) :778-779.
1970. Host plants of Myzus persicae

at the Los Angeles State and County Arboretum, Arcadia, California (Homoptera:
Aphididae ). Proc. Entomol. Soc. Wash. 72(3):294-312.

VOLUME 80, NUMBER 4 605

—. 197la. Host plants of Aphis gossypii at the Los Angeles State and County
Arboretum, Arcadia, California (Homoptera: Aphididae). Proc. Entomol. Soc.
Wash. 73(1):9-16.

—. 197lb. Host plants of three polyphagous and widely distributed aphids in the

Los Angeles State and County Arboretum, Arcadia, California, (Homoptera:

Aphididae ). Proc. Entomol. Soc. Wash. 73(2):120-131.

197lc. Host plants of Toxoptera aurantii at the Los Angeles State and

County Arboretum, Arcadia, California (Homoptera: Aphididae). Proc. Entomol.

Soc. Wash. 73(3) :324—326.

—. 1972. Host plants of aphids collected at the Los Angeles State and County
Arboretum during 1966 and 1967 (Homoptera: Aphididae). Proc. Entomol.
Soc. Wash. 74( 1) :95-120.

(HGW) 610 Rose Marie Drive, Arcadia, California 91006; (MBS)
Systematic Entomology Laboratory, IIBIII, Fed. Res., Sci. & Educ. Admin.,
USDA, Beltsville, Maryland 20705; and (LE) Los Angeles State and County
Arboretum, 301 North Baldwin Avenue, Arcadia, California 91006.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 606-610

A NEW GENUS, SPECIES AND FAMILY OF HYMENOPTERA
(ICHNEUMONOIDEA) FROM CHILE

W. R. M. Mason

Abstract—Apozyx penyai, a new genus and species, is described from
the temperate forest of Arauco province, Chile. The species has a mixture
of features of Ichneumonidae, Braconidae and Agriotypidae. It represents
a new family, Apozygidae.

A single male specimen, superficially resembling a Spathius (Braconidae)
with a second recurrent vein, collected by Luis E. Pena in Chile has long
stood in the Canadian National Collection. I delayed description in the
hope of obtaining more specimens, especially to discover the characteristics
ot the female; but now, after almost 20 years of waiting and examining
at least 20,000 Ichneumonoidea from Chile, I am convinced that the species
must be so rare that further delay is unjustifiable. I think it best to
describe this strange creature in hope that the publication will stimulate
those with opportunity to make an effort to find more specimens.

Apozyx penyai Mason, new genus and new species

This description is to cover genus and species. It seems pointless to
guess at the heirarchical levels appropriate to the various characters on
the basis of one specimen.

Etymology—tThe generic name is a Greek masculine noun' meaning
bachelor. The spelling of the specific patronym, because the diacritic “i”
is not allowed by the Code (Art. 27), is modified to produce the approxi-
mately correct pronounciation by most European language speakers.

Description —Head subcuboid, sparsely hairy, and generally shiny (Figs.
1-4); mandibles stout and short but strongly tapered, (Figs. 3 and 4)
upper tooth larger than lower; maxillary palpi with 5 articles, the most
proximal 2 apparently fused; labial palpi with 4 subequal thickened
articles; clypeus with a very prominent transverse ridge, the apical part
strongly receding (Figs. 3 and 5), apical margin concave; labrum polished
and concave, filling cavity between mandibles and clypeus, mouth thus
resembling that of a cyclostome braconid but opening lenticular, about 2
as wide as long (Fig. 4); antennae unspecialized (Fig. 9), flagellum
16-jointed, articles 2-4 longer than wide and bearing longitudinal
placodes; ocelli round and small (Fig. 2); occipital carina complete, join-
ing hypostomal well above mandible base.

Thorax (Figs. 1, 2 and 7). Pronotal structure typical of Ichneumonoidea,
that is, 2 triangular sides connected by a narrow transverse collar and

VOLUME 80, NUMBER 4 607

posterior corners touching tegulae and fore coxae; propleuron with a small
lobe overlapping pronotum at lower corner; notauli deep, crenulate, and
extending back to meet just before scutellum (Fig. 2); median lobe of
scutum broadly concave medially and strongly elevated laterally at notauli
where the scutum turns down from horizontal to vertical; scutellum apically
weakly striate and bearing a low median carina on the apical declivity
(Fig. 2); mesopleuron with complete prepectal carina and mesulcus but
without sternauli (Fig. 1); propodeum (Figs. 1 and 7) long and areolated,
with conspicuous apophyses and a strongly marked declivity behind them;
pronotum, metapleuron and propodeum mostly rugose, balance of thorax
smooth and sparsely hairy.

Legs with divided trochanters; tibial spurs short, 1: 2: 2; tarsal claws
simple; fore tibia with dense multiple rows of thickened hairs on outer
anterior side; middle tibia with a similar but weaker armature; remainder
of legs shiny and sparsely hairy except for dense hair on inner apical part
of all tibiae and on the tarsi. Hind coxae and abdomen arising from 3
separate foramina on the thorax and propodeum.

Wings (Fig. 8) with venation most similar to Braconidae but the fore-
wing with a strong 2nd recurrent; hindwing with a short, but sclerotized
discoidella and no trace of brachiella; 3 hamuli on metacarpella; basella
meeting subcostella proximad to radiella; no trace of anal lobe.

Abdomen with tergum I and sternum I solidly fused into a rigid, rugose,
tubular petiole (Figs. 1, 6 and 7); sternum I smooth and visible only
beneath anterior % of petiole; petiolar segment truncate posteriorly, thus
the sternal and tergal regions equally long; sides of tergum I apparently
[Without dissection it is impossible to decide whether the sternite ends near
the middle or extends narrowly to the posterior end of the first segment. |
meeting along midventral line of posterior % of petiole (Fig. 6); a lunule
of soft integument present at base of sternum II. Remainder of abdomen
(Fig. 1), both terga and sterna sclerotized, smooth, sparsely hairy and
evenly convex; terga II and III fused and very large; sterna overlapping
and not fused; more posterior terga and sterna freely articulated and some-
what telescoped; hypopygium small, apically emarginate and_ simple;
parameres simple, balance of genitalia difficult to observe; pygostoles
present, a little longer than wide.

Female unknown.

Holotype.— é , Pichinahuel, Cordillera Nahuelbuta, Arauco, CHILE, 10-20
Jan. 1959, Luis E. Pefia. CNC No. 15523.

Discussion

Although the affinities of Apozyx could be more confidently assessed
with a knowledge of the female and larval anatomy and the life history,
there is enough evidence in the male to show that it fits best in the

608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ryall Bl (
1s l

9

Figs. 1-9. Apozyx penyai. 1, Lateral whole view. 2, Head to scutellum, dorsal.
3, Head, lateroventral. 4, Head, anterior. 5, Theoretical median longitudinal section of
head to show arrangement of face (F), clypeus (Cl), labrum (Lbr) and mandibles
(Md). 6, Ventral side of petiole, anterior end to left. 7, Metanotum, propodeum and
petiole, dorsal. 8, Wings (scale 24 as large as other figures). 9, Antenna.

VOLUME 80, NUMBER 4 609

Ichneumonoidea. The following features are significant for this placement;
1, antennae not elbowed, i.e., the scape short; 2, flagellum with 16 articles:
3, flagellar articles with longitudinal placodes; 4, mandibles with two apical
teeth; 5, trochanters double; 6, costa and subcosta of forewing fused; 7,
large pterostigma present; 8, no anal lobe in hindwing; 9, copius venation
with two closed cells in hindwing; 10, true abdomen with 9 distinct seg-
ments; and 11, spiracles on each tergum from | to &.

The hindwing venation is of the braconid type because of the presence
of a basellan vein (2r-m) and absence of an intercubitella (3r-m). The
forewing is braconid-like because of the large 2nd cubital cell (1Rs) and
basal abscissa of cubitus (Rs+M) but the 2nd recurrent (2m-cu) vein is
never found in braconids (except rarely in freaks).

The abdomen suggests the Agriotypidae by its fused second and third
terga, by its convex and strongly sclerotized sterna and by the fused and
tubular petiole.

The petiolar structure is significant. In all generalized Ichneumonidae
and Braconidae the petiolar sternum is separate and divided into a
rigidly sclerotized anterior part and a partly membranous posterior part
that may bear one or two weakly sclerotized plates. Even in specialized
groups where the elements are fused, the posterior part of the sternite is
never as long as the tergite and always leaves a V- or U-shaped mem-
branous area at the ventral apex. In most Chalcidoidea, Cynipoidea, large
parts of the Proctotrupoidea and in Agriotypidae and Evaniidae the
petiolar tergite and sternite are fused into a tubular or ring-like structure,
whose ventral surface is about equal to the dorsal, so that there is no
apical ventral membranous area. This is the type of petiole found in
Apozyx. It raises the possibility that Apozyx may really be related to un-
known, and presumably more fully veined, ancestors of some microhy-
menopterous group. However the possibility is so speculative that I think it
better at present to place Apozyx in Ichneumonoidea where most its
characters are harmonious. It will be difficult to place it confidently until
there is far better knowledge of Apozyx and of phylogeny of Apocrita.

Nevertheless a placement must be made. The genus can be forced into
existing families only by a procrustean redefinition of family characters
that I find unacceptable. I place this genus in a new family, Apozygidae,
assigned provisionally to Ichneumonoidea.

Family Characters

Apozygidae——Mandible 2-toothed; antennal flagellum with more than
14 articles; trochanters divided. Wings present and with generalized
venation; forewing with costa and subcosta fused; stigma large; basal
abscissa of cubitus (Rs+M) present; 2nd cubital cell (1Rs) large; 2nd
recurrent (2m-cu) present. Hind wing with 2 closed cells; basellan vein

610 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(2r-m) meeting subcosta before radius; intercubitella (3r-m) absent.
Petiole tubular and with components completely fused, ventrally with no
membranous area; sternum IT and following fully sclerotized and evenly
convex when dry; metasomal spiracles 1-7 all present and approximately
equal sized.

Biosystematics Research Institute, Agriculture Canada, Ottawa, Canada,
K1A 0C6.

Footnote

‘ Apozyx is not found in most Greek dictionaries because it was used only by one
Eustathius in the 12th century.—Editor’s note (G. C. Steyskal).

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 611-615

NOTES ON NATHAN BANKS’ SPECIES OF THE MITE
GENUS CARABODES (ACARI: ORIBATEI)

Roy A. Norton

Abstract—The identity and taxonomic status of six mite species de-
scribed by Banks in the genus Carabodes is discussed. A lectotype is
selected for C. dorsalis Banks, and C. omo (Jacot) is designated a junior
subjective synonym of C. granulatus Banks.

Primarily during the period 1894-1915, the arthropodologist Nathan
Banks described more than 100 species of oribatid mites; most of these
were from eastern North America. The general quality of his descriptions
was low by modem standards, which, in conjunction with the trend in
later years toward narrowing generic concepts, necessitates reexamination
of his species. This has been done in a few instances by subsequent
oribatologists, but most are still in need of redescription and generic
recombination. The purpose of this note is to examine the identity and
taxonomic status of the six species placed by Banks (1895a, 1895b, 1896)
in the genus Carabodes C. L. Koch. I wish to thank Drs. Herbert W. Levi
(Cambridge, Massachusetts) and E. W. Baker (Beltsville, Maryland) for
providing access to Banks’ type-specimens in the Museum of Comparative
Zoology (MCZ) and U.S. National Museum (USNM) collections, respec-
tively.

Carabodes apicalis Banks and Carabodes dorsalis Banks

Two of Banks’ six “Carabodes” species are not members of the Cara-
bodidae. Jacot (1937) transferred Banks’ (1895a) species C. apicalis to
Carabodoides when the genus was first proposed; it is in need of
redescription. Most recently, this genus has been placed in the family
Anderemaeidae (Balogh, 1972).

The second species, Carabodes dorsalis Banks, 1896, was correctly trans-
ferred to Nanhermannia Berlese (family Nanhermanniidae) by Jacot
(1937). Three specimens are located in the MCZ; two are labeled “cotype”
and mounted in balsam on separate slides, one is in alcohol and is the
“suspected type” according to unpublished MCZ listings. A problem exists
because one of the two slide-mounted specimens (the ventral mount) is
not this species; it appears conspecific with Carabodes brevis Banks. It
has obviously been remounted since its deposition, but there is no record
of who mounted or labeled the specimen. The second slide (a lateral
mount) bears a correctly identified specimen; it is labeled a “cotype” in

612

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

+10 pm

C®

OO

I at
fo) =
© = p>

Pa

(@)
O

alee
(@)
O¢

1
ae,

i OF

> )

fe)

Figs. 1-2. Odontocepheus elongatus. 1, Left humeral region of specimen from
England. 2, Left humeral region of specimen from New York. Figs. 3-9. Carabodes
niger. 3, Dorsal aspect (legs removed). 4, Leg I. 5, Leg II. 6, Femur and genu of
leg III. 7, Tibia and tarsus of leg HI. 8, Femur and genu of leg IV. 9, Tibia and
tarsus of leg IV. Fig. 10. Carabodes brevis, sensillus (ss) and notogastral seta ti of
cotype. Fig. 11. Carabodes granulatus, prodorsal seta in and notogastral setae ti (with

integumental sculpturing) and pz.

VOLUME 80, NUMBER 4 613

A. P. Jacot’s handwriting and was probably originally mounted from
alcohol by him. The alcoholic specimen still carries a label in Banks’ hand-
writing. I hereby designate the single alcoholic specimen as the lectotype
of Carabodes dorsalis Banks and the laterally mounted “cotype” as a
paralectotype. Species concepts in Nanhermannia have not improved since
Hammen’s (1959) clarification of the identity of the type-species, N. nana
(Nicolet). Questions raised by Jacot (1937) and Hammen (1959) on the
synonymy of Banks’ species with species described from Europe will have
to be answered by population studies, not simply redescription of type-
specimens.

Carabodes oblongus Banks

Banks himself (1895a) was the first to indicate the need for the re-
moval of this species from Carabodes, but its present genus, Odontocepheus
Berlese, had been proposed for half a century before Johnston (1965)
effected the recombination. I have examined specimens of the type-species
of Odontocepheus, O. elongatus (Michael), from England and Scandinavia
and compared them with the type-specimen of O. oblongus (MCZ) and a
number of other specimens from the northeastern U.S. The size range of
the American mites (505-610 »m) spans the measurements of my European
specimens, and there are only two notable differences. The notogastral
setae are relatively slightly shorter in the American specimens; setae of the
series h;, ps;, ps2, and ps,, for example, do not extend posteriorly as far as
the insertion of the next seta. Also, the spine-like apophysis which extends
medially from the humeral region of the notogaster is shaped differently
in the American specimens (Figs. 1 and 2).

My European material is limited, however; and considering the variation
attributed to body length and setal shapes noted in European populations
by Perez-Inigo (1971), his synonymy of O. oblongus with O. elongatus
(apparently without the benefit of seeing American specimens) seems
justified on morphological grounds.

Carabodes niger Banks

This and the following two species are true members of the genus
Carabodes, as it is conceived at this time. Carabodes niger is one of the
largest species in the northeast. Banks’ (1895a) original estimate of
0.5 mm is misleading; 17 cotypes in the MCZ and USNM collections
ranged from 490-643 ,.m, total length (mean 580 »m). Most collections
from New York and Ohio have been from fungal fruiting bodies (especially
Polyporus spp.), but I have collected it from forest leaf litter in North
Carolina. The following is presented as a more complete diagnosis of the
species.

Integument strongly sculptured (Fig. 3), mostly with pit-like depressions

614 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

8-25 ,m in diameter. Prodorsum with weak pitting and “V” shaped ridge
between lamellae; lamellae mostly with transverse ridges instead of pits.
Dorsosejugal groove deep, broad (cervical cavity of Sellnick and
Forsslund, 1953). Seta ro smooth, attenuate; le flattened, acuminate, with
small barbs; in lanceolate, flattened, dorsally covered with small barbs,
tips usually crossing medially. Sensillus (ss) distally spatulate, with
ventrally deflexed sides; dorsal surface with small barbs. Notogaster with
distinct postero-lateral rim. Setae ta, ti, te, ms, r; and r, large, flattened,
spatulate, 2.9-3.2 times longer than broad, dorsally covered with small
barbs except along medial axis. Setae r;, pi, p2 and p, setiform, with
small barbs. Legs as in Figs. 4-9.

