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Journal

of the

New York

ENTOMOLOGICAL SOCIETY

Devoted to Entomology in General

VOLUME LXXXI1

Published by the Society
New York, N. Y.

ALLEN PRESS, INC.

V>RINTe0
u. s. *•

LAWRENCE, KANSAS

INDEX OF AUTHORS

ALEXANDER, CHARLES P. New or Little-Known Crane Flies from Iran. I
(Diptera: Tipulidae) 279

ALLEN, GEORGE E. and WILLIAM F. BUREN. Microsporidian and Fungal
Diseases of Solenopsis invicta Buren in Brazil 125

BLUM, MURRAY S. Myrmecine Trail Pheromones: Specificity, Source and
Significance 141

BROWN, F. MARTIN. Andean Larvae and Chrisalids of Dione juno andicola (Bates)
and Agraulis vanillae lucina Felder and Felder 61

BROWN, F. MARTIN. William Couper, Taxonomist-Entomologist 222

BROWN, WILLIAM L., JR. A Supplement to the Revision of the Ant Genus Basiceros
(Hymenoptera: Formicidae) 131

BUREN, WILLIAM F., GEORGE E. ALLEN, WILLARD H. WHITCOMB, FRANCIS
E. LENNARTZ and ROGER N. WILLIAMS. Zoogeography of the Imported
Fire Ants 113

CREIGHTON, WILLIAM S. and ROY R. SNELLING. Notes on the Behavior of
Three Species of Cardiocondyla in the United States (Hymenoptera: Formicidae) 82

DELFINADO, MERCEDES D. and EDWARD W. BAKER. Terrestrial Mites
of New York (Acarina: Prostigmata), I-Tarsocheylidae, Paratydeidae, and Pseudo-
cheylidae 202

ELLIOTT, NANCY B. and FRANK E. KURCZEWSKI. Seasonal variation in
Tachysphex terminatus (Smith) (Hymenoptera: Sphecidae, Larrinae) 268

EVANS, HOWARD E. Digger Wasps as Colonizers of New Habitat (Hymenoptera:
Aculeata) 259

FORBES, JAMES. The William S. Creighton Memorial Issue 66

GREGG, ROBERT E. William Steel Creighton — An Appreciation 67

HUCKETT, H. C. The Anthomyiidae and Muscidae of the Great Smoky Mountains
and Mt. Mitchell, North Carolina (Diptera) 150

MARAMOROSCH, KARL. Ovipositing of Circulifer tenellus Baker (Homoptera,
Cicadellidae) 42

MCCLUSKEY, ELWOOD S. Generic Diversity in Phase of Rhythm in Myrmi-
cine Ants 93

MCDONALD, F. J. D. Revision of the Genus Holcostethus in North America
(Hemiptera: Pentatomidae) 245

MUYSHONDT, ALBERTO. Notes on the Life Cycle and Natural History of
Butterflies of El Salvador. V. A. Pyrrhogyra hypsenor (Nymphalidae-Cato-
nephelinae) 163

NEVIN, F. REESE. A New Genus and Two New Species of Achipteriidae from New
York State (Acari: Cryptostigmata: Oribatei) 177

PECHUMAN, L. L. Two New Tabanidae from Southeastern United States (Diptera) .... 183

iii

ROLSTON, L. H. A New Genus of Pentatominae from South America, Distinguished
by the Position of its Spiracles (Hemiptera: Pentatomidae) 57

ROLSTON, L. H. and R. KUMAR. Two New Genera and Two New Species of
Acanthosomatidae (Hemiptera) from South America, with a Key to the Genera
of the Western Hemisphere 271

ROZEN, JEROME G., JR. The Biology of Two African Melittid Bees (Hyme-
noptera, Anthophoridae) 230

SCHMITT, JOHN B. The Distribution of Brood Ten of the Periodical Cicadas
in New Jersey in 1970 189

SEIFERT, RICHARD P. The Sphingidae of Turrialba, Costa Rica 45

SLATER, JAMES A. and JANE E. HARRINGTON. Tenuicoris myrmeforme : A

New Genus and Species of Myodochini (Hemiptera: Lygaeidae) 173

SNELLING, ROY R. Studies on California Ants. 8. A New Species of Cardiocondyla
(Hymenoptera: Formicidae) 76

THOMPSON, F. CHRISTIAN. The Genus Pterallastes Loew (Diptera: Syrphidae) .... 15

VAN SICKLE, DEBRA and R. M. WESELOH. Habitat Variables That Influence the
Attack by Hyperparasites of Apanteles melanoscelus Cocoons 2

VAURIE, PATRICIA. Synonymy in Sphenophorus pertinax Olivier (Coleoptera,
Curculionidae, Rhynchophorinae) 14

WHEELER, GEORGE C. and JEANETTE WHEELER. Supplementary Studies on
Ant Larvae: Simopone and Turneria 103

WILSON, EDWARD O. and ROBERT FAGAN. On the Estimation of Total
Behavioral Repertories in Ants 106

YOUNG, ALLEN M. Further Observations on the Natural History of Philaethria
dido dido (Lepidoptera: Nymphalidae: Heliconiinae) 30

YOUNG, ALLEN M. Notes on the Natural History of the Rare Adelpha Butterfly
(Lepidoptera: Nymphalidae) in Costa Rican High Country 235

BOOK REVIEWS

KLOTS, ALEXANDER B. The South Asiatic Olethreutini (Lepidoptera, Tortricidae) .

A. Diakonoff 188

KLOTS, ALEXANDER B. The Common Insects of North America. Lester A.
Swann and Charles S. Papp 213

MARAMOROSCH, KARL. Tissue Culture: Methods and Applications. Paul F.

Kruse, Jr. and M. K. Patterson 201

MARAMOROSCH, KARL. The Gunong Benom Expedition, 1967: Parts 11-13.

R. Traub 212

XV INTERNATIONAL CONGRESS OF ENTOMOLOGY 220

NEW YORK ENTOMOLOGICAL SOCIETY GUEST SPEAKERS, 1974/1975 219

PROCEEDINGS OF THE NEW YORK ENTOMOLOGICAL SOCIETY

October 2, 1973-December 18, 1973 245

Vol. LXXXII

No. 1

0 \ L^\' ( . .

rtf : i w . >v;r ^

Devoted to Entomology in General

;

The New York Entomological Society
Incorporating The Brooklyn Entomological Society
Incorporated May 21, 1968

The New York Entomological Society
Organized June 29, 1892 — Incorporated February 25, 1893
Reincorporated February 17, 1943

The Brooklyn Entomological Society

Founded in 1872 — Incorporated in 1885
Reincorporated February 10, 1936

The meetings of the Society are held on the first and third Tuesday of each month (except
June, July, August and September) at 8 p.m., in the American Museum of Natural
History, 79th St. & Central Park W., New York, N. Y. 10024.

Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00.
Members of the Society will please remit their annual dues, payable in January, to the
Treasurer.

Officers for the Year 1974

President , Dr. Daniel J. Sullivan/ S.J.

Fordham University, New York 10458

Vice-President , Dr. Peter Moller

American Museum of Natural History, New York 10024
Secretary , Dr. Charles C. Porter

Fordham University, New York 10458
Assistant Secretary , Dr. Louis Trombetta

Pelham Manor, New York 10803

Treasurer , Dr. Winifred B. Trakimas

State University of New York, Farmingdale, New York 11735
Assistant Treasurer , Ms. Joan DeWind

American Museum of Natural History, New York 10024

Class of 1974

Dr. Lee Herman

Trustees

Mr. Edwin Way Teale

Class of 1975

v: -/

Dr. Howard Topoff

■V

Dr. Pedro Wygodzinsky

Mailed May 23, 1974

The Journal of the New .York Entomological Society is published quarterly for the Society by Allen Press
Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at New/ York, New York, and
at additional mailing office.

Known office1 of publication : (Central Park West a( 79th Street, New York, New York 10024.

Journal of the

New York Entomological Society

Volume LXXXII March, 1974

No. 1

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lawrence E. Limpel
Helen McCarthy

Publication Committee

Mrs. Joan DeWind Dr. Ayodha P. Gupta

Dr. Alexander B. Klots, Chairman

CONTENTS

Habitat Variables That Influence the Attack by Hyperparasites of Apanteles
melanoscelus Cocoons Debra Van Sickle and R. M. Weseloh 2

The Biology of Two African Melittid Bees (Hymenoptera, Apoidea)

Jerome G. Rozen, Jr. 6

Synonymy in Sphenophorus pertinax Olivier (Coleoptera, Curculionidae,
Rhynchophorinae) Patricia Vaurie 14

The Genus Pterallastes Loew (Diptera: Syrphidae) F. Christian Thompson 15

Further Observations on the Natural History of Philaethria dido dido
(Lepidoptera : Nymphalidae: Heliconiinae) Allen M. Young 30

Ovipositing of Circulifer tenellus Baker (Homoptera, Cicadellidae)

Karl Maramorosch 42

The Sphingidae of Turrialba, Costa Rica Richard P. Seifert 45

A New Genus of Pentatominae from South America, Distinguished by the
Position of Its Spiracles (Hemiptera: Pentatomidae) L. H. Rolston 57

Andean Larvae and Chrisalids of Dione juno andicola (Bates) and Agraulis
vanillae lucina Felder & Felder F. Martin Brown 61

Habitat Variables That Influence the Attack by Hyperparasites
of Apanteles melanoscelus Cocoons

Debra Van Sickle and R. M. Weseloh
Department of Entomology, The Connecticut Agricultural Experiment Station,

New Haven 06504

Received for Publication September 18, 1973

Abstract: Overwintering cocoons of Apanteles melanoscelus Ratzeburg (Hymenoptera:

Braconidae), a larval parasite of the gypsy moth Porthetria dispar L. (Lepidoptera: Ly-

mantriidae), were collected from the field, and laboratory-reared cocoons were exposed in
the field to determine under what situations they would be most readily attacked by hyper-
parasites. It was found that hidden cocoons were attacked more readily than exposed
ones and that percent parasitism increased as the season progressed.

INTRODUCTION

Insect hyperparasites may be a problem in the establishment of imported
insect enemies for the control of injurious insects. As Muesebeck and Dohan-
ian (1927) point out, hyperparasites could overwhelm primary parasites before
the latter become established. Even after establishment, hyperparasites may
be important in suppressing primary parasites so that the latter are not effec-
tive. This is evidently the case for Apanteles melanoscelus Ratzeburg (Hyme-
noptera: Braconidae), a bivoltine oligophagous imported parasite of the gypsy
moth Porthetria dispar (L.) (Lepidoptera: Lymantriidae) . A. melanoscelus
overwinters as a mature larva within a cocoon that is exposed to the attack
of hyperparasites throughout late summer and autumn. The cocoons are at-
tacked by over thirty native hymenopterous hyperparasites that are mainly
in the superfamilies Ichneumonoidea and Chalcidoidea (Muesebeck and Do-
hanian, 1927). The objective of this study was to determine if habitat varia-
tion in the placement of overwintering cocoons of A. melanoscelus had any ef-
fect on the extent of hyperparasitism.

MATERIALS AND METHODS

Two procedures were used to determine the incidence of parasitism on A.
melanoscelus cocoons.

Field Collections . Approximately 300 overwintering cocoons of A. melano-
scelus were collected from trees in various forested sites in southcentral Con-
necticut during July, 1973. Each cocoon was placed individually in a 2-in.
plastic cream cup and the date of collection, height of cocoon above ground,
and extent of exposure were recorded. Cocoons were considered to be in a

New York Entomological Society, LXXXII: 2-5. March, 1974.

Vol. LXXXII, March, 1974

nonexposed situation if they were under bark flaps, in cracks, etc. Collections
were made from both dead and living trees, up to 7 ft above ground. No at-
tempt was made to record species of trees. Cocoons were kept in an incubator
at 24°C and 20 hr light/day for emergence of hyperparasites. Cocoons that
had no exit holes were dissected at the end of July to determine if they had
been hyperparasitized. Cocoons that had exit holes at the time of collection
were assumed to have formed the year before. They were classified as to
whether or not a hyperparasite had emerged by characteristics of their exit
holes. Data were converted to percentages and analyzed after transformation
to arcsines.

Placement of Lab oratory -Reared Cocoons. To exercise better control over
cocoon placement, laboratory-reared A. melanoscelus cocoons were exposed in
the field according to the following procedure.

First-instar gypsy moth larvae were paratisized by adult females of A. melano-
scelus and kept in a light chamber at 24 °C and 14 hr light/day (to induce
diapause in the A. melanoscelus larvae). The caterpillars were fed an artificial
diet until overwintering cocoons of the parasite formed. These cocoons were
placed in the field in two sites in Guilford, Conn. At each site, two dead oak
trees of about 6 in. diameter were selected. Eight cocoons were placed at each
of three heights (0, 3, 6 ft) on the trunk of each tree. Four were placed in
an exposed position and four were placed under a bark flap. Elmer’s Milk
Glue was used to hold the cocoons in place.

Cocoons were also placed in other situations as indicated below. On a living
oak tree at each site, four cocoons were placed in each of four situations at
the 6-ft level. The situations were: (1) exposed on the trunk, (2) nonexposed
under a bark flap on the trunk, (3) exposed on a small twig, and (4) exposed
on the undersurface of a leaf. Cocoons were also placed on a large 2 ft diam.
rock at each site. Four cocoons were exposed on top of the rock and four were
placed in a nonexposed position under the overhanging ledge of the rock.

Cocoons were left out for eleven days beginning July 16, 1973. They were
then collected and dissected to determine incidence of hyperparasitism.

RESULTS

At least seven different species of hyperparasites in four different families
emerged from the field-collected cocoons. The families were: Ichneumonidae,
Encyrtidae, Eurytomidae, and Pteromalidae. None were identified by species.

For the field-collected cocoons there was no difference in hyperparasitism
at the different heights. A two-way analysis of variance was performed by
grouping the data by exposure and dates collected; the results are presented
in Table 1. Variation due to dates and length of exposure was significant. Per-
cent hyperparasitism increased as the summer progressed, and the highest

New York Entomological Society

Table 1. Percent parasitism by hyperparasites of cocoons of A. melanoscelus collected

in the field

Collection Dates

Exposure

July

2, 3, 5

July
6, 9, 10

July

11-13

July

16, 18, 23

Season

1972

(old cocoons)

Exposed

Nonexposed

percent hyperparasitism was for those cocoons, formed during the previous
year, that were exposed to the attack of hyperparasites the previous summer
and autumn. Table 1 also shows that nonexposed cocoons were attacked more
readily than exposed cocoons.

For the laboratory-reared cocoons, an analysis of variance on percent para-
sitism of cocoons placed on the four dead trees failed to show any differences
in height or exposure. Analyses of variance were not run for the other situa-
tions, but it would appear that exposed cocoons were generally attacked less
by hyperparasites than nonexposed cocoons (Table 2).

DISCUSSION

Increase in percent parasitism by hyperparasites as the season progresses
has been noted by Muesebeck and Dohanian (1927), Clancy (1944), and
Schlinger (1960). As the summer progresses, the hyperparasites that attack
A. melanoscelus are evidently able to parasitize a greater and greater propor-
tion of the host population, eventually destroying up to 90 percent of them.

Table 2. Percent parasitism by hyperparasites of A. melanoscelus cocoons reared in the
laboratory and placed in the field. July 16-27, 1973

Height

Situation 0 ft 3 ft 6 ft

Four dead trees

Exposed 37 19 56

Nonexposed 44 4 50

Two living trees

Exposed on trunk 62

Nonexposed on trunk 75

Exposed on twig 62

Exposed under leaf 38

Two rocks

Exposed 38

Nonexposed 62

Vol. LXXXII, March, 1974

Of perhaps more interest is the rather unexpected result that hyperparasites
attack nonexposed cocoons more readily than exposed ones. Evidently these
hyperparasites search for hosts primarily in concealed locations. As most co-
coons of A. melanoscelus are found in nonexposed situations, especially under
bark flaps, the hyperparasites are well adapted to exploitation of this insect.

Literature Cited

Clancy, D. W. 1944. Hyperparasitization of Clausenia purpurea Ishii, an important
parasite of the Comstock mealybug. J. Econ. Entomol., 37: 450-451.

Muesebeck, C. F. W. and Dohanian, S. M. 1927. A study in hyperparasitism, with par-
ticular reference to the parasites of Apanteles melanoscelus (Ratzeburg). U.S.D.A.
Bull. 1487, 35 pp.

Schlinger, E. I. 1960. Diapause and secondary parasites nullify the effectiveness of
rose aphid parasites in Riverside, California, 1957-1958. J. Econ. Entomol., 53:
151-154.

New York Entomological Society

The Biology of Two African Melittid Bees (Hymenoptera, Apoidea)

Jerome G. Rozen, Jr.1

Received eor Publication October 18, 1973

INTRODUCTION

In terms of anatomical structures of both larvae and adults, bees belonging
to the Melittidae are diverse even though the family is small. We do not know
if this diversity is also revealed in their life histories because their biology has
been little studied. I present the following with the hope that it will augment
what is already known about their biology and that it will lead eventually to
a better understanding of the phylogeny of the family.

These observations were made on a trip in October, 1972, to the Western
Cape Region of the Republic of South Africa. Although three of the four sub-
families of melittids (Melittinae, Dasypodinae, and Ctenoplectrinae) occur in
southern Africa, these notes refer only to two species in the Dasypodinae, Capi-
cola braunsiana and Haplomelitta ogilviei. Adults of Melitta capensis Friese
(Melittinae) were also seen but nests could not be located. This species mimics
Apis mellifera Linnaeus to a remarkable extent as do species of a number of
other genera of African bees. Ctenoplectrinae apparently do not occur in the
arid regions where my observations were carried out. Mature larvae of Capi-
cola braunsiana were described in a separate paper by Rozen and McGinley
(in press). Adults and immatures collected in connection with this study are
deposited in The American Museum of Natural History.

My studies were greatly assisted by Dr. F. Christian Thompson and Mr.
Ronald J. McGinley whose companionship I enjoyed on the trip. Dr. Gerald
I. Stage, University of Connecticut, Storrs, aided in the identification of adult
bees. The research was supported by National Science Foundation Grant
GB32193.

Capicola braunsiana Friese

This species was found nesting at 67 km. east of Port Nolloth, Cape Prov-
ince, Republic of South Africa, on October 17, 1972, by Dr. Thompson. He
and Mr. McGinley assisted me in the excavation of the site between 2 and 5
p.m. on the same day, which was clear and sunny. The nesting area (Fig. 1)
was in a sandy, treeless region with low hills and numerous widely spaced

1 Deputy Director for Research and Curator of Hymenoptera, The American Museum
of Natural History, Central Park West at 79th Street, New York, New York 10024
Abstract: This paper treats the nesting biology of Capicola braunsiana Friese and Hap-
lomelitta ogilviei (Cockerell) (Melittidae, Dasypodinae) from Cape Province, Republic
of South Africa.

New York Entomological Society, LXXXII: 6-13. March, 1974.

Vol. LXXXII, March, 1974

Fig. 1. Nesting site of Capicola braunsiana, 67 kilometers east of Port Nolloth, Cape
Province, Republic of South Africa. Large-leafed vegetation is pollen plant.

Fig. 2. Nesting site of Haplomelitta ogilviei , 28 kilometers east of Velddrif, Cape Prov-
ince, Republic of South Africa. Dr. F. Christian Thompson is peering into excavation of
nest.

New York Entomological Society

desert plants, most conspicuously succulents belonging to the Mesembryanthe-
mum, sensu lato, (Aizoaceae). The gently sloping nesting site, covering an
area of approximately two meters square, was mostly barren sand with a few
plants including the pollen plant, a large, unidentified species of Mesembryan-
themum ; none of the plants shaded the nesting site appreciably. Dry loose
sand mixed with some gravel and a few stones on the surface, the soil was more
consolidated below the surface and became moist at a depth of about 12 cm.
No other species of bee was seen burrowing at the site during the course of the
brief observations but a nest of an halictid, probably Dialictus, was uncovered.

Nesting Activity: Although time did not permit an accurate counting, at
least seven nests were definitely identified and other female bees flying around
suggested that additional nests — perhaps a total of ten to fifteen — were pres-
ent within the two-meter-square area. About half the nest entrances were
adjacent to or under small stones on the surface of the ground, and the other
entrances were in the open. Tumuli were observed around some of the en-
trances but in most cases excavated material apparently was quickly blown
away.

The following information was obtained from excavating two nests (Figs. 3,
4). In each case the main tunnel, open at the surface, descended obliquely
with considerable meandering to a depth of about 13 to 15 cm. The average
rate of descent was somewhat less than 45 degrees from the horizontal. The
main tunnel, 3.0 mm. in diameter, seemed to be clogged with soil at various
intervals along the way but had some open spaces between and apparently
was open more below than near the surface. Its wall was smooth and without
a special lining. In one nest (Fig. 3), the tunnel, after reaching a depth of
about 13 cm., gave rise to a linear series of four cells, each containing a mature
larva. The tunnel then turned sharply and extended for 13 cm. in a unidirec-
tional but somewhat meandering fashion, dropping only about 2 cm. over that
length. It ended in an open cell containing a fully formed pollen ball. In the
other nest (Fig. 4) the tunnel ended in a linear series of three cells as soon as
it reached a depth of 14.5 cm. The cell closest to the tunnel was open and un-
provisioned; the other two contained pollen masses and eggs. In a third nest,
more hastily excavated than the other two, the main tunnel ran in meandering
fashion downward to a depth of 14 cm., over a distance of about 32 cm., and
ended in a linear sequence of three cells; the cell closest to the tunnel was open,
the second was closed and contained a pollen ball and presumably an egg, and
the third contained a pollen ball and an egg.

Hence, except for the single terminal cell in the first nest, all cells seemed
to be arranged in a linear series. In the first nest cells in series were separated
by a distance of 1.5 to 2.0 mm. Cells from all nests ranged in maximum length
from 7.0 to 7.5 mm. (five measurements) and in maximum diameter from

Vol. LXXXII, March, 1974

4.5 to 5.0 mm. (five measurements). They were broadly rounded at the rear
and more narrowed in front. Their long axis tilted from 30 degrees to 45 de-
grees from the horizontal and the anterior end was higher. Although cell walls
appeared to be unlined, having a dull finish, a droplet of water placed upon
the floor showed that it was waterproof while the upper part of the wall, simi-
larly treated, only retarded absorption. Closures were concave on the inside
with at best an indistinct spiral.

Only a single adult female was associated with each nest. However, the
first nest consisted of a newly provisioned cell and four cells each with a ma-
ture larva. As the female associated with the nest was fresh, she was presum-
ably not the parent of the larvae. Perhaps the nest had been occupied previ-
ously by another female.

Provisioning: The pollen plant grew profusely in the general area and was
found on the nesting site itself (Fig. 1). The bee however apparently visits a
number of species of Mesembryanthemum as it had been taken elsewhere, on
other plants. The flowers of the pollen plant opened about 2 p.m. The female
transported the pollen dry to the nest and there formed it into a perfect sphere,
which was 2.5 to 2.9 mm. (three measurements) in diameter, mealy-moist
throughout, pale green in color, and emitted no detectable odor.

Development: Several strongly curved eggs, translucent white with a shiny
chorion, were each found on top of the pollen sphere in the longitudinal verti-
cal plane of the cell. They were oriented with their anterior end toward the
front of the cell. Their anterior and posterior ends were attached to the pollen
mass while the middle looped upward. Two eggs measured 1.8 to 2.0 mm. in
length and one was 0.45 mm. maximum diameter.

No feeding larvae were found but the four predefecating mature larvae
from the first nest were oriented so that their posterior ends were at the rear
of the cells. Each started defecating within a few hours of being brought in
from the field. The feces are probably deposited, at least in part and perhaps
entirely, toward the rear of the cell. All had completed defecation by October
22, 1972, at which time pupal features could be seen through the larval integu-
ment. This fact indicates either that the species has a number of generations
per year or that unlike Melitta, Macro pis, Dasypoda, and Hes per apis it over-
winters as an adult rather than as a postdefecating larva. The larva closest
to the burrow was male, the farthest female; the sex of the other two is un-
known.

No parasitic bees were found in the vicinity of the nesting site.

Haplomelitta ogilviei (Cockerell)

This species was located first on October 15, 1972, at 28 km. east of Veld-
drif, Cape Province, Republic of South Africa. At that time the season was

New York Entomological Society

early in that neither the bees nor their pollen flowers were numerous. Conse-
quently the area was revisited on October 23, when the following observations
were made.

The site was in an extensive sandy region, the “Sandvelt,” that extends
from the mountains to the Atlantic coast. There is sufficient moisture for
wheat farming and the site had been rained on the day prior to my excavations.
The original vegetation was probably a semiarid scrub, although now the sur-
rounding area is wheatlands. The nesting site and also the pollen plant (Fig.
2 ) occurred along the sides of a paved road, where the ground surface was flat,
sandy, and covered with low herbs, including a very common yellow-flowered
composite and the rather rare purple-flowered Monop sis simplex (Linnaeus)
E. Wimmer, the pollen source of Haplomelitta ogilviei. Only two nests were
found; neither was shaded by the vegetation. The soil immediately below the
surface was moist because of the rain the previous day. The bee fauna of the
area was abundant, with Capicola, Scrapter , halictids, and associated parasitic
bees fairly common.

Nesting Activity: The two nest entrances were widely separated, one among
the pollen plants that grew most abundantly on the south side of the road and
the other 65 to 70 m. away on the north side. Neither entrance was associated
with an object, such as a stone or twig, and both entered the ground obliquely
where it was level. The low tumulus, 2.0 to 3.0 cm. in diameter, was to one
side of the entrance, the side away from the descending tunnel.

In each nest (Figs. 5, 6) the main tunnel was open, extending at first obliquel)'
downward and then descending nearly vertically. It was circular in cross sec-
tion on one nest 5.0 mm. in diameter, and on the other varying from 4.0 to 5.0
mm. in diameter. Both tunnels ended in single cells. In the first nest exca-
vated, the cell was open; in the other, closed. In each nest the diameter of
the tunnel immediately before the open cell was 4.0 mm. in diameter, some-
what less than the remaining burrow and the terminal 3 cm. of the tunnel de-
scended only very slightly. The cells were very shallow, being 4 and 6 cm. in
depth. Both tilted from the horizontal by about 30 degrees and had their an-
terior ends higher. They were comparatively short (7.0 and 8.0 mm. long) in
relation to width (6.0 mm.). Although without a visible lining, the cell wall
was somewhat waterproof when tested with a droplet of water and the wall
was slightly more rigid than the surrounding soil, an indication that the female
had applied some substance that had permeated the soil. The cell closure was
a partition of soil (mostly sand), evenly concave on the outside. On the inside
no spiral could be detected, the closure seeming to consist of a single ring. The
thickness of the closure at its middle was 1.0 to 1.5 mm.; at the periphery,
approximately 3.0 mm.

This species may normally construct only a single cell to the nest because

Vol. LXXXII, March, 1974

Figs. 3, 4. Highly diagrammatic representation of two nests of Capicola braunsiana.

Figs. 5, 6. Semidiagrammatic representation of the nests of Haplomelitta ogilviei show-
ing contents of cells. These illustrations are more precise than Figs. 3 and 4.

Fig. 7. Pollen ball and egg of Haplomelitta ogilviei.

Fig. 8. Posterior edge of mesosoma and base of metasoma of female of Haplomelitta
ogilviei , ventral view.

Fig. 9. Apex of median process of metasomal sternum I of female of Haplomelitta ogil-
viei, posterior view.

(1) the cells were very shallow; (2) each nest had only a single cell; (3) the
wings of the females from the nest were frayed (suggesting they had been ac-
tive for a considerable period); and (4) the female of the second nest, after
closing the cell, departed without first constructing either another lateral or
a cell immediately in front of the closed cell.

New York Entomological Society

Provisioning: Females, when gathering pollen from the flowers of Monop-
sis simplex, flew from flower to flower until a sufficient quantity of pollen had
been accumulated on their hind legs and then would depart for the nest. Both
females seen entering nests did so without first searching for entrances. One
female was followed for 65 to 70 m. on her return to the nest from the pollen
flowers, a path that took her across the road. Although strong winds caused
her to land occasionally, her general route was direct.

The open cell contained a pollen ball, 2.25 mm. in diameter, which was
spherical and of the same mealy-moist consistency as the provisions in the
other nest. Its small size and the fact that the cell was still open indicated that
it was a preliminary deposit of food. This species and certain panurgines (in-
cluding N omadopsis, Calliopsis, and most Perdita) are the only bees known to
me to shape preliminary provisions. The female removed the pollen from her
legs, shaped the sphere, and departed from the nest during a period of from one
to two minutes. The other cell contained a complete pale grayish yellow pollen
ball, 3.25 mm. in diameter, spherical, and evenly mealy-moist, which was not
coated with a waterproof substance. The female finished adding to and shaping
the provisions, deposited an egg, and closed the cell during a period of approxi-
mately one half-hour.

Development: The egg (Fig. 7), approximately 2.4 mm. long and 0.5 mm.
wide at maximum diameter, was elongate, curved, translucent white, and with
a shiny chorion. Its anterior end was blunter than the posterior and it was at-
tached to the pollen sphere by its ends, whereas the middle part did not touch
the food. The anterior tip of the egg was at the top of the pollen mass (Fig. 7)
and the rest of the egg was to the rear of the pollen ball. Although in general
its long axis was in the vertical longitudinal plane of the cell, the egg curved
slightly toward the rear as seen from above, a situation that probably was
atypical.

Cycles of Activity: The nests of the species were studied on a cool day that
was partly cloudy in the morning but cleared toward noon. No adults were
flying at 11:00 a.m., the first males were observed around noontime, and the
females became active at about 1:30 p.m. Activity decreased after 3 p.m.,
perhaps because the weather again turned cool and partly cloudy. Adults were
quite active during the early afternoon when the site was first discovered on
October 15. Although the above observations are obviously incomplete, these
bees may be active primarily during the early afternoon, a diurnal activity
fairly common among other South African bees.

When active the males flew close to the ground from one flower of Monopsis
simplex to another, and often rested on the ground or sat on the flowers. While
on the flowers they lay motionless across the blossom, antennae erect, and
head usually, if not always, at the edge of the corolla. On the ground, they

Vol. LXXXII, March, 1974

consistently rested with their hind legs spreading somewhat from their body
in a stereotyped pose that males of most bees assume. The somewhat inflated
hind basitarsus of the male may be a correlation with this posture. Because
females entered the center of the flower head first they could be easily dis-
tinguished from males, which merely sat across the bellshaped flower. Because
of their larger size and the fact that the hind legs were not slightly spread, fe-
males resting on the ground could be easily separated from males.

Females display a median projection (Fig. 8) on the ventral conjunctiva
between the mesosoma and the metasoma. This projection is apparently asso-
ciated with a peculiar modification of the first metasomal sternum which is
deeply emarginate posteriorly and possesses a thin median process. The process
expands slightly at the apex and gives rise to two ventrally directed, short,
apically flattened branches (Fig. 9). No observations were made that explained
the function of these adaptations.

Parasitism: Parasitic bees were not found in definite association with either
nest. Although Pseudodichroa , Sphecodopsis, and Sphecodes were captured
in the area, they seemed to be associated with solitary bees other than Haplo-
melitta. These parasitic bees and H. ogilviei have the same general body color
pattern, i.e., pitch black head and mesosoma and a deep red metasoma with
a black apex. The coloration seems to represent a wide-ranging mimetic color
pattern especially common among South African parasitic bees as well as some
nonparasitic forms.

Literature Cited

Rozen, Jerome G., Jr., and McGinley, Ronald J. Phylogeny and systematics of Melit-
tidae based on the mature larvae (Insecta, Hymenoptera, Apoidea). Amer. Mus.
Novitates. [In Press.]

New York Entomological Society

Synonymy in Splienophorus pertinax Olivier
(Coleoptera, Curculionidae, Rhyncliopliorinae)

Patricia Vaurie

American Museum of Natural History, New York City
Received for Publication October 2, 1973

Sphenophorus pertinax peninsularis Chittenden, 1905 (Jacksonville, Florida)
is not separable from nominate pertinax Olivier, 1807 (New Jersey). NEW
SYNONYMY.

Sphenophorus peninsularis , described as a species, was considered a subspe-
cies of pertinax by Vaurie (1951) on the basis of six specimens from eastern
Florida and Georgia and an additional specimen (Vaurie, 1967) from North
Carolina. These specimens differed from nominate pertinax (Virginia north
to northern New England and west to South Dakota) by having the three
pronotal vittae more widely separated and the sides ventrally coated, not shin-
ing. However, specimens seen subsequently from the intervening area of South
Carolina (1 $ from Little River, and 1 $ and 4 2 from Myrtle Beach) belong
to nominate pertinax because the vittae are not separated “by at least twice
their own width” (Vaurie, 1951, p. 163), and only two of the six specimens are
coated ventrally. I no longer have the example from Wrightsville Beach, North
Carolina, which I had considered to be peninsularis , but presumably its pronotal
vittae were widely separated. It was collected from a plant of the same genus,
Spartina, as nominate pertinax , but the species was alt erni flora instead of cyno-
suroides. Thus the range of nominate pertinax extends the entire length of
the Atlantic coast.

Another subspecies, ludovicianus Chittenden (type locality, New Orleans,
Louisiana), occurs from extreme northwestern Florida and Alabama north to
southern Missouri, west to eastern Texas. The vittae of this subspecies are
only partially separated and the bare elytral stripes are longer than those of
nominate pertinax.

Literature Cited

Vaurie, P. 1951. Revision of the genus Calendra (formerly Sphenophorus ) in the United
States and Mexico (Coleoptera, Curculionidae).

. 1967. A new Spenophorus from Arizona and distributional notes (Coleoptera:

Curculionidae) .

New York Entomological Society, LXXXII: 14. March, 1974.

Vol. LXXXII, March, 1974

The Genus Pterallastes Loew (Diptera: Syrphidae)

F. Christian Thompson

Department of Entomology, The American Museum of Natural History,
New York, New York

Received for Publication October 9, 1973

Abstract: The genus Pterallastes Loew is reviewed, its phylogeny and distribution dis-
cussed, the key to and figures of its species are given, Pseudozetterstedtia Shiraki is synon-
ymized under it, and bomboides (China) is described as a new species of it.

INTRODUCTION

The genus Pterallastes has been previously known only from the northeastern
United States, where it is represented by a single species, thoracicus Loew. The
discovery of a new Pterallastes species from the Szechuan Province of China
has prompted a review of the whole genus, the results of which are presented
below.

Loew (1863) described Pterallastes for two new species but Osten Sacken
(1875) later indicated that the two species were not congeneric and restricted
Pterallastes to thoracicus Loew, which he designated as the type species. He
erected Teuchocnemis for the other species, lituratus Loew, including also Mi-
lesia bacuntius Walker. Van der Wulp (1888), unaware of Osten Sacken’s
restriction of the generic limits of Pterallastes , described a new species of
Pterallastes , nubeculosus, from Argentina, which he stated was closely allied
to lituratus. Thus, even if van der Wulp’s statements about relationships of
his species to lituratus were accurate, his species would be assignable to Teuchoc-
nemis Osten Sacken, not Pterallastes. However, van der Wulp states the nube-
culosus has pilose eyes and this character state clearly excludes his species
from both Pterallastes and Teuchocnemis. Two other Nearctic species were
originally described in Pterallastes , perfidious Hunter and borealis Cole [ =

Acknowledgments: I would like to thank Drs. H. E. Evans and J. F. Lawrence of the
Museum of Comparative Zoology, Cambridge, for the privilege of studying the material
in their care; Drs. V. S. van der Goot of Amsterdam, Netherlands, and Prof. A. A. Stackel-
berg of the Zoological Institute, Leningrad, for the gift of the Pseudozetterstedtia uni-
color Shiraki material used in this study; Dr. G. B. Fairchild of the University of Florida,
Gainesville, for his information with regard to the types of species described by van der
Wulp; Dr. L. V. Knutson, of the Systematic Entomology Laboratory, USDA, for per-
mission to study the material in his care; and Dr. J. R. Vockeroth of the Entomology Re-
search Institute, Canada Department of Agriculture, for searching for the type of “ Pteral-
lastes” nubeculosus van der Wulp and his comments on the relationships of Pterallastes
and its related genera.

New York Entomological Society, LXXXII: 15-29. March, 1974.

New York Entomological Society

colei (Wirth)], but both belong to Helophilus ( Anasimyia ) (Curran and Fluke,
1926; Shannon, 1926; Wirth et al., 1965).

The search for the sister group of Pterallastes led to the discovery that Shir-
aki’s subgenus M allot a ( Pseudozetterstedtia ) was based on a species of Pteral-
lastes. An examination of “ Mallota ” unicolor Shiraki, the type species of
Pseudozetterstedtia, revealed that unicolor is very similar to thoracicus , the
type species of Pterallastes, and that the differences between the two species
are trivial. Thus Pseudozetterstedtia is here transferred from Mallota and
synonymized under Pterallastes.

Genus Pterallastes Loew

Pterallastes Loew, 1863:317 (also, 1864-201). Type-species, thoracicus Loew, subsequent
designation by Osten Sacken, 1875:64. Subsequent references: Kertesz, 1910:267 (cat.

citation, 3 spp. listed); Shannon, 1921:127 and 1922:31 (transfer of genus to Xylotinae),
1926:8 (descr. notes); Hull, 1949:375 (description); Wirth, Sedman, and Weems, 1965:
609 (cat. citation) .

Pseudozetterstedtia Shiraki, 1930:199 (as a subgenus of Mallota). Type species, unicolor
Shiraki by original designation. Hull, 1949:394 (descript.); Shiraki, 1968:246 (descript.).
NEW SYNONYM

Head. Higher than long; face bare, completely pollinose in male, frequently shiny medially
in female, with a low but distinct medial tubercle in male, concave in female; cheeks broad,
broader than long; facial grooves short, extending along lower third of eye margins and
only half way to bases of antennae; facial stripes indistinct, narrow, pilose; frontal promi-
nence low, at middle of head; frontal triangle of male short, from about two-thirds as
long to almost as long as vertical triangle, bare; vertical triangle of male long, about twice
as long as broad at occiput; front of female broad, only slightly longer than broad at base
of frontal prominence, slightly longer than face, with convergent sides above, only one-half
as broad at ocellar triangle as at base of frontal prominence, bare and shiny on lower third;
ocellar triangle clearly before posterior margin of eyes; eyes bare, narrowly holoptic in
males. Antennae short, about one-half as long as face; third segment orbicular; arista
bare, long, about twice as long as antennae and slightly longer than maximal facial width.

Thorax. Distinctly longer than broad, with long pile but pile not obscuring color of polli-
nosity; long yellow bristles above wings, on postalar calli, and posterior edge of mesopleura;
anterior mesopleura bare ; sternopleura with broadly separated dorsal and ventral pile
patches; posterior pteropleura bare; hypopleura including barrettes bare; metasterna under-
developed and bare; postmetacoxal bridge incomplete; metathoracic spiracle small; meta-
thoracic pleura bare; scutellum without distinct ventral pile fringe, with a few ventral
hairs laterally and in some specimens with many marginal hairs directed ventrally, usually
without apical emarginate rim; in some specimens with an indistinct and shallow apical
emarginate rim; legs simple; mesocoxae with three to four bristlelike hairs on posterior
surface; hind femora not swollen, with ventral spines, without basoventral setal patches.
Wing: marginal cell open; apical cell petiolate, with petiole short, about as long as humeral
crossvein; third vein moderately to strongly looped into apical cell; anterior crossvein
distinct beyond middle of discal cell, at outer third of discal cell, slightly oblique; anal cell
with a long and sightly curved apical petiole; apical and posterior crossveins continuous;
apical and discal cells without spurs at their apicoposterior corners.

Abdomen. Oval; first abdominal spiracle embedded in metathoracic epimeron. Male geni-

Vol. LXXXII, March, 1974

talia: Cerci simple, pilose; ninth tergum simple, bare; surstyli pilose, triangular in profile,
slightly asymmetric; ninth sternum with a ventrolateral membranous area on each side
and with a process laterad to this membranous area; lingula absent; superior lobes fused
to ninth sternum, pilose dorsobasally, produced into a long curved prong; aedeagus with
large earlike lateral lobes, with apical process short and stout.

DISCUSSION

Earlier workers considered that Pterallastes undoubtedly belonged with the
helophilines because of its looped third vein and open marginal cell. Shannon
(1921, 1922) was first to point out the true affinities of Pterallastes with the
milesine genera (Milesini = Xylotinae auctorum). While Hull (1949) recog-
nized six tribes in the Xylotinae, he did not place Pterallastes in any of them
nor did he place the genus in his key. I (1972) followed Hull’s basic arrange-
ment of genera within the Milesini {— his Xylotinae), but I made a few changes.
In my arrangement I placed Pterallastes in the Temnostoma group. I con-
sidered that the Temnostoma and Milesia groups were closely related because
both have the lateral lobes of the aedeagus well developed (synapomorphy).
I separated the Milesia group from the Temnostoma group on the basis of the
presence of well-developed metasterna and emarginate scutellar rims in most
of the genera of the Milesia group. However, this separation was not totally
satisfactory since the dichotomy is not clearcut; some genera have intermediate
conditions of the characters used, leaving the Temnostoma group as a symple-
siomorphic assemblage. The reevaluation of the phylogenetic relationships of
Pterallastes has lead me to discard my previous symplesiomorphic grouping
of the Temnostoma genera and to combine these genera with those of the Mi-
lesia group on the basis of their common possession of well-developed lateral
lobes of the aedeagus.

The sister group of Pterallastes is undoubtedly Palumbia + Korinchia. These
three genera are distinguished from all other syrphids by the presence of an
abundance of a peculiar type of long bristlelike hair above the wings, on the
postalar calli and usually along the margin of the scutellum (synapomorphy).
Also, Pterallastes , Palumbia , and Korinchia have the third vein (R4+5) looped
into the apical cell, another synapomorphous condition. While other syrphid
genera have the looped third vein character state, I consider this character
state to be convergent in all these other genera because the following characters,
among others, exclude the possibility of a close relationship with either Pteral-
lastes, Palumbia , or Korinchia : All eristaline genera have pilose metasterna
and patches of setulae on the hind femora; Rhinotropidia and Parrhyngia ,
both tropidines, have carinate faces and hind femora; Orthroprosopa, another
tropidine, has a pilose and divided metasterna; Syrittosyrphus, a milesine, has
a well-developed but pilose metasterna; Dideomimia, Salpingo gaster, Asiodidea,
and Didea, all syrphines, have bare humeri and five pregenital segments in

New York Entomological Society

PTERALLASTES

IS)

t— 1

1 — 1

•H

1— 1

.-q

pe!

•H

+->

Diagram 1. Phylogenetic relationships of and within the genus Pterallastes Loew. The
autapomorphic character states used are: 1, the presence of bristlelike pile on the mesono-
tum, the bare metasterna and looped third vein; 2, the closed and petiolate marginal cell;
3, the absence of a facial tubercle in both sexes; 4, the presence of a single asymmetric
ventral membranous area on the ninth sternum (male genitalia) ; 5, the presence of ventro-
lateral lobes on ninth sternum (male genitalia) ; 6, the bifid nature of the ventrolateral
lobe on the ninth sternum; 7-9, specialized character states of the species of Pterallastes
are discussed in the text under the respective species. Genus-group taxa are in capital
letters and species are in small letters.

the males. The phylogenetic relationships of Pterallastes are given in Dia-
gram 1.

Both Palumbia and Korinchia have petiolate marginal cells (synapomorphy).
Palumbia is distinguished from Korinchia by its lack of a facial tubercle in the
male (autapomorphy) and Korinchia is distinguished from Palumbia by the
presence of a single large ventral membranous area on ninth sternum of the
male (autapomorphy). I know of no other external or genitalic characters by
which I can distinguish these two genera. Thus I am combining Palumbia and
Korinchia ? but I have retained Korinchia as a subgenus of Palumbia. These
changes will be discussed in more detail in another paper.

Distribution and past dispersal. The known distribution of the genus Pteral-
lastes (map 1) is significantly enlarged with the addition of bomboides (China)
and unicolor (Japan). The sister group of Pterallastes is Oriental ( Korinchia )
and western Palaearctic ( Palumbia ) in distribution; the most plesiomorphic
species of Pterallastes , bomboides , is restricted to the Szechuan Province of

Vol. LXXXII, March, 1974

Map 1. Distribution of the genus Pterallastes with a diagram of its phylogenetic rela-
tionships.

China; the next most plesiomorphic species, unicolor , is restricted to Japan;
and the most derived species, thoracicus , is restricted to southeastern North
America. From these facts, the following history of dispersal in the genus Pteral-
lastes is postulated: 1) The genus arose in southern China, probably the same
area where bomboides is now found. 2) Some of the species dispersed north-
ward and eastward, with one ancestral species dispersing over the Bering land
bridge to North America. 3) During an ice age, probably the last one, the
northern species of the genus were forced to restrict their ranges. 4) Unicolor
in Japan and thoracicus in southeastern North America are the survivors of
this last episode of range restriction and extinction.

KEY TO SPECIES

1. Disc of mesonotum bright yellow to orange pollinose; 3rd vein forming a shallow

loop in apical cell (Fig. 4) ; abdomen black with fine yellowish or black pile, not

obscuring ground color 2

Disc of mesonotum dark brownish-black pollinose; 3rd vein forming a strong loop
in apical cell (Fig. 5) ; abdomen with long shaggy yellowish and reddish pile, ob-
scuring ground color on apical segments bomboides , n. sp.

2. Abdominal terga shiny, without pollinose markings; sterna yellow pilose, rarely with

a few black hairs on last sternum ; terga usually completely yellow pilose, rarely
black pilose on apical half or less of 3rd and 4th terga ; femora usually yellow pilose,
except black spinulose on ventral portion of hind femora and rarely with black
pile on dorsal edge of hind femora on apical half thoracicus Loew

New York Entomological Society

Abdominal terga with gray pollinose markings, 1st tergum all pollinose, 2nd with
two large transverse pollinose spots, 3rd with two small basal transverse spots; 4th
and 5th sterna (in female) black pilose; apical two-thirds of 4th and all 5th terga
(in female) black pilose; anterior four femora black pilose on anterior apical third;
hind femora black pilose on apical half unicolor (Shiraki)

Pterallastes thoracicus Loew

Pterallastes thoracicus Loew, 1963:317 (also, 1864:201). Type locality: U.S.A., Pennsylvania.
Types 1(5 1$ Museum of Comparative Zoology. Subsequent references: Banks, 1907:
450 (distribution rec.-Virginia) ; Kertesz, 1910:267 (catalog citation, 2 references); John-
son, 1910:770 (distribution recs.-New Jersey); Metcalf, 1913:94 (distribution recs.-Ohio),
1916:107 (distribution recs.-North Carolina); Banks et al., 1916:188 (distribution recs-
District of Columbia); Smith, 1919:273 (distribution recs.-Indiana) ; Britton, 1920:188
(distribution recs.-Connecticut) ; Metcalf, 1921:169-214, Figs. 97, 101 (male genitalia);
Wehr, 1922:157 (distribution recs.-Nebraska) ; Johnson, 1925:176 (distribution recs.-
Connecticut) ; Shannon, 1926:9 (distribution notes, type depository) ; Leonard, 1928:
800 (distribution recs.-New York); Curran, 1930:73 (distributional recs.-New York);
Brimley, 1938:353 (distribution recs.-North Carolina) ; Wirth, Sedman and Weems,
1965:609 (catalog citation, Nearctic distribution).

Male. Head: black; face silvery pollinose, with a low medial tubercle, with tubercle lower
than frontal prominence ; cheeks shiny on anterior half, whitish pollinose and pilose on
posterior half ; frontal lunule orange ; frontal triangle silvery pollinose ; vertical triangle
silvery pollinose except very sparsley pollinose on ocellar triangle, yellow pilose; occiput
silvery-white pollinose and pilose below becoming yellow on upper half. Antennae orange,
frequently with brownish tinge; third segment small, only about as large as metathoracic
spiracle; arista orange.

Thorax. Dorsum yellow pollinose and pilose, with pile of medium length except long
bristlelike hairs above wings and on postalar calli; scutellum yellow pollinose and pilose;
pleura silvery pollinose, yellowish to white pilose; squamae and plumulae white; halters
white to orangish; legs black except as follows, yellow femora tibia joints, yellowish basal
third of middle tibiae, orange middle and hind tarsi; in some specimens hind metatarsi
with brownish tinge; pile yellowish except black on front tarsi and ventral portion of tibiae
and hind femora and rarely on dorsal edge of hind femora on apical half. Wings: hyaline
or with a slight grayish tinge apically, microtrichose except bare narrowly behind anal
vein and in front of auxiliary vein; third vein with shallow loop in apical cell.

Abdomen, black, shiny, with slight metallic bluish luster under strong light; in some speci-
mens appear reddish brown under strong light; usually completely whitish yellow pilose,
rarely with black pile on apical portions of third and fourth terga (see below under dis-
cussion section). Male genitalia: Surstyli triangular, with ventral margin virtually straight,
not produced basoventrally ; ninth sternum with ventrolateral membranous area small

Figs. 1-8. Figs. 1-3. Head of Pterallastes , lateral view; 1. thoracicus Loew, male; 2.
bomboides , n. sp., male; 3. unicolor (Shiraki). 4-5. Wings of Pterallastes ; 4. thoraci-
cus Loew; 5. bomboides , n. sp. Figs. 6-8. Aedeagi of Pterallastes , lateral view. 6. thoraci-
cus Loew; 7. unicolor (Shiraki); 8. bomboides , n. sp.

Vol. LXXXII, March, 1974

New York Entomological Society

and apicomedial to lateral process, with lateral process bifid and directed apically; supe-
rior lobes only pilose dorsobasally, produced into a short broad apical prong with a single
large ventral tooth, with a few short bristlelike hairs on ventral tooth; aedeagus with lateral
lobes triangular, with apical process slender and longer than in both bomboides and uni-
color and with a single bump on ventral margin.

Female. Quite similar to male except for normal sexual dimorphism; less black pile on
legs; lower third of front shiny and upper two-thirds yellow pollinose and pilose; and
lower medial third of face frequently shiny.

MATERIAL EXAMINED

The two syntypes in the Loew Collection at the Museum of Comparative Zoology and
some 80 additional specimens of both sexes from the following states and counties: Kan-
sas (Douglas) ; Connecticut (Fairfield) ; New York (Westchester, New York City) ; New
Jersey (Bergen, Essex, Middlesex) ; Pennsylvania (Philadelphia, Montgomery, Delaware,
Westmoreland) ; Maryland (Arundel, Prince George, Montgomery, Calvert) ; District of
Columbia; Virginia (Arlington, Fairfax); North Carolina (Buncombe); and Georgia.
Some of these specimens were labeled as collected in association with the following plants:
Ceanotheus , Solidago, Viburnum nudum , and Castanea dentata. The earliest collection
recorded was 23 May (Virginia); the latest was 7 October (New York), with June and
August being the months with the most numerous records. More detailed information
about this material is available from the author.

DISCUSSION

The differences between P. thoracicus and unicolor or bomboides are dis-
cussed under the latter species. Among the material of P. thoracicus examined
there was some variation in the extent of black pile on the abdomen — ranging
from a few black hairs intermixed with yellow pile on apical portion of the
4th ( S ) or 5th ( $ ) segments to large triangular areas of solid black pile on
apical half or more of the third through fourth ($) or 5th (?) segments.

Pterallastes bomboides, n. sp.

Male. Head: black; face silvery pollinose, with a distinct medial tubercle, with tubercle
almost as high as frontal prominence; cheeks shiny; frontal lunule yellowish orange;
frontal triangle silvery pollinose; vertical triangle grayish pollinose, black pilose; occiput
silvery-gray pollinose, white pilose below becoming yellow on upper half. Antennae orange,
with brownish tinge in paratype; third segment large, larger than metathoracic spiracle;
arista orange.

Thorax. Black; dorsum dark brownish-black pollinose except narrowly silvery-gray polli-
nose laterally and on scutellum, long shaggy yellow pilose laterally and on scutellum, yel-
low and black pilose medially and almost completely black pilose behind sutures; pleura

Figs. 9-14. Figs. 9-11. Ninth tergum and associated structure, of Pterallastes , lateral
view; 9. thoracicus Loew; 10. unicolor (Shiraki) ; 11. bomboides , n. sp. Figs. 12-14.
Left surstyli of Pterallastes , lateral view; 12. thorcicus Loew; 13. unicolor (Shiraki); 14.
bomboides , n. sp.

Vol. LXXXII, March, 1974

New York Entomological Society

silvery-gray pollinose, long whitish to yellowish pilose; legs black except yellowish tips of
femora and bases of tibiae and orangish middle and hind tarsi, yellow to white pilose ex-
cept black pilose on ventral part of hind femora and apical three tarsal segments; plumulae
brown; squamae white with brownish margin and fringe; halters brown; wings hyaline
microtrichose except bare narrowly behind anal vein and in front of auxiliary vein, third
vein with a strong loop in apical cell.

Abdomen black, shiny except reddish apical half of fourth segment and genital segments;
short black pilose on medial third of second tergum ; white pilose on first tergum, sterna,
lateral third of second and lateral margins of third terga, long on lateral margins and sterna;
long yellowish red pile on third and fourth terga, obscuring ground color; short yellowish
pile on genital segments. Male genitalia: Surstyli triangular, strongly concave on ventral

margin, with a small triangular inner tooth on ventroapical margin of right surstyli ; ninth
sternum with ventrolateral membranous area large and posteromedial to lateral process,
with lateral process simple and directed ventrally; superior lobe produced into a long
slender apical prong, without teeth on ventral margin, generally pilose, with a small tuft
of hairs on ventroapical angle; aedeagus with lateral lobes broadly triangular, with apical
process stout and with an even ventral margin.

Female. Quite similar to male except for normal sexual dimorphism; lower third of front
shiny and upper two-thirds brownish pollinose and black pilose; reddish ground color of
abdomen is reduced.

MATERIAL EXAMINED

Holotype and paratype males, CHINA, Szechuan, West of Chetu Pass, near Tatsielu,
13,000 to 14,500 ft; D. C. Graham; allotype female, west of Chego Pass, July 13-18, 1923,
D. C. Graham. The holotype and allotype are in the American Museum of Natural History
collection; paratype is retained in the author’s collection.

DISCUSSION

Besides the differences mentioned in the above key, P. bomboides differs
from both P. thoracicus and unicolor in: (1) the facial tubercle is larger and
more distinct, not low and obscure; (2) the vertical triangle in male and front
and vertex in female is black pilose, not yellow or tawny; (3) the third anten-
nal segment is large, larger than metathoracic spiracle, not the same size; and
(4) the plumulae and halters are brown, not white to whitish orange. The
specific name bomboides is an adjectival form used as a substantive in the geni-
tive case and alludes to the mimetic similarity to Bombus.

Pterallastes unicolor (Shiraki)

Pseudozetterstedtia unicolor Shiraki, 1930:200. Type locality: Japan, Hokkaido, Josankei

and Sapporo; Honshu, Wakayama, Towada and Chuzenji. Types $ $ Ent. Mus., Natn.

Inst. Agric. Sci., Tokyo

Mallota unicolor : Sack, 1932:337 [descript., figs, (head)]; Stackelberg, 1950:287 (notes);

Violovitsh, 1955:350 [distr. recs. (Kuril Is.), notes]; Violovitsh, 1960:247 [distr. recs.

(Kuril Is.), notes]; Shiraki, 1968:246 (descript.).

Male. Head: black; face yellowish white pollinose, with a very low medial tubercle, with
tubercle much lower than frontal prominence; cheeks shiny black on anterior half, yellow-

Vol. LXXXII, March, 1974

ish white pollinose and pilose on posterior half ; frontal lunule brownish yellow ; frontal
triangle yellowish white pollinose ; vertical triangle yellowish white pollinose anteriorly,
slightly more brownish yellow pollinose posteriorly, tawny pilose; occiput yellowish white
pollinose and pilose below becoming more orange or tawny yellow on upper half. Anten-
nae brownish orange, black pilose; third segment small, oval, only about as large as meta-
thoracic spiracle; arista brownish orange.

Thorax, black; dorsum orange yellow to deep orange pollinose and pilose, with pile of
medium length except long bristlelike hairs above wings and on postalar calli; scutellum
the same as dorsum ; pleura more or less grayish pollinose, with meso-, ptero-, and sterno-
pleura distinctly gray pollinose, yellow pilose ; squamae and plumulae orange ; halters
white to orange; legs: black, except yellowish brown femoral-tibial joints and basal seg-
ments of middle tarsi; coxae and front four trochanters yellow pilose, with pile bristle-
like on coxae; anterior four femora whitish yellow pilose except black on anterior apical
half; anterior tibiae black pilose except for scattered yellow hairs on posterior half; ante-
rior tarsi all black pilose; middle tibiae whitish yellow pilose except black pilose on apical
ventral half; middle tarsi yellow pilose with apical segments with some black hairs inter-
mixed; hind trochanter with black setulae; hind femora whitish yellow pilose on basal
half, black pilose on apical half and with black setulae on ventral edge; hind tibiae whit-
ish yellow pilose on basal half, black pilose on apical half; hind tarsi all black pilose. Wings:
hyaline or with a slight grayish tinge apically, microtrichose except bare narrowly behind
anal vein and in front of auxiliary vein ; third vein with a moderately shallow loop in apical
cell.

Abdomen black; venter gray pollinose, white pilose on first through third sterna, with
fourth sternum black pilose; first tergum grayish pollinse and yellow pilose; second tergum
with a pair of large transverse yellowish gray pollinose spots, elsewhere brownish gray
pollinose, yellow pilose ; third tergum with narrow basal transverse yellowish gray pollinose
spots, elsewhere brownish gray pollinose except slightly shiny submedially, yellow pilose,
except usually with a few black hairs on apical margins, rarely all black pilose on
apical third; fourth tergum with yellow gray pollinose basal transverse spots simi-
lar to those on third but very narrow, black pilose except yellow pilose on baso-
lateral corners, male genitalia black pilose, sparsely grayish pollinose. Male genitalia:
surstyli triangular, slightly concave on ventral margin, without any teeth ; ninth ster-
num with ventrolateral membranous area small and apicomedial to lateral process,
with lateral process bifid and directed slightly dorsoapically ; superior lobes only
pilose dorsobasally, produced into long slender apical prong with three large ventral
teeth, with two long bristles on ventroapical margin; aedeagus with lateral lobes almost rec-
tangular in shape, with apical process more slender than in bomboides and with ventral
margin very irregular.

Female. Quite similar to male except for normal sexual dimorphism; lower third of front
shiny and upper two-thirds orangish-yellow pollinose and pilose; fifth abdominal segment
black pilose.

MATERIAL EXAMINED

JAPAN, Honshu, Iwato, 840 m, 21 July 1971, 1 $, V. S. van der Goot and J. A. W.
Lucas (FCT) .

DISCUSSION

P. unicolor is very similar to thoracicus differing principally in the structure
of the male genitalia; in the gray pollinose marking on abdominal terga; and
in the pile and pollinosity that are tawny orange, not pale yellow. Also, P.

New York Entomological Society

Figs. 15-20. Figs. 15-17. Ninth sternum and associated structures of Pterallastes , lateral
view; 15. thoracicus Loew; 16. unicolor (Shiraki) ; 17. bomboides, n. sp. Figs. 18-20.
Apical half of ninth sternum and associated structure of Pterallastes , dorsal view; 18.
thoracicus Loew; 19. unicolor (Shiraki) ; 20. bomboides , n. sp.

Vol. LXXXII, March, 1974

unicolor has more extensive black pile on the legs and abdomen than the typical
specimens of thoracicus.

“ Pterallastes ” nubeculosus van der Wulp

? nubeculosus van der Wulp, 1888:372 ( Pterallastes ). Type locality: Argentina, Prov.

Tucuman. Type $ (lost, see below). Subsequent references: Kertesz, 1910:267 (cat. cita-
tion, 1 reference); Brethes, 1907:293 (cat. citation); Fluke, 1957:1555 (cat. citation).

“Brownish-black antennae and arista, femoral apices, tibiae and tarsi rufous; eyes
strongly pilose; front, thorax scutellum and abdomen densely ocher-yellow pilose; wings
hyaline, base and costa cinereous clouded.

“Properly fitting the generic characteristics of the genus Pterallastes Loew and appears
closely allied to the North American species Pt. lituratus Loew, but nevertheless in many
respects distinct from it.

“Ground color brownish-black, shiny, with scutellum brownish red. Face pale yellowish
pollinose, the obvious facial tubercle and oral margin1 shiny, on the sides with similar colored
pile; vertical triangle very small; eyes pilose; occiput behind eyes pale yellowish pollinose,
with pile similarly colored, darker above. Antennae reddish brown, arista of a lighter color.
Front, thorax and scutellum with rather thick light ocher-yellow pile; similar pile on ab-
domen, mostly on sides and on hind edges of segment, anus curved down ventrally. Legs
blackish brown, tips of femora, tibiae and tarsi of a lighter color, almost reddish yellow;
hind femora thicken in the middle, hind tibiae curved. Halters yellow. Wings hyaline,
with base and front edge reddish gray clouded.

“One $ , prov. Tucuman, Argentina.”

DISCUSSION

“Pterallestes” nubeculosus van der Wulp has not been recognized since its
original description, which is translated above. Van der Wulp states that his
species “properly” fits the generic characteristics of Pterallastes, which were
given by Loew as “ Pterallastes forma ac figura totius corporis, praecipue capi-
tis, et pictura Myoleptam simulans, alas Helophili habetP The principal char-
acter states, which Loew was probably referring to in his description, are the
sexually dimorphic face of Myolepta and the looped third vein and open sub-
marginal cell of the helophilines. Since van der Wulp had only a male of his
species it was impossible for him to know whether his species had a sexually
dimorphic face as in Myolepta or Pterallastes but clearly it is safe to assume
that his species had the looped third vein and open submarginal cell. The com-
bination of these two character states, along with the pilose eyes, restricts the
placement of van der Wulp’s species to either Mallota or Quichuana among the
known neotropical syrphids. It is always possible that van der Wulp’s species
could represent a new genus, but 1 think this possibility is highly unlikely.

1The original text for this phrase is: “ die de duidelijke gezichtshult en den mondr ad

vrij laatp which literally translates as “the obvious facial tubercle and mouthedge leaving
free.” I have assumed this to mean that the oral margin and facial tubercle are “free” from
pollinosity.

New York Entomological Society

Unfortunately it appears that the type of nubeculosus is lost. Dr. J. R.
Vockeroth, at my request, searched for this type while examining the various
collections that contain van der Wulp material, but he was unable to find any
trace of it. In van der Wulp’s paper there is no mention of any particular col-
lection in which the material was deposited, only that the material came from
Prof. H. Weyenbergh, Jr. According to Horn and Kahle (1937:301) all the
Weyenbergh material was destroyed. Thus, if van der Wulp did return the
material to Weyenbergh, it is lost. Dr. G. B. Fairchild (in litt.) also was un-
able to find the type of another species ( Pangonia lasiophthalma ) described
in the same paper.

Literature Cited

Banks, N. 1907. Captures of Diptera. Ent. News, 18: 450.

Banks, N., Greene, C. T., McAtee, W. L., and Shannon, R. C. 1916. District of Colum-
bia Diptera. Proc. Biol. Soc. Washington, 29: 173-203.

Brethes, J. 1907. Catalogo de los Dipteros de las Republicas del Plata. An. Mus. nac.
Buenos Aires, (3)9: 277-305.

Britton, W. E. 1920. Checklist of the insects of Connecticut. Bull. State Geol. Nat.
Hist. Survey Connecticut 31, 397 pp.

Curran, C. H. 1930. Report on the Diptera collected at the Station for the study of
Insects, Harriman Interstate Park, N. Y. Bull. Amer. Mus. Nat. Hist., 61: 21-115
(1931).

, and Fluke, C. L. 1926. Revision of the Nearctic species of Helophilus and

allied genera. Trans. Wisconsin Acad. Sci., Arts, Letters, 22: 207-281, 3 pis.

Fluke, C. L. 1956-57. Catalogue of the family Syrphidae in the Neotropical region
(Diptera). Revta. Brasil. Ent., 6: 193-268, 7: 1-181.

Hull, F. M. 1949. The morphology and interrelationship of the genera of syrphid flies,
recent and fossil. Trans. Zool. Soc. London, 26: 257-408, 25 figs.

Johnson, C. W. 1910. Order Diptera. Pp. 703-814, Figs. 293-340. In Smith, J. B.,
The insects of New Jersey. Annu. Rpt. N. J. State Mus., 1909: 15-1888, 340 figs.

. 1925. Fauna of New England. 15. List of the Diptera or two-winged flies.

Occas. Papers Boston Soc. Nat. Hist., 7(15): 1-326, 1 fig.

Kertesz, K. 1910. Catalogus dipterorum hucusque descriptorum. Vol. 7, 470 pp. Lip-
siae, Budapestini [= Leipzig, Budapest].

Leonard, M. D. 1928. List of the insects of New York with a list of the spiders and
certain other allied groups. Mem. N.Y. (Cornell) Agr. Expt. Sta., 101: 1-1121

(1926).

Loew, H. 1863. Diptera Americae septentrionalis indigena. Centuria quarta. Berlin.
Ent. Ztschr., 7: 275-326 (also published in Loew, 1864: 159-210).

. 1864. Diptera Americae septentrionalis indigena. Vol. 1, 266 pp. Berolini

[— Berlin], 1861.

Metcalf, C. L. 1913. The Syrphidae of Ohio. Bull. Ohio Biol. Survey, 1: 7-122, 3

figs., 11 pis. (= Bull. 1) [= Bull. Ohio State Univ. 17:(31).]

-. 1916. A list of Syrphidae of North Carolina. J. Elisha Mitchell Sci. Soc., 32:

95-112.

. 1921. The genitalia of male Syrphidae: Their morphology, with especial refer-
ence to its taxonomic significance. Ann. Ent. Soc. Amer., 14(3): 169-214.

Vol. LXXXII, March, 1974

Osten Sacken, C. R. 1875. A list of the North American Syrphidae. Bull. Buffalo
Soc. Nat. Sci., 3: 38-71.

Sack, P. 1928-1932. 31. Syrphidae. In Lindner, E., ed., Die Fliegen der Palaeark-
tischen Region. Bd. 4, pt. 6, pp. 1-48 (1928), 49-144 (1929), 145-240 (1930), 241-
336 (1931), 337-451 (1932).

Shannon, R. C. 1926. Review of the American xylotine syrphid-flies. Proc. U.S. Natl.
Mus. #2635, 69(9): 1-52.

Shiraki, T. 1930. Die Syrphiden des Japanischen Kaiserreichs, mit Beriicksichtigung
benachbarter Gebiete. Mem. Fac. Sci. Agric. Taihoku Imp. Univ., 1 : 9-xx, 1-446,
100 figs.