In the Barneby Center, Ohio population studied, setae r;, p,, p, and p,
are somewhat longer, almost reaching the insertion of the next posterior
seta. Also, the unguinal setae (w) of all tarsi are simply scale-like, lacking
the distal attenuation present in the cotypes from Long Island.

The specific epithet nigra was used in the original description, but since

Carabodes is a masculine noun, it was later emended to niger (Banks,
1904).

Carabodes brevis Banks

The holotype (labeled “type”) is mounted in balsam and located in the
MCZ, along with an alcoholic “cotype.” As mentioned previously, the
misidentified cotype of Carabodes dorsalis belongs to this species and may
be a member of the original type-series which was wrongly labeled in later
years.

This species is generally similar to C. niger, but the sensillus is short,
strongly clavate and the central notogastral setae are much thinner, only
slightly broadened distally (Fig. 10). Seta in is similar to that of C. niger,
but slightly narrower. Setae r;, pi, p2 and p; are slightly smaller than the
central dorsal setae, but there is no strong dimorphism as in C. niger. The
dorsosejugal groove is shallower (there is no cervical cavity) and the
medial prodorsal ridge is absent. All three specimens in the MCZ are

about 500 «um in length; Banks’ (1896) original statement of 0.4 mm is
erroneous.

Carabodes granulatus Banks

A single alcoholic specimen, the holotype, is located in the MCZ. This
species can be distinguished from all other known American Carabodes
species by means of the clavate, coarsely barbed notogastral and _ inter-
lamellar setae, and the integumental sculpturing of the notogaster which
gives the impression of small, interconnected “rosettes” (Fig. 11).

VOLUME 80, NUMBER 4 615

Examination of several cotypes of Carabodes omo Jacot (1937:241)
from the USNM showed this to be a junior subjective synonym of C.
granulatus (new synonymy). The species is common in sphagnum bogs in
the northeastern U.S. and forest litter in North Carolina.

Literature Cited

Balogh, J. 1972. The oribatid genera of the world. Akademiai Kiado, Budapest.
188 pp.
Banks, N. 1895a. On the Oribatoidea of the United States. Trans. Am. Entomol.
Soc. 22:1—16.
1895b. Some acarians from a sphagnum swamp. J. N.Y. Entomol. Soc.

3:128-130.
1896. New North American spiders and mites. Trans. Am. Entomol. Soc.
23:57-77.

1904. <A treatise on the Acarina, or mites. Proc. U.S. Nat. Mus. 28:1—-114.

Hammen, L. van der. 1959. Berlese’s primitive oribatid mites. Zool. Verh., Leiden
No. 40:1-93.

Jacot, A. P. 1937. New moss-mites, chiefly midwestem—II. Am. Midl. Nat. 18:
237-250.

Johnston, D. E. 1965. <A catalogue of the determined species of Acari (excl. Ixodides )
in the collections of the Institute of Acarology. Ohio Agric. Exp. Stn., Publ.
Ser. 29. 66 pp.

Perez-Inigo, C. 1971. Acaros oribatidos de suelos de Espafia peninsular e islas
Baleares (Acari: Oribatei) III. “EOS,” Rev. Esp. Entomol. 46:263-350.
Sellnick, M. and K.-H. Forsslund. 1953. Die Gattung Carabodes C. L. Koch 1836

in der schwedischen Bodenfauna (Acar. Oribat.). Ark. Zool. 4:367—390.

S.U.N.Y. College of Environmental Science and Forestry, Syracuse, New
York 13210.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 616-625

REARING OF TEXAS TABANIDAE
(DIPTERA). I. COLLECTION, FEEDING, AND
MAINTENANCE OF COASTAL MARSH SPECIES

Patrick H. Thompson and Peter C. Krauter

Abstract —In order to study methods of rearing, 1639 females of several
species of coastal marsh Tabanidae were collected in modified Manitoba
Traps and in modified Animal Traps, augmented with CO, from dry ice,
at West Galveston Bay, Texas; five weekly collections from 18 July—
17 August, inclusive, produced the following totals of dominant species:
Tabanus nigrovittatus Macquart, 1228; T. lineola var. hinellus Philip, 349;
and T. acutus (Bigot), 62. Immediately after their collection, many fe-
males fed on warmed, citrated beef blood which was held in prophylactics
derived from animal membranes. From 26.7-39.6% of 353 females of the
three species engorged to repletion on prophylactics placed on top of trap
collection containers. Survivorship curves for one sample population
showed that mortality rates were fairly constant at all age levels for these
three species; 50% and 0% survivorships, respectively, occurred at 5 days
and 50 days for T. nigrovittatus; 4 days and 36 days, T. lineola var. hinellus;
and 9 days and 49 days, T. acutus. Comparison of collection dates of
minor species, with that from a study made in 1971, indicated that. several
inland species had greatly declined or disappeared in the 6-year interim
between 1971 and 1977.

Study of Tabanidae within the laboratory has been impeded by negative
results in rearing these insects. To the present day, scientists throughout
the world have attempted to maintain and to perpetuate many species,
mostly with refractory results. According to these researchers, major
hurdles to such success with tabanids have been caused by the reluctance
of these insects to accept blood; by their habit of indiscriminate oviposition
in cages, often under conditions not conducive to maturation of the eggs;
by the refusal of the larvae to feed or by their cannibalistic voracity,
which also hinders rearing en masse; by mortality resulting from physio-
logical disorders during ecdysis and pupation and by disease organisms
throughout development; and by the inhibitory effect of cage retention
upon reproductive behavior. To complicate this research dilemma, the
study of many species has tended to divide the disciplinary research
effort, and as a result, confused the interpretation of efforts with individ-
ual species.

In conjunction with our studies of feeding behavior and reproductive

VOLUME 80, NUMBER 4 617

physiology, rearing attempts of several species of Coastal Plain Tabanidae
were begun in 1977. Initial work with summer tabanids at inland locations
was greatly restricted by very small populations of nearly all species, even
the most abundant and successful dominants. On the other hand, popula-
tions of several species inhabiting marshes along the Gulf of Mexico were
apparently unaffected by the factors reducing populations of the inland
forms.

The Coastal Marsh Fauna

Inland from the barrier beaches and estuaries of the Texas coast line,
are intermittently flooded tidelands inhabited by a specialized fauna of
Tabanidae distinct from other faunas of the Coastal Plain found inland,
and of other groups of tideland faunas found eastward. Specifically, this
fauna is characterized by five species which are intimately associated with
coastal marshes. Tabanus eadsi Philip and T. texanus Hine share the same
range along the south Texas coast, a range extending into western Louisiana
(Thompson and Pechuman, 1970). A larger, and sometimes more prominent
species, numerically, is T. acutus (Bigot), a form occurring from West
Galveston Bay, and possibly further south, eastward along the Gulf to
coastal counties of the Florida Panhandle (Fairchild, 1937; Stone, 1938).
The distribution of T. lineola var. hinellus Philip, of T. nigrovittatus
Macquart and of their related forms extends eastward along the Gulf
Coast, and then northward along the Atlantic Coast (Philip, 1965).

A survey of marshes on the south Texas coast line in 1971 by Thompson
(1973) characterized this ecosystem and the tabanid populations then in-
habiting it, by presenting the relative abundance and seasonal ranges of
the species comprising the fauna; by providing population indices for
dominant and subdominant species throughout the season; and by de-
scribing the traps used in collecting the insects and in accumulating the
data. A concomitant study published soon thereafter (Thompson and
Gregg, 1974) described the trap modifications necessary under the rigorous
coastal weather conditions; and discussed the relative efficiency of the 2
trap types used. More significantly, this work greatly simplified the ob-
jectives of the present study.

Methods and Results

Changes in the fauna.—Initial trap collections on 18 July showed that
the population structure of the West Bay study area had changed in the
6-year interim since the previous (1971) study (Thompson, 1973) (Table
1). Although consecutive weekly population indices from 18 July through
i7 August showed that T. nigrovittatus still dominated the West Bay

618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Relative abundance of Tabanus spp. taken in five consecutive weeks
during late July and early August, 1971 and 1977, West Bay.

Percent of Population

Species 1971 1977
T. nigrovittatus Macquart" 88.5 74.0
T. lineola var. hinellus Philip 1.5 ill
T. sulcifrons Macquart 5.6 0
T. acutus ( Bigot ) 0.3 Shi
T. subsimilis subsimilis Bellardi Tha 0.2
T. texanus Hine 1.0 Onl
Remaining species” FAN aspps) 0.9 (2 spp.)
Total percent 100.0 100.0
Total flies 4,724 1,660

“Sens. lat., i.e., nominal T. nigrovittatus and T. nigrovittatus var. fulvilineis Philip
were not differentiated for the purposes of this study because of the large number of
intermediate specimens observed here.

» Species found only in 1971: Tabanus atratus F., T. cymatophorus Osten Sacken,
T. eadsi Philip, T. mularis Stone, and T. stygius Say; in both years, Tabanus lineola
lineola F. and Chrysops flavidus Wiedemann.

marshes during the summer months, 2 coastal marsh species, T. acutus and
T. lineola var. hinellus, markedly increased. In addition, inland forms
drastically declined or disappeared. No specimens of T. sulcifrons were
taken although 265 females had been trapped in 1971. Five other inland
species represented by 1-10 specimens during the same period in 1971
were not taken in 1977 (q.v., footnote “b”, Table 1). These differences
become more meaningful considering that the 1977 catches were augmented
with CO:. (The harsh winter temperatures which probably decimated
inland populations of these same species, could also have affected those at
West Bay.)

Trap characteristics—Five weekly collections, usually on several con-
secutive days in each week, were made from 18 July through 17 August.
In order to collect large numbers of Tabanus nigrovittatus et al., several
traps of 2 types were operated in areas previously shown to be productive
(Thompson, 1973). The modified Manitoba Trap (MT) and modified
Animal Trap (AT) were improved for use under the windy, open and sun-
lighted conditions of the marsh. In order to reduce heat within the col-
lection containers of the traps and thereby to reduce fly mortality and to
enable feeding by the flies held within them, the top walls or ceilings of
the containers were replaced by % in.-mesh hardware cloth.

Previous trapping experience with the coastal marsh species at West Bay
in 1971 indicated that the AT was more collective than the MT (i.e., it
collected more species in larger numbers). Also to advantage, the larger

VOLUME 80, NUMBER 4 619

size of the collection cage easily accommodated the many flies required by
the work. Finally, the AT collection container made a more suitable cage
for permanent retention of the flies in the laboratory, both because of the
dimensions and because of the nontoxic nature of the construction
materials (the styrene of MT collection containers is known to show
insect toxicity ).

Art ficial feeding in the field—Flies were fed in the field immediately
after they entered the traps, or soon thereafter, with whole beef blood
treated with sodium citrate (10 g/gal). This blood was offered to the
insects using animal membranes manufactured from lamb intestine; al-
though this material is commercially available in sheets, using the material
in the form of prophylactics allowed the membranes to be used as con-
tainers for the blood. Fourex Natural Skins! (Schmid Laboratories, Inc.,
Little Falls, New York 07424) were each filled with 30 ml of citrated
beef blood and then closed with an overhand knot near the open end.
This quantity of blood was more than ample to feed many flies and also
expose a large surface of the prophylactic across the screened ceiling of
the collection container. In the field, the blood was warmed on the hot
radiator of the vehicle which transported the collector between trap sites.

Before the blood-filled prophylactics were placed on the top of the AT
collection cage, the cage was enveloped by a corrugated cardboard box
which contained a rectangular opening, 3 in. < 8 in., centered in the ceiling
of the box. This opening was intended to serve the same function as the
aperture beneath the collection cage in the top of the trap frame—to
concentrate the insects by using a positive response to light. Then, when
trapped female flies moved to the screened cage surface beneath the
opening in the cardboard box, the prophylactics were placed above them
on the top of the cage.

Feeding periodicity and optimal time of collection—Mariculturists
working shrimp ponds in the study area offered valuable suggestions on
the feeding periodicity of tabanids in the marshes. Considering the primary
objective of the work at the time—the collection of avid flies in large
numbers, rather than the collection of data on diurnal rhythms—their sug-
gestions were most helpful. These men described predawn attacks by
annoying numbers of flies; the men had attempted, then, to avoid the
summer sun through an early work day. As we soon discovered, the
magnitude of feeding attacks and trap catches from before dawn to mid-
morning, substantiated their reports and greatly enhanced the efficiency

£ the work.

The short periods of intense biting activity were most efficiently utilized
by feeding each cage of flies after the cage was taken off the trap frame.
In this manner, flies were being trapped continuously without interruption.

620 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(Activity at dusk increased over that at mid-day but did not become as
intense as that at sunrise.) After removal of each AT cage, the opening in
the bottom of the cage—a funnel-shaped baffle made of cellulose acetate—
was sealed with wadded tissue paper, and the cage was transported to the
lab within an insulated picnic chest having convenient dimensions for this
purpose (Thermos Cooler No. 7719'; King-Seeley Thermos Co., Norwich,
Connecticut 06360). The insulating capacity of these containers was ade-
quate for safe retention of the flies below critically high temperatures, and
the insects could be returned to the laboratory in darkness, thereby prevent-
ing stress during transport.

Init‘al handling of the catch.—Af‘ter collection and return of the catch,
flies were knocked down by chilling them in a refrigerator freezer com-
partment for 10 min. or by anesthesia with N.. The former method proved
more effective for the large cages being used. Knock-down enabled
identification, and subsequent separation by species, in order to collect
data useful in subsequent trapping and to determine which species were
accepting blood and which species laid what eggs. After definitive deter-
mination of egg masses later became possible (Thompson and Holmes,
1979), trapped insects were maintained in their respective AT cages until
they died.

Feeding success—During early collections, flies were examined for the
presence of blood after anesthesia and identification. At this time, the
abdominal segments of heavily engorged specimens were distended and
rounded and the brownish integument became blackish. In some speci-
mens not feeding to repletion, the reddish coloration of fresh blood could
be recognized in the middle of the abdomen by shining the light from a
microscope lamp through it. Additional study of females which were
apparently unengorged, showed the presence of blood that was not
detected by gross examination or by light transmission; this blood was
observed on the tip of a straight pin which was used to puncture the
hindgut through the posterior abdominal segments of dead females.
Table 2 shows the results of initial (field) feeding by three coastal marsh
species taken in collections of three dates.

Engorgement and mortality—Feeding ratios were observed in three
catches but the mortality of engorged vs. unengorged insects was not re-
corded because of more pressing priorities at the time. Nevertheless, the
large percentages of engorged females which failed to survive feeding,
transportation, or anesthesia seemed unusual.

Laboratory maintenance of adults.—Flies were initially retained in half-
pint ice cream cartons in order to encourage feeding on _blood-filled
prophylactics and to confine them in a chamber where light intensity and
temperature could be controlled. Later, flies were held in cages in the

VOLUME 80, NUMBER 4 621

Table 2. Percentages of Tabanus females feeding after collection, three samples,
West Bay, 1977.*

19 July 26 July 18 Aug.
td POETS A ae Pie aes Totals
Species ate = a = ae = Fed (+)
T. nigrovittatus
Macquart” 4 2 4 3 1 19 9 (27.3%)

T. lineola var.
hinellus Philip 18 22, 38 Te 47 63 103 (39.6% )

* Sixteen of 60 T. acutus females (26.7% ) taken 9 August fed.
"Punch test with a pin showed that an additional 12 females taken on 18 August had

fed.

lab when females failed to oviposit in cartons and when the control of
light and temperature no longer seemed necessary to feeding or to oviposi-
tion. In addition, catches were no longer being anesthetized for identifica-
tion so that transfer to smaller cages was unnecessary. At this time, then,
specimens were retained in AT cages on benches in the lab. Ambient light
from two large windows in one wall determined the natural photoperiod of
the season. Thermometers and hygrothermographs operated in the lab
throughout this period recorded temperatures of 75°F +5 and RH of
65% +5, respectively. To provide any additional moisture required,
4 in. X 4 in. cotton gauze pads placed atop the cages were continuously
saturated with distilled water from an inverted pint Mason Jar above them.
Only on rare occasions was a fly observed on or near this surface; there-
fore, ambient humidity in the room, humidity which was determined in
great part by the season, was adequate under the conditions observed here.

A 10% sucrose (cane sugar) solution was provided continuously and
blood was offered daily, using the technique previously described, except
that prophylactics of blood were heated by immersing them in containers
of warm water.