. 1968. Fauna Japonica. Syrphidae (Insecta: Diptera). v. Ill, 272 pp., 47 pis.

Biogeogr. Soc. Japan, Tokyo.

Smith, M. R. 1919. A list of Syrphidae of Northern Indiana. Canad. Ent., 51: 273.

Stackelberg, A. A. 1950. [Brief survey of the palaearctic species of the genus Mallota
Meigen (Diptera, Syrphidae)]. Ent. Obozr., 31: 285-296 (in Russian.)

Violovitsh, N. A. 1955. [New and little-known hover flies (Diptera, Syrphidae) from
Kunashir Island]. Ent. Obozr., 34: 350-359, 12 figs.

— 1960. [A contribution to the knowledge of the hover flies fauna (Diptera, Syr-

phidae) of Sachalin and the Kuril Isles]. Trudy vses. ent. Obshch., 47: 217-272.

Wehr, E. 1922. A synopsis of the Syrphidae of Nebraska with descriptions of new spe-
cies from Nebraska and Colorado. Univ. Stud., Lincoln, Nebraska, 22: 119-162.

W ELLiSTON , S. W. 1886. Synopsis of the North American Syrphidae. U.S. Natl. Mus.
Bull., 31: i-xxx, 1-335 (actual date, 1887).

Wirth, W. W., Sedman, Y. S. and Weems, H. V., Jr. 1965. Family Syrphidae. In Stone,
A., C. Sabrosky, W. W. Wirth, R. H. Foote, and J. Coulsen. 1965. A catalog of
the Diptera of America North of Mexico. U.S. Dept. Agri. Handb. #276, 1696 pp.

New York Entomological Society

Further Observations on the Natural History of
Philaethria dido dido (Lepidoptera: Nymphalidae: Heliconiinae)

Allen M. Young

Department of Biology, Lawrence University, Appleton, Wisconsin 549 11

Received for Publication November 5, 1973

Abstract: Observations on the life cycle and natural history of Philaethria dido dido
(Lepidoptera: Nymphalidae: Heliconiinae) as studied in northeastern Costa Rica are

summarized. Emphasis is placed on: (1) additional descriptions of life stages, (2) a larval
food plant record ( Passiflora ^ztz/o/za-Passifloraceae), (3) developmental time in the
laboratory (37-39 days), and (4) various behavior patterns associated with oviposition
and larval development. These data and other information from the literature are dis-
cussed from the standpoint that P. dido is a specialized insect of tropical rain forests and
that it has a widespread but strongly localized geographical distribution pattern in Central
and South America.

INTRODUCTION

Owing perhaps to their apparently close evolutionary history with the
Passifloraceae and their roles in mimicry complexes, neotropical butterflies of
the subfamily Heliconiinae have received considerable attention from biologists
interested in phylogeny, ecology, and behavior (e.g., Kaye, 1917; Beebe, 1955;
Beebe, Crane, and Fleming, 1960; Crane, 1957; Emsley, 1963; 1964; 1965;
Benson, 1971; Turner, 1971; Brown and Mielke, 1972). Of the seven genera in
the subfamily, the genus Philaethria comprises a single conservative subdivision
and represents a different lineage of heliconiine evolution from the other two
subdivisions (Emsley, 1963). Since Philaethria is conservative in the sense of
exhibiting many subfamily characteristics in their most generalized form and
has very few specializations (Emsley, 1963), studies of member species in this
genus are predicted to provide more information on the general ecological and
behavioral adaptations of the subfamily as a whole. The paper of Beebe,
Crane, and Fleming (1960) gives a detailed account of the early stages and
food plants of Philaethria dido dido (Clerck) on Trinidad, and Brown and
Mielke (1972) provide similar food plant data for both P. dido dido and P.
wernickei (for two subspecies, wernickei and pygmalion) in extra -Amazonian
and Amazonian Brazil. And while P. dido has a broad geographical distri-
bution in wet tropical regions of Central and South America (Emsley, 1963;

Acknowledgments: This research is a by-product of N.S.F. Grant GB-33060, with logistic
support provided by the Costa Rican Program of the Associated Colleges of the Midwest.
I am grateful to Dr. J. Robert Hunter of A.C.M. for allowing me to conduct studies on
his property, Finca Tirimbina. Dr. Woodruff W. Benson (Rio de Janiero) kindly identi-
fied the larval food plant and also provided some of his own data on heliconiines exploit-
ing this species. Keith S. Brown, Jr. (Rio) later identified the same food plant for Heliconius
hecale.

New York Entomological Society, LXXXII: 30-41. March, 1974.

Vol. LXXXII, March, 1974

Barcant, 1970), a good deal more needs to be learned about the biology of
this interesting butterfly throughout this range. At least part of the reason
why the butterfly has not been studied in Central America stems from the
elusive habits of this insect: Many authors report that it is a high flier, pre-
ferring the canopy of virgin forests; it is seldom seen near the ground.

This paper summarizes some further observations on the biology of P. dido
dido (Fig. 1) on the Central American mainland, with an emphasis on life
cycle and natural history. The descriptions of the early stages, so well pre-
sented by Beebe, Crane, and Fleming (1960), are supplemented here with
the first illustrations of the egg, third and fourth instars, and pupa. An
egg-adult developmental time as measured for one species of Passijlora is
given for the first time. These and other aspects of natural history comprise a
new attempt to study the butterfly in Central America.

METHODS

Studies were initiated on July 1, 1973 when I made my first field record of
oviposition in P. dido in the thinned-out old secondary forest (Fig. 2) that
borders the Rio Tirimbina near La Virgen, Heredia Province, Costa Rica.
Oviposition was studied on the gentle slope of forest rising from the river
but not including the narrow strip of very disturbed young secondary growth
where another wet-forest butterfly, Parides areas mylotes, has been studied
(Young, 1973). Much of this gentle slope of old secondary forest will be
eliminated within two years in a land-clearing project to raise cattle, and it
therefore represents one habitat or part of a larger habitat of P. dido that is
endangered. This general region of Costa Rica on the Caribbean drainage of
the Central Cordillera to the west is a basal belt transitional zone between
montane and premontane tropical wet forest, and the elevation is about 225
meters.

The thinned-out condition of the forest where oviposition has been repeatedly
observed is the result of farm workers beginning to clear the land with
machetes, but this was postponed for two years when I spoke to the owner
of this land so that various ecological studies of cicadas and butterflies could
be completed. The original understory was considerably more dense than it
is now (Fig. 2), but the plant species used for oviposition by P. dido has
remained intact. Oviposition and general flying behavior of adult P. dido
were observed in this thinned-out forest by walking slowly through 200 meters
of forest, then moving about 15 meters to one side (up the slope) and repeating
this; the procedure was repeated four times giving a total of five 200-meter
transects, and the census was done on July 1-2, 1973; August 12-13, 1973;
September 1, 1973. These censuses were conducted while sampling nested
quadrats in the forest for exuviae of a large cicada ( Zammara sp.) which was
very active in the adult stage at this time.

New York Entomological Society

Eggs seen to be oviposited in the field were brought into the laboratory for
rearing studies. From eggs collected in this way during July and August, I
reared a total of eight individuals to the adult stage, keeping records on each
one for: (1) external morphology and coloration, (2) duration (days) of each
life stage, including separate instars of larvae, and (3) behavior patterns of
larvae. Each individual was reared separately in a small clear plastic bag
containing fresh clippings of the food plant; the bag was always kept tightly
tied to prevent desiccation of the food. All bags (a total of thirteen) were
kept together on a shaded shelf in a second-story apartment (Apartamentos
Miami) in San Jose and cleaned every three or four days for removal of
fecal material and excess condensation. In order to observe intraspecific
interactions among larvae, I occasionally would place three or four individuals
together in a plastic bag containing one or two cuttings of the food plant.
My interest here was to observe aggressive encounters, or lack thereof, among
larvae. All life stages were photographed at the same time that notes were
made on coloration and morphology. The adult specimens obtained from
these rearings are preserved in my permanent collection and can be made
available to interested workers upon request. Owing to the reputed high
local variation in the wing color pattern of adult P. dido (e.g., Emsley, 1963),
students of intraspecific variation in tropical insects might find such collections
useful for systematic studies. Pupal shells have also been preserved from
this study.

RESULTS

Life Cycle

Since there exists one excellent text account of the life stages of P. dido
(Beebe, Crane, and Fleming, 1960), it would be redundant to describe the
stages as seen in the present study. Rather, I refer the interested reader to the
account of Beebe, Crane, and Fleming, and wish only to supplement those
observations with figures of the egg, third instar, fourth instar, and pupa
(Fig. 3) — stages described but not pictured in Beebe, Crane, and Fleming.
I also present some observations on color differences in the fifth instar be-
tween Costa Rica and Trinidad and give specific developmental time data
for P. dido (not given in Beebe et al., 1960), contrasting this developmental
time to the general pattern offered by Beebe et al.

Beebe et al. (1960) report that the head of fifth instar is bright orange,
but for the Costa Rican individuals studied the head is clearly beige and
slightly shiny. At the base of each of the two head scoli there is an irregularly
shaped black spot not given in Beebe et ah, and the portions of the body
described in Beebe et ah as being white are pale green throughout the fifth
instar in the Costa Rican P. dido studied. The supralateral thoracic scoli
in the Costa Rican fifth instars do not have black tips and the basal portion

Vol. LXXXII, March, 1974

Fig. 1. An adult male Philaethria dido dido (dorsal view). This is one of the individuals
obtained from eggs collected at Finca Tirimbina, La Virgen, Heredia Province, Costa Rica,
July 1973.

of each is distinctly orange while the shaft is red. This does not occur in
Trinidad P. dido. Furthermore, the sublaterals of abdominal segments in
Costa Rica are greenish-white with faint black tips, and not the white-orange-
black pattern as seen on Trinidad (Beebe et al., 1960). The longest scoli on
the Costa Rican fifth instars are 8-9 mm long.

Beebe et al. (1960) also state that the usual duration of the egg stage
in Trinidad heliconiines (including P. dido) is about four days. For Costa
Rican P. dido at about 23 °C and humid confined bags, the egg stage lasts
seven days. The entire larval period is about nineteen days, with the first
instar lasting three days, the second three days, the third three days, the
fourth about four days, and the fifth about six days.

Beebe et al. (1960) state that pupae of the “group A type,” which includes
(in addition to P. dido) Agraulis vanillae, Dione juno, Dryadula phaetusa ,
Dry as julia , and Heliconius doris, are generally brown in color. I figure the
pupa of P. dido here (Fig. 3F) to point out its distinct mottled color pattern
as seen in Costa Rica. The pupa is about 30 mm long and resembles a piece
of broken-off, rough tree bark, being mottled boldly in various shades of
brown and gray. It is perhaps one of the most cryptic of heliconiine pupae.

New York Entomological Society

Fig. 2. The thinned-out old secondary forest habitat at the edge of transitional-zone
rain forest at Finca Tirimbina where female P. dido dido oviposits on Passiflora vitifolia
(Passifloraceae) vines that grow along the ground and over rocks and tree stumps.

The pupa for Costa Rican P. dido lasts about eleven days for the male and
twelve or thirteen days in the female. Beebe et al. do not give a typical
duration figure for the pupal stage.

From these considerations, the overall egg-adult developmental time for
P. dido in laboratory culture in Costa Rica is 37 to 39 days. Some discrepancies
between the present study of P. dido and the larval developmental time gen-
eralizations of Beebe et al. (1960) for heliconiines include: (1) the first and
second instars of three days as opposed to two days, (2) the fourth instar of
four days, and (3) the usual duration of the fifth instar of six days instead of
the usual five days. The overall larval period noted by Beebe et al. is twelve
days as opposed to the nineteen days found here for P. dido.

Larval Food Plant

The plant used for oviposition is Passiflora vitifolia and it occurs at the
study site as a large, sprawling vine on the ground, logs, and tree stumps.
The vine does not go into the canopy here, but it is difficult to say if this has

Vol. LXXXII, March, 1974

Fig. 3. Some life stages of P. dido dido in Costa Rica. (A) egg, (B) third instar larva,
(C) fourth instar larva showing one aspect of scoli distribution and coloration, (D) fourth
instar larva showing another aspect of scoli distribution and coloration, (E) fifth instar,
dorsal view, and (F) the pupa, lateral view.

New York Entomological Society

always been the case owing to the selective thinning out of trees by farmers.
But the large size of the sprawling vine near the ground is indicative of
successful thriving in this zone of the forest environment. The leaves are
thick and with rough texture. I have not found other heliconiines on this
species of Pas si flora in the study site, but at Rincon de Osa, a lowland trop-
ical wet-forest site on the southern Pacific coast of Costa Rica, Dr. Woodruff
W. Benson (pers. comm.) has found P. dido and several other heliconiines
feeding on this plant. It is a species of Pas si flora that has clearly been exploited
by heliconiines at different stages in the evolutionary development of the sub-
family. It appears to be a favorite food plant for the subfamily at wetter low-
land sites in Costa Rica and other regions of southern Central America.

The larvae of all instars for P. dido feed primarily on the older and larger
leaves of P. viti folia as studied in the laboratory. But this may also be true
in the wild since eggs are laid on older leaves (see below).

Oviposition Behavior

Some authors (e.g., Barcant, 1970; Brown and Mielke, 1972) have com-
mented that P. dido is a high flier over treetops in forests. But oviposition
clearly sometimes occurs near the ground as seen in the present study. For
all of the times I observed oviposition, it occurred in sunny weather, either in
the morning or early afternoon. The typical flight pattern of female P. dido
would be to appear suddenly in the lower portion of the thinned-out forest
and make several attempted ovipositions before actually laying an egg. In
several instances, the female would actually grasp a tendril or leaf with wings
fluttering for stability, but an egg was not laid. I followed one female on
July 2 and it was almost twelve minutes before an egg was laid, despite
several intervening attempts at oviposition. The pattern is somewhat frus-
trating to the observer since it almost appears as if the Passi flora species in
question was not acceptable to P. dido.

The bright yellow large egg is laid either on the ventral side of an older
leaf of P. vitifolia, or else on a dead tendril. In a total of fourteen eggs
actually observed to be oviposited, nine were on leaves and the remaining five
on dead tendrils. The female flies very erratically between ovipositions or
attempted ovipositions, suddenly darting up into the canopy and then coming
back down to the Passi flora. A single female will remain in the same general
area where the Passi flora is growing for as long as 25 minutes in my ex-
perience. Out of a total of probably five different females of P. dido observed
ovipositing during July and August, three of these were very fresh individuals,
and could not have been more than a day or two out of the chrysalis. It is
especially easy to distinguish very young or fresh adults in P. dido since, as
noted by Emsley (1963), wing color fades very rapidly in this species.
Females of P. dido may therefore be mated almost immediately after emerging

Vol. LXXXII, March, 1974

from their pupae. It is not known, however, if males wait near female pupae
for future mates, as noted for Ornithoptera (Papilionidae) species (Borch
and Schmid, 1973).

Larval Behavior

The first instar larva devours its empty eggshell, and during subsequent
molts, larvae also eat their castoff exuviae. These two behavior patterns have
not been previously reported for P. dido by Alexander (1961a). In terms of
feeding on leaf tissue, the larvae exhibit the channeling behavior reported by
Alexander (1961a) for Dione. The larvae of P. dido do not remove fecal mater-
ial with the jaws as described in Alexander (1961a) for the larvae of several
Heliconius. The resting behavior, not studied for P. dido by Alexander, is very
variable in the laboratory, but usually involves the larva facing the direction of
feeding; often a larva assumes a hooked or “J” position as noted by Alexander
for Heliconius isabella , H. Melpomene, and H. ricini.

Weaving is also well expressed in the construction of silken pathways along
stems and ventral sides of leaves. Disturbed larvae are very mobile for several
minutes, and they walk very fast. As in Heliconius erato (Alexander, 1961a),
the larvae of P. dido are clearly asocial and very aggressive and there are no
signs of any gregarious behavior (including social defecation) of the type so
evident in Dione juno (Alexander, 1961a; Muyshondt, Young, and Muyshondt,
1973). Individual larvae do not share the same leaf without fighting, as seen
in laboratory culture. Another aspect suggesting that P. dido is truly a solitary
species is the complete lack of synchrony among larvae for feeding and resting.
From these observations, I suspect that the larvae of P. dido are considerably
more aggressive and asocial than originally predicted by Alexander, thus being
closer to the behavior patterns of H. erato than to H. melpomene or Dione.

DISCUSSION

The above observations are intended to supplement what has been already
determined of the life cycle and natural history of P. dido as studied in
Trinidad (Beebe, Crane, and Fleming, 1960; Alexander, 1961#; Emsley,
1963; Barcant, 1970) and Brazil (Brown and Mielke, 1972). This report
concerns the butterfly in Central America where I am sure several researchers
have reared the species in the past.

One of the interesting characteristics of the distributional biology of P. dido
is the apparently widespread but strongly localized occurrence of the butterfly
in the rain forests of Central and South America. The species is susceptible
to regional restriction by land barriers: Brown and Mielke (1972) comment
that in extra-Amazonian Brazil this species and several other heliconiines
are restricted to tropical regions by high southeastern coastal mountains but
that it and P. wernickei are sympatric over the lower and middle Amazon

New York Entomological Society

Basin and along the eastern coast of Brazil (as far south as Rio de Janeiro).
In Costa Rica, the distributional pattern is also molded by the Central
Cordillera and confines the butterfly to all of the Caribbean lowlands and
southwestern Pacific wet lowlands. The butterfly seldom occurs above
400-meters elevation in Costa Rica, although an occasional adult is seen at
elevations as high as 900 meters. The butterfly is also rare in lowland
Guanacaste Province, where there is a strong but variable dry season each year.

Various authors (Emsley, 1963; Barcant, 1970; Brown and Mielke, 1972)
have emphasized that P. dido inhabits forest clearings and edges of rain forests.
The same pattern is seen in Costa Rica, although studies are lacking from
the canopy in the interior of forests. Certainly one larval food plant, P.
vitifolia, is abundant in thinned-out forest at their edges; the distributional
pattern of such plants and the spectrum of Pas si flora species exploitation by
P. dido are the major determinants of habitat selection for oviposition and
larval development.

The literature suggests that there may exist considerable specificity for
certain species of Passiflora in P. dido. Beebe, Crane, and Fleming (1960)
noted that although the usual food plant in Trinidad is P. lauri folia, oviposition
(I am equating oviposition with correct larval food plant) also occasionally
occurs on P. cyanea. Brown and Mielke (1972) report that the preferred food
plant is P. mucronata in extra-Amazonian Brazil and that larvae refuse P.
alata and P. speciosa, even though both of these are very closely related to
P. vitifolia, the food plant in Costa Rica (this paper and Woodruff Benson,
pers. comm.), and also in Colombia and Panama (Brown and Mielke). Of
other Brazilian Passifloraceae, P. dido also refuses P. violacea, P. jileki, and
Tetrastylis ovalis (Brown and Mielke, 1972). Such food plant specialization
in P. dido in Brazil could have resulted from an evolutionary divergence in
food plant exploitation brought about by sympatry with P. wernickei, which
has been observed (Brown and Mielke, 1972) to feed on other species of
Passiflora not used by P. dido. But clearly other heliconiines might also have
exerted some ecological pressure for food plant specialization. However,
this is apparently not the case in Costa Rica where several heliconiines exploit
P. vitifolia at least on the Osa Peninsula. Another important factor to consider
is the relative ease with which some species of Passiflora can be exploited as
larval food plants over others. For example, Heliconius hecale and Agraulis
vanillae are at least two other heliconiines found together on P. vitifolia in
Guanacaste, Costa Rica (Allen M. Young, pers. obs.). And it is known that
H. hecale exploits this species over much of Central America and Colombia and
as far as Ecuador, along with many other heliconiine species (Keith S. Brown,
Jr., pers. comm.). The abundance of this vine and the size of individuals may
provide a nonlimited food source for many heliconiines, especially if many of
these butterflies have low average fecundities per female (e.g., see Labine,

Vol. LXXXII, March, 1974

1968, for Heliconius erato egg production). Furthermore, if the local complex
of heliconiines exploiting one or a few species of Passifloraceae contains some
genera with high dispersal tendencies in the adult stage (see Benson, 1971,
for comments concerning generic patterns of heliconiine dispersal tendencies),
then this would also lessen the local exploitation of single patches of the vines,
a consideration especially important if a given local species of Pas si flora is very
dispersed itself. Although I have not found many other heliconiines on P.
vitifolia at Tirimbina, this is because systematic searches have not yet been
conducted, although I suspect that the herbivore load would be reasonably simi-
lar to that observed by Benson on the Osa Peninsula (to be reported by him in
a forthcoming paper).

The traditional question of palatability that shrouds ecological and evolu-
tionary approaches to the Heliconiinae is interesting to consider for Philaethria
butterflies, owing to the generalized separate lineage from other members of
the subfamily (Emsley, 1963). If it is assumed that the genetic and physi-
ological adaptations for withstanding toxic compounds derived from plant
tissue are a derivative or advanced evolutionary trait in butterflies (Brower
and Brower, 1964), the question then arises whether a generalized genus like
Philaethria has the ability to develop unpalatability. Certainly, the rather
convincing resemblance between this butterfly and the presumably palatable
mimic Victorina ( Metamorpha ) stelenes (Nymphalidae: Nymphalinae) sug-
gests that P. dido is a Batesian model in this interaction (Brower and Brower,
1964), although (1) P. dido is absent in El Salvadore where V. stelenes is very
abundant, and (2) local abundance of V. stelenes exceeds that of P. dido in
young secondary fields in northeastern Costa Rica (Young, 1972). The
ability of several of the more primitive heliconiine genera, such as Agraulis
to feed on P. vitifolia suggests that Philaethria dido may be palatable, if primi-
tive or generalized genera are unable to develop detoxication systems, as sug-
gested recently by Benson (1971). But since Philaethria represents a separate
lineage of heliconiine evolution from these other genera, a physiological di-
vergence might have occurred with respect to detoxication systems: Philaethria
may have evolved them while the clustered lineage of Agraulis, Dione, Dryadula,
and Podotricha (Emsley, 1963) might not have achieved this, allowing toxic
materials to pass out of the gut in fecal pellets. It is clear that the question of
palatability, or lack thereof, is still very open in Philaethria and warrants fur-
ther study.

Beebe, Crane, and Fleming (1960) have gone into considerable detail on the
comparative analysis of life stages among many heliconiines, including P. dido.
But I do wish to point out some variations in coloration seen in the fifth instar
larva and adult from Costa Rica. It is difficult at the present to attach
significance to color differences in the fifth instar between Costa Rica and
Trinidad since presumably the same subspecies {dido) applies to both regions.

New York Entomological Society

But in the adult series from Tirimbina, the marginal row of light green spots
on the dorsal side of each hindwing (Fig. 1) is considerably smaller than in
the adult figured in Beebe, Crane, and Fleming (1960). The difference is
very stable in all reared adults, and it is also consistent with wild-caught
specimens from Tirimbina and nearby Finca La Selva (8 km) that I have
obtained over the past five years. Color differences in immatures are difficult
to evaluate since they may reflect contemporary ecological specializations
and have little or nothing to do with evolutionary history. Brown (1972)
comments that the color pattern of older larvae for Heliconius hermatheria
converges on that of P. dido and he interprets this as ecological specialization.

Borrowing from the recent discussion of Benson (1971), it is interesting to
cast life cycle and natural historical data for P. dido in terms of a pattern of
ecological adaptation. Single oviposition, highly aggressive larvae, and ap-
parent food plant specialization are ethological and ecological mechanisms
that reflect increased dispersal tendencies of the adult population. From
selection pressures favoring a noncohesive adult population, it is also predicted
that the opportunity for communal roosting to evolve in P. dido would also be
very low. In my experience and surely in that of other researchers, P. dido
adults occur at very low densities in tropical forests, even though a given
population might be strongly localized in a region (Brown and Mielke, 1972,
point out the latter). I believe that the natural historical observations dis-
cussed here and in previous papers are consistent with a non-home-ranging and
non-viscous (Benson, 1971) adult population structure for P. dido at edges
and clearings of tropical rain forests in Central and South America. Local-
izations of populations of P. dido are predicted to be determined primarily by
local topographic effects and by a variety of other natural and perhaps man-
made land barriers. If topographic barriers are few in a region or become
modified by man, we might expect zones of overlap where two or more
different forms might co-occur. This would account for the confusion in the
literature concerning sympatric populations of P. dido dido and P. dido
wernickei or other varieties on the Central American mainland, while other
populations contain only one form, as discussed in Emsley (1963). Especially
near breaks in mountains and low hills, we might expect considerable local
variety in color pattern, but along the more distal coastal regions, uniformity
of the sort encountered in Costa Rica at Tirimbina and La Selva would be
expected.

Literature Cited

Alexander, A. J. 1961a. A study of the biology and behavior of the caterpillars, pupae,
and emerging butterflies of the subfamily Heliconiinae in Trinidad, West Indies.
Part I. Some aspects of larval behavior. Zoologica, 46 : 1-24.

Alexander, A. J. 19616. Part II. Molting, and the behavior of pupae and emerging adults.
Zoologica, 46: 105-122.

Vol. LXXXII, March, 1974

Barcant, M. 1970. “Butterflies of Trinidad and Tobago.” London: Collins, 314 pp.

Beebe, W. 1955. Polymorphism in reared broods of Heliconius butterflies from Surinam
and Trinidad. Zoologica, 40: 139-143.

Beebe, W., Crane, J., and Fleming, H. 1960. A comparison of eggs, larvae, and pupae
in fourteen species of heliconiine butterflies from Trinidad, West Indies. Zoologica,
45: 111-154.

Benson, W. W. 1971. Evidence for the evolution of unpalatability through kin selection
in the Heliconiinae (Lepidoptera: Nymphalidae) . Amer. Nat., 105: 213-226.

Borch, H., and Schmid, F. 1973. On Ornithoptera priamus caelestis Rothschild, demo-
phanes Fruhstorfer and boisduvali Montrouzier (Papilionidae) . J. Lep. Soc., 27:
196-205.

Brower, L. P., and Brower, J. V. Z. 1964. Birds, butterflies, and plant poisons: A study
in ecological chemistry. Zoologica, 49 : 137-159.

Brown, K. S., Jr., and Mielke, O. H. H. 1972. The Heliconians of Brazil (Lepidoptera:
Nymphalidae). Part II. Introduction and general comments, with a supplementary
revision of the tribe. Zoologica, 57 : 1-40.

Crane, J. 1957. Imaginal behavior in butterflies of the subfamily Heliconiinae: Changing
social patterns and irrelevant actions. Zoologica, 42: 135-146.

Emsley, M. G. 1963. A morphological study of imagine Heliconiinae (Lep.: Nymphali-
dae), with a consideration of the evolutionary relationships within the group. Zoo-
logica, 48: 85-130.

Emsley, M. G. 1964. The geographical distribution of the color pattern components of
Heliconius erato and Heliconius melpomene, with genetical evidence for a systematical
relationship between the two species. Zoologica, 49: 245-286.

Emsley, M. G. 1965. Speciation in Heliconius (Lep.: Nymphalidae): Morphology

and geographic distribution. Zoologica, 50: 191-254.

Kaye, W. J. 1917. A reply to Dr. Eltringham’s paper on the genus Heliconius. Trans.
Ent. Soc. London, 1916: 149-155.

Labine, P. A. 1968. The population biology of the butterfly Euphydryas editha. VIII.
Oviposition and its relation to patterns of oviposition in other butterflies. Evolution,
22: 799-805.

Muyshondt, A., Young, A. M., and Muyshondt, A., Jr. 1973. The biology of the but-
terfly Dione juno huascama (Nymphalidae: Heliconiinae) in El Salvador. Jour. New
York Entomol. Soc., 81 : 137-151.

Turner, J. R. G. 1971. The genetics of some polymorphic forms of the butterflies Heli-
conius melpomene (Linnaeus) and H. erato (Linnaeus). II. The hybridization of
subspecies of H. melpomene from Surinam and Trinidad. Zoologica, 56: 125-157.

Young, A. M. 1972. Interactions of Philaethria dido (Heliconiinae) and Victorina stelenes
(Nymphalinae) at Stachytarpheta flowers in Costa Rica: Evidence against mimetic
association. Act. Biol. Venez., 8: 1-17.

Young, A. M. 1973. Notes on the life cycle and natural history of Parides areas mylotes
(Papilionidae) in Costa Rican premontane wet forest. Psyche, 80: 1-22.

New York Entomological Society

Ovipositing of Circulifer tenellus Baker
(Homoptera, Cicatlellidae)

Karl Maramorosch

Institute of Microbiology, Rutgers University, New Brunswick, N. J. 08903
Received for Publication November 15, 1973

In the course of experiments with leafhopper vectors of certain plant disease
agents, such as viruses and mollicutelike organisms (Maramorosch, 1969),
an observation was made concerning oviposition by Circulifer tenellus Baker,
the beet leafhopper that transmits the agent of sugar beet curly top disease.
Groups of 10 adult male and female leafhoppers were routinely confined to
sugar beet plants in small cages, fastened to either the upper or the lower leaf
surface by clip cages (Maramorosch, 1951) or by magnetically attached
cages (Kaloostian, 1955). The latter were modified sometimes so as to provide
adequate aeration through a cylinder made of Saran monofilament plastic
screen (Fig. 1). Irrespective of the type of leaf cage used, only the upper
or lower leaf surface was accessible to the feeding insects. This “limited
access” feeding differed from the usual methods in which stock culture or
disease agent-carrying insects are given free access to all aboveground parts of a
test plant.

Frequently during the summer months gravid beet leafhopper females
deposited eggs in leaf tissues while confined to beet plants in small cages.
Surprisingly, eggs were deposited in such a manner that nymphs never hatched
on the side on which the females were confined. Whenever the cages were
attached to the upper surface of leaves (Fig. 1), the eggs were found pro-
truding from the lower surface (Fig. 2). When insects were placed on the
lower leaf surface, their eggs were seen on the upper surface only (Fig. 3).
The number of eggs found on the lower surfaces seemed to exceed the number
deposited on the upper ones, but no statistical analysis was made to ascertain
whether the difference was significant.

In a few instances leaf cages containing gravid females were left attached
for as long as three to five weeks without disturbing the insects. In such
instances nymphs that hatched from deposited eggs began to feed on the side
opposite the caged adults. Some nymphs managed to squeeze through occasional
narrow gaps between the leaf surface and the bottom part of the clip cage
and they would occasionally appear on other parts of a test plant. Once free
to move, such first and second instar nymphs would become potential sources
of greenhouse contamination.

To prevent the escape of progeny nymphs and accidental greenhouse
contamination, exposed leaves were marked by punched holes. After the

New York Entomological Society, LXXXII: 42-44. March, 1974.

Vol. LXXXII, March, 1974

Fig. 1. Two insect cages, magnetically attached to leaves. Upper (left) cage is of cellu-
lose nitrate tubing, with a Saran monofilament screen on top. Lower (right) cage is made
entirely of Saran monofilament screen, with cotton plug on top to insert insects. The bottom
of each cage, resting on the leaf surface, is covered by a 15 dernier nylon screen.

Fig. 2. When caged insects were confined to the upper leaf surface, eggs were protrud-
ing from the lower surface.

Fig. 3. Single leafhopper egg, protruding from upper surface of a leaf; in this instance
a gravid female was confined to the lower surface of the leaf.

New York Entomological Society

removal of insect cages, usually within the first ten days of test feeding,
the marked leaf portion was cut off and destroyed before first instar nymphs
began to hatch. Whenever gravid females had to be confined to plants in
leaf cages for periods exceeding three to four days, the cages were transferred
from one area to another, or from leaf to leaf, and the exposed portion con-
taining deposited eggs was excised and discarded. This procedure did not
prevent successful inoculation of plants with viruses or mollicutelike agents
(Maramorosch et al., 1962) since these disease agents were rapidly transported
through phloem elements to other parts of the plant.

A probable explanation of the observed hatching of nymphs on the leaf
surface opposite that of female confinement was the length of the ovipositor
and the depth of penetration (Muller, 1942). It seems less likely, though
not inconceivable, to assume that the females were making a deliberate attempt
to place their eggs in such a manner as to ensure that their progeny would
not hatch within the limited area of their own “prison confinement.” The
first, purely mechanistic, explanation seems the more plausible.

Forcing oviposition by means of leaf cages within a limited area of a leaf
has also been advantageous for the rapid collection of leafhopper eggs, used
as the source of embryonic material for insect tissue culture (Hirumi and
Maramorosch, 1964).

Literature Cited

Hirumi, H. and Maramorosch, K. 1964. Insect tissue culture: use of blastokinetic stage
of leafhopper embryo. Science, 144: 1465-1467.

Kaloostian, G. H. 1955. A magnetically suspended insect cage. J. Econ. Entomol., 48:
756-757.

Maramorosch, K. 1951. Handy insect- vector cage. Jour. New York Entomol. Soc., 59:
49-50.

Maramorosch, K. (ed.). 1969. “Viruses, Vectors, and Vegetation.” John Wiley-Inter-

science, New York. 666 pp.

Maramorosch, K., Martinez, A. L., and Maisey, S. 1962. Translocation of aster yellows
virus in aster plants. Phytopathology, 52: 20.

Muller, H. J. 1942. Uber Bau und Funktion des Legapparates der Zikaden (Homoptera
Cicadina). Z. Morphologie u. Okoologie der Tiere, 38: 534-629.

Vol. LXXXII, March, 1974

The Spliingidae of Turrialba, Costa Rica

Richard P. Seifert

Department of Ecology and Evolution, State University of New York, Stony Brook,

New York 11790

Received for Publication October 19, 1973

Abstract: A five-month study of the Sphinigidae at Turrialba, Costa Rica, was made.

A total of 565 specimens of 66 species was collected, of which Errinyis ello was the most
common. A monthly record of these data is presented. Selected data from other Costa
Rican sites are included; they bring the total number of species discussed to 80. The
known distribution of ten species is extended.

Statistical data on the times of flights are given for the Turrialba species, particularly
for those that are common. Differences in temporal activity of closely related species
may be the result of competition.

INTRODUCTION

In most regions of the New World Tropics, there is a paucity of knowledge
about insect communities. Even in the families of Lepidoptera where the tax-
onomy is well known, distributional data in general are very limited and collec-
tions covering a considerable time span at a single locality are few. In this
study, I focus on the seasonal abundance of the hawk moths (Sphingidae)
within the vicinity of the Interamerican Institute of Agricultural Sciences near
Turrialba, Costa Rica, during a five-month period from January through
May of 1967. Certain species discussed here were collected only in Costa
Rican localities other than Turrialba. I have included these species to pro-
vide broader knowledge of the Costa Rican fauna. However, this paper gives
a complete record of sphingids from Turrialba only.

Field Site

The Interamerican Institute of Agricultural Sciences (09'45" North Lati-
tude; 83'38" West Longitude; 602 meters elevation) is located 45 km. south-

Acknowledgments : I should like to thank Robert Hunter and Leslie Holdridge for

their help throughout this project, as well as Kenneth Christiansen and O. R. Taylor who
read drafts of this paper and contributed valuable comments. It gives me particular plea-
sure to thank W. L. Staudinger, who provided encouragement and constant help, and
William Sieker, who helped identify many of the difficult specimens and was never too
busy to answer my myriad questions. This research was sponsored by Grinnell College
and by the Associated Colleges of the Midwest. The Tropical Science Center, the Inter-
american Institute of Agricultural Sciences, and the Universidad de Costa Rica allowed
use of their facilities. The typing was done by Florence H. Seifert and Eileen Fischer.

New York Entomological Society, LXXXII: 45-56. March, 1974.

New York Entomological Society

east of San Jose. The average yearly temperature, rainfall, and relative humid-
ity of this area are 22.4°C, 2547.5 mm., and 87.5 percent, respectively. This
area is classified by the Holdridge system (Holdridge, 1964) as Tropical Pre-
montane Wet Forest. An important feature of the study site is the large area of
primary vegetation near the institute. Two rivers, the Rio Turrialba and the
Rio Reventazon, join near the study area, and the land along the edges of
these rivers is largely primary forest. The study area is surrounded on three
sides by these forests; the fourth side faces pasture and coffee fields. The
site is situated in a small valley and probably drew moths from the surround-
ing hills.

Fourteen fluorescent lights were situated approximately three meters off
the ground along the edges of the forest. One additional fluorescent light was
set up on a white reflecting wall looking into the forest at a height of ten
meters. Two black lights were installed at about two meters height facing the
forest. The lights were situated in a manner so that each light could be ob-
served at least once within a one-hour period and the time of collection of
most moths was recorded. The perimeter of this study site was about three
kilometers and collections were generally made from 7:00 p.m. until 2:00 A.M.,
approximately fifteen nights each month.

RESULTS AND DISCUSSION

Annotated List of Species

A total of 80 species of Sphingidae was covered in this report, of which 565
specimens of 66 species were collected in Turrialba. The most abundant spe-
cies in the Turrialba collection was Erinnyis ello, which is the most common
species of Sphingidae in the New World tropics. Amply terus gannascus , Erin-
nyis oenotrus, A grins cingulata, and P achylia resumens were the next most
common species, in order of abundance. Tables 1 through 5 list by subfamily
the Sphingidae collected in Costa Rica. Except where otherwise noted, the
species were collected at Turrialba. Numbers of specimens per month are
given as well as the relative abundances. The terms for abundance are those
used by Fleming in his 1947 paper on the Sphingidae of Rancho Grande. They
are as follows:

Abundant 101 or more specimens

Common 11 to 100 specimens

Occasional 5 to 10 specimens

Rare 2 to 4 specimens

Unique Only 1 specimen

Seventeen species of the subfamily Acheron tiinae were collected through-
out Costa Rica during the study, of which 92 specimens of thirteen species

Agrius cingulata (Fabricius) Common

Neococytius cluentius (Cramer) Collected only at San Jose

Cocytius duponchal (Poey) Common

Cocytius anteus medor (Stoll) Unique

Vol. LXXXII, March, 1974

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Amply terus gannascus (Stoll) Common

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Amply terus donysa (Druce) Collected only at Vara Blanca

Pseudosphinx tetrio (Linnaeus) Unique

Isognathus rimosa inclitus (W. H. Edwards Unique

New York Entomological Society

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Vol. LXXXII, March, 1974

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New York Entomological Society

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Vol. LXXXII, March, 1974

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Hyles lineata (Fabricius) Collected only at Liberia

New York Entomological Society

were found in Turrialba and four species, Amphimoea walkeri , Neococytius
cluentius, Manduca corallina, and M. dilucida were taken at secondary sites
(sites other than Turrialba). The Acherontiinae represented 16 percent of
the total sample from Turrialba. Agrius cingulata and Manduca jlorestan
were the most common species of this subfamily.

Only four species of the subfamily Ambulicinae were collected in Costa
Rica, three of which were found in Turrialba. One species, Amply pterus donysa,
was collected at a secondary site. All of the species of Ambulicinae found at
Turrialba were common, totaling 57 specimens, or nearly 10 percent of the
samples.

The Sesiinae is the largest subfamily of hawk moths in the New World
tropics with about 100 described species. Thirty-five species were taken in
Costa Rica, with twenty-eight being taken at Turrialba. At Turrialba, 294
specimens were collected, representing nearly 54 percent of the total number
of specimens collected. Erinnyis ello was the most common hawk moth at
Turrialba, representing 49 percent of the Sesiinae collected.

Only thirty-nine specimens and eight species of the subfamily Philampeli-
nae were collected in Costa Rica, all of which are represented in Turrialba.

Of the sixteen species of the subfamily Choerocampinae captured in Costa
Rica, fourteen were represented at Turrialba, yielding 83 specimens. Xylo-
phanes pluto and Xylophanes porcus continentalis were the only species that
were common at Turrialba where they represented 41 percent of the Choero-
campinae collected.

RANGE EXTENSIONS

This study extends the known range of ten species of Sphingidae (Draudt,
1931; Rothschild and Jordan, 1903; Mosser, 1939; and Cary, 1951). Of
these species, five were known from Panama and the extension is therefore
not surprising. Manduca hannibal was believed to exist from southern Brazil
to Panama. Manduca pellenia was known from records of its capture in Pan-
ama, Colombia, Venezuela, and Mexico. Pachylia darceta was believed to be
distributed from Panama to Bolivia and Peru. Pachygonia drucei has pre-
viously been recorded only from Panama and Honduras and Nyceryx eximia
was recorded only from Panama. Cautethia spuria was evidently considered
to be endemic to Mexico. Manduca dilucida was recorded from Mexico to
Honduras. Eumorpha phorbus, which was found at Turrialba, was previously
considered to be distributed from Venezuela to northern Brazil and Eumorpha
capronnieri , which was also found in the collection at Turrialba, was recorded
from sites in Venezuela, Surinam, Ecuador, Peru, and the Amazon Valley.
Nyceryx magna, collected at Turrialba, had been previously recorded only in
Peru and Ecuador.

Vol. LXXXII, March, 1974

Analysis of Temporal Activity for the Turrialba Specimens

An estimate of the period of flight activity for each species of moth was
made by recording the time of capture for each individual at Turrialba. Each
light was visited at least once and often twice each hour from 7:00 p.m. to
2:00 a.m. Table 6 gives a summation per half-hour of the number of speci-
mens of twenty-two species collected throughout the study. The final column
in Table 6 sums all individuals collected after 1:30 a.m. Since collecting was
only occasionally continued after 2:00 a.m., further late groupings yield little
information. The table includes all species that I have considered common (and
that, therefore, provide a sufficient amount of data to be examined), plus
four additional species that show significant nonrandom flight times. The flight-
time distribution of each species was tested for randomness of temporal ac-
tivity, using a Kolmogorov-Smirnov one-sample test. The level of significance
of each test is given in the table using standard statistical notation (n.s. = not
significant; * = significant at 0.05 level; ** = significant at 0.01 level). The
remaining forty-four Turrialba species showed no significant departure from
randomness; for this reason and because their sample sizes are small, data
from these species were not included in the table. Three of the common species
in Table 6 each yielded a computed D, the Kolmogorov-Smirnov statistic,
close to the appropriate critical values of D, indicating that nonrandom flight
times probably occur for these species and would be shown if the sample size
were larger. These species were Cautethia spuria , Eumorpha triangulum , and
Xylophanes porcus. The computed D’s for these species are, respectively, .346,
.322, and .353, while the critical values are, respectively, .375, .328, and .361.

The time of activity may be a resource subdividable by the Sphingidae.
Certain species are using highly discrete time periods for activity (presumably
feeding) while others are acting as generalists with respect to the time resource
and are flying randomly throughout 7:00 p.m. to 2:00 a.m. To elucidate this,
Table 7 gives a representation of the times of activities of the eighteen species
of sphingids that I consider common as well as the four additional species
that show significant nonrandom temporal activity.

Using a Kolmogorov-Smirnov two sample test, six pairwise comparisons
were made between common species of the same genus to determine if closely
related species exhibited statistically significant differing times of flight, in-
dicating effects of competition. These six pairs are: Manduca occulta and
M. florestan ; Amply terus gannascus and A. ypsilon ; Erinnyis ello and E.
oenotrus ; P achylia ficus and P. resumens ; Eumorpha anchemolus and E . tri-
angulum ; and Xylophanes porcus and X. ceratomioides. The Erinnyis, Eu-
morpha, and Xylophanes pairs were nonsignificant, indicating that the ran-
dom flight times of one of the species for each pair had a mean value near the
mean of the nonrandom pair member. This does not, of course, invalidate

Table 6. Per half-hour summary of collection of twenty-two species of Sphingidae collected over a five-month period at Turrialba. The
data for each species were subjected to a Kolmogorov-Smirnov one-sample test ; the level of significance for each test is given following
the appropriate species using standard statistical notation.

New York Entomological Society

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Xylophanes pluto,n s'
Xylophanes porcus,n s'
Xylophanes ceratomioides*

Table 7. Temporal activity periods of twenty-two species of Sphingidae collected at Turrialba; the solid lines indicate activity times for
each species. The notation following each species is the same as that in Table 6. The species whose names are followed by n s- are consid-
ered, with the possible exception of Cautethia spuria , Eumorpha iriangulum, and Xylophanes porous , to fly throughout the night.

Vol. LXXXII, March, 1974

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New York Entomological Society

the fact that for these species pairs, competition apparently has resulted in one
generalist and one specialist with respect to the time resource.

The remaining three pairs were significant at the .01 level indicating high-
resource dividing between closely related species. In the Manduca and Eu-
morpha pairs this apparently indicates that the mean time of flight for the
random member is different from that of the nonrandom member. In the case
of the Amply terus pair, both species show nonrandom temporal activity. We
are then faced with the interesting conclusion that each species shows a non-
random and separate activity period, thus indicating high competitive effects.

It should be pointed out that any nonrandom temporal flight activity may
be associated with corresponding synchronous nonrandom flower opening by
food sources. This type of flowering system occurs in the tropics and yields
a system whereby long-distance pollinators, such as euglossine bees (Janzen,
1971) and sphingids (personal observation and Janzen, 1971), promote out-
crossing, thus allowing individuals of plant species to reproduce even at great
distances from their nearest neighbors.

Literature Cited

Cary, M. M. 1951. Distribution of Sphingidae (Lepidoptera: Heterocera) in the An-

tillean-Caribbean region. Trans. Am. Ent. Soc., 77: 63-129.

Draudt, M. 1931. Sphingidae in Sietz, A., Macro-Lepidoptera of the world. Kernen,
Stuttgart, 6: 839-900 and plates 90-98.

Fleming, H. 1947. Sphingidae (moths) of Rancho Grande, North Central Venezuela.
Zoologica, 30: 133-145.

Holdridge, L. R. 1964. Life-zone ecology. Tropical Science Center, San Jose, 124 pp.
Janzen, D. 1971. Euglossine bees as long-distance pollinators of tropical plants. Science,
171: 203-205.

Mosser, O. 1939. Enumeracion de los esfingidos Mexicanos. Anales Escuela National
de Cienc. Biologicas, 1: 407-495.

Rothschild, W., and Jordan, K. 1903. A revision of the Lepidopterous family Sphingi-
dae. Novit. Zool., 9. 972 pp. and 67 plates.

Vol. LXXXII, March, 1974

A New Genus of Pentatominae from South America, Distinguished by
the Position of Its Spiracles (Hemiptera: Pentatomidae)

L. H. Rolston

Department of Entomology, Louisiana State University,

Baton Rouge, Louisiana 70803

Received for Publication November 30, 1973

Abstract: Caonabo, new genus, and C. casicus , new species, are described from Brazil. In
this genus the spiracles are located near the lateral margins of the abdomen, while the paired
trichobothria associated with each spiracle remain in a submarginal position. The spatial
relationship of the spiracles and trichobothria appears unique among genera of Pentatominae.

INTRODUCTION

This new genus is apparently unique among members of Pentatominae in
having the spiracles located near the lateral margin of the abdominal sternites
while the paired trichobothria are caudad and mesad of each spiracle in the
submarginal position usual for the subfamily. A similar spatial relationship
between the trichobothria and spiracles exists in several pentatomoid families
(Acanthosomatidae, Dinidoridae, Tessaratomidae, and Urostylidae) but among
pentatomids only in Phyllocephalinae according to Ruckes (1962).

Caonabo , n. g.

Juga and tylus subequal in length; antennae five-segmented, basal segment not surpassing
apex of head; bucculae moderately developed, weakly toothed at anterior limit, then per-
current, terminating truncately at base of head near distal end of first rostral segment.
Anterolateral margins of pronotum rounded vertically, anterior angles contiguous with
eyes. Frena extending along basal half of scutellum. Costal angles of coria surpassing apex
of scutellum.

Prosternum and metasternum nearly flat; median low carina on mesosternum diminish-
ing posteriorly to obscurity. Metathoracic ostiole auriculate. Femora unarmed, tibiae
weakly sulcate on superior surface, tarsi three-segmented. Spiracles located near lateral
margin on second through fifth visible abdominal sternites, laterad and cephalad of paired
trichobothria, these somewhat diagonally transverse with one trichobothrium entad and
cephalad of other (Fig. 2). Abdomen lacking median tubercle or spine.

Male with one pair of lateroventral thecal appendages and greatly developed median
penal lobes (Figs. 4 to 6).

Type species: Caonabo casicus , new species.

Caonabo casicus, n. sp.

Light brown to castaneous above, generally grading to black on humeri, brownish yellow
beneath; occasionally entirely fuscous or black. Length of body, 7.4 to 10.7 mm.

Head usually slightly longer than wide across eyes, 1.6 to 1.9 mm wide, 1.7 to 2.1 mm
long. Lateral margins weakly concave above antenniferous tubercles, strongly elevated

New York Entomological Society, LXXXII: 57-60. March, 1974.

New York Entomological Society

Fig. 1. Dorsal aspect, female. Fig. 2. Lateral margin of abdominal sternite; spiracle (s) ;
trichobothrium (tr). Fig. 3. Pygophore, dorsal aspect; carina (ca) ; elevated margin of
genital cup (e) ; median process (m) ; paramere (pa); proctiger (pr). Fig. 4. Theca and
related structures, dorsal aspect; conjunctiva (c). Fig. 5. Same, lateral aspect; thecal
process (tp). Fig. 6. Same, ventral aspect; penisfilum (p). Fig. 7. Genital plates; 8th
paratergite (pt 8); 9th paratergite (pt 9). Fig. 8. Spermatheca. Fig. 9. Distal portion
of spermatheca; spermathecal bulb (sb). Figs. 10 to 12. Variations in right paramere.

Dimensional lines equal 0.5 mm.

toward apex. Juga usually a little shorter than tylus, each jugum and tylus separately
rounded at apex. Punctation moderately strong and rather uniformly arranged, entirely
fuscous or with castaneuos to concolorous punctures on and about tylus and vertex. An-
tennae uniformly yellowish brown to pale castaneous ; basal two segments subequal in
length, next two segments each about twice as long as basal segment, distal segment longest;
length of segments, 0.4 to 0.6; 0.5 to 0.6; 0.8 to 1.0; 0.9 to 1.0; 1.2 to 1.4 mm. Apex of
rostrum usually falling between mesocoxae and metacoxae, occasionally shorter, not sur-
passing mesocoxae, or longer, reaching between metacoxae

Humeri strongly produced, subacute to acute, somewhat elevated, directed obliquely for-
ward (Fig. 1). Anterolateral pronotal margins concave from dorsal view, rough but not
crenulate, usually slightly tuberculate at anterior angles; posterolateral margins convex,
rough, usually pale. Disk traversed by strong ruga about midway between base and apex.
Cicatrices indistinct, their posterior margin delineated by a transverse ruga swollen on each

Vol. LXXXII, March, 1974

side of meson into an irregular callous, this often accented posteriorly by a dense patch of
black punctures. Punctation stronger than on head, ranging from dark brown to black,
coalescing into irregular lines on humeri. Width across humeri, 4.9 to 6.0 mm, length at
meson, 1.9 to 2.5 mm.

Scutellum usually a little longer than wide at base, 2.6 to 3.3 mm wide; sides weakly
concave. A small pale callous, indifferently to clearly delineated, present in basal angles.
Disk often rough, sometimes with submarginal impressions converging beyond distal end
of frena. Punctation as strong as on pronotum, many punctures forming short irregular
lines on basal disk.

Coria more finely and regularly punctate than scutellum; posterior margin extending
posterolaterad in slight arc from scutellum; membrane frosty, venation weakly differentiated.
Connexiva moderately exposed, black with border of each segment draped in brownish
yellow; posterior angle of each segment markedly produced, resulting in strongly serrate
connexival margin.

Punctation beneath humeri especially dense, strong, black, continuing as submarginal
band along thorax and along abdomen mesad of trichobothria (Fig. 2) ; punctation in broad
marginal band on abdomen concolorous, shallow, dense, with trichobothria located about
midway in this band; punctation elsewhere on head, thorax, and disk of abdomen mostly
black, moderately dense, variable in size, irregular in distribution. Evaporatorium on each
side rugose, extending from between coxae laterad about halfway from orifice to lateral
margin of metapleuron.

A segment of anterior margin of genital cup elevated on each side of median process;
entad of this segment a carina running along lateral wall; posterior margin of genital cup
produced on each side as short oblique process located caudad of parameres and cephalad
of large impression in broad posterior border of pygophore (Fig. 3). Proctiger longitudinally
impressed, elevated at distal extremity as pale median crest, this semicircular from lateral
view. Distal portion of parameres variable in form (Figs. 10 to 12). A pair of thecal ap-
pendages located ventrolaterally (Figs. 4 to 6). Median penal lobes longer and wider than
theca, without a discernible division. Penisfilum lying on median plane, pigmented basally
except along midline, flagellate and hyaline distally. Conjunctiva with median lobe.

Apical angles of 8th paratergite acute, narrowly rounded (Fig. 7) ; 9th paratergite longi-
tudinally impressed, deeply so toward base; spermathecal bulb ovoid (Figs. 8 and 9).

Types. Holotype, male, labeled Museum Leiden, Nova Friburgo, Estado do Rio, 900 m.
1-1946, Wygodzinsky. Deposited in Rijksmuseum van Natuurlijke Historie, Leiden, Nether-
lands.

Paratypes. 9 males, 5 females. Same data as holotype (2 $ $ , $ Rijksmuseum) ; Vigosa,
Minas Gerais, Brazil, 25-IV-33, E. J. Hambleton ($ U. S. Nat. Mus.) ; (a) Rio Ver-
melho, S. Cath., Brazil, Apr. 1947; ( b ) A. Mailer Coll., Frank Johnson, Donor ($, $
Am. Mus. Nat. Hist.); Cacador, S. Catarina ($ authors coll.); 12.2.73 Parana, Bocaiuva
do Sul ($ Univ. Fed. do Parana); (a) Stieglmayr, Rio Gr. do Sul; ( b ) Brit. Mus. 1955-
16 (3 $ $, 3 $ $ Brit. Mus. Nat. Hist.).

Distribution. Brazil, in states of Rio Grande do Sul, Santa Catarina, Parana, Rio de Janeiro,
and Minas Gerais.

DISCUSSION

Caonabo seems related to a South American species group, currently placed
in the genus Euschistus, whose members have a pair of lateral appendages on
the aedeagus. In at least some species of this group the conjunctiva is eversible

New York Entomological Society

and the appendages clearly pertain to this structure rather than arising on or
near the posterior margin of the theca as elsewhere in this and neighboring
genera. In Caonabo the conjunctiva is not eversible but the appendages arise
within the theca, their origin, whether on the theca or on the conjunctiva, ob-
scure. If these conjunctival and thecal appendages are homologous, Caonabo
may represent a transitory stage from the former to the latter condition.

Acknowledgments: Messr. W. R. Dolling of the British Museum (Natural History),

R. C. Froeschner of the United States National Museum, A. R. Panizzi of the Universidade
Federal do Parana, P. H. van Doesburg of the Rijksmuseum van Natuurlijke Historie, and
P. Wygodzinsky of the American Museum of Natural History loaned the specimens upon
which these descriptions are based.

Literature Cited

Ruckes, H. 1961 (1962). The diagnostic value of trichobothria in pentatomid taxonomy.
Verh. XI Internat. Kongr. Entomol. (Wien), 1 : 35-37.

Vol. LXXXII, March, 1974

Andean Larvae and Chrysalids of Dione juno andicola (Bates) and
Agraulis vanillae lucina Felder & Felder

F. Martin Brown

6715 So. Marksheffel Road, Colorado Springs, Colorado 80909

Received for Publication November 19, 1973

Abstract: The mature larva and pupa of Dione juno andicola Bates are described from
Banos, Ecuador, and compared with those of other subspecies of juno. The differences are
sufficient to cast doubt upon the assignment of andicola to the species juno. The egg and
five larval stages of Agraulis vanillae lucina Felder & Felder are described from Banos,
Ecuador. Although the imago of lucina is quite different from those of other subspecies
of vanillae, the larval stages support assignment of lucina to vanillae.

INTRODUCTION

While I was going through old papers trying to make manageable the accu-
mulations from three offices I found the following notes of observations made
in 1938 in Ecuador. I sent them at that time to the late Dr. John A. Comstock
and do not recall that he ever published them. At this time I have added to
the original manuscript comparisons with the mature larvae and pupae of the
species as observed in other areas.

Dione juno andicola Bates

Two full-grown larvae of this Andean taxon were collected October 9, 1938, on the trail
from Banos, Tungurahua, Ecuador, to Runtun — a high hill just south of the town. They
were making silk patches on the heavy leaves of maguey, which obviously is not their
food plant. One specimen was preserved in alcohol (and dispatched to Comstock) and the
other allowed to pupate and emerge for determination.

Mature larvae. Length 3.0 cm., greatest diameter 4.5 mm. The ground color is dark olive
brown, almost black. This is almost obliterated by a mosaic of dark burnt-orange spots.
The anal plate and head are black. Segment Tl bears two short subdorsal scoli, T2 has
two long lateral scoli and T3 two long subdorsal and two short lateral scoli. Each ab-
dominal segment except the last bears six short scoli. These are arranged in subdorsal,
lateral, and sublateral rows. The lateral pair is missing on the last segment. All of the scoli
and the spines that adorn them are black.

Beebe, Crane, and Fleming (1960, text Fig. 5 A) show a caterpillar of Dione juno juno
(Cramer) from Trinidad, and Muyshondt, Young, and Muyshondt (1973, Fig. 2A) show
D. juno huascama Reakirt caterpillars from Salvador with the scoli on the thorax not promi-
nently different in length from those on the abdominal segments. This suggests to me that
andicola may not be a subspecies of juno but a valid, albeit cryptic, species. Trinidadian
juno (Beebe, Crane, and Fleming, 1960, p. 129) is described as “body velvety dark brown
to almost black with small, paired spots, brownish-yellow to brown-orange. On middle
part of body the arrangement is very regular, in three series. . . . Thus structurally and in
coloring andicola differs from juno juno in the fifth instar.

New York Entomological Society, LXXXII: 61-64. March, 1974.

New York Entomological Society

Salvadorian huascama is colored more like andicola. Muyshondt et al. (1973, p. 141)
states: “In the fifth instar the coloration is distinctive, being a mottled light brown.’’ This
is in accord with andicola so far as coloring is concerned. The scoli on huascama are like
those on juno juno.

Pupa. Length 2.23 cm., greatest depth 0.89 cm., greatest width (at wing flanges) 0.67 cm.
Highly cryptic, marbled black and cream with the black predominant. It hangs pendant
from a tuft of silk. It is strongly keeled with a deep thoracic arch in the dorsum. All the
organs of the head are studded with dull red-brown warts. The inner margins of the fore-
wings form thick, dirty white flanges on the sides. The outer margins of the wings are
decorated with fine black lines forming “Ts” at the ends of the nervules. The first three
abdominal segments are decorated with subdorsal warty ridges and a wart above the
black stigma. The ventrum of these segments is covered by the wing cases. Segments 4-6
bear large, subdorsal, warty prominences at their caudal margins and, centrally, at the an-
terior margin, have smaller, deep cream-colored warts. Segment 7 bears a subdorsal pair
of small, red-brown warts. Segment 8 and the cremaster are covered with small red-brown
warts. On the sides of segments 4-7 the creamy white marbling dominates the coloring.
The ventrum of segments 4-6 is a creamy pink that is reminiscent of a patch of mold.

The specimen pupated during the night of October 11 and emerged at 10:30 a.m. on
October 25.

Beebe et al. (1960, pi. XIV, Fig. 82) and Muyshondt et al. (1973, Fig. 2B) suggest that
the shapes of the pupae of juno, huascama, and andicola are essentially the same. This
probably is a generic feature. Both juno juno and huascama are described as brown or dark
brown in color. Thus andicola1 s strongly mottled coloring is quite different.

Until proven wrong I will consider andicola Bates to be a full species in the genus Dione.

Agraulis vanillae lucina Felder & Felder

This seems to be the most common representative of the family in Banos during October.
The imagoes vary considerably, some of them being very dark on the underside, others much
lighter and in that respect like Agraulis vanillae. For a while I wondered if this was moneta
(Huebner). The larvae, however, are markedly different from those of Dione. Michener
(1942) found what have been accepted to be valid generic differences between moneta and
vanillae. As he pointed out, lucina is quite unlike vanillae in markings and some might hold
it a different species. Oviposition: The only females observed ovipositing were very much
battered. Oviposition took place only in bright sunshine. The eggs were laid singly on vari-
ous parts of a Passiflora vine (species not determined). Some were on the leaves, some on
the stem, others on the tendrils and buds. One female was observed on the first sunny after-
noon after almost a week of rain and dull weather. During a half-hour period she laid
twenty-three eggs on the upper side of a leaf near its tip. This is the only time such an oc-
currence was observed. Unfortunately for me and posterity, a pet parrot discovered the
batch of eggs before I tried to collect them! Egg: Subconical, lemon-yellow in color, 0.9
mm. high and 0.4 mm. in greater diameter. The sides are sculptured with 14 ridges. Between
these ridges the surfaces are pitted with elliptic depressions.

These eggs were somewhat smaller than those of vanillae vanillae (Linnaeus) (Beebe et
al., 1960, p. 117) and with several fewer vertical ridges. The number of ridges and the
coloring probably have little taxonomic value at species level since, within the Heliconiidae,
species are known with highly variable eggs.

First instar. The larva at eclosion is 2.2 mm. long with a head capsule about 0.3 mm.
across. It grew to between 4 and 5 mm. before making its first moult. The head is black

Vol. LXXXII, March, 1974

with scattered long, black spines. The body is dirty olive gray with a little white mottling.
T1 bears 16 black spines of which 10 terminate in little knobs. Such knobbed spines (or
setae) are highly diagnostic for Agraulis. T2 and T3 each bear 14 spines of which 8 are
knobbed. Each of the abdominal segments bears 10 knobbed spines. The anal plate bears
4. The 6 dorsal spines of T1 are set in a black patch.

Second instar. The larvae at this stage are about 7 mm. long and the head capsule is 0.6
mm. across. In general the insect appears as in the first instar with increased white mottling
and each of the spines set in a small, subconical, brown papule, except the dorsal spines of
T1 in the black patch.

Third instar. The larvae of this stage are about 15 mm. long and the head capsule 1.1 mm.
wide. The ground color is purple brown. There are broad dorsal and lateral stripes of
yellow which are finely set with black hairs. Each segment bears three pairs of black scoli.
The head, legs, and anal appendages are black. The head bears in addition to short black
spines two rather large coronal scoli. As the time for the third moult approaches, the yellow
dorsal stripe breaks up in each segment into a “T” with 5 dots, 3 over the crossbar and
one on each side of the stem.

Fourth instar. During this stage the larvae attain 25 mm. length and the head capsule
is 2.0 mm. wide. The markings and decoration are as in the third instar with one great
exception: The dorsal stripe is divided into three parallel stripes broken at the septa.

Fifth instar. The largest specimens of this instar measured 51 mm. long but the average
was about 45 mm. The head capsules were close to 3.5 mm. wide. The ground color is
nearly puce. The dorsal stripe is dull orange yellow to canary yellow and patterned as in
the 4th instar. The lateral stripes are creamy white to pale canary yellow. These are oc-
casionally tinged with purple toward the close of the instar. The dorsal stripes are some-
times edged with gray-white. The coronal scoli are prominent.

Pupa. Length 23 mm., greatest depth 9.6 mm., greatest width 6.4 mm. The pupa is sus-
pended from a pad of silk. The shape of the pupa is not much different from typical
vanillae as described by Beebe et al. (1960, p. 147, Figs. 71, 72). Compared with juno,
the pupa of lucina has the keel and laterally compressed thoracic process but much more
prominent. The color is dull rust brown with a few streaks of darker brown on the wing
cases. The entire surface is finely rugous. The abdominal processes are not as bold as
those on juno nor are they as warty.

There appears to be very liitle difference between the larvae of vanillae vanillae from
Trinidad noted by Beebe et al., and those of the “unvam7Jae”-looking subspecies
lucina. The longitudinal stripes on lucina appear to be broader and more continuous. The
knobbed hairs of the lst-instar larvae are highly characteristic of vanillae. D’Almeida’s
(1922, p. 126) description of the larvae of the Brazilian subspecies maculosa (Stichel) dif-
fers only in minor points of coloration from the Andean subspecies.

Literature Cited

D’Almeida, R. Ferreira. 1922. “Melanges Lepidopterologiques,” R. Friedlander & Sohn,
Berlin.

Beebe, William, Crane, Jocelyn and Fleming, Henry. 1960. A comparison of eggs,
larvae and pupae in fourteen species of Heliconiine butterflies from Trinidad, W.I.
Zoologica, 45: 111-154, xvi plates, 111 figures.

Michener, Charles D. 1942a. A generic revision of the Heliconiinae (Lepidoptera, Nym-
phalidae). American Museum Novitates, No. 1197, 8 pp., 17 figs. 9 October 1942.

New York Entomological Society

. 1942 b. A review of the subspecies of Agraulis vanillae (Linnaeus). Lepidoptera,

Nymphalidae. American Museum Novitates, No. 1215, 5 pp., 31 December 1942.
Muyshondt, Alberto, Young, Allen M., and Muyshondt, Alberto, Jr. 1973. The
biology of the butterfly Dione juno huascama (Nymphalidae: Heliconiinae) in El

Salvador. Jour. New York Entomol. Soc., 81: 137-151, September 1973.

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American Museum of Natural History, New York 10024

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Fordham University, New York 10458

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The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press
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Known office of publication: Waksman Institute of Microbiology, New Brunswick, New Jersey 08903.

A A:

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Journal of the

New York Entomological Society

Volume LXXXII June, 1974 No

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Waksman Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lawrence E. Limpel
Helen McCarthy

Publication Committee

Dr. Kumar Krishna Dr. Ayodha P. Gupta

Dr. James Forbes, Chairman

CONTENTS

The William S. Creighton Memorial Issue James Forbes

William Steel Creighton — An Appreciation Robert E. Gregg

Studies on California Ants. 8. A New Species of Cardiocondyla (Hymenoptera :
Formicidae) Roy R. Snelling

Notes on the Behavior of Three Species of Cardiocondyla in the United
States (Hymenoptera: Formicidae) .... William S. Creighton and Roy R. Snelling

Generic Diversity in Phase of Rhythm in Myrmicine Ants ... Elwood S. McCluskey

Supplementary Studies on Ant Larvae: Simopone and Turneria

George C. Wheeler and Jeanette Wheeler

On the Estimation of Total Behavioral Repertories in Ants

Edward O. Wilson and Robert Fagen

Zoogeography of the Imported Fire Ants William F. Buren,

George E. Allen, Willard H. Whitcomb, Frances E. Lennartz, and Roger N. Williams

Microsporidan and Fungal Diseases of Solenopsis invicta Buren in Brazil

George E. Allen and William F. Buren

A Supplement to the Revision of the Ant Genus Basiceros ('Hymenoptera:
Formicidae) William L. Brown, Jr.