Longevity —Under the conditions described above, survivorship of speci-
mens taken in one sample is shown in Fig. 1. This figure shows that the
rate of mortality was fairly constant at all age levels for the three species
considered. This curve corresponds to Slobodkin’s Type III curve (Slobod-
kin, 1962), a plot line becoming straight when the ordinate scale is
logarithmic. The relative regularities of the graph lines for these three
species reflect, in great part, the corresponding sizes of the sample popula-
tions; with Tabanus nigrovittatus, a species represented by 410 specimens
in this sample, the mortality rate was more constant than with the less
numerous species.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

100

Tabanus nigrovittatus

410 22

al) (0) Xe)

90

BO

70
4 GO Tabanus lineola var. hinellus
o
Se s.0 ) PEELS
>
-
s
n

percent

Tabanus acutus

3199

9900009000000000

survivorship in days

VOLUME 80, NUMBER 4 623

Conclusions

The techniques and materials used here for collecting, feeding and
maintaining Tabanus nigrovittatus, T. lineola var. hinellus and T. acutus
will enable further research in rearing these species. Many references
considering collection devices for these and other coastal marsh species of
eastern North America have been cited by Axtell (1976). Bearing collect-
ing methodology in mind, the present work and that conducted 6 years
before (Thompson, 1973), show that the AT and the MT are highly
effective for attracting and trapping T. nigrovittatus in large numbers. The
efficiency of collecting methods for the less numerous species considered
here, such as the potential of the MT for taking T. lineola var. hinellus
(Thompson and Gregg, 1974), needs further research.

Feeding of many species of Tabanidae on a great variety of vertebrate
hosts has been attempted primarily in conjunction with animal disease
transmission experiments. These studies, exhaustively reviewed by Krinsky
(1976), have usually dealt with small numbers of insects—numbers neces-
sary only to effect the transfer of pathogens from carriers to a small number
of uninfected susceptibles. Because of this priority, rather than the en-
gorgement of large numbers of flies intended for egg production, these
studies offer little quantitative information on feeding methods and _ host
acceptance. Moreover, fundamental studies of the basic biology of some
species, studies dealing with engorgement and oviposition per se, such as
those of Hafez et al. (1970), Jones and Anthony (1964), Roberts (1966),
Schwardt (1936), Singh (1967), Webb and Wells (1924) and Wilson
(1967), have not offered much insight into the relevant factors which
stimulate feeding under conditions of retention and confinement. Also,
these studies reported poor feeding results or did not present quantitative
information on the numbers of flies exposed to the hosts or the percentages
of those numbers feeding. In addition, papers reporting prominent success
with artificial methods are very rare. Hafez et al. (1970) noted that
Tabanus taeniola Palisot de Beauvois females, in contrast to other Egyptian
species, fed to repletion on citrated calf blood. During studies of blood
meal volume and digestive enzymes, Thomas and Gooding (1976)
noted that several species of Chrysops and Hybomitra accepted warmed
defibrinated beef blood through Silverlight membranes.

Results with beef blood-filled prophylactics used in this study were
markedly negative or positive. Approximately 25-50 adult females of

Fig. 1. Survivorship of three coastal marsh tabanids, taken at West Galveston Bay,
July and August, 1977.

624 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Tabanus fuscicostatus Hine, T. lineola lineola, T. melanocerus Wiedemann,
T. proximus Walker, T. subsimilis sub:imilis and T. sulcifrons each re-
jected this combination while the same numbers, or more, of T. mularis,
T. abactor Philip and the three coastal marsh species considered here,
readily accepted it. In most instances, the insects had come to traps baited
with dry ice; and at least some individuals of each of these species were
offered blood-filled prophylactics on the tops of trap collection containers
very soon after being taken. These data, as well as those of Thomas and
Gooding (1976), suggest membranes hold potential success in feeding this
group of Diptera, as they have for tsetse species and mosquitoes. On the
other hand, results with free blood could be much less favorable; none of
the species that we exposed to citrated beef blood on cotton, including
Hybomitra lasiophthalma, fed.

In studies of the longevity of several hundred caged Tabanus nigrovit-
tatus females in the laboratory in Delaware, Olkowski (1966) expressed
survivorship in “mean fly days” (12.3 days for T. nigrovittatus ). Compar-
ing his data with ours is not very meaningful because he excluded 43% of
his population sample because these females survived less than 6 days.
Secondly, he presents no expression of variation in this population.

In any event, the survivorship that we observed in this the present
work, as the collecting and feeding results that we also experienced, will
enable study of further consecutive events in the life histories of these
species.

Acknowledgments

We gratefully acknowledge the help of Mr. Joseph W. Holmes, Jr. for
feeding and maintaining tabanids; of the Drs. B. J. Cook of this laboratory,
Roger Meola of Texas A&M University and L. L. Pechuman of Cornell
University for their comments on the manuscript; also of the Messrs. B. F.
Hogan and G. E. Spates for their comments; and of the Messrs. Hoyt Hol-
combe and Jim Herrold, Brazoria County Mosquito Control District,
Angleton, Texas, for their observations on fly feeding periodicity.

Literature Cited

Axtell, R. C. 1976. Coastal. horse flies and deer flies (Diptera: Tabanidae). pp.
415-445. In L. Cheng (ed.) Marine Insects. North-Holland Publishing Co.

Fairchild, G. B. 1937. A preliminary list of the Tabanidae (Diptera) of Florida. Fla.
Entomol. 19:58-63.

Hafez, M., S. El-Ziady and T. Hefnawy. 1970. Studies on the feeding habits of
female Tabanus taeniola P. de B. (Diptera: Tabanidae). Bull. Entomol. Soc.
Egypt 54:365-376.

Jones, C. M. and D. W. Anthony. 1964. The Tabanidae (Diptera) of Florida. U.S.
Dep. Agric. Tech. Bull. No. 1295. 85 pp.

VOLUME 80, NUMBER 4 625

Krinsky, W. L. 1976. Animal disease agents transmitted by horse flies and deer flies
(Diptera: Tabanidae ). J. Med. Entomol. 13:225-275.

Olkowski, W. 1966. Biological studies of salt marsh tabanids in Delaware. MS
thesis, Univ. Delaware. Mimeo, 107 pp.

Philip, C. B. 1965. Family Tabanidae. pp. 319-342. In A. Stone et al. A Catalog
of the Diptera North of Mexico. USDA Handbook No. 276. 1696 pp.

Roberts, R. R. 1966. Biological studies on Tabanidae I. Induced oviposition. Mosq.
News 26:435—-438.

Schwardt, H. H. 1936. Horseflies of Arkansas. Arkansas Agric. Exp. Stn. Bull. No. 332.

Singh, S. 1967. Life-history and biology of Tabanus rubidus. Res. Bull. Punjab Univ.
Sci. 19:113—122.

Slobodkin, L. B. 1962. Growth and Regulation of Animal Populations. Holt. 184 pp.

Stone, A. 1938. The horseflies of the Subfamily Tabaninae of the Nearctic Region.
U.S. Dep. Agric. Mise. Publ. No. 305. 171 pp.

Thomas, A. W. and R. H. Gooding. 1976. Digestive processes of hematophagous
insects. VIII. Estimation of meal size and demonstration of trypsin in horse flies
and deer flies (Diptera: Tabanidae). J. Med. Entomol. 13:131-136.

Thompson, P. H. 1973. Tabanidae (Diptera) of Texas. I. Coastal marsh species,
West Galveston Bay; incidence, frequency, abundance, and seasonal distribu-
tion. Proc. Entomol. Soc. Wash. 75:359-364.

Thompson, P. H. and E. J. Gregg. 1974. Structural modifications and performance
of the modified Animal Trap and the modified Manitoba Trap for collection
of Tabanidae (Diptera). Ibid. 76:119-122.

Thompson, P. H. and J. W. Holmes, Jr. 1979. Rearing of Texas Tabanidae (Diptera).
II. Oviposition and larval development of coastal marsh species. Ibid. In press.

Thompson, P. H. and L. L. Pechuman. 1970. Sampling populations of Tabanus
quinquevittatus Wiedemann about horses in New Jersey, with notes on the
identity and ecology. J. Econ. Entomol. 63:151—155.

Webb, J. L. and R. W. Wells. 1924. Horse-flies: biologies and relation to western
agriculture. U.S. Dep. Agric. Dept. Bull. No. 1218. 36 pp.

Wilson, B. H. 1967. Feeding, mating, and oviposition studies of the horse flies
Tabanus lineola and T. fuscicostatus (Diptera; Tabanidae). Ann. Entomol. Soc.
Am. 60:1102-1106.

Veterinary Toxicology and Entomology Research Laboratory, Fed. Res.,
Sci. and Educ. Admin., USDA, College Station, Texas 77840.

Footnote

*This paper reports the results of research only. Mention of a commercial or
proprietary product does not constitute a recommendation for use by the U.S. Depart-
ment of Agriculture.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 626-654

THE GENUS TYCHIUS GERMAR (COLEOPTERA:
CURCULIONIDAE): LARVAE AND PUPAE OF SOME
SPECIES, WITH EVALUATION OF THEIR
CHARACTERS IN PHYLOGENETIC ANALYSIS?

Wayne E. Clark, Horace R. Burke and Donald M. Anderson

Abstract—Larvae of eight species and pupae of five species of North
American Tychius are described. Of these, only the larvae of T. picirostris
(Fabricius) and T. stephensi Schonherr, have been previously described.
Larvae of the European T. quinquepunctatus (L.) and T. flavicollis
Stephens, also described, are compared to larvae of the North American
species and to other European species known from previously published
descriptions. Larval and pupal characters appear to support some group-
ings previously postulated on the basis of morphological characters of
adults and host plant relationships. However, because of the lack of
structural complexity in known characters of Tychius immatures, and the
apparent lack of major differences between many of the species in these
stages, it appears that the larval and pupal characters do not provide
strong evidence of phylogenetic relationships. Superficial similarity due to
parallelism or convergence appears likely in some cases. The hypothesis
of polyphyly of the genus Miccotrogus Schonherr is not refuted by charac-
ters of the immature stages of some of the species formerly assigned to
that taxon.

Since revision of the North American species of Tychius (Clark, 1971),
the senior author has continued studies of members of that genus and of
the related genus Sibinia. A classification of the subfamily Tychiinae
(Clark et al., 1977), including the genera Tychius and Sibinia, provides
a framework for research on the group. Relationships of North American
Tychius to members of the much larger Old World, primarily Palearctic,
Tychius fauna are discussed by Clark (1976, 1977). A revised classification
of native North American Tychius and host and distributional records for
these are also included in the 1977 paper. Clark and Burke (1977) present
notes on life histories and habits of native North American Tychius and
review biology of world members of the genus which consists of more than
300 nominate species, several of which are of economic importance (see
Muka, 1954; Hoffmann et al., 1963; Yunus and Johansen, 1967; Nasredinov,
1975). Similar information on members of the genus Sibinia is presented
by Clark (1978a).

Most published descriptions and illustrations of immature stages of
members of the genus Tychius are not sufficiently detailed to be taxo-

VOLUME 80, NUMBER 4 627

nomically useful. Exceptions are those of T. quinquepunctatus (L.) by
Grandi (1916) and Scherf (1960), and T. flavus Becker by Servadei
(1947). Scherf (1964) also assembled information from numerous sources
on bionomics and immature stages of Curculionoidea, including several
species of Tychius, of Central Europe. These and other publications (see
Burke and Anderson, 1976) have been searched for taxonomic characters
of Tychius larvae and pupae, and these characters are compared with the
ones revealed by our own observations. In this paper we describe larvae
of 6 of the 12 known native North American species of Tychius and pupae
of 4 of these, larvae of 2 European species introduced into North America
and the pupa of one of these, and larvae of 2 other European members of
the genus. The usefulness of the larval and pupal characters described
in recognizing monophyletic groups within the genus has also been
analyzed.

Materials and Methods

As Ahmad and Burke (1972) and Burke and Anderson (1976) pointed
out, systematic studies of immature Curculionidae, as well as of other
Coleoptera, have not progressed as rapidly as studies based on adults.
This is due in part to the relative difficulty encountered in obtaining
specimens of the immature forms and the more time-consuming procedures
necessary to prepare larvae for study. Timing is the critical factor in ob-
taining Tychius larvae and pupae. Larvae suitable for taxonomic study
are available for only a short period when fruits of their hosts, members of
the legume subfamily Papilionoideae, are mature. Once infested fruits are
found, however, larvae may be obtained relatively easily by placing the
pods in porcelain pans. When larvae emerge from infested pods they may
be transferred to a mixture of sand and peat moss in widemouth glass jars.
The larvae tunnel into the substrate, pupate and complete their develop-
ment to the adult stage there (see Clark and Burke, 1977).

Larvae of 8 and pupae of 5 of the 14 species of Tychius known to
occur in North America have been collected. Larvae and pupae were
preserved in 70% ethyl alcohol; a few reared adults were also preserved
with the immatures to facilitate subsequent identification. When larvae
were collected but pupae and adults were not subsequently reared, identifi-
cation was based upon association with adults taken on plants from which
the larvae were obtained.

Larvae of some species were obtained for study through loans from the
Ohio State University (OSU), the U.S. National Museum (USNM), and
from Dr. C. A. Johansen, Washington State University (collections are
referred to in the text by the abbreviations in parentheses); specimens
collected by Clark are deposited in the collections of the Department of

628 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Entomology, Texas A&M University (TAM) and in the USNM. For some
of the borrowed material, larvae and pupae were identified by association
with accompanying adults, even when there was no direct evidence that
the adults were reared from larvae from the same lot. When no adults
were present, reliance was placed upon determination labels with the im-
matures, even though there was usually no indication as to how these
determinations were made.

Pupae and entire larvae were examined in alcohol, but examination of
most Jarval characters required that the specimens be slide-mounted. Ex-
cept for a few modifications, slides were prepared by the procedures
described by Ahmad and Burke (1972). It was found to be easier to re-
move the mouthparts from the head capsule before removing the latter from
the body. Mounting and examination were also easier when the 2 sides
of the larval skin remained broadly intact mid-dorsally than when the two
halves were completely separated. Hoyers was used as the mounting
medium. Drawings were made by procedures outlined by Ahmad and
Burkey(Glg72)).

Anderson (1947) is followed for terminology of larval characters. Where
applicable, pupal terminology proposed by Burke (1968) is used. In the
text the setae on only 1 side of the body are described.