Myrmicine Trail Phermones: Specificity, Source and Significance

Murray S. Blum

. 2

103

106

113

125

131

141

New York Entomological Society

The William S. Creighton Memorial Issue

The sequence of circumstances that led to the publication of this issue of the
Journal demonstrates the desire of friends and associates to recognize the achieve-
ments of William Creighton and to provide for him a fitting memorial. At the
end of last summer the news of Dr. Creighton’s death was received by Dr.
Howard Topoff, President of the New York Entomological Society. A few weeks
later at an Executive Committee meeting of the Society the decision was made
to solicit papers from myrmecologists for a memorial issue of the Journal. Prep-
arations were just begun when a letter was received from Dr. Roy Snelling of Los
Angeles, California, advising the Society that he and Dr. E. O. Wilson were col-
lecting papers and asking if the New York Entomological Society would sponsor
a memorial issue. The two projects were combined.

William Creighton joined the New York Entomological Society shortly after
he came to the Biology Department of the City College of New York in 1931. In
his early membership his main service to the group was as a speaker at meetings.
From time to time he would spend an evening presenting some aspect of his
research. During the 1950’s he became an active worker in the Society. In 1957
he was elected Vice-President. Later he joined the Publication Committee, and
in 1960 he undertook the Editorship of the Journal. He held this post for a year
and a half until his early retirement from C.C.N.Y. It was a change in the retire-
ment regulations of the New York City colleges that gave him the opportunity to
retire early and to begin a period of vigorous research in the taxonomy and the
biology of the ants in the southwestern region of the United States.

The New York Entomological Society is proud to have had William S. Creigh-
ton as a member and as a Society worker and is grateful for the contributions he
has made to the field of myrmecology. To his memory this issue of the Journal
is affectionately dedicated.

James Forbes

New York Entomological Society, LXXXII: 66. June, 1974.

Vol. LXXXII, June, 1974

William Steel Creighton — An Appreciation

Robert E. Gregg

Department of Biology, University of Colorado, Boulder, Colo. 80302
Received for Publication January 7, 1974

The accompanying photograph of Dr. Creighton is an excellent likeness,
and while there seem not to be many such available, this one has been
graciously furnished by his wife, Martha. I am indebted to her for this and
also for certain biographical information without which this account could
not be written.

Dr. Creighton was born April 3, 1902, in Philadelphia, Penn., the son of
John Harvey and Ethel Steel Creighton, and died July 23, 1973, at Alexandria
Bay, New York. He received a bachelor’s degree from Roanoke College, Vir-
ginia, in 1924, his M.S. degree from Princeton in 1926, and the D.Sc. degree
from Harvard in 1930. During his attendance at Princeton he carried out
research on the luminescence mechanism of fireflies, but, in 1926, while
working with Dr. Frank E. Lutz of the American Museum of Natural His-
tory, he became interested in the study of ants. Dr. William Morton Wheeler,
one of the leading authorities on ants, who was at the American Museum,
was transferred to Harvard University, so Creighton moved to Boston in
order to continue his interests and his training under Professor Wheeler. The
two men cemented a lasting friendship and an academic rapport that was
often reflected in Creighton’s sincere and high regard for his mentor by his
references to “Uncle Bill Wheeler.” The combination of their efforts was
to have a very significant effect on the course of myrmecology; it brought
a maturity to the study of ants in North America that was salutary. Never-
theless, Creighton did not hesitate to disagree with Wheeler nor to correct
his own mistakes when scientific accuracy so demanded, and this is quite
evident to those who peruse the now standard volume on North American
Formicidae.

In 1931, Dr. Creighton joined the faculty of the Department of Biology
at the City College of New York and gave continuous and dedicated service
for 31 years, retiring as professor emeritus in February 1962. His associates
have prepared a mimeographed resolution of respect, and I have the per-
mission to quote from this statement, as no more fitting tribute to his capaci-
ties as a teacher could be made.

“His chief devotion . . . was to his teaching and to his students. A grandson of
a preacher, he had a touch of the pulpit in his proselytizing approach to biology. He
was deeply committed to demonstrating to his students the intellectual and emotional
rewards of the pursuit of the science of biology. This commitment was especially evident
by his preference for teaching our freshman general biology courses. These are clearly

New York Entomological Society, LXXXII: 67-75. June, 1974.

New York Entomological Society

the most difficult courses to teach successfully, and here, he felt, was where future
biologists could be made. He believed that, course content and syllabus aside, first and
foremost we must instill in our students a respect and enthusiasm for the subject. The
communication of information comes easily as a consequence. He was a stirring lecturer,
and always managed to project a personal involvement in the subject matter — that
magical feeling of wonder about life, and curiosity about how it works. It was a
dull mind indeed that could not be stimulated with a sense of excitement by one

of his lectures.

In his last several years prior to retirement, Professor Creighton saw major changes
in his field. The advent of molecular and biochemical approaches to biology took many
by surprise. Not so Bill Creighton. He revised and up-dated his teaching materials
constantly, but always with the basic principle of projecting his personal enthusiasm

to the students. In this way, he found that he could demand high standards of per-
formance, and his students gave him their best willingly. His patience with students

and their questions seemed inexhaustible, yet there were times when he erupted easily,
especially when faced with sloth or with bureaucratic red tape. Under such conditions
he spoke his mind clearly and unequivocally.

Bill’s colleagues found in him a staunch friend and a frank critic. He had a delightfully
earthy sense of humor, and had the remarkable facility for separating fraud from truth,
and doing so with devastating clarity.”

Vol. LXXXII, June, 1974

Professor Creighton’s interests and research contributions were in the field
of ant biology, especially the systematics of the Formicidae. In collaboration
with Wheeler he began a study of the taxonomy of ants in North America that
was to summarize the diverse knowledge that had already accumulated, and
to provide in a single publication a guide to the identification and other
aspects of the taxonomy of all our native species. Before this effort was far
along, Professor Wheeler died (1937), and Creighton was left the task of
completing the study. That he did so with outstanding success is now history.
He enjoyed unlimited access to the Wheeler Collection, part of which was
in the American Museum in New York, and part in the Museum of Compara-
tive Zoology at Harvard in Cambridge, both of which contained a large
number of type specimens, and otherwise authentically determined material
identified by Wheeler himself and such renowned authorities in Europe as
Emery and Forel. In addition, he availed himself of the very substantial
holdings in the United States National Museum in Washington, through the
cooperation of Dr. Marion R. Smith, who was then in charge of these speci-
mens. Creighton also built a personal collection which contained numerous
types presented to him by Wheeler, by other ant students, and types resulting
from the descriptions of new species for which Creighton was the original
author. But more than this, Bill Creighton amassed a superb collection based
upon years of field work and direct acquaintance through automobile travel
with virtually all sections of the country. Thus he reinforced his study of
cabinet specimens by coming to know the geographical and ecological condi-
tions in which his species existed. He always insisted that those who did no
field work were under serious handicaps and could not possibly understand
the full significance of the biology of the forms being studied. Although he
did not travel widely over the world, Creighton did acquire experience at
the Soledad field station in Cuba in the earlier years of his life and, in
later years, after the publication of his book, extended his journeys to many
parts of Mexico, particularly the nontropical areas. And, finally, he culti-
vated opportunities for augmenting his collection by exchanging specimens
and making gifts to support the work of colleagues, too numerous to mention
separately.

Museum work and laboratory work were indispensable tools, as they are
for all taxonomists, but Bill Creighton was a first-class naturalist as well,
and his knowledge of distributional ecology, vegetation, and the names of
plants (as well as animals), was truly remarkable. He knew of the manifold
complexities and interrelationships that exist in the natural world and that
this matrix must not be ignored if one hoped to gain a satisfactory compre-
hension of the distribution, and the causes thereof, of the particular group of
organisms in which he was interested. As far as practicable, his collecting
of specimens always entailed the gathering of relevant habit and habitat notes
associated with ants, and he was very cognizant of the role of altitude in

New York Entomological Society

affecting the occurrence of living things. Especially for mountainous dis-
tricts, elevation became a conspicuous inclusion on his labels. Meticulous
attention to essential details was not foreign to him. A glimpse of the in-
tensity and thoroughness of his field work is revealed by his willingness
to lie prone in the dirt in search of minute species, his efforts literally to
leave no stone unturned, his pursuit of the arboreals (a most difficult type
of ant to find and bag) even when it necessitated tearing off rotten branches
and injuring his hands in the bargain. He visited type localities in an effort
to locate additional topotypes, although in the areas he ransacked for speci-
mens he was not always successful. Rare species were a challenge, and he
had the perseverance to track them in likely as well as unlikely places.

The work of revising the taxonomy of the entire North American ant fauna
developed into an enormous undertaking. He soon realized that far-reaching
changes would be necessary to modernize the nomenclature and that the job
would go well beyond what he and Wheeler had originally visualized. Many
species had been described utilizing subtle and confusing differences in color
and inconsequential variations in size. To Creighton, these criteria (par-
ticularly color) as separatory characters were suspect, and he succeeded in
many instances in proving his point by demonstrating that the supposed

differences were inconstant over long series of specimens, or could be found
among a series of individuals obtained from a single nest! The work en-
tailed not only a critical re-examination of specimens representing numerous
taxa but also a painstaking review of the voluminous literature on the subject,
both foreign and domestic, containing original descriptions of ants, as well
as a penetrating analysis of what the authors said or did not say about
the insects they studied. The result of such investigations was extensive

synonymy of many well-known taxa commonly accepted by previous writers.
Although this was unavoidable in the light of new knowledge and although
some cherished names had to be sacrificed on the altar of scientific accuracy,

Creighton strove sincerely to salvage and conserve every ant name possible,

and as he would put it, “refrain from upsetting too many apple carts.” To
serve the concept of stability in nomenclature, he exercised the privilege of
the “first reviser” to sustain a name by not adhering strictly to the law of
priority, where to do so would only weaken the status of the taxon in ques-
tion. However, this action was rare indeed.

Of even greater magnitude was the problem of complicated quadrinomial
nomenclature that had gradually built up in myrmecology over the years and
that reached its culmination in Emery’s treatment of the world ant fauna
in the several fascicules of the Genera Insectorum, the last of which appeared
in 1925. Subsequent to this, new ideas concerning the nature of species
came into clear focus through the work of Ernst Mayr and others who
showed that species should be regarded each as living and evolving popula-
tions of individuals that are for the most part biologically (reproductively)

Vol. LXXXII, June, 1974

isolated from one another. In the course of their evolution, it was thought,
these populations developed through stages of divergence that could be de-
tected and recognized as subspecies, and these subspecific populations could
be designated by formal names added to the basic binomial, thus producing
the widely used trinomial nomenclature. But only this one infraspecific cate-
gory seemed to be objective and defensible, varieties and forms then having
to be discarded. In the midst of this atmosphere Creighton carried out his
revision, and he adopted the practice of employing binomials, as usual, and
the trinomial wherever the facts, particularly distributional data, justified
its use. For it was believed that subspecies arise as geographic or topographic
isolates and that by definition they must replace each other geographically
to be valid. Accordingly, wherever a species appeared to be composed of
two or more structurally different and essentially allopatric populations,
and especially if there could be found any evidence of intergradation among
them in intermediate localities, Creighton treated them as subspecies. Forms
that were unquestionably distinct, with constant nonintergrading traits, how-
ever minute, he regarded as full species. In given cases, forms that were
thought of as separate species were, on review, reduced to subspecies; those
that had been subspecies or even varieties, if they qualified, were necessarily
raised to species rank. The varietal category itself was abandoned. Thus in
one great consistent effort he effectively swept aside the troublesome varietal
rank, recognized the validity of any previous taxon as a full or subspecies
(if the data supported such action), described new species when their popula-
tions conformed to the refined criteria of the biological species concept, and
in the end simplified and enormously improved the taxonomy of American
ants. This achievement was and will remain a milestone in the history of
myrmecology. Since then new forms are not given varietal status, most
being described as species and some as subspecies. Interestingly, the bio-
logical validity of the latter category also has been challenged since Creighton
completed his studies. His work was entitled, “The Ants of North America,”
and was published as Volume 104, 585 pp., 57 plates, 1950, in the Bulletin
of the Museum of Comparative Zoology, at Harvard College.

I came to know Professor Creighton personally in 1950, just after the
publication of his treatise on American ants. There had been a few previous
exchanges of letters, but there was no indication he might be spending a
year at the University of Colorado. The chairman of our department, Dr.
Gordon Alexander, had met Creighton at Princeton, and the Creightons and
Alexanders were close friends. So when a year’s leave of one of our staff
created a temporary vacancy, Bill Creighton was invited to fill it. His
inimitable capacities as a teacher showed themselves again, judging from
the testimony of students, and he seemed to fully enjoy the experience as
well. My opportunities to consult with him at length, for he was always
most generous with his time, were invaluable. Our myrmecological conversa-

New York Entomological Society

tions were almost a daily occurrence, but I must confess the benefits flowed
nearly completely in my direction. For one who was (prior to that time)
necessarily accustomed to identifying ants with limited access to scattered
publications, the help of irregular correspondence, and the aid of occasional
exchanges of specimens, this association was a bonanza. He brought with him
numerous type specimens of North American ants, and we were able to make
many direct comparisons with ants collected in Colorado. All this, together
with the newly published Ant Book which was worked over in minute detail,
gave tremendous impetus to the study of Colorado ants. It may be safely
asserted also that this book opened the way for a much clearer understanding
of our native ants than ever before, as it has stimulated the publication of
many significant investigations by myrmecologists in widely separated por-
tions of the United States and Canada.

Toward the end of the academic year, Dr. Creighton was awarded a Gug-
genheim Fellowship, and with this recognition of his accomplishments, he
was able to turn his attention to another aspect of American ant biology.
He had already traveled extensively in many parts of the United States,
including the Rocky Mountain States and the Pacific Coast areas. He now
looked to Mexico, whose ant populations had been so little studied and so
sparsely collected that it was a country with almost unlimited possibilities.
He penetrated far south in Mexico but found the nontropical territories ex-
ceedingly rich as well as the tropical, and decided to confine the most
concentrated efforts of field work to the former. His letters about the various
localities visited were full of enthusiasm, and frequently contained descriptions
of people, places, or events made vivid by a style distinctively his own.
The jeep the Creightons used was ingeniously fitted out as a traveling labora-
tory, even to the addition of a rigid support for a binocular microscope.
Martha designed a monogram for the vehicle showing the queen of a leaf-
cutting ant! They camped out and traveled Mexico the hard way, and on
a very rugged road high in the Sierra San Pedro Martir of Baja, California,
they nearly came to grief. Today very many localities are easily accessible
by fine paved highways, and the numbers of good motels are increasing steadily.

After Bill’s retirement from City College, the Creightons spent summers
at their modest but charming home on Tar Island in the St. Lawrence near
the exit from Lake Ontario. The house was improved for comfort and a
room added. By his own labor, the boat dock was rebuilt and heavy rocks
were fashioned into a sea wall to avert erosion from annual changes in the
level of the river. Most enjoyable of all was the development of the grounds,
complete with rock garden, and a collection of dwarf evergreens from nu-
merous localities, in which the Creightons took great and justifiable pride.
For their winters they returned to the community of La Feria, Texas, in the
valley of the Rio Grande. Here Bill became interested in studying the be-
havior of ants. Long mild seasons and newly acquired leisure time, as well

Vol. LXXXII, June, 1974

as ideal ant subjects, contributed to his success, and a series of papers flowed
from his pen. During this period also a latent ability showed up for making-
excellent, finely shaded, pen-and-ink drawings of certain ant species. He
did not lose his concern for ant systematics, however, for he continued to
visit Mexico for specimens, and some months before his death had suggested
that I accompany him to one of the localities near the eastern Sierra. Un-
fortunately, this could not be carried out. Studies on Cardiocondyla and on
Colobopsis are scheduled to appear posthumously.

He was a member of a number of scientific societies, Sigma Xi, the New
York Entomological Society, Georgia Entomological Society, and he was
honored by election as a Research Associate to the American Museum of
Natural History. He seemed to have a constitutional aversion to politics in
science, preferring to make his influence felt strictly through the publication
of research papers on the subject to which he was so deeply devoted.* One
could be certain always of his candor and of his absolute sincerity. His advice
was trustworthy wisdom, but with it all he retained an enviable humility.
Whether consciously or not, he was aware that science is greater than men.

To one who has known Bill Creighton as friend and colleague for the
past twenty-three years, his passing is an irreparable loss. He has left an
indelible imprint on my efforts in myrmecology, as well as on the contribu-
tions of many others. In an exchange of correspondence with him, letters
in excess of 225 have been accumulated and preserved. Our collection of ants
has received important and crucial additions, some of them types, thanks
to his generosity and willingness to share in mutually advantageous exchange.

He married Martha Patterson, of Cranford, New Jersey, in 1930, and
although there were no children, he, together with Martha’s constant help
and companionship, has given us a legacy of solid scientific achievement and
unimpeachable integrity. In the words, again, of his associates during his
long tenure at the “College” in the City of New York, he was “a unique combina-
tion of scholar, teacher, and powerful personality. His death leaves a hiatus
in the hearts of his friends that will never be filled.”

Bibliography

Creighton, W. S.

1927. The slave raids of Harpagoxenus americanus. Psyche, 34:11-29.

1928. Notes on three abnormal ants. Psyche, 35:51-55.

1928a. A new species of Thaumatomyrmex from Cuba. Psyche, 35:162-166.

1929. New forms of Odontoponera transversa. Psyche, 36:150-154.

1929a. Further notes on the habits of Harpagoxenus americanus. Psyche, 36:48-50.

1930. A review of the genus Myrmoteras. Jour. New York Entomol. Soc. 38 : 177—191 .
1930a. The New World species of the genus Solenopsis. Proc. Amer. Acad. Arts Sci.

66:39-151.

1932. A new female of Acamatus from Texas. Psyche. 39:73-78.

* See accompanying bibliography.

New York Entomological Society

1933. Cyathomyrmex, a new name for the subgenus Cyathocephalus Emery. Psyche,
40:98-100.

1934. Descriptions of three new North American ants with certain ecological observa-
tions on previously described forms. Psyche, 41:185-200.

1935. Two new species of Formica from western United States. Amer. Mus. Nov.,
No. 773, 8 pp.

1937. Notes on the habits of Strumigenys. Psyche, 44:97-109.

1938. On formicid nomenclature. Jour. New York Entomol. Soc. 46:1-9.

1939. A generic re-allocation for Myrmoteras kuroiwae. Jour. New York Entomol.
Soc. 47:39-40.

1939a. A new subspecies of Crematogaster minutissima with revisionary notes con-
cerning that species. Psyche, 46:137-140.

1940. A revision of the North American variants of the ant Formica ruja. Amer.
Mus. Nov., No. 1055, 10 pp.

1940a. A revision of the forms of Stigmatomma pallipes. Amer. Mus. Nov., No. 1079,

8 pp.

1945. Observations on the subgenus Rhachiocrema with the description of a new
species from Borneo. Psyche, 52:109-118.

1950. The ants of North America. Bull. Mus. Comp. Zool. 104:1-585.

1950a. Polyhomoa Azuma, a synonym of Kyidris Brown. Psyche, 57:93-94.

1951. Studies on Arizona ants 1. The habits of Camponotus ulcerosus Wheeler, and
its identity with C. bruesi Wheeler. Psyche, 58:47-64.

1951a. Studies on Arizona ants 2. New data on the ecology of Aphaenogaster huachucana
and a description of the sexual forms. Psyche, 58:89-99.

1952. Studies on Arizona ants 3. The habits of Pogonomyrmex huachucanus Wheeler,
and a description of the sexual castes. Psyche, 59:71-81.

1952a. Studies on Arizona ants 4. Camponotus ( Colobopsis ) papago , a new species
from southern Arizona. Psyche, 59:148-162.

1952b. Pseudomyrmex apache , a new species from the southwestern United States.
Psyche, 59:131-142.

1953. New data on the habits of the ants of the genus Veromessor. Amer. Mus.
Nov., No. 1612, 18 pp.

1953a. New data on the habits of Camponotus ( Myrmaphaenus ) ulcerosus Wheeler.
Psyche, 60:82-84.

1953b. A new subspecies of Xenomyrmex stolli from northeastern Mexico. Amer. Mus.
Nov., No. 1634, 5 pp.

1953c. The rediscovery of Leptothorax silvestrii (Santschi). Amer. Mus. Nov., No.
1635, 7 pp.

1954. Additional studies on Pseudomyrmex apache. Psyche, 61:9-16.

1955. Observations on Pseudomyrmex elongata Mayr. Jour. New York Entomol.
Soc., 63:17-20.

1955a. Studies on the distribution of the genus Novemessor. Psyche, 62:89-97.

1956. Notes on Myrmecocystus lugubris Wheeler and its synonym, Myrmecocystus
yuma Wheeler. Amer. Mus. Nov., No. 1807, 4 pp.

1956a. Studies on the North American representatives of Ephebomyrmex. Psyche, 63:
54-66.

1957. A study of the genus Xenomyrmex. Amer. Mus. Nov., No. 1843, 14 pp.

1957a. A revisionary study of Pheidole vasliti Pergande. Jour. New York Entomol.

Soc. 65:203-212.

Vol. LXXXII, June, 1974

1963. Further studies on the habits of Cryptocerus texanus Santschi. Psyche, 70:133-
143.

1963a. Further observations on Pseudomyrmex apache. Amer. Mus. Nov., No. 2156,
4 pp.

1964. The habits of Pheidole ( Ceratopheidole ) clydei Gregg. Psyche, 71:169-173.

1965. Studies on southwestern ants belonging to Camponotus , subgenus Myrmo-
brachys. Amer. Mus. Nov., No. 2239, 9 pp.

1965a. The habits and distribution of Macromischa subditiva Wheeler. Psyche, 72:28 2-
286.

1966. The habits of Pheidole ridicula Wheeler, with remarks on the habit patterns
in the genus Pheidole. Psyche, 73:1-7.

1967. Studies on free colonies of Cryptocerus texanus Santschi. Psyche, 74:34-41.

1967a. Living doors. Nat. Hist., 76:71-73.

19676. Ant, In Encyclopedia Brittanica, 2:1-4.

1969. Studies on Camponotus ( Myrmaphaenus ) andrei Forel. Amer. Mus. Nov., No.
2393, 6 pp.

1971. New data on the distribution and habits of Leptothorax ( Nesomyrmex ) wilda.
Jour. Georg. Entomol. Soc., 6:207-210.

Creighton, W. S., and Tulloch, G. S. 1930. Notes on Euponera gilva (Roger).
Psyche, 37: 71-79.

, and Crandall, R. H. 1954. New data on the habits Myrmecocystus melliger

Forel. Biol. Rev., C.C.N.Y. 16:2-6.

, and Gregg, R. E. 1954. Studies on the habits and distribution of Cryptocerus

texanus Santschi. Psyche, 61:41-57.

, and Gregg, R. E. 1955. New and little-known species of Pheidole from the

southwestern United States and northern Mexico. Univ. Colo. Stud., Ser. Biol.,
No. 3, 46 pp.

, and Creighton, M. P. 1959. The habits of Pheidole militicida Wheeler. Psyche,

66:1-12.

, and Nutting, W. L. 1965. The habits and distribution of Cryptocerus rohweri

Wheeler. Psyche, 72:59-64.

, and Snelling, R. R. 1966. The rediscovery of Camponotus ( Myrmaphaenus )

yogi Wheeler. Psyche, 73:187-195.

and . 1974. Notes on the behavior of three species of Cardiocondyla

in the United States. Jour. New York Entomol. Soc., 82:82-92.

Wheeler, W. M., and Creighton, W. S. 1934. A study of the ant genera Novomessor
and Veromessor. Proc. Amer. Acad. Arts Sci., 69:341-387.

New York Entomological Society

Studies on California Ants. 8. A New Species of
Cardiocondyla (Hymenoptera: Formicidae)

Roy R. Snelling

Natural History Museum of Los Angeles County, Los Angeles, California 90007
Received for Publication January 7, 1974

Abstract: A new species of introduced ant, C. ectopia , is described and figured, based

on material from Orange and Los Angeles counties. All three castes are included and
the species is compared to the other four species known to occur in the United States.

Cardiocondyla is an Old World genus of approximately 30 species, about half a dozen
of which are regularly transported by commerce into new areas. Four species have
been introduced into the eastern United States (Smith, 1944) ; all seem to be firmly
established in Florida. No species has previously been reported from California. This
seems surprising since the three most commonly transported species are common in the
Pacific region.

The first California specimens to come to my attention were collected by R. J. Hamton
at his home in Long Beach, Los Angeles Co., in 1967. During the following year, speci-
mens were collected by K. C. Stephens in Downey and Artesia, L. A. Co. Specimens
from Tustin, Orange Co., were collected in 1970 by A Mintzer, and the author found
the species in his yard at Seal Beach, Orange Co., in 1972.

I have been unable to associate this species with any previously described name. In
order to discuss this species in the following paper, I am describing the ant as new.
Hopefully the correct name, if the species is previously described, can be determined
at a later date.

Cardiocondyla ectopia Snelling, n. sp.

DIAGNOSIS

Worker (among species in North America) with shallow but distinctly impressed meta-
notal suture ; antennal scape failing to attain occipital margin by about apical breadth ;
propodeum with a pair of short, triangular denticles ; promesonotum slight shiny, ir-
regularly roughened and with shallow, obscure punctures; petiolar node, from above,
slightly longer than wide ; anterior border of postpetiolar node slightly concave. Female
and male: see DISCUSSION (Figs. 1-5).

WORKER Measurements (Figs. 1, 2). HL 0.55-0.60 (0.60); HW 0.43-0.48 (0.47); SL
0.40-0.44 (0.44) ; WL 0.65-0.71 (0.68) ; PW 0.30-0.33 (0.33) mm.

Head distinctly longer than wide, Cl 76-81 (79), longer than scape, SI 90-97 (92);
in frontal view, sides nearly straight, a little convergent above; occipital margin straight,
corners fully rounded. Median lobe of clypeus high, weakly carinate laterally, apical
margin shallowly concave. Scape short of occipital margin by about its maximum thick-
ness, less than length of second antennomere. Eye large, with 11-14 facets in greatest

Acknowledgments: I wish to thank R. J. Hamton, A. Mintzer, and K. C. Stephens
for the gift of material of C. ectopia. Important sexual material of other species was
loaned by D. R. Smith, United States National Museum. The figures were prepared
by Ruth A. DeNicola.

New York Entomological Society, LXXXII: 76-81. June, 1974.

Vol. LXXXII, June, 1974

Figs. 1-4. Cardiocondyla ectopia. 1. Worker, dorsal view; 2. Same, lateral view
3. Male, dorsal view; 4. Same, lateral view. Figures by Ruth Ann DeNicola.

New York Entomological Society

diameter, removed from mandibular insertion by 0.58-0.80 (0.80) times its greatest
diameter. Mandible quinquedentate.

Thorax slender, PW 0.44-0.49 (0.49) X WL. Pronotum, from above, with rounded
humeri. In profile, metanotal suture broadly, shallowly impressed. Propodeum spinose,
spines stout, about as long as basal width; distance between apices of spines about three
times their length.

Anterior peduncle of petiole slightly longer than height of node; node in profile dis-
tinctly longer than high ; node, from above, a little longer than wide ; peduncle with

anteroventral tooth. Node of postpetiole about twice wider than that of petiole; from
above, 1.24-1.50 times wider than long, lateral margins strongly convex; anterior margin
straight or slightly concave.

Integument. Front of head slightly shiny, finely reticulate and with obscure fine punc-
tures; median line obscure. Supraclypeal area pol'shed, shiny. Median lobe of clypeus
slightly shiny, with several irregular, fine longitudinal rugulae. Sides and venter of
head shiny and sparsely punctate, reticulae faint.

Thoracic dorsum a little shinier than front of head, faintly reticulate and with shallow
punctures. Pronotal side shiny, with sparse, fine punctures. Sides of mesopleura and

propodeum moderately shiny; closely, finely striatopunctate. Petiolar node moderately

shiny, finely sparsely punctate ; anterior peduncle dull, closely punctate. Node of
postpetiole moderately shiny, with sparse, obscure, fine punctures. First gastric tergite
shiny, with sparse, fine piligerous punctures.

Vestiture. Pubescence everywhere fine, appressed, as usual in genus. Clypeal margin with
three long, erect hairs; mandibles with a few long, decumbent hairs; apical gastric seg-
ments with a few long, decumbent hairs.

Color. Head brownish ferruginous, lighter anteriorly; thorax, petiole and postpetiole light
ferruginous to yellowish; gaster blackish; antenna and legs light ferruginous to yellowish.

FEMALE Measurements (Fig. 5). HL 0.59-0.63; HW 0.47-0.50; SL 0.43-0.44; WL
0.84-0.89; PW 0.40-0.42; Wing 2.00-2.07 mm.

Head shape similar to that of worker, Cl 78-82. Scape short of occiput by about its
greatest thickness, less than length of second antennomere ; proportionately shorter than
that of worker, SI 88-91. Eye large, removed from base of mandible by 0.58-0.67 times
its greatest length. Lateral ocelli about V3 smaller than median ocellus, separated by
about five times their diameters. Clypeus and mandible as in worker.

Thorax slender, WL 0.47-0.49 times WL. Pronotal humeri weakly angulate. Basal
face of propodeum about as long as posterior face ; spines triangular, length slightly less
than greatest width, distance between apices about three times their length.

Petiole and postpetiole as in worker.

Integument. Head as in worker. Pronotal humeri, mesoscutum and scutellum slightly
shiny, coarsely reticulopunctate. Side of pronotum, pleurae and side of propodeum
shinier than dorsum, finely longitudinally striatopunctate. Basal face of propodeum
closely, finely punctate, posterior face shiny, finely transversely striate. Petiole and
postpetiole as in worker. First gastric tergite similar to that of worker, but punctures
relatively coarser.

Fig. 5. Cardiocondyla ectopia. Female, dorsal view. Figure by Ruth Ann DeNicola.

Vol. LXXXII, June, 1974

New York Entomological Society

Vestiture. As in worker.

Color. Light brownish ferruginous, thorax, petiole, postpetiole, and appendages lighter;
gaster blackish. Wings whitish, veins and stigma pale yellowish.

MALE Measurements (Figs. 3, 4). HL 0.51-0.54 (0.53); HW 0.48-0.50 (0.48); SL
0.38-0.39 (0.38) ; WL 0.61-0.63 (0.63) ; PW 0.30-0.31 (0.30).

Head slightly longer than broad, Cl 91-93 (91), distinctly longer than scape, SI 78-
79 (79) ; in frontal view, sides of head and occipital margin nearly straight, occipital
corners broadly rounded. Median lobe of clypeus short, weakly carinate at sides, margin
slightly concave. Antenna 12-segmented, apical club one-segmented ; scape short of
occiput by a little less (5:6) than its greatest thickness, about length of second antenno-
mere. Eye small, separated from mandibular insertion by 0.88-1.00 (1.00) times its
greatest diameter. Ocelli absent. Mandible quinquedentate, apical tooth massive, pre-
apical tooth larger than basal teeth.

Thorax slender, PW 0.48-0.50 (0.48) times WL, broadest at humeri. Humeri right-
angular; pronotum and mesonotum abruptly declivitous laterally. Metanotal suture
impressed. Basal face of propodeum distinctly longer than posterior; spines short, tri-
angular, about as long as greatest width, apices separated by slightly more than twice
length.

Node of petiole, from above, a little broader than long; in profile, longer than high;
peduncle with anteroventral tooth. Node of postpetiole about twice wider than that of
petiole, 1.4-1. 5 times wider than long, sides strongly convex from above.

Integument. Head shiny, smooth to slightly roughened between sparse, fine, shallow,
piligerous punctures; clypeus moderately shiny, with obscure median carinula; sides and
venter duller, integument more roughened. Promesonotum shiny, with sparse, fine punc-
tures; side of propodeum smooth and shiny, with sparse fine punctures; pleurae similar,
but weakly striatopunctate on lower half. Basal face of propodeum shiny, with sparse,
fine punctures; side similar, but obscurely striatopunctate below. Nodes of petiole and
postpetiole moderately shiny, with sparse, fine punctures. First tergite smooth and
shiny between scattered fine, piligerous punctures.

Vestiture. As described for worker.

Color. Head, thorax, petiole, and postpetiole pale yellowish; clypeus, mandible, thoracic
sutures, and pleurae more reddish ; gaster light brownish ; vertex with obscure pale
brownish spot; appendages pale reddish yellow.

Holotype worker, allotype male; 17 female, two male and 282 worker paratypes: Seal
Beach, 25', Orange Co., Calif., 17-24 July 1972 (R. R. Snelling, No. 72-9), in Natural
History Museum of Los Angeles County, except one female and two worker paratypes
in each of the following: American Museum of Natural History, Museum of Comparative
Zoology, and United States National Museum.

The specific name is from Greek, ektopios, strange or out of place, alluding to the
alien origin of this species.

DISTRIBUTION

Although certainly of Old World origin, this species is presently known only from
southern California. In addition to material from the type locality, specimens from the
following localities have been studied: Long Beach, Los Angeles Co., various dates (R.
J. Hamton; LACM, RJH) ; Downey, Los Angeles Co., 6 June 1968 (K. C. Stephens;

Vol. LXXXII, June, 1974

LACM) ; Artesia, Los Angeles Co., 22 Aug. 1968 (K. C. Stephens; LACM) ; Tustin,
Orange Co., 6 June 1970 (A. Mintzer; LACM).