Descriptions
Genus Tychius Germar

Larva—Body (Fig. 8): Elongate, slightly curved; integument rugulose
in large species, smooth in smaller ones; larger species with transverse rows
of fine asperities on ventral portions of thoracic and Ist few abdominal
segments. Head (Figs. 1-7): Free, wider than long, narrowed slightly
anteriorly; anterior ocelli present. Antenna with membranous basal article
bearing a blunt to elongate, acute accessory appendage and several minute
sensory processes. Hypopharyngeal bracon distinct. Frontal sutures distinct
throughout length, incomplete anteriorly. Epicranial suture less than % as
long as head capsule. Frons bearing 3 pairs of setae, setae 1 and 2 absent,
seta 3 short to minute, seta 4 long, but shorter than seta 5; 2 pairs of frontal
sensilla, | pair between frontal setae 3 and 4, the other anterior, lateral, or
posterior to seta 3. Dorsal epicranial seta 1 minute or long, setae 4 and 5
long, seta 3 short, located on or slightly posterior to frontal suture between
setae 1 and 4, seta 2 absent; 1 sensillum located between dorsal epicranial
setae 4 and 5 on each side, another just posterior to seta 1. Lateral
epicranium with 2 pairs of setae, seta 1 short, seta 2 longer; 1 pair of
sensilla located anterior to seta 1. Ventral epicranium with 2 pairs of short
setae. Posterior epicranium with 2 or 3 pairs of minute setae and 2 pairs of

VOLUME 80, NUMBER 4 629

Figs. 1-3. Dorsal views of head capsules of larval Tychius. 1, T. sordidus. 2, T.
lineellus. 3, T. quinquepunctatus.

sensilla. Clypeus wider than long, with 2 pairs of minute setae and 2
sensilla, each 1 of which is located between or slightly distad of clypeal
setae on each side. Labrum (Figs. 1-3) with 3 pairs of setae, seta 3 shorter
than seta 2; labral sensilla present, median labral sensillum present or
absent. Epipharynx (Fig. 5) with 2 anterolateral setae on each side, 6

630 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 4-7. Mouth parts of Tychius sordidus. 4, Left mandible, dorsal view. 5,
Epipharynx. 6, Labium, ventral view. 7, Right maxilla, ventral view.

anteromedian setae, and 4 medium spines; 2 pairs of epipharyngeal sensilla
present. Labral rods short, stout, widely separated. Mandible (Fig. 4)
with 2 apical teeth, 2 mandibular setae and 1 sensillum. Maxillary palpus
(Fig. 7) consists of 2 articles; apical article with 1 sensillum and an apical
cluster of minute papillae; basal article shorter than apical article, bearing
1 seta, 2 sensilla. Labium (Fig. 6), palpus consists of 1 article; premental
sclerite with anterior and posterior median extensions obsolete; prementum

VOLUME 80, NUMBER 4 631

Figs. 8-11. Lateral views of larval Tychius. 8, T. sordidus. 9, T. stephensi (thoracic
and Ist adbominal segments). 10, T. tectus (thoracic and Ist abdominal segments).
11, T. tectus (thoracic and Ist abdominal segments ).

with 1 pair of long setae and 1 pair of sensilla; glossa with 1 or 2 pairs of
short to minute setae and 2 or 3 pairs of medial sensilla, and 1 sensillum
located near base of each labial palpus; postmentum with 2 or 3 pairs of
setae, seta 1 short or absent, seta 2 long, seta 3 shorter than 2. Thorax
(Figs. 8-11): Not narrower than abdomen. Pronotum with 7-8 minute to
long setae and 2 minute setae on anterolateral margins. Thoracic spiracle
unicameral or bicameral. Prodorsum and epipleural lobe of mesothorax
and metathorax each with 1 long seta. Pleural area of prothorax with 2

632 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

20mm ;

Fig. 12. Pupa of Tychius sordidus, ventral view on left, dorsal view on right.

long setae, and with 3 minute setae on extreme anteromedian margin; pleural
area of mesothorax and metathorax each with 1 long seta. Sternum of each
thoracic segment with 1 moderately long seta and 1 minute seta. Abdomen
(Figs. 8-11): 8 pairs of lateral, unicameral spiracles. Segments I-VII each
with 3 dorsal folds, prodorsal fold more prominent than postdorsal; segment
VIL with 2 dorsal folds. Segments I-VII each with 1 long prodorsal seta
and 2-3 postdorsal setae. Epipleurum with 1 long and 1 shorter seta.
Pleural area with 1 long and 1 shorter seta. Pedal areas each with | long
seta. Eusternum with 2 long setae. Segment IX with 1 postdorsal seta, 1
short and_1 longer epipleural setae, 1 short and 1 longer pleural setae and
1 short and 1 longer eusternal setae. Sternellum distinct. Anus terminal,
anal folds bearing a few minute setae.

Pupa (Fig. 12).—Head: Distirostral setae absent. Two pairs of basirostral
setae, 1 pair located just distad of ocular area, the other directly over the
point of antennal insertions, proximal pair usually longer than distal pair,
each borne on summit of rounded to conical tubercle. One pair of in-
terorbital setae, each seta borne on summit of conical tubercle; interorbitals
longer and stouter than basirostrals. Supraorbital setae absent. One or

VOLUME 80, NUMBER 4 633

2 pairs of frontal setae present, each seta borne on summit of conical
tubercle; frontals about same length as interorbitals. Pronotum: 1 pair
of anteromedian setae; 2 pairs of anterolateral setae; 1 pair of discal setae
located directly posterior to anteromedian setae, sometimes more widely
separated than anteromedians; 1 pair of posteromedian setae, and 1 or 2
pairs of posterolateral setae. Mesonotum and Metanotum: Anteronotal
setae absent. One to 3 pairs of posteronotal setae; when 2 or more pairs
present, setae on each side close together, borne on separate tubercles or
on common tubercle; 3rd seta when present not borne on_ tubercle.
Abdomen: Anterotergal setae absent. One to 3 pairs of discotergal setae,
each seta borne on summit of conical tubercle, tubercles becoming larger
posteriorly. One pair of laterotergal setae on terga 1-7 or 8, each seta
borne on summit of low, indistinct tubercle. Segment 9 with a pair of
short, sharply pointed, widely separated posterior processes. Abdomen
devoid of ventral setae. Femora: Bearing 1 or 2 setae, or without setae;
when only 1 seta present it is borne on outer face near apex, 2nd seta if
present borne on dorsal surface near apex.

Material examined.—The larval description is based upon examination
of larvae of T. sordidus LeConte, T. lineellus LeConte, T. tectus LeConte,
T. semisquamosus LeConte, T. soltaui Casey, T. prolixus Casey, T. quin-
quepunctatus, T. stephensi Schonherr, T. picirostris (Fabricius) and T.
flavicollis Stephens, and published descriptions of T. flavus (Servadei,
1947; Scherf, 1964), T. meliloti Stephens, and T. crassirostris Kirsch (Scherf,
1964). The pupal description is based on examination of pupae of T.
sordidus, T. tectus, T. soltaui, T. prolixus and T. picirostris and on published
descriptions of T. quinquepunctatus (Scherf, 1964), T. flavus (Servadei,
1947; Scherf, 1964) and T. meliloti and T. crassirostris (Scherf, 1964).

Discussion—kKnown Tychius larvae are distinguished from larvae of
Sibinia sulcatula (Casey) (Rogers et al., 1975), and S. sodalis Germar
(Scherf, 1964), by the following combination of characters: Dorsal
epicranial seta 1 long (except T. picirostris); dorsal epicranial seta 3 short,
located on or slightly posterior to frontal suture between dorsal epicranial
setae 1 and 4; epipharynx with 2 instead of 3 sensory pores; mandible with
2 rather than 3 apical teeth; abdominal prodorsal seta long; spiracular seta
2 long. Known pupae of Tychius members can be distinguished from the
pupa of S. sulcatula by the number of posterolateral pronotal setae; 1 or 2
in Tychius, 4 in S. sulcatula. Scherf's (1964) description of the pupa of
S. sodalis does not clearly indicate the number of posterolateral setae.

The larvae and pupae of the species listed above under the material
examined heading are each described below; characters cited in the generic
description are omitted. The diagnostic characters of the larvae are listed
in Table 1, those of the pupae in Table 2.

634 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Summary of selected larval characters showing distribution among some

species of Tychius.

) al) g ~n ® 3 = 3 2 oy
3 = a” > 2 ‘S = = = rz) S 8 2) iS
Larval Character eel soos s = SS.) SoS eae
States See S88, 8 io Syne Se wath. eee
Dorsal short xX
epicranial
seta 1 long X Xe Ee ee i es X
Median present XT SXVex xX Xe EXO xX
labral
sensillum absent Xx
Dorsal 3 X XX
malar
setae 4 Xe We XOX OX EX te
Sensilla 1 xX > Se 5), a, Gee, G Py DS ed, «
labial
palpus 2; X DG 2K
Pairs Ht XS EEX Xe XO XT “GX X
glossal
setae 2, X x Pex
Long 5 ke ek Dk
pronotal
setae 4 XV KOE
Thoracic unicameral x xe OS
spiracle bicameral XRF XS XE CREEK AT MX HOXG X
Thoracic ® Xo XG
postdorsal
setae 3 XN ENS OG TOXIN 1 EXCTLIOX: XX
Alar short Kent MEXIAIIXS OU EIKO ES” UXO E XEROX
setae long XxX
Epipleural — short Ke PX XT XE NO ee CNG
seta
2 long xX
Pleural 1 xX
setae 2 TPXGRE NCHS WXEL EXT SEX EE NGERIONG

Tychius sordidus LeConte

Figs. 1, 4-8, 12
Tychius sordidus LeConte, 1876:217.

Larva—Body: 4.6-6.6 mm long (40 larvae); shining, yellow. Head
(Fig. 1): Uniformly dark yellowish brown; truncate posteriorly; width of

VOLUME 80, NUMBER 4 635

Table 2. Summary of selected pupal characters showing distribution among some
species of Tychius.

ae)
3 Sees aa z s = a
kg USM ete Seon: anual ole ts tire
Pupal Character SS SES ESS 3 Suen 5 = 2
States Oe SS aS er ee
Anterolateral 2 Keg fXh pee Xie ea WX X
setae of pronotum
(pairs ) 3 ? ? iF
Posterolateral 1 X iF
setae of pronotum
( pairs ) 2 MEX ke x OX
Posteronotal 1 xX Xx
setae of pronotum
( pairs ) 1-2 DX FXO OK
Discotergal 1 DS RS, Maa X X
setae of abdomen
( pairs ) 2 X P
Femoral 1 X X On ex X
setae wy) xX xX XxX

head capsule 0.82-0.94 mm (8 larvae). Ocellar areas lacking subcutaneous
pigment. Accessory appendage of antenna robust, sides rounded. En-
docarina more than % as long as frons. Dorsal epicranial seta 1 long.
Median labral sensillum present. Malar area of maxilla (Fig. 7) with 5
ventral and 4 dorsal setae. Labial palpus (Fig. 6) with 1 sensillum; glossa
with 1 pair of minute setae; postmental seta 1 present. Thorax (Fig. 8):
Pronotum with 5 long setae, 2 moderately long setae, 1 short seta and 2
minute anterolateral setae. Spiracle bicameral, air tubes each with 5 to 8
annuli. Postdorsum of mesothorax and of metathorax each with 3 long
setae. Pedal area of each thoracic segment with 3 long and 3 minute to
moderately long setae. Sternal seta slightly shorter than ventralmost pedal
seta of the same segment. Abdomen (Fig. 8): Postdorsum with 3 long
setae of subequal length. Spiracular seta 1 long, seta 2 shorter. Epipleural
seta 1 slightly shorter than seta 2. Pleural seta 2 slightly shorter than seta 1.

Pupa (Fig. 12)—Length: 3.9-5.0 mm (25 pupae). Head: Distalmost
pair of basirostral setae absent in some specimens. Pronotum: 2 pairs of
posterolateral setae. Mesonotum and Metanotum: 1-3 pairs of posteronotal
setae. Abdomen: 2 pairs of discotergal setae on terga 1-5; terga 6 and 7
often each with 3 pairs of setae, tergum 8 usually with 1 pair, but some-
times with 2 pairs of discotergals. Legs: Femur with 1 seta or without a
seta.

636 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Material examined.—Fifty larvae (40 entire, 10 slide-mounted speci-
mens, TAM, USNM), which emerged from pods of Baptisia leucophaea
Nutt. var. laevicollis (Gray) Small, collected 10 May 1973, at College
Station, Brazos County, Texas, and 25 pupae (TAM, USNM), laboratory
reared from larvae from the same pods, all determined by association with
reared adults.

Discussion.—Tychius sordidus occurs in the eastern and midwestern
United States. Its known hosts are all members of the plant genus Baptisia
(Clark, 1971; Clark and Burke, 1977). The larva is distinguished from that
of other members of the genus by the following combination: Alar area of
each thoracic segment with 1 long seta; mesothorax and metathorax each
with 3 long postdorsal setae of subequal length. The pupa is distinguished
as follows: Femoral setae 1 or none; posteronotal setae 1-3 pairs; dis-
cotergal setae 2 pairs.

Tychius lineellus LeConte
Fig. 2

Tychius lineellus LeConte, 1876:217.

Larva.—Body: 5.7-6.8 mm long (3 larvae); opaque, pale yellowish
white. Head (Fig. 2): Uniformly dark brown; emarginate posteriorly;
width of head capsule 0.74-0.854 mm (5 larvae). Ocellar areas with sub-
cutaneous pigment. Accessory appendage of antenna moderately long,
sides slightly rounded. Endocarina more than % as long as frons. Dorsal
epicranial seta 1 long. Labral setae 1 and 2 subequal in length. Median
labral sensillum present. Malar area of maxilla with 5 ventral and 4 dorsal
setae and 1 ventral sensillum. Labial palpus with 2 sensilla; glossa with 1
pair of minute setae; postmental seta 1 present. Thorax: Pronotum with
5 long setae, 2 shorter setae and 1 minute seta, in addition to 2 minute
anterolateral setae. Spiracle bicameral, air tubes with 5 to 8 annuli.
Postdorsum of mesothorax and of metathorax with 3 setae, seta 3 long,
seta 2 very short, seta 1 intermediate in length. Pedal area of each thoracic
segment with 3 long and 3 minute to moderately long setae and a few
sensilla. Sternal seta slightly shorter than ventral-most pedal seta of the
same segment. Abdomen: Postdorsum with 3 setae, setae 1 and 3 slightly
shorter than seta 2. Spiracular seta 1 long, seta 2 slightly shorter. Epipleural
seta | slightly shorter than seta 2. Pleural seta 2 slightly shorter than seta 1.

Pupa.—Unknown.

Material examined.—Eight larvae (3 entire, 5 slide-mounted specimens,
TAM, USNM), which emerged from pods of Lupinus leucophyllus Dougl.,
collected 6 July 1968, 5 miles east of Springville, Utah County, Utah,
determined by association with adults collected on the plants.

Discussion —Tychius lineellus occurs in the western United States and

VOLUME 80, NUMBER 4 637

adjacent portions of Canada. Its known hosts are members of the plant
genus Lupinus (Clark, 1971). The larva is distinguished from larvae of
other species of Tychius by the following combination: Labial palpus with
2 sensilla; mesothorax and metathorax each bearing 3 setae, setae 1 and
2 short, seta 3 long. It also has a sensillum on the malar area of the
maxillary palpus, as does the Old World T. quinquepunctatus. Adult
characters do not indicate a close relationship between T. lineellus and
T. quinquepunctatus, however.

Tychius tectus LeConte
Fig. 10

Tychius tectus LeConte, 1876:217.
Tychius mixtus Hatch, 1971:355.

Larva.—As described for T. sordidus, except as follows. Body: 5.5-
5.8 mm long (8 larvae); pale yellowish white to yellow. Head: Uniformly
light yellowish brown; posteriorly truncate; width of head capsule 0.60-
0.67 mm (4 larvae). Ocellar areas lacking subcutaneous pigment. Accessory
appendage of antenna short, sides rounded. Endocarina distinct, more
than % length of frons. Dorsal epicranial seta 1 long. Labral seta 1 longer
than seta 2. Median labral sensillum present. Malar area of maxilla with
5 ventral and 4 dorsal setae. Labial palpus with 1 sensillum; postmental
seta 1 present. Thorax (Fig. 10): pronotum with 5 long setae, 2 moderately
long setae, and | minute seta, in addition to 2 minute anterolateral setae.
Spiracle bicameral, air tubes with 4 to 6 annuli. Postdorsum of mesothorax
and of metathorax with 3 setae; setae 1 and 3 long, seta 2 minute. Pedal
area of each thoracic segment with 3 long setae, 1 short to minute seta, and
1 sensillum. Sternal seta long, subequal in length to ventral-most seta of
pedal area of same segment. Abdomen (Fig. 10): Postdorsum with 3
setae, setae 1 and 2 longer than seta 3. Spiracular seta 1 long, seta 2 very
short. Epipleural seta 1 much shorter than seta 2. Pleural seta 2 much
shorter than seta 1.

Pupa.—The pupa of T. tectus differs from that of T. sordidus in its
smaller size (length 2.9-3.4 mm) and by possession of the lowermost pair
of basirostral setae, 2 femoral setae, 1 or 2 pairs of mesonotal and metanotal
posteronotal setae and | pair of discotergal setae on terga 1-5.

Material examined.—Twelve larvae (8 entire, 4 slide-mounted specimens,
TAM, USNM), from pods of Astragalus distortus var engelmanii (Sheldon)
Jones, collected 5 May 1972, at College Station, Brazos County, Texas and
3 pupae (TAM, USNM), laboratory-reared from other larvae taken from
the same plants at the same time, all determined by association with
reared adults.

Discussion.—Tychius tectus is widely distributed throughout the Great

638 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Plains and Rocky Mountain regions of the western United States and
adjacent portions of Canada (Clark, 1971 and 1977). Its hosts, members
of the papilionoid tribe Astragaleae, include species of Astragalus in pri-
marily Eurasian groups, members of the same genus in autochthonous
American groups, and species of Oxytropis, a primarily Eurasian genus
closely related to Astragalus (Clark, 1977). The larva is distinguished from
larvae of other members of the genus as follows: Mesothorax and meta-
thorax each with 3 postdorsal setae, setae 1 and 3 long, seta 2 short;
typical abdominal segments also with 3 postdorsal setae, setae 1 and 2
long, seta 3 short. Distinguishing characters of the pupa are listed above.