DISCUSSION

The worker of C. ectopia cannot be run out in the key by Smith (1944) since it
fails to agree fully with either alternative of the first couplet. In that of Creighton
(1950) it will go to C. emeryi Forel. Workers differ from those of C. emeryi by the
broader head, longer oculomandibular distance, concave anterior clypeal margin, striato-
punctate pleurae, broader propodeal spines, and less compressed petiolar node. From
C. nuda (Mayr), C. ectopia is readily separable by the shorter oculomalar distance,
rounded humeri, striatopunctate pleurae, and impressed metanotal suture. In C. venustula
Wheeler and Mann, the clypeus is more massive, the pleurae punctate only, the pro-
podeal spines are reduced to minute tubercules and the antennal scape fails to reach
the occipital margin by less than its greatest thickness. In C. wroughtoni Forel, the
node of the petiole is broader, the anterior margin of the postpetiole is distinctly con-
cave, the pronotal humeri are subangular and the propodeal spines are longer.

Males in this genus are very poorly known and the few descriptions are meaningless,
especially those of the ergatoid males. These usually have been compared to the workers.
Normal, winged males are produced by C. emeryi. This same species also has modified
ergatoid males in which the antennae are 11-segmented, the mandible is unusually long
and slender, without a dentate cutting margin, the anterior margin of the clypeus is
deeply emarginate and with lateral angulations, and the mesonotum has a transverse
gibbosity. An ergatoid male similar to that of C. ectopia is produced by C. nuda but
the description of that form by Forel (1904) is too general to be useful. No males of
C. venustula or C. wroughtoni have been available, nor have they been described.

The female of C. venustula has the propodeal spines reduced to denticles, the nodes
of petiole and postpetiole are sharply reticulopunctate, and the sides of the thorax
are longitudinally rugulose. Those of C. wroughtoni and C. emeryi also have rather
coarsely and closely punctate petiolar and postpetiolar nodes, the anterior margin of
the postpetiole is concave in dorsal view, the petiolar spines are about thrice longer than
wide and the sides of the pronotum are uniformly contiguously punctate. The female
of C. ectopia is most similar to that of C. nuda , but the sides of the pronotum are
shinier, with irregularly spaced punctures and longitudinal rugulae, rather than uniformly
closely punctate. In C. nuda the piligerous punctures of the first tergite are very fine,
hardly exceeding the diameter of the hairs arising from them. In that species, too, the
oculomandibular distance is about half the maximum eye length, a little longer in C.
ectopia.

Literature Cited

Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool., 104:1-
585.

Forel, A. 1904. Miscellanea Myrmecologiques. Rev. Suisse Zook, 12:1-52.

Smith, M. R. 1944. Ants of the genus Cardiocondyla Emery in the United States.
Proc. Entom. Soc. Wash., 46:30-41.

New York Entomological Society

Notes on the Behavior of Three Species of Cardiocondyla in the
United States ( Hymenoptera : Formicidae)

William S. Creighton1
La Feria, Texas 78559
AND

Roy R. Snelling

Natural History Museum of Los Angeles County, Los Angeles, California 90007
Received for Publication January 7, 1974

Abstract: The behavior, both in the field and in observation nests, of three species of

Cardiocondyla are described: C. emeryi and C. nuda were studied in Texas and C. ectopia
in California. Observations include data on length of immature stages, foraging habits, foods
utilized, reactions with other ant species and mating behavior. The data are summarized
and compared against observations made by E. O. Wilson on C. venustula and C. emeryi.

Although the observations presented in this paper are based on a limited num-
ber of colonies, we believe that they will substantially augment the habit data
now available for three of the five species of Cardiocondyla which occur in the
United States. When M. R. Smith monographed our representatives of Car-
diocondyla in 1944, four species were known to occur in the United States,
restricted to southern Florida. At that time Dr. Smith opined that subsequent
field work would turn up records in other parts of the southern United States.
His view has been amply confirmed. The senior author has been able to study
colonies of C. emeryi Forel and C. nuda (Mayr) at La Feria, Texas, and the
junior author has studied colonies of C. ectopia Snelling at Seal Beach, Cali-
fornia. Moreover, the colonies have been favorably placed for continuous
observation. It is the lack of continuous observation that has limited previous
accounts of the habits of these species. In the many times that C. emeryi has
been taken in the field, few of these encounters have permitted a protracted
study of the colonies. Indeed, in many instances no nest was found and ob-
servations were based on the behavior of strays or foragers at food sources. As
far as we have been able to determine, no one has attempted to study these tiny
ants in observation nests. This is not surprising since they are so small that
they are difficult to confine. They can escape through minute apertures and
frequently do so. We have been able to study all three species in observation
nests. In emeryi the observations extended over a period of a year and a half.
The observations on the free nests covered a considerably longer period.

1 Died 23 July 1973. The sections on C. emeryi and C. nuda were in unfinished manuscript,
completed by the junior author.

New York Entomological Society, LXXXII: 82-92. June, 1974.

Vol. LXXXII, June, 1974

Cardiocondyla emeryi Forel

The senior author took stray workers of emeryi near La Feria, Texas, as early
as 1964, but extensive search failed to reveal the nest. This was particularly
frustrating, since it was clear that the nest must be situated in an area that was
less than twenty-five feet square. Although this nest was never found, its
workers continued to forage for the ensuing three years. At that time the colony
either moved to another area or came to an end. Similar difficulties marked
the second nest of emeryi. In early December of 1970, foragers of emeryi
were found on a concrete sidewalk in front of the senior author’s cottage in
La Feria, Texas. The opportunity for observing these foragers could scarcely
have been more favorable. They were close at hand and as long as they kept
to the sidewalk there was little possibility of losing sight of them. Yet despite
daily inspections it was not until nearly two months later that the nest was
discovered. The entrance was a tiny, circular opening about one millimeter in
diameter in the soil at the edge of the walk. Originally this entrance was com-
pletely concealed by a heavy overgrowth of grass. When the nest was excavated,
three dealated females and thirty-four workers were secured. These were placed
in a small Janet nest for further observation. It later became apparent that
only a part of the colony had been taken, for, after a few days, a new nest
entrance was opened up and foraging began again. This provided a good
opportunity for checking the actions of the captive workers against those of
their free nest mates. These observations were continued over a period of
eighteen months, for the captive colony proved to be hardy and comparatively
easy to handle.

The transfer of the females to the observation nest did not interrupt their
egg laying, and subsequent events showed that this proceeds all year long. The
egg is a stubby oval, approximately 0.3 mm. long and 0.17 mm. wide. When
about to lay an egg, the female turns her abdomen under the thorax until she
can touch the tip of it with her mandibles. As a rule the female grasps the egg
in her mandibles as it emerges but occasionally the emerging egg will be seized
by one of the workers. The eggs, which appear to be unusually sticky, are
collected in packets which are often shifted about by the workers.

No attempt was made to maintain the captive colony at a constant tempera-
ture, although, for the most part, the temperature ranged between 60°-70°F.
The following data are, therefore, mainly useful in showing the relative length
of the various stages. Twelve days after the egg is laid the larva emerges. As
the larva increases in size, its anterior end becomes more and more bent until
finally the mandibles are in close proximity to the swollen ventral body wall.
Twenty-three days after the larva has hatched, the meconium is voided and the
larva transforms to a pupa four days later. Six days after transformation, the
eyes and the mandibular teeth of the pupa begin to show pigmentation, and by
the end of two weeks the pigmentation is general. Transformation to the adult

New York Entomological Society

occurs a day or two later. As the pigmentation is well advanced by this time,
the callow period lasts only a day or two.

The data just presented may be summarized as follows:

Since the observations on which the above figures are based were made during a
period from January 18 to March 15, it is probable that during the warmer
months of the year the rate of development of emeryi is more rapid.

It was soon apparent that the workers in the captive colony would accept
anything edible that was put in the food chamber. Moreover, there was no
indication of preference for a particular kind of food. Honey-dew and nectar
elicited just as vigorous feeding responses as did the tissues and body fluids of
other insects. On the other hand, there were some interesting points in the
mechanics of feeding. It is not to be expected that an ant as small as emeryi
could easily cut to pieces heavily sclerotized insect body wall. But the ants
seemed particularly inept at this sort of activity. They often failed to cut up
the soft body wall of termites, although they would “mine” termites extensively
when the body wall was torn open. They had better success with lightly
sclerotized insects such as mosquitoes, crane flies, and may flies and usually
managed to cut them apart although the process was a slow one. When this
was done, pieces of the dissected victim were sometimes carried into the brood
chamber and fed to the larvae, but most of the time the larvae were fed by
regurgitation. This also seemed to be true of the free colony. During two years
of observation, only a few foragers were seen to carry anything back to the
nest. Two of these foragers were relieved of their burdens which, in each case,
consisted of a much macerated bit of soft insect body wall.

Most of the insects placed in the food chamber of the captive colony were
killed before they were given to the ants, but on three occasions living victims
were offered to them. Prudence demanded that these be small insects which
would not be able to disrupt the colony. The living victims used were collembola
(snow fleas), the small nymphs of the woolly apple aphis, and the first instars
of a pentatomid bug. The reaction of the emeryi workers was essentially the
same for all three. The victims were immobilized either by having the head
crushed or by having the appendages bitten off, whereupon their body fluids
and softer tissues were cleaned out. While the ants attacked their victims
energetically, there was no evidence that any of them were stung. If the captive
colony was not liberally supplied with food, the workers would eat the brood.
When they did so, it was usually the pupae which were eaten.

To summarize the above, it appears that emeryi is omnivorous. It is prob-

Egg to larva
Larva to pupa
Pupa to adult
Egg to adult

12 days
27 days
16 days
55 days

Vol. LXXXII, June, 1974

ably predatory on small, soft-bodied insects and almost certainly scavenges the
remains of larger ones. Solid food is rarely brought back to the nest and food
transfer in the colony mainly involves regurgitated liquid foods.

Although the captive colony kept the brood chamber fairly clean, it was
remarkably slack about the rest of the nest. Dead members of the colony
which had been cut to pieces, together with bits of insects which had served as
food, would be dropped at random in any part of the nest except the brood
chamber. At times so much of this refuse accumulated in the passages that
the workers had difficulty in getting through them. This led to a serious mold
problem; since the refuse was not concentrated in a kitchen midden, mold easily
spread throughout most of the nest. This made frequent cleaning necessary.
It was soon found that the best way to do this was to chill the ants to immobility.
This was done many times without deleterious results, a rather remarkable
response on the part of a species which is regarded as a tropicopolitan tramp.

The foraging activities of the free colony showed a number of puzzling fea-
tures. About the only clear-cut controlling factor was temperature, for foraging
would not occur unless the surface temperature was 70°F (21°C) or higher.
Beyond this there was little to indicate what factors were involved in the
foraging pattern. The foragers emerged singly from the nest entrance at widely
separated intervals. Even under optimum conditions at least fifteen minutes
intervened between the emergence of one worker and that of its follower. As
a result, there was no concentration of foragers around the nest entrance, since
the forager was well away from the nest entrance by the time the next one
emerged from it. A secondary result was that no more than a dozen foragers
(often less) were outside the nest at once.

On leaving the nest the forager might start off in any direction and the course
which it followed was extraordinarily crooked. It was exceptional for a forager to
move for more than three or four centimeters in a straight line. Moreover, they
often doubled back over their previous course. This same random, tortuous
course marked the return to the nest and here there was even stronger evidence
of lack of orientation, for the returning forager would often overshoot the nest
entrance even though it passed close by it. As already noted, few returning
foragers carried solid food to the nest, and it seemed possible that the erratic
return course to the nest might be an indication that no food had been found.
In order to test this, grains of sugar were placed on the sidewalk fifteen centi-
meters from the nest entrance. The foragers carried the sugar grains back to the
nest in their mandibles. But it was exceptional to find that one of these ob-
viously food-burdened foragers returned directly to the nest. Instead they con-
tinued on their erratic courses and were equally inept at finding the nest entrance.
It may be recalled that when Dr. E. O. Wilson (1959) published his observations
on the foraging of C. venustula , he reported that the foragers proceeded from
nest entrance to food source over a straight course. Moreover, they would

New York Entomological Society

stick to this course even when this involved surmounting obstacles which could
have been avoided by slight course deviations. Dr. Wilson believed that this
behavior was due to the fact that the foragers orient themselves by sight. It is
hard to see how such an explanation could apply to the tortuous courses char-
acteristic of emeryi. On the other hand, it is equally hard to suggest what, if
anything, controls the random foraging of emeryi. But it is clear enough that
this sort of foraging gives no chance for recruitment by tandem running. Over
many months of observation, no tandem running was ever observed in emeryi.

The foragers of emeryi often met other species of ants on the sidewalk. Most
of the dozen or so species which foraged on the sidewalk posed no particular
threat to the emeryi foragers, but two of them, Solenopsis geminata (Fabricius)
and Pheidole dentata Mayr, are aggressive and carnivorous, and it seemed
likely that encounters with these two species would be extremely hazardous for
the emeryi workers. It was a surprise, therefore, to discover that in such encoun-
ters the two larger species exhibited a marked avoidance reaction to emeryi.
When such encounters occurred, the emeryi forager stood still while the other
ant scrambled away. This was particularly noticeable in P. dentata minors which
seemed to go into a near panic in the presence of an emeryi forager. Since the
size disparity rules out any possibility that the foragers of geminata and dentata
were trying to avoid an attack by the emeryi worker, it can only be supposed
that despite its small size the worker of emeryi possesses a highly effective
repellent pheromone. This would also explain how emeryi is able to nest in close
proximity to flourishing colonies of 5. geminata and P. dentata.

Cardiocondyla nuda (Mayr)

It now seems clear that the senior author was mistaken in treating Forel’s
variety minutior as a subspecies in 1950. At that time there were few long
nest series of nuda available for study; hence it was not certain how the single
nest series which had yielded workers of the typical nuda and others of the
variety minutior ought to be handled. Subsequent studies have shown that the
above situation is normally encountered in any long nest series of nuda. It fol-
lows that minutior must be treated as a synonym of nuda , as shown by Wilson
and Taylor (1967).

In April 1972, several nests of nuda were found in a brick sidewalk about
seventy-five yards away from the nest of emeryi described earlier in this paper.
These were built in the thin layers of soil which had pressed up between the
bricks. On April 14 one of these colonies was excavated and installed in an
observation nest. The colony consisted of two dealated queens and thirty-eight
workers. Since emeryi had shown itself to be easily adaptable to life in an
artificial nest, no difficulty was anticipated in the observation nest of nuda.
Actually the nuda colony proved to be far more difficult to handle.

At first the nuda colony seemed to be doing well. Both queens laid eggs and

Vol. LXXXII, June, 1974

the workers carried out their usual nest activities. But at the end of two weeks,
the rate of egg laying declined. Ultimately both queens ceased to produce eggs.
By this time some had been in the nest a month, and it might have been
expected that larvae would have been present. However, no egg ever hatched.
About the end of May, both queens were cut to pieces by the workers and the
colony expired.

It seems worth noting that during the brief duration of this nuda colony,
the observation colony of emeryi was bringing much brood to maturity. Since
this seems to indicate that nest conditions were satisfactory, considerable effort
was made to assure that the nuda colony received identical treatment. The
two Janet nests were kept in contact to minimize temperature differences.
Their humidity was, as far as possible, kept at the same level and the same
food was given each. Since the emeryi colony survived until the summer of
1972, at which time the last queen died, the obvious conclusion must be that
nuda requires nest conditions different from those which satisfy emeryi.

Better results were secured from observation on the free colonies of nuda.
The foragers leave the nest singly but as they emerge more frequently than do
those of emeryi there are usually more of them outside the nest. They forage
somewhat more rapidly than do the workers of emeryi and the courses they
follow, while by no means straight, are far less tortuous than those of emeryi.
It is rare for a forager of nuda to double back on its own course. The result
is that their progress between nest entrance and food source is more direct.
They also seem to have less difficulty finding the nest entrance on their return.

Cardiocondyla ectopia Snelling

A single polydomous colony was found in the junior author’s front yard in
mid- July 1972. The subcolonies from northeast to southwest were designated
A, B, C. Subcolony B was situated 2.6 m. SW of A; C was about 3 m. S of B.
The entrances of A and B, both between bricks set into the soil, were fully
exposed and marked by piles of debris. That of C, although similarly marked
by debris, was more difficult to discover because it was partially concealed by a
dense mat of Euphorbia serpens.2

Foraging activities of this species have been observed intermittently for about
one year. Other than modifications of diurnal activity, which seem directly
related to temperature, activity appears to be uniform. During the cooler
months, November to March, there is little surface activity. Daytime tempera-
tures often are not sufficiently high to prompt activity, or the duration of
suitable temperature levels is too short.

During July 1972 the colonies were watched whenever circumstances permitted.
At five-minute intervals during observation periods, two temperature readings

2 Determined by R. Gustafson, Natural History Museum of Los Angeles County.

New York Entomological Society

were made — one of the ambient temperature at a level of six feet, the other of
the surface of the asphalt driveway. At this time of the year, worker ants
emerge from the nest and begin foraging when the ambient temperature is at
19°-20°C; the asphalt surface, in full sun, is at about 24°C. On one date in
July, the ambient reading at 1100 hrs. was 26°C and workers were no longer
moving onto the exposed surfaces. For workers from colonies A and B, access
to food resource areas was across a distance of about 0.3 m. When surface
temperatures reached as high as 42 °C at 1100 hrs., there was no foraging
activity by individuals from these colonies. Workers from C continued to forage,
though at a greatly reduced density, since they had direct access to the patch
of Euphorbia.

On another day, however, by 1100 hrs. the ambient temperature was at
23° C, the surface at 36°C. Because of cloud cover, the surface temperature
had been at that level for nearly an hour. From 1000-1045 hrs., ants were
active on the paved surfaces, but at about 1045 began to abandon these surfaces.
On other days, when surface temperatures never exceeded 35°C, there was a
noticeable lessening of activity after 1100 hrs., so it would appear that time of
day is, at least partially, a controlling factor in the foraging pattern of this
species. However, notes by the junior author record some surface activity as
late as 1925 hrs.

Foragers traveled at least 6 m. from the nest in search of food. The small
size of the ants made observation difficult once the workers reached food source
areas amid the plants in the yard. Attempts to determine foraging distances
were frustrated by two factors: the reluctance of the ants to accept bait of any
sort and the competition of the much larger Argentine ant Iridomyrmex humilis
(Mayr). The latter species quickly discovered and monopolized baits.

Individuals which foraged in the mats of Euphorbia were observed with some
success. Many proceeded directly to the flowers and took nectar. Several
seconds (3-21, average 7.4) were spent at each blossom. After a period of up
to 35 min. the forager returned to the nest with distended gaster. Other workers
wandered about, picking up bits of soil, fragments of plant fiber, and pieces of
dead arthropods. Once an acceptable item was discovered, it was transported
back to the nest. Fragments collected were so small that no attempt was made
at specific identification.

From the onset of foraging until about 1030 hrs., the Cardiocondyla workers
foraged throughout the area, even though I. humilis frequently utilized the same
areas. After about 1030, however, most areas were abandoned by the smaller
species. An exception was the patch of Euphorbia adjacent to nest C. This
resource was worked until about 1300 hrs., after which time only Argentine ants
were to be found on it. Since this pattern was consistent, it seems safe to assume
that the foraging period of the Cardiocondyla regularly ends in early afternoon.
Occasional encounters between the two species were uncommon. As a rule, both

Vol. LXXXII, June, 1974

species retreated from the point of encounter. Often, however, the Cardiocondyla
would continue along its original course or was but slightly diverted. The
Iridomyrmex behaved in a very erratic fashion, and usually left the point of
encounter rapidly, and at a course highly divergent from its original course. Less
often, the Cardiocondyla worker would stop with gaster slightly elevated, head
lifted and directed forward, with spread mandibles.

Workers departing from the nests were observed often. Most proceeded
singly in a very irregular but basically unidirectional mode. There were many
turns and divergences for no apparent reason. Return from the foraging area
did not always reverse the outward course. In fact, seldom was this so, for
often the ant would head back to the nest from a point in the foraging area
fully a meter from where she entered it. The return was frequently much
less direct than the departure, involving more divergences from the straight
line and much back-tracking. Once within 0.2 m. of the nest the ant seemed
more certain of her direction and would head more or less directly toward the
nest entrance. Even so, misses were frequent and some search was necessary.

From the above we may perhaps deduce that orientation is partially solar-
directional and partially a matter of visual recognition within a limited area.
There may also be a distance recognition factor, as suggested by a series of
tests. An individual which discovered a bait placed in the driveway was marked
on the gaster. After feeding, it returned to the nest, 1 m. distant. A period of
almost 15 min. elapsed before the marked ant appeared, heading toward the
bait. The path between the ant and the bait was washed with ethyl alcohol
and the bait displaced 5 cm. to one side. The worker proceeded across the
washed area with only momentary hesitation. She stopped when she reached
a point about 1.5 cm. from where the bait had been and began searching for
several centimeters in all directions. Ultimately the new location was dis-
covered. This experiment was repeated, with similar results, several times with
different individuals. When the bait was displaced as much as 15 cm., it usually
was not relocated for the ants would not search so far from the known locus.

Although most foragers depart singly from the nest, tandem running is fre-
quent in this species. Tandems almost invariably consist of a pair of ants,
rarely three. The pattern is as described by Wilson (1959) for venustula.
By working with marked individuals in an observation colony, it was possible
to discover that the leader of the tandem pair was guiding the follower to a
previously discovered food source and that the follower was recruited. The
follower, in turn, would recruit another individual once she returned to the nest.
When the lead ant arrived at the bait, she would immediately begin feeding. The
follower, after searching for a few seconds, would discover the bait and also begin
to feed.

On 23 July 1972 colony A was excavated and placed in a Janet nest. The
colony consisted of eight dealate females, two alate females, about 75 workers,

New York Entomological Society

and two males. Brood was not counted but estimated at 55 larvae and 15 pupae.
About five hours after removal to the artificial nest, mating between one of
the males and one of the alate females was observed. The female was motionless,
antennae slightly extended, head vertical, mandibles slightly open. The male
mounted the female, parallel with her body, and began to nip at the top of the
head of the female, then rapidly jerked his head up and down along the front
of the female’s, apparently effecting contact with her mouthparts. The jerking
movement of the male was excited and rapid, appearing almost violent, and
lasted about 15 sec. After this, he backed along the dorsum of her body,
curled the gastric tip under that of the female, establishing genital contact.
Genitalic contact lasted about five sec., after which the male returned to the
forward position, cleaned the gastric apex, followed by renewal of the entire
procedure. The entire sequence of activities was repeated three times within a
five-min. period. After the last repetition, the male completed his cleaning
procedure, then remained motionless. At this time the female began to walk
about and the male eventually fell off. At some time during the following day
this female shed her wings and became indistinguishable from the others. The
other female was not observed to mate; on 17 August she was not to be found
among the colony residents.

Santschi (1907) stated that males of nuda “var. mauritanica ” assisted in
moving brood. We have no observations to indicate that the male of ectopia
practices such remarkable behavior.

Males of this species are ergatoid, hence wingless, so mating flights do not
take place. Mating, of necessity, occurs within the nest. But, does the mating
take place between individuals born within the same colony, hence potentially
brother and sister? If so, does the female always, or only occasionally, shed
her wings and remain with her colony? Or does she shed her wings and migrate
with part of the worker force to establish a new colony by budding? Does she
sometimes fly after mating either to (a) found a new colony or (b) become
adopted into a neighboring nest? Or does she fly forth from her parent nest still
virgin, become adopted into a neighboring colony, and mate with a male there?

Unfortunately, no answer as yet can be given to the above questions. We
incline to the last-mentioned alternative, however. Adoption into another colony
seems to be a very simple matter. Workers and females from different nests
have been introduced into the observation colony without evidence of animosity;
the new ants acted as if they were a part of the colony. One alate female in
colony A at the time of its capture was not observed to mate, nor did she shed
her wings. On 17 August she disappeared. No remains were found in the
midden. From this we assume she escaped by flight. It is assumed that she
attempted to locate another colony.

In our view the pre-mating flight-adoption-mating alternative seems most
logical, even though there is no firm evidence for it. A mated female represents a

Vol. LXXXII, June, 1974

considerable reproductive potential, far more than does an unmated female.
Furthermore, mating with a male from a different colony is genetically more
sound.

Females do not emerge en masse for flight. Rather, they emerge singly and
take flight over a period of an hour or more. On 17 July 1972 females flew
from colony C as follows: 0905 — 1 2; 0906 — 1$; 0907 — 12; 0908 — 22 2;
0910—12; 0915—12; 1018—12; 1023—1 2. On 23 July 1972: 1010—1 2;
1012 — 12; 1015 — 12; 1018 — 12; 1035 — 1 2. On 7 August 1972 one female
emerged from C at 0930. On 19 February 1973 a single female flew from C at
1400, air temperature about 23 °C, as was true for all of these (observed range:
22.7°-24.1°C).

DISCUSSION

Wilson (1955) summarized the natural history of venustula as he observed
it in Puerto Rico. He found that the colonies were polydomous and that the
populations were low, probably not in excess of two hundred workers. Nest
entrances were small and surrounded by debris. Foraging occurred mostly
during the middle part of the day. Tandem running was observed and presumed
to be a highly evolved form of recruitment. Tandem running was also noted
to occur in emeryi in Puerto Rico.

Our observations, based on emeryi, nuda , and ectopia , tend to corroborate
Wilson’s conclusions. The siting of nests seems to be very similar for all four
species, and the entrances are concealed by miscellaneous debris. Foraging of
the three species which we studied takes place mostly during the midmorning to
midafternoon period, at temperatures above 19°C, and ceases when the ambient
temperature reaches 26°C.

Colonies of ectopia appear to be polydomous, as in venustula , with the com-
ponents up to six meters apart. Multiple queens seem to be normal in all
species. Males of ectopia are wingless and mating takes place within the nest.
Alate females emerge singly and fly quickly. Mating possibly occurs before
emergence or it may be that the females seek adoption in a neighboring nest and
mate there.

Curiously, although Wilson reported tandem running of emeryi, the senior
author found no examples of such behavior in this species at La Feria, Texas.
The foraging pattern of this ant is highly erratic and it may be that tandem
recruitment is not common in this species. This recruitment technique was
not observed in nuda. The two species should be more thoroughly studied.

Better results were obtained for ectopia. Departing workers pursue an erratic
pattern when searching for food; but once a large source, requiring more ants,
is located, the foragers return more or less directly to the nest. Here, another
worker is recruited and led back to the food source, again more or less directly.
Additional ants are recruited in the same manner, if necessary.

New York Entomological Society

Wilson described his experiments with tandem pairs of venustula. Similar
experiments with ectopia produced similar results. From these data it seems
clear that chemical trails are not laid down and orientation to a food source
is probably based on solar position and distance. Since there is no chemical
trail to indicate direction, it follows that the discoverer must lead its nestmates
to the food source. The leader of the tandem pair evidently releases an excitant
pheromone by which the follower is led. It seems possible that this chemical is
so volatile it functions for only a short distance and that only a single individual
can follow it. Hence tandems consist only of a single pair.

The ants are small and can carry only small arthropods or fragments of larger
ones. Relatively large items, such as can be exploited only by large numbers,
probably are seldom available. These are likely effectively taken over by larger
ants ( Pheidole , Solenopsis, Iridomyrmex, etc.) or by those which recruit in
large numbers ( Monomorium , Wasmannia, Iridomyrmex, etc.). While individ-
uals of Cardiocondyla can apparently repel individuals of Iridomyrmex, it is
unlikely that the colonies can effectively compete against such numerically
superior species at more bountiful food sources. Further studies of these and
other Cardiocondyla should investigate food resource utilization as compared
to other ants in the same foraging area.

Literature Cited

Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. Harvard,
104: 1-585.

Santschi, F. 1907. Fourmis de Tunisie capturees en 1906. Rev. Suisse Zool., 15: 305-334.
Smith, M. R. 1944. Ants of the genus Cardiocondyla Emery in the United States. Proc.
Entom. Soc. Wash., 46: 30-41.

Snelling, R. R. 1974. Studies on California ants. 8. A new species of Cardiocondyla.
Jour. New York Entomol. Soc., 82: 76-81.

Wilson, E. O. 1959. Communication by tandem running in the ant genus Cardiocondyla.
Psyche, 66: 29-34.

Wilson, E. O. and Taylor, R. W. 1967. The ants of Polynesia. Pac. Insects Monog., 14:
1-109.

Vol. LXXXII, June, 1974

Generic Diversity in Phase of Rhythm in Myrmicine Ants

Elwood S. McCluskey

Departments of Physiology and of Biology, Loma Linda University,

Loma Linda, California 92354

Received for Publication February 11, 1974

Abstract: Few comparative studies in functional biology have been made at the genus

level. In the tribe Myrmicini, the latest morning hour the workers are aboveground was
compared for 58 species in 9 genera and the hour of mating flight for 40 species in 13 genera.
In each case there is more difference in phase of rhythm among than within genera. When
the possible influence of season, altitude, latitude, average rainfall, and average tempera-
ture is statistically removed by analysis of covariance, the generic diversity remains sig-
nificant. This evidence suggests a taxonomic explanation of the diversity (as opposed to a
strictly ecological or geographical explanation).

INTRODUCTION

Relatively few comparative studies in functional biology have been made at
the genus level. In the ant tribe Formicini time of day both of mating flight and
of worker foraging is much more alike from species to species within a genus
than among genera (McCluskey, 1973). This correlation of behavior with
taxonomic grouping suggested the value of studying more groups of ants. The
present paper considers the tribe Myrmicini (broad sense) in another subfamily
(Myrmicinae) . It is again based on literature records.

DESCRIPTIVE COMPARISONS

Mating Flight

Figure 1 presents the midpoint flight hour for each species of each genus
where records are available for at least two species. It can be seen first that the
generic means range from 0700 to 1800 and second that the species flight hours
tend to be similar within each genus. This generic diversity was tested by the
circular distribution method of Watson and Williams (1956; cf. Batschelet

Acknowledgments: Field studies of rhythms in this group were made by Dr. Creighton,

most notably his paper of 1953. He gave me personal encouragement when I visited his
home in 1956 at an early stage in my research. I thank P. Yahiku for help with statistical
analysis; L. Brand, B. Buttler, I. Fraser, and B. Neufeld for reading the manuscript; and
R. Snelling for determining Pheidole pilifera pacifica and Solenopsis xyloni. Part of my
inspiration for developing explanatory comparisons was the paper by Waterman (1961).
Computer time for the project was supported in part by NIH, Division of Research Resources,
Grant RR-00276. For the analysis of covariance, UCLA Health Sciences Computing Facility
made its BMDX General Linear Hypothesis program available; this was run at the Loma
Linda University Data Processing Facility. G. McCluskey aided with programing.

New York Entomological Society, LXXXII: 93-102. June, 1974.

New York Entomological Society

A. M.

hours

0 2 4 6 8

10 12

Genus

Veromessor f *f

Messor

(F) . F F

PnnoNQHYRMry

F F

F F . F

* = MEAN
HOUR

SOLENOPSIS

F F F * F F F

NYRMEA

F F F * F

Aphaenqgaster

F *

F (F)

Pheidole (f)

F F * F

Fig. 1. Flight hours (all recognizable Daylight Time records were converted to Standard
Time). Each F represents for one species the midpoint between earliest and latest literature
records of flight; () indicate the most fragmentary records. Each asterisk shows the mean
of the species midpoints for a particular genus. Following are the species and literature
sources represented, including single -species records for 6 genera not plotted on the graph.
Where personal communication (person, com.) is the source, the hour precedes name.
ATOPOMYRMEX : mocquerysi (Wheeler 1922). APHAENOG ASTER: megommatus

(Smith 1963), pythia (Saunders 1969), treatae (Talbot 1966). CAREBARA : junodi

(Wheeler 1922). CAREBARELLA : bicolor (Kempf 1969). LEPTOTHORAX : mon-

jauzei (Cagniant 1968). MESSOR: capitatus and structor (Delage 1968, Meyer 1927),

semirufus concolor (Mursaloglu 1957). MYRMICA: laevinodis (Donisthorpe 192 7),

lobicornis fracticornis (Kannowski 1959), ruginodis (Beare 1913; Brian & Brian 1955;
Donisthorpe 1927), sabuleti americana (Kannowski & Kannowski 1957), schenki emeryana
(Medler 1958; Talbot 1945, 1965). PHE1DOLE : bicarinata (1600, W. L. Brown pers.

com.), creightoni (Gregg 1955), megacephala (Illingworth 1933, 1935; Williams 1935), nari
and sp. #10591 (Kusnezov 1962), sitarches (Wilson 1957), pilifera pacifica (1530, E. A.
McCluskey pers. com.). POGONOMYRMEX : badius (Van Pelt 1953), barbatus (Wheeler
1910, 1917), calif ornicus (Michener 1942), imberbiculus (Wheeler 1917), maricopa (Cole
1968), occidentalis (Nagel & Rettenmeyer 1973), rugosus (1515, F. Taylor pers. com.).
SOLENOPSIS: angulatus and sp. # 10576 and sp. #10577 (Kusnezov 1962), invicta (Markin
et al. 1971), molesta (Mallis 1941; Talbot 1966; Wilson & Hunt 1966), richteri or saevissima ?
(Kusnezov 1962; Rhoades & Davis 1967), xyloni (1815, McCluskey unpublished; Wheeler &
Wheeler 1973). STENAMMA: brevicorne (Kannowski 1958). TETRAMORIUM : caespitum
(0700, G. C. & J. Wheeler pers. com.). VEROMESSOR : andrei (McCluskey 1963), per-
gandei (0730, McCluskey unpublished). An annotated table giving the details of support for
Figs. 1 and 2 is available from the author.

Vol. LXXXII, June, 1974

1965, but only for a two-sample case) : Fq-i,N-q — [ (N - q) (Ri - R) ]/[ (q - 1)

( N - Rl) ] =2.80 and P < .02. (N = 40 species, q— 13 genera, R * refers to the
combination vector for all the species in each genus, and R refers to the com-
bination vector of all genera.)

Worker Surface Activity

Whereas mating flights usually occur at a particular season for a given
species, the workers come out of the nest over several seasons. In order to
compare the various species most directly, I attempted to use only summer records
from clear days. Since many of the literature records are incidental or otherwise
fragmentary, the single item of information most useful for comparison was
the approximate lateness of the hour the workers stay out of the nest in the
morning.

In Fig. 2 a nocturnal species is indicated by an X representing out until
“Dawn”; a species which stays out until the sun hits the nest is placed under
“Sunshine”; etc. An intermediate time is indicated by an X between two
adjacent columns. Every genus was included where records are available for at
least three species.

The mean generic hour is seen to range from soon after the sun hits the nest
(Aphaenogaster) to late morning (Monomorium) ; and there is a noticeable
grouping of species. Arbitrarily scoring “Dawn” as 5 am, “Sunshine” as 7 am,
“Midmorn” as 10 am, and “Midday” as 1 pm, Watson and Williams’ test in-
dicates significant diversity (P < .001).

EXPLANATORY COMPARISONS

How might this generic diversity in phase of rhythm be explained? The data
are too limited to answer an ultimate question such as whether the time relations
are adaptive. But it is possible to ask preliminary questions. Is there a
relationship at the genus level between phase and such gross measures of en-
vironment as altitude, latitude, average temperature, average rainfall, or season?
(Current examples of studies at the species and microhabitat level indicating
sensitivity to the environment include Bernstein, 1971; Levins et al., 1973; and
Whitford, 1973).

For each observation locality cited I estimated the altitude (range, 0-2300
m), latitude (3°-55°), average temperature (10°-32°C) and total rainfall
(2-75 cm) for the appropriate season (using mainly Hammond’s Comparative
World Atlas, 1963, and Nystrom’s World Rainfall maps).

Analysis of covariance permitted consideration of the regression of the de-
pendent variable on a number of independent variables (co variates) simul-
taneously. For the workers the latest-hour-out was used as the dependent
variable, and altitude and latitude, or temperature and rain, as covariates. The
generic diversity again appeared highly significant, even though possible altitude,

New York Entomological Society

Genus

DAWN

SUNSHINE

MIDMORN

MIDDAY

Aphaenogaster

(X)

(x)

Veromessor

(x) (x)

Pheidole

(X)

X *

Manica

* X

liESSOR

* = MEAN HOUR

P0G0N0MYRMEX

(x)

Myrmica

X *

Monomorium

Fig. 2. Worker surface activity. Each X represents one species and shows its nearest
approach to midday. See text and also legend for Fig. 1. The following are represented:
APHAENOG ASTER: ashmeadi & floridana (Van Pelt 1958; Whitcomb et al. 1972), fulva
(Park et al. 1931), longiceps (Brown 1955), megommatus (Cole 1966), pallida (Bernard
1968), rudis picea and tennesseensis (Park & Strohecker 1936), splendida (Tohme 1969),
treatae (Talbot 1953, 1966). MANIC A: bradleyi, hunteri, and mutica (Wheeler & Wheeler

1970) , rubida (Reichle 1943). MESSOR : aegyptiacus (Delye 1968; Sheata & Kaschef

1971) , alexandri and orientalis (Tohme 1969), arenarius (Delye 1968, 1971), barbarus
(Buxton 1924; Forel 1928; Pickles 1944; Wheeler & Creighton 1934), capitatus (Delage
1968), caviceps (Delye 1964, 1968, 1969), semirufus (Bodenheimer & Klein 1930; Mursaloglu
1957). MONOMORIUM : chobauti (Delye 1968, niloticoides and venustum (Tohme 1969),
salomonis (Delye 1968; Kemp 1952). MYRMICA: lobicornis fracticornis (Dondale et al.

1972) , rubra (Reichle 1943), ruginodis and scabrinodis (Brian 1955), sabuleti americana

Vol. LXXXII, June, 1974

Table 1. Analyses of Covariance. The contribution of rainfall and temperature might
heavily overlap that of altitude and latitude as variables; therefore a second analysis, shown
in (), was done, with rain and temperature replacing altitude and latitude as covariates

Source of Variation

Workers (latest hour out) (58 species, 9 genera)

Genera

12.7 (12.7)

5.5 (5.4)

<.001

(<•001)

Covariates

3.7 ( 3.7)

1.6 (1.6)

(ns)

Altitude

(or rain)

2.1 ( 5.4)

.9 (2.3)

(ns)

Latitude

(or temp.)

5.9 ( 1.4)

2.5 ( .6)

(ns)

Error

2.3 ( 2.3)

Flights (deviation from midday) (40 species, 13 genera)

(With

season

as months’ deviation

from August 1)

Genera

8.7 (8.2)

2.5 (2.3)

<.05

«.05)

Covariates

5.4 (4.1)

1.6 (1.1)

(ns)

Altitude

(or rain)

.7 (2.9)

.2 ( .8)

(ns)

Latitude

(or temp.)

9.2 (2.9)

2.7 ( .8)

(ns)

Season

.0 (8.2)

.0 (2.3)

(ns)

Error

3.4 (3.6)

(With

season

as months’ deviation

from June 21)

Genera

8.5 (7.4)

2.6 (2.1)

<.05

«.io)

Covariates

7.0 (4.2)

2.2 (1.2)

(ns)

Altitude

(or rain)

.4 (2.2)

.1 ( .6)

(ns)

Latitude

(or temp.)

11.8 (1.6)

3.7 ( .5)

(ns)

Season

4.6 (8.5)

1.4 (2.4)

(ns)

Error

3.2 (3.6)

latitude, average temperature and rainfall effects had been statistically removed
by the analysis of covariance (see Table 1). The same was true if the possible
complicating variables of the tropics were eliminated by performing the analysis
for only the temperate zone species (which were the majority).

(Dondale et al. 1972; Talbot 1946, 1953), schenki emeryana (Talbot 1965). NOVOMESSOR :
albisetosus (Wheeler & Creighton 1934), cockerelli (Wheeler & Creighton 1934; Whitford
1973), manni (Kannowski 1954). PHEIDOLE : megacephala (Carnegie 1960; Greenslade
1972; Levins et al. 1973; Steyn 1954), morrisi (Van Pelt 1958), ridicula (Creighton 1966),
saxicola (Wheeler 1922), sculpturata and/or crassinoda and sp. A and sp. Q (Kemp 1952),
xerophila (until 0630, R. Bernstein pers. com.). POGONOMYRMEX : badius (Carlson &

Gentry 1973; Golley & Gentry 1964; Van Pelt 1953, 1966), barbatus (Box 1960; McCook
1879), californicus (Cole 1932; Michener 1942; Whitford 1973), desertorum (Whitford
1973), magnacanthus (Cole 1968), maricopa (La Rivers 1968), occidentalis (Headlee &
Dean 1908; McCook 1882; Stevens 1965; Wheeler & Wheeler 1963), owyheei (Cole 1934b;
Willard & Crowell 1965), rugosus (until 0745, R. Bernstein pers. com.; Whitford 1973).
VEROMESSOR: andrei (Creighton 1953; McCluskey 1963), juliana (Creighton 1953),

lariversi (Cole 1963, 1966), lobognathus (Cole 1963; Wheeler & Wheeler 1959, 1963),
pergandei (Cole, 1934a, 1963; Creighton 1953; Tevis 1958; Went et al. 1972; Wheeler &
Creighton 1934), smithi (Cole 1963, 1966).

New York Entomological Society

A similar search was made for an explanation of the diversity in flight timing.
Flights, unless near midday, are generally either morning or afternoon, rather
than bimodal like worker activity and like the environment. To relate both
morning and afternoon flight times similarly to the middle of the “environ-
mental” day, I used the difference between the hour of midday and the hour
of either morning or afternoon flight as the dependent variable for regression
studies. “Midday” was arbitrarily defined as 1300, since that is closer than
noon to the hottest time of day according to the thermometer. Also 1345 divides
the day into two equal halves with respect to the number of species flying.
Further, 1307 is the average center of the midday hours avoided by workers;

I calculated this from the 12 species (4 genera) where the records cited (legend
of Fig. 2) are complete enough to show the worker bimodal activity pattern.

Season was included as an additional covariate, because flight records were
used (Fig. 1) no matter what the season, rather than just summer records as
for the workers. Season was measured two ways: as the difference between the
date of the cited observation and either August 1 (to represent the average
“heat center” of the summer) or June 21 (with the longest dawn- or dusk-to-
midday interval). (Southern Hemisphere records were converted by 6 mo.)

Taking distance of flight hour from midday as the dependent variable, analy-
sis of covariance shows the generic difference in timing to remain significant
after removal of the effects of the covariates (Table 1). This is the more
noteworthy because the morning-vs. -afternoon difference between genera is
ignored in the choice of the dependent variable as simply the time from midday.

CONCLUSION

The records displayed here indicate generic diversity both of worker phase
and of flight phase of rhythm beyond the effects of altitude, latitude, average
temperature, average rainfall, and season. This is not to suggest that no rela-
tion with such variables would be found locally (e.g., temperate latitudes only),
or in a microclimatic study, or at the species level. Nevertheless, at the genus
level the evidence suggests a taxonomic explanation of the diversity (as opposed
to a strictly ecological or geographical explanation).

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Vol. LXXXII, June, 1974

103

Supplementary Studies on Ant Larvae: Simopone and Turneria 1

George C. Wheeler and Jeanette Wheeler

Laboratory of Desert Biology, Desert Research Institute,

University of Nevada System, Reno> 89507

Received for Publication January 7, 1974

Abstract: This study supplements our “Ant Larvae: Review and Synthesis” (1974). The

larvae of Simopone n. sp. and Turneria sp. (near dahli ) are described and figured and each
genus is characterized. Simopone is definitely cerapachyine but quite distinct from the
larvae of other known genera of the subfamily. Turneria is typically dolichoderine but
readily distinguished from other genera of the subfamily by the tail and the shape and
location of the dorsal bosses.

SUBFAMILY CERAPACHYINAE

The larva of Simopone is definitely cerapachyine. Its profile is myrmecioid like that of the
other four known genera (Cerapachys, Eusphinctus, Lioponera , and Phyracaces) . The
mandible is sui generis and we must establish for it a new monotypic rubric usimoponoidT
The small size of the mouth parts is also distinctive for the genus. In our general key for ant
larvae Simopone runs to 49 b in company with Cerapachys, Eusphinctus , and Phyracaces ;
from these it can be distinguished by the shape of the mandibles. Its index of specialization
(see our 1974) is 24; that of the subfamily is 22. [The most specialized ant larvae —
Leptanillinae — have an index of 35, while the Ponerinae are less specialized with 17.
The index for the family as a whole is 22.]

Genus SIMOPONE Forel

Body myrmecioid; head on anterior end; anus ventral. Body hairs sparse and minute.
Head suboctagonal ; antennae large; mouth parts small. Mandibles with about 8 teeth on
medial border.

Simopone n. sp. (Fig. 2). Length (through spiracles) about 3 mm. Shape myrmecioid
(i.e., elongate and rather slender; curved ventrally; without a differentiated neck; diameter
decreasing only slightly from AV to anterior end) ; leg vestiges present as small papillae ;
anus ventral. Head on anterior end. Segmentation indistinct. Spiracles small; ten pairs.
Entire integument spinulose, the spinules minute and in arcuate rows, rows forming a
reticulate pattern on venter of T1 ; isolated and coarse, or minute and in short rows, else-
where. Body hairs sparse and minute (0.013-0.025 mm. long) ; unbranched, smooth and
slightly curved, most numerous on AX. Cranium suboctagonal; occipital border sinuate;
mouth parts small. Antennae rather large, slightly raised ellipsoids with 3 sensilla, each
of which bears a spinule. Head hairs few, minute (about 0.004 mm. long), unbranched,
smooth, and slightly curved. Labrum bilobed, about 3 times as wide as long; each lobe with
4 minute sensilla on each ventrolateral surface ; posterior surface of each lobe with about 7
sensilla near the middle in a longitudinal row. Mandibles small; subtriangular ; without
a blade; with the apex slightly curved medially and with about 8 minute to large teeth
on distal % of convex medial border. Maxillae apparently adnate ; palp a slightly raised

1 Hymenoptera : Formicidae .

New York Entomological Society, LXXXII: 103-105. June, 1974.

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Text figure 1. Turneria sp. (near dahli) . la. Head in anterior view, X88; b. Left
mandible in anterior view, X397; 2c. Body hair, X 2 12 ; d. Body in side view, X28. Text figure
2. Simopone n. sp. 2a. Head in anterior view, X101; b. Left mandible in anterior view,
X314; 1 c. Body hair, X667; d. Larva in side view, X28.

cluster of 5 sensilla; galea represented by 2 sensilla with a spinule each. Labium with a few
short transverse rows of minute spinules on the anterior surface ; palp represented by a
cluster of 5 sensilla; an isolated sensillum between each palp and the opening of the
sericteries; the latter a short transverse slit.

Material Studied: 4 larvae from Ghana: New Tafo (Akim), 29 XI 1970, B. Bolton;
courtesy of Dr. W. L. Brown.

SUBFAMILY DOLICHODERINAE

The larva of Turneria is so typically dolichoderine that it does not disturb in the least the
nearly perfect homogeneity of the subfamily. Nevertheless it is distinct from all other
dolichoderine genera. In our key to all ant larvae (1794) it would run to “5 la. Boss or
bosses dorsal .... Forelius, Froggattella and Iridomyrmex It can be distinguished from
those three genera by its tail and the shape and location of the bosses.

The index of specialization (see our 1974) for Turneria is 27, while that for the subfamily
is 24. [For the most specialized ant larvae — the Leptanillinae — the index is 35, while the
Ponerinae are less specialized with 17. The index for the family as a whole is 22.]

Vol. LXXXII, June, 1974

105

Genus TURNERIA Forel

Body dolichoderoid but with 2 middorsal doorknob-shaped tubercles (1 on T3 and 1
on AIV) ; AIX and AX narrowed and turned ventrally as a stout tail. Body hairs un-
branched, smooth and spike-like. Labrum subtriangular, broadest dorsally. Mandibles
dolichoderoid.

Turneria sp. (near dahli) (Fig. 1). Length (through spiracles) about 1.9 mm. Body
dolichoderoid (i.e., short, stout, plump, and nearly straight, with both ends broadly rounded;
anterior end formed by the enlarged dorsum of the prothorax; head ventral, near anterior
end; no neck; segmentation indistinct) ; AIX and AX narrowed abruptly and bent ventrally
as a small tail; on the dorsum of each T3 and AIV a middorsal doorknob-shaped boss.
Anus on anterior surface of tail. Spiracles small; those on AI greatest in diameter, on AVIII
vestigial. Entire integument spinulose, the spinules minute and in short transverse rows.
Body hairs sparse, short (0.005-0.025 mm. long), unbranched, smooth and spike-like, longer
and more numerous on the dorsal surface. Cranium subtrapezoidal with corners rounded;
mouth parts small. Each antenna with 2 or 3 sensilla, each of which bears minute spinule.
Head hairs few, minute (0.006-0.013 mm. long) and spike-like. Labrum subtriangular in
anterior view ; anterior surface with 2 minute sensilla ; ventral border with 2 sensilla each
on a slight elevation; posterior surface with 6 small sensilla medially and with a few
arcuate rows of minute spinules laterally. Mandibles small, feebly sclerotized, dolichoderoid
(i.e., basal portion inflated and narrowed abruptly to the distal portion, which is slender
and sharp-pointed; no medial teeth); with a few short ridges at base of apical tooth.
Maxillae small, apex rounded, appearing adnate ; palp represented by a cluster of 5 (1
encapsulated and 4 with a spinule each) sensilla; galea a low knob with 2 sensilla, each
with a minute spinule. Each labial palp represented by a cluster of 4 (1 encapsulated and
3 with a spinule each) sensilla; an isolated sensillum between each palp and the opening
of the sericteries; the latter a short slit between the tips of the maxillae. Hypopharynx
densely spinulose, spinules arranged in subtransverse rows, rows grouped in 2 subtriangles
which have their bases near middle.

Material Studied: numerous larvae from Espiritu Santo, New Hebrides, E. O. Wilson,
7-13 Jan. 1954; courtesy Dr. W. L. Brown.

Literature Cited

Wheeler, G. C. and Wheeler, J. 1974. Ant larvae: review and synthesis. Mem. Entom.

Soc. Washington (in press) .

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On the Estimation of Total Behavioral Repertories in Ants

Edward O. Wilson

Museum of Comparative Zoology, Harvard University, Cambridge, Mass. 02138

AND

Robert M. Fagen

Division of Engineering and Applied Physics, Harvard University, Cambridge, Mass. 02138
Received for Publication January 7, 1974

Abstract: The total behavioral catalog size of Leptothorax curvispinosus workers in a

nest environment has been estimated by means of the Fagen-Goldman method of fitting
frequency data to negative binomial and lognormal Poisson distributions. The worker
repertory is characterized by a smaller number of rare behaviors in comparison with
vertebrate repertories. This trait makes the preparation of an adequate ethogram much
less time-consuming. The behavior of a partially bilateral worker-male gynandromorph is
described and the estimation method used to show that its repertory is probably inter-
mediate in size between those of full workers and males. The limitation of worker be-
haviors to the worker (as opposed to male) appendages suggests a bilateral as opposed to
diffuse control of movement by the gynandromorph’s brain. The advantages and difficulties
of the estimation technique are then discussed.

INTRODUCTION

The listing of behavioral repertories to produce “ethograms” is an essential
first step in the comparative study of behavior. But it is also one of the most
time-consuming. Studies of single bird species commonly last hundreds of hours,
while a few primate projects have consumed a thousand observation hours or
more over a period of years. Even at this level, there has been no systematic
way of judging how nearly complete the ethogram has become, and ethologists
have ordinarily relied on unaided intuition in choosing the time to stop. Re-
cently Fagen and Goldman (1974) proposed a method for estimating the total
size of behavioral categories by fitting frequency data of behavioral acts to
one or both of the most general distributions likely to be appropriate, namely the
lognormal Poisson and negative binomial.

The present article examines the application of this technique to two castes
of the ant genus Leptothorax and considers its general strengths and weaknesses
for insect studies. The method has also been used to evaluate the repertory of
a rare gynandromorph discovered in a colony of L. curvispinosus.

Acknowledgments: We are grateful to Dr. Mary Talbot for supplying live colonies of

Leptothorax and to Dr. Arnold M. Clark for advice on the study of gynandromorph be-
havior. The research was supported by funds from National Science Foundation grant
number GB-40247.

New York Entomological Society, LXXXII: 106-112. June, 1974.

Vol. LXXXII, June, 1974

107

Table 1. Relative frequencies of behavioral acts by workers and a gynandromorph of the
ant Leptothorax curvispinosus and by males of L. duloticus. ( N , total number of behavioral

acts recorded in each column)

Behavioral Act

Leptothorax
curvispinosus
workers
(N = 1962)

L. curvispinosus
gynandromorph
(N = 45)

L. duloticus
males
(N = 65)

1. Self-grooming

0.2370

0.7333

0.6462

2 . Antennal tipping

0.0122

3. Allogroom worker

0.0428

0.0667

4. Allogroom queen

0.002 S

Brood care:

5. Carry egg

0.0153

6. Lick egg

0.0255

7. Carry larva

0.1264

8. Licking larva

0.1804

9. Assist larval ecdysis

0.0056

10. Feed larva solid food

0.0336

11. Carry pupa

0.0122

12. Lick pupa

0.0484

13. Assist eclosion of adult

0.0082

14. Lay egg

0.0025

Regurgitate :

IS. With larva

0.0775

0.0222

16. With worker

0.0642

0.1778

0.3538

17. With queen

0.0138

18. Fight queen or workers

0.0092

19. Lick wall of nest

0.0138

20. Forage

0.0291

21. Feed on honey

0.0056

22. Feed on solid

0.0173

23. Carry dead insect

0.0025

24. Carry dead nestmate

0.0025

25. Carry live nestmate

0.0015

26. Handle nest material

0.0041

27. Stridulate

0.0061

TOTALS

1.0

METHODS

Colonies of Leptothorax were collected by Dr. Mary Talbot at the E. S.
George Reserve, near Pinckney, Michigan. Some consisted of pure L. curvi-
spinosus, with curvispinosus queens, others of curvispinosus enslaved by the
rare parasitic species L. duloticus, the latter containing duloticus queens. The
colonies were maintained in narrow glass tubes moistened by cotton wool at
one end and left open at the other. The workers were allowed to forage freely
out of the tubes and onto the floor of small, steep-sided containers. The con-
tainers were small enough in turn (9 X 15 cm on the side by 6 cm deep) to be

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placed on the stage of a dissecting microscope. As a consequence the entire
worker populations of colonies, consisting of 20 to 100 workers, could be
monitored simultaneously. Behavioral catalogs were constructed and frequencies
of each behavior accumulated by scanning back and forth for as long as an hour
or more in continuous sessions. By this means it was possible to record all of
the discrete behavioral acts displayed by virtually every worker. Observation
periods were scattered according to convenience from 8 in the morning to one
or two hours past midnight. Over this span no differences in level or pattern
of activities were noted. Nor were any expected, since the internal nest environ-
ment remained essentially constant.

Behavioral repertories and their frequency distributions did not differ sig-
nificantly between pure curvispinosus colonies and those mixed with duloticus.
Consequently, in order to obtain as large a sample size as possible, counts were
taken from two curvispinosus colonies that had been especially well analyzed
in connection with a separate study of ant slavery (see Wilson, 1974), one pure
and the other enslaved. When a worker-male gynandromorph eclosed in one
of the pure colonies during the course of the study, it was closely monitored
during its short life. Simultaneously, duloticus males in a mixed colony were
monitored; curvispinosus males were not available at this time for quantitative
study, but earlier studies had shown that the repertories, if not the frequency
distributions, were identical. The data were then analyzed by the method of
Fagen and Goldman.

RESULTS

Worker repertory. The behavioral catalog and frequency data are presented in
Tables 1 and 2 and Fig. 1. The estimates given were based on a fit of the data
to the negative binomial distribution. Similar results were obtained with the
lognormal Poisson distribution. In the case of L. curvispinosus workers, the
estimated total repertory size is 29, with a 95 percent confidence interval of
[27,35]. The sample coverage, defined as 2 Ph where pi is the probability of

performance of each observed act i, is much greater, being 99.95%. This very
high value means that the still missing behaviors have an aggregate probability
of 0.0005.

Male repertory. The repertory of the L. duloticus males in the nest was ex-
tremely limited, and the two behaviors observed were not far from equiprob-
ability. As a result the estimated repertory is identical to the observed repertory,
a remarkable result in view of the small number of data utilized.

Male-worker gynandromorph. The observed repertory falls far short of the
estimated repertory, especially the upper limit of the 95 percent confidence
interval, a result that accords well with our intuitive feeling during the period

Vol. LXXXII, June, 1974

109

Table 2. Catalog and estimated total repertory of two castes and a worker-male gynandro-
morph of Leptothorax. Estimates were obtained by fitting the data of Table 1 to
a negative binomial distribution

No. of acts
observed

No. of
kinds of
behaviors
observed
(observed
repertory
size)

Estimated

total

repertory

size

Estimated
95% con-
fidence inter-
vals, total
repertory
size

L. curvispinosus
workers

1962

[27,35]

L. duloticus
males

[ 2,2 ]

L. curvispinosus
gynandromorph

[ 4,27]

of observation. Because opportunities seldom arise for the observation of living
gynandromorphs, further notes on this one individual will now be given. Data
were taken on an almost daily basis from the time the ant was discovered as
a one- or two-day-old callow until its death eleven days later. The total observa-
tion time was six hours.

The gynandromorph was the size of a small worker. The body behind the
head was covered preponderantly by worker exoskeleton (easily distinguished
by its yellow as opposed to blackish brown coloration). The only male portions
were the left lateral edges of the pro- and mesothoraces and the left fore and
middle legs. The two legs were mostly useless, ordinarily being carried folded
beneath the body. We gained the impression that these two appendages, which
were longer and more slender than their worker counterparts, were under
the control of the worker part of the central nervous system. (Presumably the
thoracic ganglia consisted of worker tissue.) The foreleg often moved in a
nearly normal fashion down to about the level of the metatarsus, where a worker
leg would have ended, but the terminal segments kept folding under the meta-
tarsus when the leg was moved forward and down.

The division of the head was exactly bilaterally symmetrical. So precise
was the line of demarcation that the median ocellus was half developed — or
the male side. It is a reasonable supposition that the division extended througl
the brain. The behavior of the gynandromorph proved to contain an inter
esting mixture of male and worker elements, as follows.

1. Level of activity. The gynandromorph was quite inactive and spent most
of its time resting in one position. In this respect it much more closely resembled
males of the same age than workers. When nearby workers discovered honey
and began to rush excitedly in and out of the tube nest while regurgitating to

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ABUNDANCE RANGE

Figure 1. Frequency distributions of the 2 7 observed behaviors of Leptothorax curvi-
spinosus workers, 111 behaviors of playing children, and 120 behaviors of rhesus monkeys.
The mode has clearly emerged in the ants and children, indicating that most kinds of be-
haviors have been cataloged. This is particularly true of the ants, in which relatively few
rare categories have so far been discovered. (Human and rhesus data from Fagen and
Goldman, 1974.)

each other, the gynandromorph did not participate, a male-like rather than
worker-like characteristic.

2. Location. The gynandromorph spent over 90 percent of its time at all
hours of the day standing or walking around slowly within one cm of the nest
entrance. This was a position sometimes taken by young workers but seldom
if ever by males, which preferred to remain deep in the nest and especially near
the brood. Males showed a circadian increase in activity, sometimes walking all
the way out of the nest and attempting to escape from the foraging arena be-
tween about 9 pm and 1 am. Males from wild colonies were captured at lights at
11 pm. Together, these data indicate that nuptial flights are conducted at night.
No such circadian rhythm was noted in the gynandromorph.

3. Alio grooming. On three occasions the gynandromorph was observed to
groom workers, a behavior commonly seen in workers but not in males. Sig-
nificantly, the worker antenna was employed for orientation much more than
was the male antenna during these bouts. On another occasion the gynandro-
morph regurgitated with a larva, another behavior characteristic of workers but
not of males. The worker antenna was used to investigate and the worker fore

Vol. LXXXII, June, 1974

111

tarsus to stroke the larva; the corresponding male appendages were not em-
ployed. Although the allogrooming responses were worker-like, they occurred
less frequently than in full workers.

4. Antennal posture and general orientation. The male antenna was held in a
more extended position than the worker antenna; the postures of both were
typical of the caste they represented. When the gynandromorph investigated a
worker nestmate (as opposed to grooming it), both antennae were used equally.

5. Investigation of solid food. The gynandromorph was seen to explore a
fragment of moth thoracic muscle being eaten by a larva, a behavior common in
workers but not seen in males. During this brief episode only the worker
antenna was used.

In summary, the gynandromorph displayed a mixture of male and worker
traits. Its actual and estimated total repertory sizes were intermediate between
those of males and workers. The estimation technique indicates that a smaller
fraction of the total repertory was observed than in the case of the full males
and workers. The worker behaviors were also displayed less frequently than in
full workers. When the gynandromorph behaved as a worker, it used its worker
antenna primarily or exclusively, suggesting a bilateral separation of effector
control at the level of the central nervous system as opposed to a mixed control.
This correspondence between anatomical and behavioral mosaicism is con-
sistent with earlier findings on Drosophila and Habrobracon (Manning, 1967;
Stern, 1968; Hotta and Benzer, 1972).

DISCUSSION

Let us next consider the strengths and weaknesses of the Fagen-Goldman
catalog estimation method with special reference to ants and other insects.
The obvious advantage of the technique is that it improves unaided intuition
without forcing any new, unsupportable assumptions on the analysis. It is
possible to judge more precisely the point of diminishing return during the
preparation of ethograms.

This point came surprisingly early in the case of the ants. After only 51
hours of observation, during which 1,962 separate acts were recorded, the mode
of the frequency curve emerged and the estimated sample coverage attained
99.95 percent. Thus the effort required to secure a nearly complete repertory
seemed to be a full order of magnitude less than in the vertebrates. This result
implies that comparative ethological studies can proceed much more rapidly
in ants and other insects.

A reason of considerable potential biological interest exists for this relative
tractability of ant studies. This is the scarcity of rare acts compared with
common acts (see Fig. 1). In other words, whatever ants do they do rather
frequently; few if any rare behaviors exist to surprise the investigator in the
late stages of a study. We conjecture that the small size of the ant brain

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precludes the storage of responses that are not used commonly. As one of us
has pointed out previously (Wilson, 1971), a characteristic of behavior in
social insects is the repeated use of the same communicative signals and re-
sponses in different contexts to achieve various purposes.

There are two disadvantages of the method which we do not regard as par-
ticularly serious. The first is the probability that the repertories and frequency
distributions change in different contexts. It remains for the biologist to define
those contexts and to repeat the analysis within them. In the case of ants
distinguishable contexts are not only finite but also probably quite limited in
number. By far the greatest part of an ant’s life is conducted in the homeostatic
environment of the nest interior. Thus the lifetime sample coverage in the
present study, defined as the cumulative probability of all behavior for all
contexts, was probably very high in spite of the fact that it was limited to one
environment. We suggest that the following list might exhaust the remaining
contexts for the worker caste: extended foraging periods; major disturbances
of the nest, including invasion by alien colonies, flooding, and overheating;
emigration to a new nest site; and assisting during the initiation of nuptial
flights on the part of the reproductive forms.

The second difficulty in repertory estimation is the arbitrariness of the
definition of the behavioral act. One observer might see three distinct neuro-
muscular patterns where another sees only one. Thus “foraging” as defined in
the present study could easily be broken down into several acts. This is
essentially a problem of language, and different observers can solve it by a
straightforward mapping procedure. One observer’s acts a , b, and c will be
recognized as comprising the second observer’s act a\ the first observer’s act h
will be seen as representing the second observer’s acts m and n\ and so forth.
No great difficulty should occur when the same species is considered or closely
related species are compared. Serious conceptual problems might exist, how-
ever, when an attempt is made to compare the size and frequency characteristics
across radically different species.

Literature Cited

Fagen, R. M. and Goldman, R. N. 1974. Behavioral repertory size estimation. {In Fagen,
R. M. 1974. Theoretical bases for the evolution of play in animals. Ph.D. Thesis,
Harvard University, Cambridge, Mass, xvi -f- 255 pp.)

Hotta, Y. and Benzer, S. 1972. Mapping of behaviour in Drosophila mosaics. Nature,
240: 527-535.

Manning, A. 1967. Genes and the evolution of insect behavior. In J. Hirsch, ed.,
“Behavior-Genetic Analysis.’’ McGraw-Hill, New York. Pp. 44-60.

Stern, C. 1968. “Genetic Mosaics and Other Essays.” Harvard University Press, xiv +
185 pp.

Wilson, E. O. 1971. “The Insect Societies.” Belknap Press of Harvard University Press,
x -f 548 pp.

Wilson, E. O. 1974. Leptothorax duloticus and the beginnings of slavery in ants.
Evolution, in press.

Vol. LXXXII, June, 1974

113

Zoogeography of the Imported Fire Ants* 1

William F. Buren,2 George E. Allen,2 Willard H. Whitcomb,2
Frances E. Lennartz,3 and Roger N. Williams4

Received for Publication February 14, 1974

Abstract. The present known ranges of the imported fire ants Solenopsis richteri and
S. invicta in North America and South America are shown. Hypothetical answers are given
to the questions of how far the species will spread in North America, why both species
first became established in the Mobile, Alabama, area, why S. invicta has an extremely
elongate, narrow, north-south range in South America, and why it is absent from areas
of South America which appear ecologically favorable.

Key words: Solenopsis, richteri, invicta , ranges, homelands.

Buren (1972) recognized two species of imported fire ants in the United
States, the black imported fire ant, Solenopsis richteri Forel, and the red im-
ported fire ant, S. invicta Buren. Southernmost Brazil, Uruguay, and Argentina
are the homelands of S. richteri (Creighton, 1930; Wilson, 1952; Buren, 1972;
and authors) and the state of Mato Grosso, Brazil, has been proposed as the
homeland of S. invicta (Buren, 1972; Allen, et al., 1974). S. richteri is thought
to have been imported into the Mobile, Alabama, area as early as 1918 (Creigh-
ton, 1930) or perhaps even as early as the turn of the century (Lewis, 1951).
A secondary spread of this species into the area near Starkville, Mississippi,
probably by means of dirt ballast via railroad transport, may have occurred as
early as 1935 to 1940 (Wilson, 1951). The black imported fire ant slowly in-
creased its range in this northeastern area of Mississippi and by 1968 had
occupied an area approximately 135 miles long (Tupelo to Meridian, Miss.,
personal records) and with eastern extensions into western Alabama (near
Aliceville, Cochrane, Pickensville, and Ethelsville, and Vernon, Sulligent, Win-
field, and Carbon Hill), plus an isolated record at Rogersville. This is the only
known area in the United States where S. richteri , the original imported fire ant,
is still extant.

The profound behind-the-scenes influence of Dr. William S. Creighton in the

Acknowledgments: The authors wish to thank Miss Debbie Brandt for drawing the maps

and Dr. Murray S. Blum for kindly permitting us to reference information in a personal
communication from the late Dr. William S. Creighton. This research was supported in large
part by Cooperative Agreement 12-14-100-10, 952(33), Agriculture Research Service, U.S.
Department of Agriculture, with the University of Florida.