The larvae and pupae of most of the other American Astragaleae-as-
sociated Tychius are either apparently identical to those of T. tectus, or
are distinguished only by minor, in some cases variable characters (these
are enumerated in the discussions of T. semisquamosus, T. soltaui and
T. prolixus, which follow). The close similarity of these species to each
other supports the contention that the Astragaleae-associated Tychius form
a monophyletic (possibly paraphyletic) group (Clark, 1977).

Tychius semisquamosus LeConte

Tychius semisquamosus LeConte, 1876:217.
Tychius lamellosus Casey, 1892:418.
Tychius squamosus Hatch, 1971:356.
Tychius intermixtus Hatch, 1971: 356.

Larva.—tThe larva of T. semisquamosus has subcutaneous pigment on
the ocellar areas and fine asperities on the thoracic and abdominal sterna
but cannot otherwise be distinguished from that of T. tectus.

Pupa.—Unknown.

Material examined.—Sixteen larvae (12 entire, 4 slide-mounted speci-
mens, TAM, USNM), from pods of Astragalus beckwithii T. & G., collected
1 June 1972, 2.5 miles southeast of Provo, Utah County, Utah, determined
by association with adults collected on the plants.

Discussion—Tychius semisquamosus occurs in the Rocky Mountain re-
gion of the western United States and adjacent portions of Canada, and in
southern California (Clark, 1971 and 1977). Its known hosts are all species
of Astragalus in native American groups (Clark, 1977).

Tychius soltaui Casey

Tychius soltaui Casey, 1892:416.
Tychius hirsutus Clark, 1971:30 (replacement name for T. hirtellus LeConte,
1876:218; not Tournier, 1873).

VOLUME 80, NUMBER 4 639

Larva.—The larva of T. soltaui is indistinguishable from that of T. tectus,
except for possession of subcutaneous pigment on the ocellar areas. It is
distinguished from the larva of T. semisquamosus only by the lack of
conspicuous asperities on the thoracic and abdominal sterna.

Pupa.—the pupa of T. soltaui differs from that of T. tectus by having
1 instead of 2 femoral setae.

Material examined.—Thirty-one larvae (20 entire, 11 slide-mounted
specimens, TAM, USNM), which emerged from pods of Astragalus
nuttallianus A., D.C., collected 10 April 1973, 0.7 miles southeast of San
Diego, Duval County, Texas, and from pods of A. emoryanus (Rydb.)
Cory, collected 31 May 1973, 35 miles south of Alpine, Brewster County,
Texas, determined by association with reared adults.

Discussion.—Tychius soltaui occurs in the western United States and
adjacent portions of Canada (Clark, 1971) and in the state of Durango,
Mexico (Clark, 1977). Its known hosts are species of Astragalus in native
American groups (Clark, 1977). The relationship of T. soltaui to the other
American species of Tychius associated with Astragalus was considered in
the discussion of T. tectus.

Tychius prolixus Casey
Tychius prolixus Casey, 1892:419.

Larva.—The larva of T. prolixus is indistinguishable from that of T.
tectus. It is distinguished from larvae of T. semisquamosus and T. soltaui
by the absence of subcutaneous pigment from the ocellar areas.

Pupa.—The pupa of T. proxilus differs from that of T. tectus only in
possession of 2 instead of 1 pair of meso- and metathoracic posteronotal
setae. Some of the specimens examined have 1, but most have 2 femoral
setae, as in T. tectus.

Material examined.—Twenty-one larvae (11 entire, 10 slide-mounted
specimens, TAM, USNM), which emerged from pods of Astragalus
utahensis (Torr.) T. & G., collected 1 June 1972, at the mouth of Rock
Canyon, northeast of Provo, Utah County, Utah, and from pods of A.
wootonii Sheldon, collected 30 May 1973, 6.5 miles east of the junction of
Highways 505 and 166, Jeff Davis County, Texas, and 15 pupae (TAM,
USNM), reared from larvae from the A. utahensis lot, all determined by
association with reared adults.

Discussion.—Tychius prolixus occurs in southern California, Nevada,
Utah, Arizona and western Texas (Clark, 1971). Its known hosts are
species of Astragalus which belong to native American groups (Clark,
1977). Significance of characters by which T. prolixus and other Astragalus-

640 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

associated species are distinguished from each other was considered in the
discussion of T. tectus.

Tychius quinquepunctatus L.
Fig. 3

Curculio 5-punctatus L. 1758:383.
Tychius 5-punctatus Germar, 1817:340.

Larva.—Body: 4.8-6.5 mm long (8 larvae); shining, yellow. Head
(Fig. 3): Uniformly dark brown; rounded posteriorly; width of head
capsule 1.23-1.30 mm (2 larvae). Anterior ocellar areas with minute spot,
or without subcutaneous pigment. Accessory appendage of antenna robust.
Endocarina more than % as long as frons. Dorsal epicranial seta 1 long,
located well posterior to frontal suture. Posterior epicranium with 3 pairs
of minute setae. Median labral sensillum present. Malar area of maxilla
with 5 ventral setae, 4 dorsal setae, and 1 ventral sensillum. Labial palpus
with 1 sensillum; glossa with 2 pairs of minute setae; postmental seta 1
present. Thorax: Pronotum with 5 long setae, 2 shorter setae, and 1 minute
seta in addition to 2 minute anterolateral setae. Spiracle bicameral. Post-
dorsum of mesothorax and of metathorax each with 3 setae, of which seta
1 is long, seta 2 shorter, seta 3 longer than 1. Pedal area of each thoracic
segment with 3 long and 2 short setae. Sternal seta long, subequal in
length to ventral-most pedal seta of same segment. Abdomen: postdorsum
with 3 long setae of subequal length. Spiracular seta 1 long, seta 2 shorter.
Epipleural seta 1 longer than seta 2. Pleural seta 1 longer than seta 2.

Pupa (from Scherf, 1964, Fig. 238) —Length: 5-5.3 mm. Head: 2 pairs
of frontal setae. Pronotum: Anterior-most pair of anterolateral setae
located beside anteromedian setae, 2nd pair displaced posteriorly, behind
subapical constriction of pronotum; 2 pairs of posterolateral setae present.
Mesonotum and Metanotum: each with only | pair of posteronotal setae.
Abdomen: Terga 1-8 each with 1 pair of discotergal setae. Legs: Femora
each with 2 setae.

Material examined.—Ten larvae (8 entire, 2 slide-mounted specimens,
USNM), labeled “Tychius quinquepunctatus, 209, Erlangen Rosenhauer,”
were examined. Identity of these specimens cannot be verified as there are
no associated adults.

Discussion.—This European species reportedly has hosts in the papilio-
noid genera Lathyrus, Phaseolus, Pisum and Vicia (Scherf, 1964). The
larva is distinguished from larvae of other known species of Tychius by the
following combination: Glossa with 2 pairs of minute setae; malar area of
maxilla with a ventral sensillum; postdorsum of mesothorax and of meta-

VOLUME 80, NUMBER 4 641

thorax with 3 setae, seta 1 long, seta 2 short, seta 3 longer than 1. The
pupa has 2 femoral setae, 1 pair each of mesonotal and metanotal setae,
and terga each with | pair of discotergal setae.

Larvae and pupae of T. quinquepunctatus have more in common with
larvae and pupae of T. sordidus, T. lineellus, and the Astragaleae-associated
species (T. semisquamosus, T. tectus, T. soltaui and T. prolixus) than with
those of the clover seed weevils (T. stephensi and T. picirostris) and that
of T. flavus (Table 1). The only other species of Tychius known to have
a sensillum on the malar area of the larval maxilla is T. lineellus; T.
picirostris is the only other known species with only | pair of mesonotal
setae. Adult characters do not indicate that T. quinquepunctatus is closely
related to either of these, however, and it is likely that both similarities are
due to convergence.

? Tychius flavicollis Stephens
Tychius flavicollis Stephens, 1831:56.

Larva—As described for T. tectus, except as follows. Body: White.
Head: Width of head capsule 0.52 mm (1 larva); rounded posteriorly.
Endocarina 7% length of frons. Dorsal epicranial seta 1 long, located
posteriorly distant from frontal suture. Mandibular setae long. Thorax:
pronotum with 5 long setae and 3 short setae, in addition to 2 minute
anterolateral setae. Pedal area of prothorax with 3 long and 2 short setae;
pedal areas of mesothorax and metathorax each with 3 long setae and 3
short setae. Abdomen: Postdorsum with 3 setae, setae 1 and 3 short, seta
2 long.

Pupa.—Unknown.

Material examined.—Two larvae (1 entire, 1 slide-mounted specimen),
labelled Schonebeck, a.d. Elbe 21-8-1931, C. Urban, (USNM) were ex-
amined.

Discussion—No adults are associated with the specimens described
here, so their identity is questionable. The specimens were apparently
transmitted to the USNM from Urban, their collector. Urban (1935) re-
ported finding larvae of T. flavicollis, along with larvae of T. meliloti, in
the fruit of Melilotus alba Lam. Perris (1877), however, cited by Hoffman
(1954) and Scherf (1964), stated that larvae of T. flavicollis develop in
seeds of Lotus corniculatus L. Lengerken (1941) stated that T. meliloti
develops in galls on the leaves of Melilotus macrorhiza Pers., and Hoffmann
(1954) adds M. alba and M. officinalis (L.) to the list of hosts of that
species.

There is indirect evidence that the larva described above as T. flavicollis

642 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

is not that of a Trifolieae-associated species, and is therefore possibly
correctly associated with the species Franz (1942) called T. flavicollis.
Franz considered T. flavicollis to be related to T. kulzeri Penecke, T. cin-
namomeus Kiesenwetter and T. cretaceus Kiesenwetter. Tychius cretaceus
was listed as synonym of T. cinnamomeus by Hoffmann (1954) who listed
Dorycnium suffruticosus Vill. as the host of the weevil. The genera
Dorycnium and Lotus belong to the papilionoid tribe Loteae (Hutchinson,
1964). Since closely related species of Tychius usually have closely related
hosts, it is likely that Franz’s T. flavicollis is a species associated with the
plant genus Lotus. The tribes Loteae and Trifolieae are considered to be
closely related (Hutchinson, 1964). Adult characters, however, indicate
that T. flavicollis is also closely allied to the Trifolieae-associated Tychius
(specimens of T. flavicollis as well as specimens of Trifolieae-associated
species determined by Franz himself have been examined). The larva
described above, however, shows greater affinity to larvae of Astragaleae-
associated Tychius, than to those of the Trifolieae-associated T. picirostris,
T. stephensi and T. flavus (see discussions of these below). Clark (1977)
noted that T. flavicollis resembles the American Astragaleae-associated T.
liljebladi Blatchley.

Tychius stephensi Schonherr
Fig. 9

Curculio picirostris Fabricius, 1787:101.

Curculio tomentosus Herbst, 1795:278 (not Olivier, 1790:536).
Tychius stepheni Schonherr, 1836:412.

Tychius stephensi Schoénherr. Stephens, 1839:229.

Larva—As described for T. tectus, except as follows. Body: 2.74—
3.59 mm long (8 larvae); opaque, white. Head: Width of head capsule
0.41-0.48 mm (5 larvae). Ocellar areas with subcutaneous pigment. Dorsal
epicranial seta 1 long. Clypeal setae moderately long. Median labral
sensillum present. Malar area of maxilla with 5 ventral setae and 3 dorsal
setae. Labial palpus with 2 sensilla; glossa with 2 pairs of minute setae.
Thorax (Fig. 9): Pronotum with 4 long setae, 2 slightly shorter setae and
2 minute setae, in addition to 2 minute anterolateral setae. Spiracle uni-
cameral, air tube with 4.or 5 annuli. Postdorsum of mesothorax and of
metathorax with 2 setae, seta 1 moderately long, seta 2 longer. Pedal area
of each thoracic segment with 3 long setae. Abdomen (Fig. 9): Post-
dorsum with 2 setae, seta 1 long, seta 2 shorter. Pleuron with 1 long and
1 short seta.

Pupa.—Unknown.

VOLUME 80, NUMBER 4 643

Material examined.—Six slide-mounted and numerous whole specimens
labelled “Minnesota, St. Paul, 25 June, 1948, A. Peterson, ex heads red
clover’ (OSU), determined by association with adults preserved with
larvae, were examined. These are the specimens upon which Peterson
(1951) based his description of the larva of T. stephensi.

Discussion.—Introduced from Europe into North America, T. stephensi
develops in the fruit of red clover, Trifolium pratense L. It is known as
the red clover seed weevil in America. Although it has been called T.
stephensi (sometimes T. stepheni) in America, in Europe it is known as
T’. tomentosus Herbst. As Clark (1971) pointed out, neither name is strictly
correct since the types of both are conspecific with the type of Curculio
picirostris Fabricius, 1787. Schonherr (1825) misidentified Fabricius’ C.
picirostris and applied the name picirostris to the species commonly known
in Europe and America as Miccotrogus picirostris. Since Miccotrogus is
now in synonymy with Tychius, the appropriate combination is Tychius
picirostris (Fabricius), following Clark (1976). A proposal to the Inter-
national Commission on Zoological Nomenclature suggesting a solution is
in preparation.

The larva of T. stephensi is distinguished from the other known Tychius
larvae as follows: Thoracic spiracle unicameral, postdorsum of mesothorax
and of metathorax, as well as of each abdominal segment, with 2 setae. The
larva of T. picirostris is distinguished from that of T. stephensi by the
bicameral thoracic spiracle, absence of the median labral sensillum, the
short dorsal epicranial seta 1, possession of a single pleural seta on the
abdominal segments, only 2 long pleural setae on the mesothorax and meta-
thorax, and generally shorter thoracic and abdominal setae throughout.

Larvae of at least 2 other species of Tychius with hosts in the papilionoid
tribe Trifolieae, T. flavus and T. meliloti, also have unicameral thoracic
spiracles. The larvae of these do not differ from the larva of T. stephensi
in any known characters. If the close resemblance of the larvae of the
Astragaleae-associated species described above to each other is any in-
dication of the way closely allied species of Tychius may be expected to
differ, the larvae of T. stephensi, T. flavus and T. meliloti may in fact be
difficult, if not impossible, to distinguish. Larvae of all 3 of these also
have in common with T. picirostris most observed characters, including
several (2 sensilla on labial palpus, 4 long pronotal setae, 2 thoracic and
abdominal postdorsal setae and 3 dorsal setae on malar area of maxilla)
not shared by the other known Tychius larvae. These shared characters
support the contention that the Trifolieae-associated Tychius form a mono-
phyletic group. Some of the members of this group, including T. picirostris,
were formerly assigned to Miccotrogus (see Clark, 1976).

644. PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Tychius picirostris (Fabricius )
Fig. 11

Miccotrogus picirostris (Fabricius). Schonherr, 1825:583, and subsequent
authors, based on misidentification of Curculio picirostris (Fabricius,
WPS Ne

Tychius picirostris (Fabricius). Clark, 1976.

Larva—As described for T. tectus, except as follows. Body: 1.78-
2.81 mm long (8 larvae); opaque, white. Head: Width of head capsule
(.43-0.44 mm (2 larvae). Ocellar areas with subcutaneous pigment. Dorsal
epicranial seta 1 minute. Clypeal setae minute. Median labral sensillum
absent. Malar area of maxilla with 5 ventral setae and 3 dorsal setae. Labial
palpus with 2 sensilla; glossa with 2 pairs of setae, without sensilla. Thorax
(Fig. 11): Pronotum with 4 long setae, 2 slightly shorter setae, and 2
minute setae, in addition to 2 minute anterolateral setae. Spiracle bicameral,
air tubes of unequal length. Postdorsum of mesothorax and of metathorax
with 2 setae, seta 1 short, seta 2 longer. Pedal areas of mesothorax and of
metathorax with 2 long setae and 1 shorter seta. Abdomen (Fig. 11):
Postdorsum with 2 setae, seta 1 long, seta 2 short. Pleuron with 1 moderately
long seta.

Pupa.—Length: 1.14-1.28 mm. Head: 2 pairs of frontal setae, 1 pair
small, each seta borne on small tubercle located slightly anterior and mesal
to larger pair of tubercles which bear much stouter, longer setae. Pronotum:
Anterior-most pair of anterolateral setae located beside anteromedian setae,
2nd pair displaced posteriorly to apical % of pronotum; 1 pair of postero-
lateral setae. Posteromedian setae more widely separated than discal
setae. Abdomen: Terga 1-8 each with 1 pair of discotergal setae. Legs:
Devoid of setae.

Material examined.—Ten larvae (8 entire, 2 slide-mounted specimens
TAM, USNM) and 3 pupae (TAM, USNM) collected in eastern Washing-
ton or western Idaho, by Yunus and Johansen, determined (by Johansen)
by association with adults reared from white clover seeds, were examined.

Discussion—This European species, like T. stephensi, has been intro-
duced into North America where it is a pest of clover seed. It is called
the clover seed weevil. Its biology was studied by Yunus and Johansen
(1967). Hosts in North America are white clover, Trifolium repens L.,
and alsike clover, T. hybridum L. Problems of the nomenclature of this
species and the red clover seed weevil are reviewed above in the discussion
of T. stephensi.

The larva of T. picirostris is distinguished from the other known Tychius
larvae as follows: Thoracic spiracle bicameral; median labral sensillum
and glossal sensilla absent; and pedal area of mesothorax and of metathorax

VOLUME 80, NUMBER 4 645

each bearing 2 long setae and | shorter seta. The pupa is distinguished by
possession of 1 pair of posterolateral pronotal setae, 2 pairs of posteronotal
setae on the meso- and metathorax, 2 pairs of discotergal setae on the
abdomen, and absence of femoral setae.

Comparisons of the larva of T. picirostris and of the related T. stephensi,
T. flavus and T. meliloti and hypotheses concerning relationships of these
to each other and to other Tychius are set forth above in the discussion of
T. stephensi.

Tychius flavus Becker
Tychius flavus Becker, 1864:488.

Larva.—Servadei (1947) provided fairly good illustrations and descrip-
tion of the larva of T. flavus, from which the following diagnosis is taken:
Length 2.5-3.0 mm; head brown, endocarina % as long as frons; dorsal
epicranial seta 1 long; median labral sensillum present; malar area of
maxilla with 3 ventral setae, and 9 (?) dorsal setae; postmental seta 1
present; pronotum with 6 long setae (this probably indicates 4 long and
2 slightly shorter setae); all spiracles unicameral; postdorsum of meso-
thorax and that of metathorax with 2 setae; pedal area of each thoracic
segment with 3 long setae; postdorsum of typical abdominal segments each
with 2 setae.

Scherf’s (1964) description and illustrations of the larva of T. flavus are
apparently adapted from those of Servadei (1947). These are not as
complete as those of Servadei, however, and some inconsistencies are
noted. For example, Scherf illustrated an extra frontal seta, and omitted
the median labral sensillum. Servadei’s description and illustrations are
more in line with our observations on members of the genus Tychius. Un-
fortunately, neither Scherf’s nor Servadei’s descriptions are complete
enough to permit some important comparison (e.g. relative setal lengths,
number of sensilla on the labial palpae and glossa).

Pupa.—Scherf (1964, Fig. 244) also reproduced Servadei’s (1947)
illustration of the pupa of T. flavus (which unfortunately included only a
ventral view) and gave a brief description. From these the following
diagnosis is derived: Length 2.0-2.5 mm; rostrum devoid of setae; 1 pair
of frontal setae present, interocular and supraocular setae absent; ap-
parently 3 pairs of anterolateral pronotal setae present; legs each with
2 femoral setae.

The identity of the specimens examined by Scherf and Servadei has not
been verified.

Discussion—This European species develops in pods of Medicago sativa
L. (Scherf, 1964). It is a serious pest of seed production of this important

646 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

forage crop in Europe. Its biology was studied by Tanasijevic (1955) and
by Ionescu et al. (1965).

The only character mentioned by Servadei (1947) by which the larva
of T. flavus differs from the larvae of the other Tychius with unicameral
thoracic spiracles (T. picirostris and T. meliloti) is the number of ventral
setae on the malar area of the maxillary palpus, which Servadei (1947:159,
Fig. XVII) depicted as no fewer than 9; we observed not more than 5
ventral malar setae in the species examined. Servadei’s (1947) description
of the pupa is too brief to be of value in making comparisons.

Tychius meliloti Stephens
Tychius meliloti Stephens, 1831:55.

Larva.—Scherf (1964) provided a brief description, without illustration,
of the larva of T. meliloti. This agrees with our generic description but,
aside from showing that the thoracic spiracles are unicameral, it is not of
further taxonomic value.

Pupa.—sScherf’s (1964) description of the pupa of T. meliloti omits
important diagnostic characters but does indicate that it has the following:
Length 2.5-2.8 mm; rostrum with 1 pair of basirostral setae and 1 pair of
interocular setae; head with 1 pair of frontal setae; pronotum with 6 pairs
of setae; abdomen apparently with 2 pairs of discotergal setae. No mention
is made of femoral setae.

Discussion.—Urban (1935) reported that this European species develops
in fruit of Melilotus alba and M. officinalis. Scherf (1964) also listed M.
altissimus as a host.

The unicameral larval thoracic spiracle and the host association indicate
that T. meliloti is closely allied to 2 of the other Trifolieae-associated species,
T. stephensi and T. flavus, as considered in the discussion of T. stephensi.
The pupa is apparently like that of T. picirostris in possessing only a single
pair of posterolateral pronotal setae.

Tychius crassirostris Kirsch
Tychius crassirostris Kirsch, 1871:48.

Larva.—Mik (1885) and Scherf (1964) presented brief descriptions and
illustrations of the larva of T. crassirostris. These agree with our generic
description, but do not contain information diagnostic at the species level.

Pupa—Mik (1885) illustrated a ventral view of the pupa of T. cras-
sirostris. Scherf (1964) reproduced the figure and from it wrote a brief
description. According to Scherf the pupa is 2.8-3.4 mm long. The figure
indicates that the setae on the head are as illustrated for T. sordidus

VOLUME 80, NUMBER 4 647

(Fig. 12), except that the lowermost basirostral seta is actually “distirostral”
in position. One pair of frontal seta are present. The pronotum apparently
has 3 pairs of anterolateral setae, the abdomen only a single pair of
discotergal setae. The legs each bear 1 femoral seta.

Discussion —Mik (1885) and Urban (1935) give accounts of the life
history and habits of this European species whose larvae develop in galls
on the leaves of Melilotus alba. Scherf (1964) lists other Melilotus as
well as Medicago hosts.

The larva of T. crassirostris is poorly known. The pupa is evidently
like those of T. flavus and T. meliloti in possessing 3 pairs of anterolateral
pronotal setae, and like that of T. picirostris in having only a single pair
of posterolateral pronotal setae.

Taxonomic Relationships

Although only a small proportion of the total number of species of
Tychius (13 out of more than 300) are known from the immature stages,
some general statements concerning taxonomic relationships based on
characters of these stages can be made. Characters of the larvae and pupae
of some species appear to support some groupings indicated by adults and
host plant relationships. A comprehensive evaluation of character states of
the adults for recognition of monophyletic groups in Tychius has yet to be
carried out, however; and we have not attempted to determine relative
apomorphy-—plesiomorphy in the characters of the relatively small sample
of immatures studied.

Distributions of character states of larvae and pupae diagnostic at species
level in Tychius are presented in Tables 1 and 2, respectively. Diagnostic
characters for larvae include differences in relative lengths of setae on the
head capsule, thorax and abdomen, presence or absence of setae and
sensilla on the mouthparts, and numbers of chambers in the thoracic
spiracles. Presence or absence of pigment on the ocellar areas, and presence
or absence of asperities on the sternal areas of the thoracic and abdominal
segments also appear to distinguish some species, but others exhibit varia-
tion in these features. Pupae of species of Tychius differ from each other
in numbers of pairs of anterolateral and posterolateral pronotal setae,
nesonotal and metanotal setae, discotergal abdominal setae, and in presence
or absence and numbers of femoral setae. Some of these pupal characters
are variable within the small samples examined.

Trifolieae-associated species——The most distinctive grouping indicated
by larval characters includes T. stephensi, T. picirostris, T. flavus and T.
meliloti, all of which have hosts in the papilionoid tribe Trifolieae
(Trifolium, Melilotus and Medicago). The existence of a monophyletic
Trifolieae-associated group within the genus Tychius was postulated by

648 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Clark (1976). The known larvae of members of this group are distin-
guished as follows: Malar area of maxillary palpus with 3 dorsal setae;
labial palpus with 2 sensilla; glossa of labium with 2 pairs of setae;
pronotum with 4 long setae; mesothorax and metathorax each with 2
postdorsal setae. Some of these character states are shared by other
Tychius larvae, but none have them in the same combination. Only 1
Trifolieae-associated species, T. picirostris, is known in sufficient detail in
the pupal stage to permit meaningful comparisons. The pupa of this spe-
cies, and apparently that of T. meliloti, are distinguished from other
Tychius pupae by possession of only | pair of posterolateral pronotal setae.

The larva of T. picirostris differs from known larvae of other Trifolieae-
associated Tychius in the short dorsal epicranial seta 1, absence of the
medial labral sensillum and short abdominal pleural seta 1. It is also
unique among Trifolieae-associated species in possession of bicameral
thoracic spiracles, a feature which it shares with all known non-Trifolieae-
associated Tychius larvae.

Tychius flavicollis—Adult characters and host relationships indicate that
the other Tychius whose immature stages are known represent at least 5
different infrageneric groups (see Clark, 1976 and 1977). These groups are
more homogeneous in the immature stages than the Trifolieae-associated
group, however (Tables 1 and 2). The adult of T. flavicollis resembles
adults of Trifolieae-associated species, but it also resembles some adults
of a group of species associated with members of the papilionoid tribe
Astragaleae, most notably the American T. liljebladi (see Clark, 1977).
The larva of T. flavicollis, however, does not differ from larvae of
Astragaleae-associated species in any of the characters studied (the pupa
is unknown). The host of T. flavicollis, Lotus corniculatus, belongs to
Loteae, a papilionoid tribe which Hutchinson (1964) considered to be more
closely related to Trifolieae than to Astragaleae.

Tychius quinquepunctatus——On the basis of adult characters, T. quin-
quepunctatus appears to be closely allied to species assigned to the genera
Apeltarius Desbrochers and Xenotychius Reitter. The larva of T. quin-
quepunctatus differs from larvae of Astragaleae-associated Tychius only in
possession of 2 rather than 1 pair of setae on the glossa of the labium. It
differs from the larvae of T. sordidus and T. lineellus only in characters
which it shares with Astragaleae-associated species. It has 2 pairs of glossal
setae on the labium, like Trifolieae-associated Tychius. The pupa of T.
quinquepunctatus also shows affinity to Trifolieae-associated species in
possession of a single pair of posterolateral pronotal setae. Characters of
the adults of T. quinquepunctatus and of Trifolieae-associated species,
however, do not indicate that the 2 groups are closely related. Known
hosts of T. quinquepunctatus belong to the genera Pisum and Vicia, mem-

VOLUME 80, NUMBER 4 649

bers of the tribe Vicieae, a tribe which Hutchinson (1964) considers to be
closely allied to the tribe Trifolieae.

Astragaleae-associated species—Monophyly of this group whose mem-
bers occur in North America and Eurasia was postulated by Clark (1977)
on the basis of adult characters and host relationships. Members of the
group with known immature stages, T. tectus, T. semisquamosus, T. soltaui
and T. prolixus, have nearly identical larvae. They differ from each other
in presence or absence of pigment on the ocellar areas and presence or
absence of asperities on the sternal areas of the thorax and abdomen. These
differences are minor and of doubtful taxonomic value. The pupa of T.
soltaui has 1 instead of 2 femoral setae, but it is otherwise indistinguishable
from known pupae of other Astragaleae-associated species, which in turn
are indistinguishable from each other.

Tychius lineellus and Tychius sordidus——Adult characters indicate that
these North American species represent 2 distinct but related species
groups (Clark, 1977). Larvae of the 2 species differ in several respects.
The larva of T. lineellus differs from all known Tychius larvae, except those of
Trifolieae-associated species, by possessing 2 sensilla on the labial palpus.
This similarity is probably the result of convergence; adult T. lineellus in
no way resemble adults of Trifolieae-associated species, and the hosts of
T. lineellus (species of Lupinus, tribe Lupineae) are not closely related to
the Trifolieae. The head of the larva of T. lineellus (Fig. 2) also has
several distinct features not listed in Table 1, namely the head capsule is
deeply emarginate posteriorly, the accessory appendage of the antenna is
long and slender and dorsal epicranial setae 1 and 3 are relatively short
(cf. T. sordidus Fig. 1). In adult characters T. lineellus appears to be more
closely allied to the European T. venustus (Fabricius) than to any
American Tychius (see Clark, 1977). The larva of T. sordidus differs from
that of T. lineellus and larvae of Astragaleae-associated species in the rela-
tively long setae on the alar area of the thoracic segments and on the
epipleural folds of the abdominal segments. The pupa of T. sordidus is
unique among known Tychius pupae in possessing 2 pairs of discotergal
setae on abdominal terga 1-8.

Status of Miccotrogus Schoénherr.—One of the species whose immatures
are described herein, T. picirostris, along with several other European
Tychius, were previously assigned to a separate genus, Miccotrogus Schén-
herr, because they have 6 instead of 7 antennal funicular articles. The North
American T. prolixus, although never formally assigned to Miccotrogus,
also has 6 funicular articles. After examining adult morphological characters
and host relationships of several Old World and American species, Clark
(1976) concluded that Miccotrogus was polyphyletic, and he placed that
name in synonymy with Tychius. The immatures of T. picirostris and of

650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

T. prolixus have nothing in common which would justify their separation
from other Tychius examined. We conclude, therefore, that Clark’s (1976)
hypothesis of polyphyly is not refuted by characters of the immatures
described herein. It should be noted, however, that although immatures of
T. prolixus differ only in minor respects from those of other Astragaleae-
associated Tychius, the larva and pupa of T. picirostris are distinguished
from other Trifolieae-associated Tychius by several features (see discussion
of Trifolieae-associated species ).

Conclusions

It is often claimed that characters of the immature stages of weevils and
other insects should or do contribute to knowledge of phylogeny (see
Emden, 1957; Ahmad and Burke, 1972; Pfaffenberger and Johnson, 1976).
It is true that the immature stages provide a set of attributes which
theoretically have as much potential for revealing monophyletic groups as
do adult structures. Some authors have noted the lack of congruence be-
tween classifications based on characters of immatures and those based on
adults (see Rohlf, 1963; Pfaffenberger and Johnson, 1976). It seems to us
that any 2 systems (e.g. morphological, karyotype, behavioral, biochemical,
etc.) may differ to the extent that purely phenetic classifications based on
them will be incongruent. However, we feel that what is needed in sys-
tematic studies is that all systems be analysed, the evidence drawn from
each synthesized into a unified system, preferably a phylogenetic one. This
is the essence of Hennig’s (1966) holomorphological method. The attri-
butes of different developmental stages should be considered of equal
value in phylogenetic analysis to the extent that similarities are not the
results of parallelism or convergence.

We have observed larvae and pupae of several members of the subfamily
Tychiinae (those treated herein, those of Sibinia sulcatula (Rogers et al.,
1975), and of other species of Sibinia which will be described elsewhere,
and those of Neotylopterus pallidus (LeConte) (Clark, 1978b), as well as
immatures of members of several other subfamilies (including published
accounts by Emden, 1938; 1952; Ahmad and Burke, 1972; and others listed
by Burke and Anderson, 1976). For the most part, character states of these
have yet to be rigorously analysed with the express intent of recognition of
synapomorphies. Instead, analyses have been of the sort presented in the
previous section; i.e., recognition of groups on the basis of subjectively
weighted similarity. From the present study of tychiines and from our
general familiarity with curculionid immatures, it is expected that the
search for synapomorphy, at least at and below the generic level, will be
fraught with difficulty. This is because attributes observed up to now, and
upon which most studies, including the present one, are based (differences

VOLUME 80, NUMBER 4 651

in numbers, positions and relative lengths of setae, variation in numbers of
chambers in the spiracles) are, for the most part, of the sort that one
worker (Schlee, 1969) referred to as “simple features poor in structure and
having only a few details.” It is generally agreed that with such characters
it may be difficult to distinguish similarity due to common ancestry from
similarity resulting from parallel or convergent development.

Nevertheless, we do not advocate abandonment of the search for
synapomorphies in the larvae and pupae of tychiines and other weevils. It
is quite possible that patterns and combinations of the kinds of characters
examined in the present study will prove to be of greater value in deter-
mining relationships at higher levels. Other systems such as internal
anatomy (see May, 1967 and 1977) are also likely to contribute useful
information. Furthermore, the host and other biological information usually
obtained while collecting immatures has already been shown to be of value
as evidence of phylogenetic relationships in the Curculionidae.

Acknowledgments

Some of the information presented here was part of a Ph.D. dissertation
submitted by the senior author to the Department of Entomology, Texas
A&M University. This information was supplemented and re-evaluated by
the senior author during his tenure as Post-Doctoral Fellow in the Entomol-
ogy Department, Smithsonian Institution. Some of the specimens studied
were provided by C. A. Triplehorn, Ohio State University, and C. A.
Johansen, Washington State University. We also thank W. N. Mathis and
D. R. Whitehead for reading and critiquing parts of early drafts of the
manuscript.

Literature Cited

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Anderson, W. H. 1947. <A terminology for the anatomical characters useful in the
taxonomy of weevil larvae. Proc. Entomol. Soc. Wash. 49:123—-132.

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37(2):477-493.

Burke, H. R. 1968. Pupae of the weevil tribe Anthonomini (Coleoptera: Curcul-
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Burke, H. R. and D. M. Anderson. 1976. Systematics of larvae and pupae of American
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Casey, T. L. 1892. Coleopterological Notices, IV. Ann. N.Y. Acad. Sci. 6:359-712.

Clark, W. E. 1971. <A taxonomic revision of the weevil genus Tychius Germar in
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versity Science Bulletin, Biological Series. Vol. 13, No. 3. Brigham Young Uni-
versity Press, Provo, Utah. 39 p.

652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

——. 1976. Review of genus-group taxa included in the genus Tychius Germar
(Coleoptera: Curculionidae). Entomol. Scand. 7:91-95.

—. 1977. North American Tychius: new synonymy and observations on phy-
logeny and zoogeography (Coleoptera: Curculionidae). Entomol. Scand, 8(4):
287-300.

1978a. The weevil genus Sibinia Germar: natural history, taxonomy, phy-
logeny, and zoogeography, with revision of the New World species (Coleoptera:

Curculionidae). Quaest. Entomol. 14(2):91-387.

———. 1978b. Notes on the life history, and descriptions of the larva and pupa of
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Clark, W. E. and H. R. Burke. 1977. The curculionid genus Tychius Germar: nat-
ural history and coevolution with leguminous host plants. Southwest. Entomol.
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Clark, W. E., D. R. Whitehead and R. E. Warner. 1977. Classification of the weevil
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TES?

Emden, F. I. van. 1938. On the taxonomy of Rhynchophora larvae (Coleoptera ).
Trans. R. Entomol. Soc. London 87: 1-37.

———. 1952. On the taxonomy of Rhynchophora larvae: Adelognatha and Alophinae
(Insecta: Coleoptera). Proc. Zool. Soc. London 122:651—795.

———. 1957. The taxonomic significance of the characters of immature insects. Ann.
Rev. Entomol. 2:91—106.

Fabricius, J.C. 1787. Mantissa insectorum, Vol. I. Hafniae. 348 p.

Franz, H. 1942. Vorarbeiten zu einer Monographie der Tychiini (Coleoptera:. Cur-
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Taxon. Entomol. Berlin-Dahlem. 9:104—-133, 182-205, 242-266.

Germar, E. F. 1817. Miscellen und Correspondenzennachrichten. Mag. Entomol.
(Germar) 2:339-341.

Grandi, G. 1916. Contributo alla conoscenze die costumi e delle metamorfosi del
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Agrar. R. Scuola Sup. Agr. Portici 10:103-119.

Hatch, M. H. 1971. The beetles of the Pacific Northwest, part V: Rhipiceroidea,
Sternoxi, Phytophaga, Rhynchophora, and Lamellicornia. Univ. Wash. Press,
Seattle. vii-xiv, + 622 p.

Hennig, W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana.

263 p.
Herbst, J. F. W. 1795. In Herbst and K. G. Jablonsky, Natursystem aller bekannten
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Hoffmann, A. 1954. Faune de France, Coléoptéres Curculionides II. Vol. 59. Leche-
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Hoffmann, A., P. Jourdheuil, P. Grison, M. Chevalier, J. Steffan, J. Cuille, A. Villardebo
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A. S. Balachowsky, Entomologie appliquee a l’agriculture, Tome, I Coléopteéres,
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Hutchinson, J. 1964. The genera of flowering plants, (Angiospermae ) Dicotyledones
I. Clarendon Press, Oxford. v—vii + 516 p.

Ionescu, M., L. Mitru and I. Tucra. 1965. Combaterea gargaritei Tychius flavus
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Kirsch, T. F. W. 1871. Tychius crassirostris Kirsch n. sp. Berl. Entomol. Z. 15:48.

LeConte, J. L. 1876. Tribe XI. Tychiini, p. 211-219, In J. L. LeConte and G. H.
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Lengerken, H. 1941. Von Kafern erzeugte Pflanzengallen. Entomol. Blatt. 37:145-
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Linnaeus, C. 1758. Systema Naturae, ed. 10, vol. 1. Holmiae. 823 p. (original not
seen ).

May, B. 1967. Immature stages of Curculionidae, 1. some genera in the tribe Arau-
cariini (Cossoninae ). N.Z. J. Sci. 10(3):644-660.

———. 1977. Immature stages of Curculionidae: larvae of the soil-inhabiting weevils
of New Zealand. J. R. Soc. N.Z. 7(2):189-228.

Mik, J. 1885. Zur biologie von Tychius crassirostris Kirsch. Wien. Entomol. Z. 4:
289-292, + Taf. IV.

Muka, A. A. 1954. The biology of the clover head weevil, Tychius stephensi Schon-
herr, with some notes on control. Unpublished Ph.D. Dissertation, Cornell Uni-
versity, Ithaca, N.Y. Summary in: Diss. Abstr. 15:12—13.

Nasredinov, Kh. A. 1975. A brief review of the weevils (Coleoptera, Curculionidae )
of southern Tadzhikistan. Entomol. Rev. 54:40-52.

Olivier, A. G. 1970. Entomologic, ou histoire naturelle des insectes, avec leurs
caractéres génériques et spécifiques, Coléoptéres, Vol. 2. Paris.

Perris, E. 1877. Larves de Coléoptéres. Deyrolle, Paris. 590 p.

Peterson, A. 1951. Larvae of Insects. Part II Coleoptera, Diptera, Neuroptera, Si-
phonaptera, Mecoptera, Trichoptera, Ist ed. Edward Bros., Ann Arbor. 146 p.

Pfaffenberger, G. S. and C. D. Johnson. 1976. Biosystematics of the first-stage larvae
of some North American Bruchidae (Coleoptera). U.S. Dept. Agric. Tech. Bull.
1525. 75 p.

Rogers, C. E., W. E. Clark and H. R. Burke. 1975. Bionomics of Sibinia sulcatula
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Scherf, H. 1960. Zur morphologie und biologie der metamorphosestudien einiger an
Lathyrus vernus lebender Coleopteren aus den Gattungen Bruchus, Apion und
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Schlee, D. 1969. Hennigs principle of phylogenetic systematics, an “intuitive,

statistico-phenetic taxonomy’? Syst. Zool. 18:127—134.

Schonherr, J. C. 1825. Tabula synoptica familiae curculionidum. Isis von Oken 5:
581-588.

1836. Genera et species curculionidum .. ., Tom. 3, pars. 1. Roret, Parisiis.
505 p.

Servadei, A. 1947. Il Tychius flavus Becker (IV contributo delle leguminose forag-
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Stephens, J. F. 1831. Illustrations of British entomology . . ., Mandibulata, vol. 4.
Baldwin and Cradock, London. 366 p.

1839. A manual of British Coleoptera or beetles. Longman, Orme, Brown,
Green and Longmans, London. xii + 443 p.

Tanasijevic, N. 1955. Morfologia i razvice surlasa Tychius flavus Beck. Zast. Bilja.
29:3-33.

654 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Tournier, H. 1873. Observations sur les especes Europeennes et circumeuropeennes
de la tribu des Tychiides. Ann. Soc. Entomol. Fr. 3(5):449-522.

Urban, C. 1935. Tychius meliloti Steph. und Tychius crassirostris Kirsch. Entomol.
Blatt. 31:24-29.

Yunus, C. M. and C. A. Johansen. 1967. Bionomics of the clover seed weevil, Mic-
cotrogus picirostris (Fabricius), in southeastern Washington and adjacent Idaho.
Wash. Agric. Exp. Stn. Tech. Bull. 53. 16 p.

(WEC) c/o Department of Entomology, Smithsonian Institution, Wash-
ington, D.C. 20560 (now at Department of Zoology and Entomology, Auburn
University, Auburn, Alabama 36830; (HRB) Department of Entomology,
Texas A&M University, College Station, Texas 77843; and (DMA) Sys-
tematic Entomology Laboratory, Fed. Res., Sci. and Educ. Admin., USDA,
c/o U.S. National Museum, Washington, D.C. 20560.

Footnote

* Technical contribution TA No. 14248. Department of Entomology, Texas Agricultural
Experiment Station, College Station, Texas 77843.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, p. 655

NOTE

ON THE USE OF PECTINATE MAXILLARY SPINES TO SEPARATE
STENONEMA AND STENACRON
(EPHEMEROPTERA: HEPTAGENIIDAE)

Jensen (1974. Proc. Entomol. Soc. Wash. 76:225-228) removed the
Interpunctatum group from Stenonema and described it as a new genus,
Stenacron. His diagnosis of the new genus incorrectly states that the
nymphs of Stenacron have “pectinate spines on the maxillae ... which are
absent in Stenonema.” Several species of Stenonema also have pectinate
spines on the crowns of the maxillae. For example, the nymphs of the
Stenonema terminatum (Walsh) group possess comb-like spines (Fig. 1)
very similar to those found on nymphs of Stenacron and Heptagenia. Use
of this character to separate the two genera should, therefore, be dis-
continued.

Fig. 1. Pectinate spines on the crown of the maxilla of Stenonema terminatum ares.

Philip A. Lewis, Biological Methods Branch, Environmental Monitoring
and Support Laboratory, U.S. Environmental Protection Agency, Cincinnati,
Ohio 45268.

656 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 656-657

NOTE

PARATROPES BILUNATA (ORTHOPTERA: BLATTIDAE):
AN OUTCROSSING POLLINATOR IN A NEOTROPICAL
WET FOREST CANOPY ?

Ongoing controversy and speculation surround discussions of how canopy
plants outcross. Scant studies are available which show cockroaches to be
pollinators (Proctor, M. and Yeo, P. 1972. The Pollination of Flowers.
Taplinger Pub. Co., New York) and there are no observations on cockroaches
as pollinators of canopy plants. I report here on such activity by Paratropes
bilunata (Saussure and Zehntner) (identification by F. W. Fisk at Ohio
State University, who holds the specimen) in an undisturbed Lowland wet
forest at Finca La Selva, near Puerto Viejo, Heredia Province, Costa Rica. A
new method of tree-climbing was used to gain access to the region of the
canopy where the observations were made (Perry, Biotropica, in press).

Little is known about the biology of P. bilunata or the genus. Biolley
(1900. IV Ortépteros recogidos en Costa Rica desde 1890 4 1900. Informe
Nacional Costa Rica. Pp. 40-49.) did note that some Paratropes species are
found inactive during the day under the leaves of certain small trees and
can be confused with beetles of the family Lycidae. This confusion may
be in part due to a striking coloration which is characteristic of many spe-
cies in the genus, including P. bilunata.

At 1200 hours on 24 June 1976 a P. bilunata was seen walking in full
sunlight 34 m above the ground on an inflorescence of Dendropanax
arboreus (L.) Dec. & Planch. Rey. Hortic. (Araliaceae), an arboyeal
epiphyte within the crown of a Dipteryx panamensis ( Pitt.) Record & Mell
(Leguminosae ). It flew successively to three nearby inflorescences over a
period of several minutes without returning to a previously visited area. An
attempt to net it failed, after which there were no new sightings for the
remainder of the observation period which lasted one hour.

Subsequently, at 1030 hours, on 25 June, a P. bilunata was again seen
visiting the same group of inflorescences. Its behavior was similar to the
previous individual. This time, however, the cockroach was caught. Again,
no other P. bilunata were seen for the duration of the observations which
lasted for about an hour. Finally, at 1400 hours, on 3 July another P.
bilunata was seen at the same location after it behaved similarly to the
above.

Interestingly, no P. bilunata (adult or nymph) were ever seen at any
place other than the inflorescences and proximal leaves of D. arboreus,
even though numerous inflorescences of Dipteryx were present nearby.

VOLUME 80, NUMBER 4 657

The exposed condition of the anthers and stigmas of this plant allows P.
bilunata to be a likely pollinator, of which the plant has many. More
important, the behavior of P. bilunata together with the total absence of a
resident population in the canopy area and the temporal distribution of
sightings suggest that this species may range between a number of
Dendropanax individuals and thus could be an outcrosser of this canopy
epiphyte.

The total length of the pronotum and tegmina of the specimen is 25 mil-
limeters. Two large cream-yellow spots are on the anterior lateral portion
of a totally black pronotum. These spots are connected by a very narrow
band along the anterior medial edge of the pronotum. The tegmina, when
closed, are symmetrically colored. Starting laterally and going medially the
tegmina are marked with first a reddish-brown longitudinal band, the
anterior portion of which bears a cream-yellow spot. A black band is
medial to this, followed by a reddish-brown axial band. Precisely how this
patterning relates to the observations is not clear but since P. bilunata was
diurnally active and visible to potential predators such as birds some
significance may be found.

Donald R. Perry, Department of Biology, University of California at Los
Angeles, Los Angeles, California 90024.

PROC. ENTOMOL. SOC. WASH.
80(4), 1978, pp. 657-658

NOTE

CEROPLASTES DENUDATUS, JUNIOR SYNONYM
OF C. RUSCI (HOMOPTERA: COCCOIDEA: COCCIDAE)

The syntypes of Ceroplastes denudatus Cockerell (1893. Entomologist
26:82) from Antigua agree closely with specimens of Ceroplastes rusci (L.)
(1758. Syst. Nat. Ed. 1:456) as currently recognized. Some syntypes of C.
denudatus have spiracular setae in a complete marginal row between the
anterior and posterior spiracular depressions on each side. Interspersed
between these setae in the median part of the row are few bristlelike setae.
The other syntypes, however, are similar to C. rusci in having two or more
bristlelike setae separating the anterior and posterior spiracular setae.

According to Lindinger (1936. Entomol. Zaharb. 45:154), C. denudatus
Green (nec Ckll) (1923. Bull. Entomol. Res. 14:88) from Madeira is equal
to C. rusci (L.) Sign., thus implying that the C. denudatus Cockerell
determined by Green was different from Cockerell’s species. However,
Green (op. cit. 94) based his determination on type-material of C.

658 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

denudatus. Therefore, I conclude that only one species was involved and
that C. denudatus Cockerell is a junior synonym of C. rusci as currently
recognized (NEW SYNONYM).

Ceroplastes rusci is distinguishable from other species of Ceroplastes in
North America by the following morphological characters: Dorsal pores
predominately bilocular, few simple or trilocular; medio-dorsal clear area
present; spiracular setae bullet-shaped, confined to margin laterad of spirac-
ular furrow, in 3 rows; antenna 6-segmented; leg with tibio-tarsal
articulatory sclerosis, claw digitules of equal size.

The lectotype of C. denudatus here designated is a young adult female
mounted from the dry, type-material and labeled as follows: Left label
“Ceroplastes denudatus CkIl, Type, Antigua, Barber Coll., Ckll. Coccidae
104, 6111, from Cockerell, Jan. 3. 94°; right label “mounted from type-
material, LECTOTYPE designated by S. Nakahara.” Eleven adult, female
paralectotypes on five slides and five first instar paralectotypes on one
slide with same data; type slides and unmounted paralectotypes with same
data in USNM. The original description of C. denudatus does not indicate
the collection date. Whereas the description was published in 1893, “Jan. 3.
94” apparently represents the date the type material was received from
Cockerell by the USDA.

Sueo Nakahara, Plant Protection and Quarantine, Animal and Plant
Health Inspection Service, U.S. Department of Agriculture, Beltsville,
Maryland 20705.

BOOK REVIEW

Entomofauna Cubana, Tomo III. Subclase Polyneoptera. 1976. Fernando de
Zayas. 29.2 cm spine. 130 pp., 119 figs. (In Spanish.) Instituto
Cubana del Libro, La Habana, Cuba. No price marked; said by Librarian,
Instituto de Zoologia, Havana, in letter to reviewer, to be available in ex-
change for similar books.

This is a general, illustrated book that has numerous keys and summarizes
well the insect groups covered. It probably will be most useful to local
Cuban students, but it will be consulted generally by specialists. Volumes
I and II of this series have not appeared. The groups in Vol. III comprise
the Polyneoptera, so named by Martynov in 1923 and essentially accepted
as an arrangement of orthopteroids (s. 1.) by Rohdendorf (1961) and
Bei-Bienko (1962). Included are Orthoptera (s. ].), Dermaptera, Isoptera,
Plecoptera, Embioptera and Zoraptera. No Plecoptera are known from
Cuba. For Zoraptera there has been an uncertain record of a wing found
in Quarantine, and now de Zayas records an unidentified species collected

VOLUME 80, NUMBER 4 659

from logs in Oriente and at the old experiment station in Santiago de las
Vegas. At present, the cockroaches, earwigs, termites and embiids are
represented by modern taxonomic reviews.

An index, unfortunately without page numbers, lists 179 species and
various additional genera of orthopteroids considered as Cuban inhabitants.
Coverage appears to be relatively complete. One small mantid, Acontista
cubana from Cuba, and a_ beautiful black spotted yellow cricket,
Hygronemobius histrionicus from the Isle of Pines, are described as new,
the types to be deposited in the Instituto de Zoologia. Their descriptions
are very sparse, but there are good habitus figures of both. In fact,
illustrations are a strong feature. Of the 179 named species noted, 106
(including 11 of the 12 listed genera of earwigs) are represented by habitus
figures, and there are additional figures of details. Several figures, including
those of the cricket Nemobiopsis gundlachi Bol. and the grasshopper
Nichelius fuscopictus Bol., are apparently the first published habitus figures
of those species.

The author is a veteran Cuban entomologist and is said to have the
largest private collection in Cuba. He studied in Louisiana as a young man
and later consulted collections in Washington, D.C. Although he has
published on Lepidoptera and other orders, Coleoptera are his chief in-
terest. A 443-page volume on Cuban Cerambycidae, dated 1975, is an im-
portant comprehensive monograph; it is not of the current series.

Unhappily, the Polyneoptera volume is marred by misspellings and by
errors in the bibliography, perhaps due to language problems with the
printer and to gaps in available library sources. For instance, Walker ap-
pears as “Walter”; a work on termites by Snyder and Miller is credited to
Rehn and Hebard; and Stephen C. Bruner of Cuba, Lawrence Bruner of
Nebraska, and Carl Brunner von Wattenwyl of Vienna have been confused.

All in all, however, de Zayas has produced an attractive and helpful piece
of work that merits much use, and I am delighted that major fruits of his
long career are appearing.

Ashley B. Gurney, Cooperating Scientist, Systematic Entomology Labora-
tory, Fed. Res., Sci. and Educ. Admin., USDA, c/o U.S. National Museum,
Washington, D.C. 20560.

BOOK REVIEW

Insect Flight. R. C. Rainey, editor. Royal Entomological Society of London,
Symposium 7. John Wiley & Sons, New York, 287 pp. Cost: $47.00.

This volume is the product of the seventh biennial symposium of the
Royal Entomological Society. The purpose of this series of symposia and

660 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

their published reports, is to provide a topical summary of extant knowledge
concerning aspects of entomological research. Earlier symposia have been
concerned with insect polymorphism (1961), reproduction (1964), be-
havior (1966), abundance (1968), ultrastructure (1970) and insect/plant
relationships (1973).

The text of the volume on insect flight consists of 12 contributed papers.
The topics are as pertinent to insect flight as the authors are notable to
insect flight research. Chapters and authors include: J. W. S. Pringle
“The muscles and sense organs involved in insect flight’, B. Mulloney
“Control of flight and related behaviour by the central nervous systems of
insects’, W. Nachtigall “Wing movements and the generation of aerodynamic
forces by some medium-sized insects”, T. Weiss-Fogh “Energetics and
aerodynamics of flapping flight: a synthesis”, R. C. Rainey “Flight be-
haviour and features of the atmospheric environment”, Elizabeth Betts
“Forecasting infestations of tropical migrant pests: the Desert Locust and
the African Armyworm’, R. J. V. Joyce “Insect flight in relation to problems
ef pest control”, G. W. Schaefer “Radar observations of insect flight”, M.
Lindauer “Foraging and homing flight of the honey-bee: some general
problems of orientation”, C. G. Johnson “Lability of the flight system: a
context for functional adaptation”, R. J. Wootton “The fossil record and
insect flight”, and V. B. Wigglesworth “The evolution of insect flight”.

Individual contributions range from 13-42 pages, with most chapters
14-22 pages long; thus, the volume is balanced. Each chapter has its own
terminal question-and-answer section developed at the symposium and its
own bibliography. There are 646 references on insect flight in this volume
with titles from 1974.

It is difficult to provide a comprehensive review of this work because it
has multiple authorship and each author addresses a different aspect of
insect flight. In this context the book appears somewhat disjointed. How-
ever, the editor has done an excellent job in minimizing the problems as-
sociated with multiple-author books and at the same time enabled the text
to flow smoothly through the uniform application of words.

In short, I believe this book will be useful to anybody studying insect
flight. However, because of its cost, it probably will be prohibitively ex-
pensive to purchase by the general reading audience.

Gordon Gordh, Division of Biological Control, Department of Entomol-
ogy, University of California, Riverside, California 92502.

VOLUME 80, NUMBER 4 661
SOCIETY MEETINGS

849th Regular Meeting—March 2, 1978

The 849th Regular Meeting of the Entomological Society of Washington
was called to order by President Sutherland at 8:00 PM on March 2, 1978,
in the Ecology Theater of the National Museum of Natural History. Thirty-
three members and 16 guests were present. The minutes of the February
meeting were read by Joyce Utmar and were approved.

Membership chairman Utmar read for the first time the names of the
following new applicants for membership:

R. M. Hendrickson, Jr., USDA, ARS, 501 S. Chapel St., Newark, Delaware.
H. Pearson Hopper, 3713 35th Street, NW, Washington, D.C.
Alvin Mark Wilson, 4221 Metzerott Rd., College Park, Maryland.

Ted Spilman suggested that the meeting announcement should indicate
where the meeting is to be held. President Sutherland announced that the
Washington Academy of Science will have their awards dinner on March
16th. President Sutherland reported that a committee composed of Chris
Thompson, Doug Miller, Ted Spilman and Doug Sutherland will review
the Society’s By-laws for any needed change.

The main speaker for the evening was Mr. Arnold Mallis, Extension
Entomologist, Retired, Pennsylvania State University. Mr. Mallis spoke on
“Recollections of Some Western Entomologists,” and told of how his interest
in the biographies of American Entomologists developed. Afterwards, he
discussed his recollections of professors and associates he had as a student
at the University of California, Berkeley.

Notes and Exhibitions

John Neal showed a sesiid pest (Synanthodon rhododendri) of rhododen-
dron shrubs that were being marketed in the area.

T. J. Spilman described the structure and function of the mandibles of
larvae of Dirrhagofarsus lewisi, a false click beetle (Eucnemidae), as com-
pared with those of larvae of ordinary beetles. Illustrations and models of
the mandibles were also used.

John Kingsolver reported on recent difficulties in sending dead insects
from France to the United States, apparently due to their interpretation of
the Endangered Species Act.

The meeting was adjourned at 9:20 PM, after which pie and punch were
served as refreshments.

Wayne N. Mathis, Recording Secretary

662 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

850th Regular Meeting—April 6, 1978

The 850th Regular Meeting of the Entomological Society of Washington
was called to order by President Sutherland at 8:00 PM on April 6, 1978,
in the Ecology Theater of the National Museum of Natural History.
Twenty-six members and 10 guests were present. The minutes of the March
meeting were read and approved.

Membership chairman Utmar read for the first time the names of the
following new applicants for membership:

Ahmad Marwan Chawkat, 400 Arundel Rd., Dunkirk, Maryland

Italo Currado, Istituto di Entomologia dell'Universita, Via Guiria 15,
10126, Torino, ITALY

Kenneth L. Deahl, USDA, Vegetable Laboratory, Bldg. 004-Rm 210,
Washington, D.C.

Neil J. Lamb, Military Ent. Information Service/AFPCB, Washington,
DiC.

Lynda L. Richards, Insect Zoo, NHB 101, Smithsonian Institution, Wash-
ington, D.C.

Patricia A. Zungoli, 3416 Tulane Drive, Hyattsville, Maryland

Joyce Utmar announced that the annual banquet will be held at Fort
Lesley McNair on Thursday, June Ist. Tickets are available.

President Sutherland introduced both speakers for the evening. The first,
Mr. Tom Shortino, USDA, Insect Physiology Laboratory, Beltsville, Mary-
land, spoke on “Bioassay diets for insects.” Mr. Shortino passed around
various culture media, some with developing insects, and described
techniques used in the production of insect diets. Dr. A. B. Gurney,
Cooperating Scientist, Systematic Entomology Laboratory, USDA, was the
second speaker. He addressed the topic of “Photographs and biographical
notes of early entomologists.” Dr. Gurney presented slides and provided
biographical sketches of many notable entomologists, mostly from the late
19th and early 20th centuries.

Notes and Exhibitions

A. B. Gurney passed around the new common names list that was recently
published by the Entomological Society of America. Our president, D.
Sutherland, chaired the committee responsible for it. Dr. Gurney and
others commented on the historical development and format of the list.

President Sutherland announced that the Missouri Botanical Garden will
publish a listing of North American plants entitled “A provisional checklist
of species for flora of North America.”

Mr. Nelson Beyer of the U.S. Fish and Wildlife Service reported that his
agency will be surveying the moths and butterflies of the Washington, D.C.

VOLUME 80, NUMBER 4 663

area for assay. He inquired if there were members within the area who

would be willing to let them black-light in their yards.
The meeting adjourned at 9:54 PM, after which punch and cookies were

served.
Wayne N. Mathis, Recording Secretary

664 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

XVI INTERNATIONAL CONGRESS OF ENTOMOLOGY

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August 3-9, 1980
FIRST ANNOUNCEMENT

The 16th International Congress of Entomology will be held in Kyoto the ancient
capital of Japan, 3-9 August, 1980, under the sponsorship of the Science Council of
Japan, the Entomological Society of Japan, the Japanese Society of Applied Entomology
and Zoology and the Japan Plant Protection Association. Sessions will be held in the
excellent modern facilities of the Kyoto International Conference Hall. All sessions and
printed materials will be in English.

Scientific program.—The tentative program of the 16th Congress will include plenary
sessions, symposia, regular sessions, poster sessions, film sessions and informal meet-
ings. The Congress will cover the following fields of Entomology: Systematics,
Morphology, Physiology, Biochemistry, Ecology and Population Dynamics, Genetics,
Developmental Biology, Behavior, Agricultural and Forest Entomology, Toxicology,
Pathology, Stored Product Insects, Medical and Veterinary Entomology, Sericulture,
Apiculture, Biological Control, Integrated Control, Pesticide Development, Manage-
ment and Regulation, Social Insects, Acarology and others.

Participation.—All participants are required to register for one of the three member-
ships: full member, associate member or student member. Registration forms will
be provided in the Second Announcement which will be mailed in June 1979.

Social and associates programs.—An interesting social program will be proposed for
all participants. Also a number of tours and local activities will be arranged for the
associate members.

Exhibition.—It is planned to display scientific equipment, insect specimens, and
books and periodicals on Entomology.

Travel and hotel accommodation.—Japan Travel Bureau (JTB) has been appointed
as the official travel agent of the Congress. Hotel accommodation and travel within
Japan will be arranged by JTB.

Notice.—Anyone wishing to participate in the Congress should send his or her
name, address and field of interest (see list under Scientific Program) to the following
address:

XVI International Congress of Entomology
c/o Kyoto International Conference Hall
Takara-ike, Sakyo-ku

Kyoto 606, Japan

PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

MISCELLANEOUS PUBLICATIONS

Cynipid Galls of the Eastern United States, by Lewis H. Weld _ $ 5.00
Cynipid Galls of the Southwest, by Lewis H. Weld 3.00

Mocha papersconyreruiid. Salis <2 8h et ee 6.00
Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman 1.00

Unusual Scalp Dermatitis in Humans Caused by the Mite Dermato-
Mince aMlanr ser narein se 6. te ke ee Le el ee

A Short History of the Entomological Society of Washington, by Ashley

RRO RTE Eee eet see eS PO ary tO ol ee a a a 1.00

Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae),
Pp SeOn Min SECY SRAM Oe nt 6 Ee ee eS deere ee 1.50

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

No. 1. The North American Bees of the Genus Osmia, by Grace
Sandhouse. 1939 __. gS. eee ERP OE DGS Ser Ro tL Pe $15.00

No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga,
yoann Go Bayi. 1949) 3 8 ee 15.00

No. 3. The Nearctic Leafhoppers, a Generic Classification and Check
fastby-Padk WalsonvOman. 1949) 2 - 25:00

No: 4... A eae of the Chiggers, by G. W. Wharton and H. S. Fuller.

No. 5. A Classification of the Siphonaptera of South America, by Phyllis
Pasta, UO were ee ce NAS Re. be eee DEO

No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace
Po Murdoch:and Hirosi. Takahasi,, 1969) 2 15.00

No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and
Jeanette Wheeler. 1976 - i ee NS a Se Bey ag dS ee

Back issues of the Proceedings of the Entomological Society of Washington are
available at $15.00 per volume to non-members and $9 per volume to members of
the Society.

Prices quoted are U. S. currency. Dealers are allowed a discount of 10 per cent
on all items, including annual subscriptions, that are paid in advance. All
orders should be placed with the Custodian, Entomological Society of Wash-
ington, c/o Department of Entomology, Smithsonian Institution, Washington,
D.C. 20560

CONTENTS

(Continued from front cover)

A new genus, species and family of Hymenoptera (Ichneumonoidea) from
Chile W. R. M. MASON

A synopsis of Neotropical Eleleides Cresson (Diptera: Ephydridae)
W. N. MATHIS

A new genus near Canaceoides Cresson, three new species and notes on their
classification (Diptera: Canacidae) W. N. MATHIS and W. W. WIRTH

New species, records and key to Texas Liposcelidae (Psocoptera)
E. L. MOCKFORD

Notes on Nathan Banks’ species of the mite genus Carabodes (Acari: Oribatei)
R. A. NORTON

Aphids of sunflower: Distribution and hosts in North America (Homoptera:
Aphididae) C. E. ROGERS, T. E. THOMPSON and M. B. STOETZEL

A new species of Hylemya from Wyoming (Diptera: Anthomyiidae)
G. CG. STEYSEKAL

Rearing of Texas Tabanidae (Diptera). I. Collection, feeding, and maintenance
of coastal marsh species P. H. THOMPSON and P. C. KRAUTER

Additional aphid-host relationships at the Los Angeles State and County Arboretum
(Homoptera: Aphididae)
H. G. WALKER, M. B. STOETZEL and L. ENARI

Studies on the genus Forcipomyia. V. Key to subgenera and description of a
new subgenus related to Euprojoannisia Bréthes (Diptera: Ceratopogonidae)
W. W. WIRTH and N. C. RATANAWORABHAN

NOTES:

On the use of pectinate maxillary spines to separate Stenonema and Stenacron
(Ephemeroptera: Heptageniidae) PL AS LEWIS
Ceroplastes denudatus, junior synonym of C. rusci (Homoptera: Coccoidea:
Coccidae) S. NAKAHARA
Paratropes bilunata (Orthoptera: Blattidae): An outcrossing pollinator in a Neo-
tropical wet forest canopy? Do Re Brno

BOOK REVIEWS:

Insect Flight (R. C. Rainey, editor) G. GORDH
Entomofauna Cubana, Tomo III. Subclase Polyneoptera (Fernando de Zayas)
A. B. GURNEY

SOCIETY MEETINGS

ANNOUNCEMENT: XVI International Congress of Entomology
TABLE OF CONTENTS, VOLUME 80

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