1 Florida Agricultural Experiment Station Journal Series No. 5327.

2 Department of Entomology and Nematology, Univ. of Florida, Gainesville, Fla. 32611.

3 Department of the Interior, Washington, D.C.

4 Ohio Agriculture Research Station, Wooster, Ohio.

New York Entomological Society, LXXXII: 113-124. June, 1974.

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development of the two imported fire ant species concept is not apparent from
the literature. Creighton (1930) had suggested that the then recognized sub-
species of Solenopsis saevissima (F. Smith) were more than usually distinct
and might one day need to be recognized as separate species. He privately
held firm in this view even though later authors (Wilson, 1952; Ettershank,
1966) synonymized all but one of these taxa under saevissima. Dr. Creighton
was the first myrmecologist in recent years to recognize that the black imported
fire ant, identical with the original Mobile population, was still present in the
United States. This was done in a personal letter (April, 1968) to Dr. Murray
S. Blum after identifying some specimens from Tupelo, Mississippi. Dr.
Creighton was also unstinting in his private encouragement and advice to the
senior author in his taxonomic studies on the fire ants. The authors are indeed
pleased that Dr. Creighton lived to see his 1930 viewpoints vindicated.

S. invicta appears to have invaded the United States in the Mobile, Alabama,
area some time between 1933 and 1945, possibly between 1933 and 1941. This
time span seems reasonably certain because Creighton (personal communication
to Miss Lennartz, 1973) was actively collecting in the Mobile area and along
the Gulf Coast until 1933 and found only S. richteri , whereas the first authentic
specimens of S. invicta were not captured until 1945 (Buren, 1972), although
Wilson (1951) believed he may have seen the “light phase” imported fire ant
in the dock area at Mobile in 1941. In any case the new invader was quickly
successful in expanding its territory, both by mating flight dispersal and by
man’s agency (Markin, et al., 1971; Culpepper, 1953). This species is now the
dominant formicid in a very large area of the southern United States, with large
infested areas in North Carolina, South Carolina, Georgia, Florida, Alabama,
Mississippi, Louisiana, and Texas, plus a smaller area in Arkansas. Some of the
early history was obfuscated by the unfortunate confounding of the two species
and listing as a single taxon (as Solenopsis saevissima richteri Forel or as S.
saevissima [F. Smith] by numerous authors).

While it is difficult to be certain which species is being discussed, it seems
reasonable to suggest that the early reports about the spread of the fire ant up
through the late 1930s and early 1940s probably apply to S. richteri. M. R.
Smith (in an unpublished report, 1949) records S. richteri from several localities
in Mobile County and one in Baldwin County in Alabama in 1931. By 1937
it had been seen in several localities in Jackson County in southern Mississippi.
By 1947, Clay Lyle had found a large isolated population around Artesia,
Mississippi, a small railroad stop east of Starkville. Another isolated population
was found near Meridian, Mississippi. From specimens collected by E. O.
Wilson, it is known that S. richteri still existed along with invicta in the Mobile
area and at Foley, Alabama, in the late 1940s. During the 1950s, however,
richteri was becoming sparse or appeared to be eliminated from many of its
southern areas, and only invicta remained (Wilson and Brown, 1958).

Vol. LXXXII, June, 1974

115

The existence of two species rather than one makes it certain that two separate
importations are involved and leads to the question of why both importations
were in the Mobile, Alabama, area. One of us (Lennartz, 1973) has shown
that no single imported commodity (Brazil nuts, quebracho, coffee, rubber,
mahogany, etc.) can be definitely associated with the importation of S. invicta.
Anemochore or hydrochore dispersal seems out of the question. It can only be
stated that the species must have been aboard shipping from South America and
came ashore in an unknown manner. If an established colony were aboard ship
and happened to have a wedding flight involving both males and females while
in port, then the mated females hypothetically could have flown ashore and
established a number of colonies. To hypothesize this method, however, it also
seems necessary to suppose that the biotic factors ashore were favorable for this
type of invasion. Whitcomb, et al. (1973), believe that 99 percent or more of
S. invicta females are destroyed by predation by other ants, other animals,
and by other biotic and abiotic factors in north Florida during and after mating
flights and during colony founding. With an annual production of circa 97,000
females per acre (Whitcomb, et al., 1973) this mortality may not be able to halt
the spread of S. invicta from heavily infested areas but might be a serious impedi-
ment to the establishment of the species by a few females flying ashore from a
wedding flight initiating aboard a ship in port.

It may be postulated, therefore, that the biotic conditions at Mobile at the
time of the invicta invasion were somehow favorable to this species. Again we
are led to the question of why both species of Solenopsis were first established at
Mobile. Why not one of them at New Orleans? It is known (United States
Shipping Board Report, 1926-1936) that New Orleans received more shipping
from South America than Mobile during this period.

Our hypothetical answer is rooted in what we can piece together of the history
of several ant invasions in southern United States. It seems reasonably estab-
lished that S. richteri arrived in Mobile about 1918 or perhaps even earlier and
that by 1928 was common there although it was not as numerous as invicta
was to become approximately 20 years later. It is also known that another South
American ant, Iridomyrmex humilis Mayr, the Argentine ant, became established
in southern United States, probably first at New Orleans, before the turn of the
century (Foster, 1908). By 1913 (Newell and Barber, 1913) and continuing to
the early 1940s (personal observations), this ant had become overwhelmingly
abundant at New Orleans and had completely eliminated all other ant species
in its held territory.

It is doubtful that queens of 5. invicta could have established new nests during
those years at New Orleans. At Mobile, however, it is possible that S. richteri
was keeping /. humilis in partial check. A hypothesis (reported in Wilson,
1951) that /. humilis had pushed the S. richteri population north of Mobile
during the early 1920s seems doubtful to us, but in any case it is known

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(Creighton, 1930) that the S. richteri population was back in place by 1928 and
subsequently.

Possibly S. richteri not only helped to prevent I. humilis from reaching massive
population levels at Mobile but was also having some effect on the abundance
of native ants, in particular the native fire ants S. geminata (Fabr.) and S.
xyloni McCook. However, S. richteri never fully occupied the territory available
to it at Mobile as invicta was to do later and as richteri itself was to do later
in northern Mississippi. Thus, in our view, the success of the initial invasion
by invicta may have been caused by a “preconditioning” of the area by the
original imported fire ant richteri in a manner which helped to alleviate some
of the competition and predation from native ants and from /. humilis, while
at the same time leaving an ecological niche partially open, a niche which
invicta was to find eminently suitable for exploitation. These factors may even
partially explain the early explosive buildup of invicta in the Mobile area.
Northern United States seaports such as New York, which receives even more
shipping from South America than either New Orleans or Mobile, can be excluded
from consideration because of the obvious abiotic factor of winter severity.

The present areas of infestation of S. richteri and invicta in the United States
are shown in Fig. 1 . These areas are based on the identification of approximately
600 nest collections, plus data as given by the United States Department of
Agriculture (Markin, et al., 1972). A few isolated locality records for S. invicta
are known from farther south in Florida than shown. It is impossible to guess
the eventual boundaries of the richteri infestation, but as the range of richteri
probably extends from approximately 30° to 38° south latitude in South America,
a more northward extension of the range of richteri in the United States could
reasonably be expected, possibly northward into Tennessee and Kentucky. Con-
trol and eradication efforts, if continued, may negate or strongly modify this
projection. The rate of expansion of the territory of S. richteri appears to be
slow.

The northward progression of S. invicta, on the other hand, after a period of
extremely rapid expansion well documented by various authors (see especially
Wilson and Brown, 1958; Adkins, 1970), seems to have reached close to a
northern limit, except for minor local enclaves. We believe this may be due
mainly to winter kill conditions. S. invicta is a species in which hibernation
apparently does not occur. Examinations of nests in near freezing or freezing
temperatures (personal observations) reveal that the ants are up in the tumulus
at about the same depth as in more favorable temperatures. Only in hot, dry
conditions will the ants be down in the nest out of the tumulus. At Atlanta,
Georgia, about on the northern boundary of the range, based on observations
over a four-year period, the species is not abundant and its limited population
appears to be maintained with difficulty. A few colonies in favorable situations,
such as on southern slopes fully exposed to the winter sun, achieve fair size, but

Vol. LXXXII, June, 1974

117

almost all new colonies which arise during the summer do not appear to survive
the winters. This contrasts strongly with the abundance of the species, prior to
eradication and control programs, only approximately 100 miles south of
Atlanta.

S. invicta is expanding its range to the west. The species has been taken as
far west as San Antonio, Texas. There seems no doubt but that our previous
prediction (Buren, 1972) regarding its possible establishment in the cities and
favorable localities in the southwest eventually could come true. We know of
no reason why S. invicta could not become established in the southwestern
cities where S. xyloni is now common, displacing the latter as it has in the south-
eastern states. The native desert fire ant, Solenopsis aurea Wheeler, is small in
size and lives in small colonies, and it seems inconceivable that this species could
offer any resistance to the spread of S. invicta in those southwestern ecological
niches where the latter could colonize. The distribution of invicta in the south-
west could be expected always to remain sporadic, along canals, in irrigated
fields, in watered lawns, etc. We would not expect it to become established
in actual desert situations. It is possible that if it ever reached California, the
species could become a pest there in irrigated areas, displacing /. humilis as it
apparently now has done almost completely at New Orleans and other south-
eastern areas.

The hypothesis that winter kill is limiting the northward expansion of S. invicta
seems reasonable and we can think of no other explanation which fits the data

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as well. A previous tentative hypothesis that the northern boundary of the
S. invicta range could be influenced by the range of Lasius neoniger Emery, a
northern predator and competitor (Bhatkar, et al., 1972), or by a number of
northern predators no longer seems reasonable to us. There is no northern ant
species or series of species known to us which has a cohesive range that would
fit in its southern limit the relative smoothness of the northern limit of S. invicta.
As far as we are aware, Lasius neoniger is absent or rare in the southern Great
Plains (northern Texas and Oklahoma) where 5. invicta has not penetrated any
farther north than in the southeastern states. Lasius neoniger is not present
or is probably rare in Atlanta, Georgia, also, where, as previously stated, the
S. invicta population appears to be in difficulty. From data given by Wilson
(1955), L. neoniger appears to be very sporadic in the southern states. About
all that may be postulated in regard to the biotic factors in this question is that
the abiotic factor of winter kill from freezing temperatures possibly weakens
the colonies sufficiently so that they are more subject to competition or preda-
tion from native ants, if they are not completely killed initially. It seems likely
that the severity of winter kill is roughly proportional to the depth to which
the soil becomes frozen. Freezes up to four inches, which can occur at Atlanta,
possibly often kill most of the workers and brood of a colony and sometimes the
queen. The combined biotic and abiotic factors are probably especially harsh
on incipient colonies and thus there can be little or no population buildup or
spread.

The South American ranges of S. richteri and invicta are shown in Fig. 2.
Actual locality records of S. invicta seen and identified by the senior author are
marked as well as the range postulated from these data. A large number of
individual nests have been sampled at some of these localities. The differences
between the shapes of the ranges in North and South America are striking for
S. invicta, which has an enormously long north-south range in South America
with only a narrow east-west distribution, whereas in North America the main
axis is east-west. The combined biotic and abiotic factors which enforce these
distributional differences are not fully understood and deserve further study.

The range of S. richteri shown represents our “guesstimate” for this species.
We know that it occurs in southern Rio Grande du Sul, Brazil, probably
throughout Uruguay, and south an unknown distance into Argentina. We have
not seen specimens from Bahia Blanca, but this city is the type locality of
Solenopsis quinquecuspis Forel, a species found in parapatric associations with
S. richteri in Uruguay (Buren, 1972), and it seems reasonable that the two
species would have fairly similar range extensions. The western limits of the
range of S. richteri are not known, but we have seen no Solenopsis specimens
which can be identified as S. richteri from Cordoba, from northwestern Argentina
(provinces of Jujuy, Salto, Tucuman, Formosa, or Chaco), or from Paraguay.
Therefore, our estimation of the range of 5. richteri is considerably less extensive

Vol. LXXXII, June, 1974

119

Homeland Areas of Fire Ants

60' • 50°

So/enopsis invicta ^ 5, invicta collection sites

So/enopsis richteri national boundaries

Figure 2

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than that given by Wilson (1952). Buren (1972) made similar remarks on the
range of 5. interrupt a Santschi.

The northernmost record of S. invicta in South America is Porto Velho,
Rondonia Territory, Brazil, and the southernmost record is near Resistencia,
Chaco, Argentina, a distance of about 3,000 km. This compares with 1,345 miles
or 2,250 km. in North America, the distance between the easternmost records of
invicta in North Carolina and San Antonio, Texas. The width of the invicta
range in South America appears to be relatively narrow and if exemplified by
the distance between Corumba and Coxim, Mato Grosso, is only about 350 km.
wide and possibly even considerably less wide in its southern arm into Argentina
and Paraguay and its northern arm into the Amazon drainage along the Guapore
River. Most of the available records are from localities which fringe the
Pantanal (large flood plain [60,000,000 to 90,000,000 hectares] of the head
waters of the Paraguay River), and although the interior of this area has not
been sampled there seems little doubt but that the species occurs in favorable
locales throughout the Pantanal. Otherwise we could not expect it to be so
uniformly distributed around the periphery. The Pantanal has been proposed
(Allen, et al., 1974) as the probable original homeland of 5. invicta and this
hypothesis still appears reasonable to us. Hydrochore dispersal via the well-
known phenomena of massing together and floating downstream during flooding
(Lennartz, 1973) could easily account for the far south and far north popula-
tions of invicta along the Paraguay and Guapore rivers, respectively.

The western extensions of the range of S. invicta are not known but we believe
them to be rather limited. None of the Solenopsis material captured so far in
Bolivia can be identified as this species. We would expect it to occur in
easternmost Bolivia, however, since portions of the Pantanal extend into this
country.

Why a vigorous species such as invicta has not penetrated farther to the east
of its present area remains an ecological mystery about which we can only make
guesses. The species has not yet been taken in the state of Sao Paulo and has
not been found east of Rondonopolis or from Campo Grande eastward in Mato
Grosso. Other species in the S. saevissima complex have been found (Allen,
et al., 1974) in these areas, so it is obvious that the areas are not entirely
insalubrious to Solenopsis. (These species are presently under taxonomic study
by the senior author.) However, in effect, invicta has not been found either in
the cerrado area to the east of the Pantanal, where Allen et al. (1974) and
Lennartz (1973) have postulated that a lack of moisture during the prolonged
dry season might halt its progress, or even in what would seem to be favorable
limited areas along streams and rivers more than a short distance (approxi-
mately 50 to 100 kilometers) from the Pantanal. If the species is “at home” and
fairly abundant in the flood plain of the Paraguay River, why, apparently, is it
absent from the flood plain of the Parana River, which joins the Paraguay near

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121

where invicta has been captured in Argentina? And if present there why could
it not move thence into favorable areas of the state of Sao Paulo? Other species
of Solenopsis have been collected along the Parana River, but not S. invicta.

It is easy to postulate that a combination of abiotic and biotic factors en-
forces these territorial limits without knowing the exact parameters or how
they act precisely. One can logically postulate in a general way that invicta
needs more soil moisture than certain other species in the saevissima complex
and, therefore, is at a competitive disadvantage with these species in the
campo cerrado and thus has not been able to expand eastward out of the
Pantanal region.

Another hypothesis is that competitive action by other ants, possibly species
of the genus Pheidole, may be of importance in limiting the spread of 5. invicta.
Pheidole is a large and predominant genus in the neotropics, with nearly 400
taxa (Kempf, 1973). Various species are numerous both in the forested areas
and in the campo cerrado, where Solenopsis of the saevissima group is rare or is
limited to ecologically disturbed areas. On the other hand, in our observations
on the fringes of the Pantanal, Pheidole spp. do not seem very common in this
area, possibly due to the annual flooding which Solenopsis can withstand by
massing together and floating but which, perhaps, Pheidole cannot.

Mutual exclusiveness in the ranges of ants has not been studied in depth but
is known to occur. See, for example, the remarks of Levins and Heatwole (1973)
on the mutually exclusive ranges of Solenopsis geminata (Fabr.) and Pheidole
megacephala (Fabr.) on islands in the West Indies, and also those of Buren
(1968) on the mutually exclusive ranges of Conomyrma bicolor Wheeler and
Crematogaster larreae Buren in the deserts near El Paso, Texas. In each case,
the range limitations were due to a combination of abiotic and/or biotic factors.

It is the ground-patroling activities of Pheidole which are suspected of being
inimical to Solenopsis through efficient detection and attacks on the newly mated
queens after wedding flights. The queens, unlike the workers, do not sting or
effectively defend themselves. A number of adverse factors, such as the
postulated attacks on the queens by Pheidole workers, lack of soil moisture for
long periods of the year, and, perhaps, predation on incipient colonies by maraud-
ing ants (Dorylines) could severely limit even a vigorous species.

A time factor must also be considered. It is reasonable to suggest that a cer-
tain time must elapse before any two or more species which are in competition can
come to equilibrium in the territory occupied. This time could be relatively short
in a case where one species is clearly more aggressive than others or very long
where the species are more or less evenly matched. Erickson (1972) has in-
vestigated the displacement of Pogonomyrmex calif ornicus Buckley by Irido-
myrmex humilis and finds that this has proceeded at the rate of about 100 meters
per year in the old field studied. Other studies concerning the displacement
of various ants by I . humilis have been comparable. The encroachment upon

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native ants by Solenopsis invicta has been much more rapid, however, and
occasionally may have been as much as 5 miles per year (Wilson and Brown,
1958). Even where very rapid, minor enclaves or pockets of the lesser species
are likely to remain. For instance 5. invicta appears to be having difficulty be-
coming predominant in parts of the central sandy uplands of Florida and S.
geminata has remained the predominant ant in Alachua County and other
locales of this region, in spite of the fact that S. invicta is found there
sporadically. 5. geminata also remains in small areas at Tall Timbers Research
Station north of Tallahassee, Florida, where invicta has otherwise claimed
exclusive usage of certain territory especially favorable to it, such as along the
mucky shoreline of Lake Iamonia, where the water table is very close to the
soil surface.

Where the several species are nearly evenly matched, the distribution patterns
either can become sympatric, as in the case of invicta and one or more unknown
Solenopsis species in the Pantanal, and in the case of S. blumi Buren, interrupta
Santschi, and quinquecuspis Forel in Uruguay or can assume parapatric patterns,
as in the case of richteri and quinquecuspis in Uruguay (Buren, 1972).

Ants, just as many other insects and other animals, have highly differing
ranges. There are examples of ants with extremely extensive ranges such as
the holarctic ranges of Camponotus herculeanus (Linne) and Formica jusca
Linne or the extensive neotropical range of Paraponera clavata (Fabr.). These
contrast with the very limited ranges of such species as Crematogaster opuntiae
Buren (cholla cactus associations in the sonoran desert of southern Arizona),
C. navajoa Buren (pinyon pine-juniper-grasslands of northern Arizona and
southern Utah), and Discothyrea testacea Roger (in fern areas, coastal plains
of the Carolinas and Georgia). In the case of species with extensive ranges, a
long time span of existence as stable species seems to be the only explanation.
In the case of species with small, limited ranges, it can be postulated, however,
that their existence as separate species has either been relatively short (possibly
the case with Crematogaster navajoa and opuntiae since the arid desert condi-
tions of the southwest are relatively recent and the two species have not spread
out of their small ranges to other ecologically similar areas), or they may be
ancient, impoverished relict species, or they may have very restrictive crypto-
biotic habits (as may be the case with Discothyrea testacea).

In South America the evidence is that the rain forests have not always occupied
the extensive area now occupied but, owing to severe continent-wide drought
conditions, have periodically retreated into isolated enclaves, the latest period
only 2,600 years ago and the period previous to this only 11,000 years ago.
The isolated enclaves are thought to have contributed to the complexity of
speciation seen in the hylean forests. For a review of this subject see Vanzolini
(1973) and Vuilleumier (1971). We submit, however, that if the hylean forests
withdrew into enclaves during periods of severe drought, then very probably

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123

other moist areas in South America were severely limited also, including the
Pantanal. If Solenopsis invicta speciated in the recent geologic past within the
Pantanal during a period of great isolation, then it follows that its range would
have been severely limited, and our tentative hypothesis is that the species
has not had time since these periods, considering the many biotic and abiotic
factors mitigating against its spread, to reach all the areas which might be
ecologically favorable to it. In North America, however, following its chance
introduction, its spread was almost unbelievably rapid and unhampered by the
factors which are operative in South America. The progress of the co-invader
S. richteri does not seem conspicuously successful in the areas of the United
States that it has invaded, but it can be wondered how it would have fared
if it had reached the southern Great Plains, perhaps fairly similar to its homeland
pampas.

Literature Cited

Adkins, H. G. 1970. The imported fire ant in the southern United States. Ann. Assoc.
Amer. Geograph., 60(3) : 578-592.

Allen, G. F., Buren, W. F., Williams, R. N., DeMenzes, M., and Whitcomb, W. H.
1974. The red imported fire ant Solenopsis invicta : Distribution and habitat in

Mato Grosso, Brazil. Ann. Ent. Soc. Amer., 67(1): 43-46.

Bhatkar, A., Whitcomb, W. H., Buren, W. F., Callahan, P., and Carlysle, T. 1972.
Confrontation behavior between Lasius neoniger (Hymenoptera: Formicidae) and

the imported fire ant. Environ. Ent., 1(3): 274-279.

Buren, W. F. 1968. A review of the species of Crematogaster, sensu stricto, in North
America (Hymenoptera, Formicidae). Part II. Descriptions of new species. J.
Georgia Entomol. Soc., 3(3): 91-121.

Buren, W. F. 1972. Revisionary studies on the taxonomy of the imported fire ants. J.
Georgia Entomol. Soc., 7(1) : 1-26.

Creighton, W. S. 1930. The new world species of the genus Solenopsis (Hymenoptera,
Formicidae). Proc. Amer. Acad. Arts Sci., 66(2): 39-151, 8 plates.

Culpepper, G. H. 1953. Status of the imported fire ant in the southern states in July,
1953. U.S. Department of Agriculture, Bureau of Entomology and Plant Quarantine.
Special Report E-867, Washington, D.C.

Erickson, J. M. 1971. The displacement of native ant species by the introduced Argentine
ant Iridomyrmex humilis Mayr. Psyche, 78(4): 257-266.

Ettershank, G. 1966. A generic revision of the world Myrmicinae related to Solenopsis
and Pheidologeton (Hymenoptera: Formicidae). Aust. J. Zool., 14: 73-171.

Foster, E. 1908. The introduction of Iridomyrmex humilis into New Orleans. Jour. Econ.
Ent., 1(5): 289-293.

Kempf, W. W. 1972. Catalago abreviado das Formigas da Regiao Neotropical (Hym.

Formicidae). Studia Entomologica, 15(1-4): 3-343.

Lennartz, F. E. 1973. Modes of dispersal of Solenopsis invicta from Brazil to continental
United States — A study in spatial diffusion. Unpublished master’s degree thesis.
Univ. of Florida.

Levins, R. and Heatwole, H. 1973. Biography of the Puerto Rican Bank: Introduction
of species onto Palominitos Island. Ecology, 54: 1056-1064.

Lewis, L. F. 1951. Investigation in the biology of the imported fire ant. In U.S. Depart-
ment of Agriculture Quarterly Report, April-June 1951.

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Markin, G. P., Dillier, J. H., Hill, S. O., Blum, M. S., and Hermann, H. R. 1971.
Nuptial flight and flight ranges of the imported fire ant Solenopsis saevissima richteri
(Hymenoptera: Formicidae). J. Georgia Entomol. Soc., 6(3): 145-156.

Markin, G. P., Ford, J. H., Hawthorne, J. C., Spence, J. H., Davis, J., Collins, H. L.,
and Loftis, C. D. 1972. The insecticide Mirex and techniques for its monitoring.
U.S. Department of Agriculture, Animal and Plant Health Inspection Service, 81-3.
Newell, W. and Barber, T. C. 1913. The Argentine ant. U.S. Department of Agriculture,
Bureau of Entomology, Bull. no. 122.

Report on the volume of water-borne foreign commerce of the United States by ports
of origin and destination. 1926-1936. U.S. Shipping Board, Bureau of Research,
Maritime Records Division, Washington, D.C.

Smith, M. R. 1949. Report on imported fire ant investigations. U.S. Department of
Agriculture, Bureau of Entomology and Plant Quarantine. Unpublished report, July
26, 1949.

Vanzolini, P. E. 1973. Paleoclimates, relief, and species multiplication in equatorial forests.
In “Tropical Forest Ecosystems in Africa and South America: A Comparative Review.”
Smithsonian Institution Press.

V uilleumier , B. S. 1971. Pleistocene changes in the fauna and flora of South America.
Science, 173: 771-780.

Whitcomb, W. H., Bhatkar, A., and Nickerson, J. C. 1973. Predators of Solenopsis
invicta queens prior to successful colony establishment. Environ. Ent., 2(6):
1101-1103.

Wilson, E. O. 1951. Variation and adaptation in the imported fire ant. Evolution, 5:
68-79.

Wilson, E. O. 1952. O complexo Solenopsis saevissima na America do Sul (Hymenoptera:
Formicidae). Mem. Inst. Oswaldo Cruz., 50: 49-68.

Wilson, E. O. 1955. A monographic revision of the ant genus Lasius. Bull. Mus. Comp.
Zool., 113(1) : 1-199, 2 plates.

Wilson, E. O. and Brown, W. L. 1958. Recent changes in the introduced population of
the fire ant Solenopsis saevissima (Fr. Smith). Evolution, 12: 211-218.

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125

Microsporidan and Fungal Diseases of Solenopsis invicta
Buren in Brazil* 1

George E. Allen and William F. Buren

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

Received for Publication February 14, 1974

Abstract: The first record of a microsporidan infection in the family Formicidae is
presented. The organism, a Thelohania sp., was isolated from Solenopsis invicta Buren
colonies in Brazil in 1973. Microsporidan spores were also observed in three other species
of the S. saevissima complex. M etarrhizium anasopliae is also reported from S. invicta and
the leaf-cutting ant Atta sexdens rubropilosa.

DISCUSSION

Although the Formicidae has been one of the most extensively studied families
of insects, our knowledge of pathogens associated with the ant group is one of
the most deficient areas in insect pathology. Many of the pathogens described
from ants were isolated from a small number of individuals since “epizootics,”
such as those that occur in the Lepidoptera, are rarely observed.

All levels of association, ranging from symbiotic to parasitic relationships, can
be found between microorganisms and various ant groups. The association of
members of the Tribe Attini and their respective fungal symbionts is a well-
known phenomenon and is discussed in detail by Wheeler (1907) and Weber
(1972). Another well-documented relationship is that of the Laboulbeniomycetes
fungi and the various insect orders including Formicidae. This group includes
predominately obligate parasites which seem to have little or no effect on the
well-being of their hosts (Benjamin, 1973). According to Smith (1946), Formica
is the most common ant genus associated with members of the Laboulbenio-
mycetes, especially the genus Laboulbenia. For further information the reader
is referred to an excellent review by Benjamin (1971) which includes a summary
of the studies of Thaxter and others on the Laboulbeniomycetes.

A brief review of known pathogen involvement with the ant group is sum-
marized in Table 1. An interesting disease of Formica ruja Linne in Western
Siberia has been attributed to the fungus Alternaria tenuis Nes. (Dlusskii, 1967).
The course of epizootics of the disease is vividly described by Marikovsky
(1962) and is the first report of Alternaria as an insect pathogen. There is,

Acknowledgments: We gratefully acknowledge the assistance of Mr. Ed Hazard, Insects

Affecting Man Research Laboratory, United States Department of Agriculture, Gainesville,
Florida, for verifying the Thelohania sp.

1 Florida Agricultural Experiment Station Journal Series No. 5326.

New York Entomological Society, LXXXII: 125-130. June, 1974.

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however, reason to question the identity of the fungus, since members of the
genus Alternaria have been reported only as common plant pathogens.

Several species of the fungus Cordyceps have been reported as “pathogens”
of ants (Mains, 1948; Petch, 1932; and Van Pelt, 1958). However, McEwen
(1963) raises the question of pathogenicity of Cordyceps, noting the lack of
detailed accounts of pathological conditions in infected hosts.

Mains (1948) discusses two species of fungi as possible conidial or imperfect
stages of bicolored species of Cordyceps pathogenic to ants. Included in this
group are the genera Stilbum and Hymenostilbi.

In addition, the cosmopolitan fungal pathogens Metarrhizium anasopliae
(Metchnikoff) Sorokin and Beauveria bassiana (Bals.) Vuill. have been described
from ants (Steinhaus and Marsh, 1967). These two fungi appear to be very
important ant pathogens in South America.

To date, no virus diseases in ants have been reported. However, Steiger et al.
(1969) have observed “virus-like” particles in cell lines derived from the
cephalic ganglionic center of Formica lugubris Zetterstadt. As in the case of
viruses, pathogenic protozoa have not been reported in the Formicidae. Several
groups, especially the microsporida, are important pathogens of other families in
the order Hymenoptera. Nosema apis, the cause of “Nosema disease” in the
honey bee, Apis mellifera, is an excellent example.

diseases associated with the Solenopsis saevissima COMPLEX

Our knowledge of ant pathogens is no doubt related to the “economic im-
portance” given to this group of insects. Generally speaking, no ant species
is ranked as a “major” pest of agricultural crops, man, or animals. An exception
to this is the “red imported fire ant,” Solenopsis invicta Buren, which in recent
years has been the target of an extensive research program in the southeastern
United States.

Buren (1972) proposed a taxonomic model for the Solenopsis saevissima com-
plex and showed that there are two species of imported fire ants in the United
States (S. richteri Forel and S. invicta Buren), each from different homelands
within South America. In 1971, the authors organized and coordinated a 17-
day trip through western Brazil and established as the homeland of S. invicta
certain areas of Mato Grosso, Brazil (Allen et al., 1974). One of the objectives
of the trip was to isolate pathogens of the ant. The following is a report of
primary pathogens isolated from specimens collected during the 1971 trip and
later collections made by the senior author in February 1973 in and around the
city of Cuiaba, Mato Grosso.

Microsporida

During a taxonomic examination of the 1973 collections, the junior author ob-
served subspherical “cyst-like” bodies in the gasters of alcohol-preserved workers

Vol. LXXXII, June, 1974

127

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Atta sexdens rubropilosa M. anasopliae Allen (unpublished)

UNKNOWN Beauveria densa Placed in synonymy with B. Leatherdale (1958)

bassiana by de Hoog, 1972

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New York Entomological Society

Fig. 1. Octanucleate sporonts (S) and spores within a sporont membrane (Sp) of
Thelohania sp. in S. invicta workers, Giemsa smear, X 2,200.

of 5. invicta. Histological examination showed the bodies were not cysts but
rather masses of microsporida spores enclosed within fat body-cell membranes.
These structures were not found in living ants examined later, even though both
sporonts and spores were present.

The microsporidan isolated from living workers of S. invicta was Thelohania
sp. Giemsa-stained smears showed that the octonucleate sporonts produce eight
spores enclosed in a sporont membrane (Fig. 1). These characteristics place the
microsporidan in the genus Thelohania Henneguy. Spores are pyriform with
average fixed spore measurements of 3.4 fx X 2.0 /x. Schizonts of the micro-
sporidan were also observed in Giemsa-staned smears of adult workers and
queens of S. invicta. The primary site of infection was the fat body. To the
best of the authors’ knowledge, this is the first report of a microsporidan infection
in the family Formicidae.

Every S', invicta colony sampled in 1973 showed a high infectivity rate. No
mounds were evident and the colonies were found only after digging around
large rocks, cement pillars, and other protective objects. Infected colonies
appeared to have lower than normal populations and noticeable loss of vigor
and pursuit when disturbed.

The potential of the Thelohania sp. as a biological control agent of S. invicta

Vol. LXXXII, June, 1974

129

in the United States can only be speculated at this point. The genus is well
known and the associations of many of its species and their insect hosts have
been described (Kudo, 1924; Weiser, 1961; Kellen et al., 1965; Chapman
et al., 1966).

Microsporida spores were also observed in collections made in 1971 (Allen
et al., 1974) from several localities, one of which appeared to be a Nosema.
Collections of S. invicta from Cuiaba and Porto Velho were infected at the time
of collection as well as three other species of the S. saevissima complex from
Cuiaba, Mato Grosso, Campo Grande, Porto Manga, and Corumba. Porto
Velho is located in the Territory of Rondonia, which borders the state of Mato
Grosso to the northwest. For a map depicting the location of these localities
the reader is referred to Buren et al. (1974).

Fungi

The fungus Metarrhizium anasopliae was isolated from S. invicta workers and
Atta sexdens rubropilosa Forel queens collected during the 1971 trip. This
cosmopolitan pathogen is a well-known entomogenous fungus (Steinhaus and
Marsh, 1962; Charles, 1941; and Leatherdale, 1958) which attacks a wide range
of insect hosts. It was also reported from S. saevissima richteri Forel ( = S.
richteri Forel) in Uruguay (Steinhaus and Marsh, 1967).

Metarrhizium anasopliae reportedly attacks only the queens of A. sexdens
rubropilosa in Brazil, where it is known as “queens disease” by the local citizens.
The fungus may also attack worker ants, but these are not observed because
the infected individuals probably leave the colony when infection is apparent,
as is the case with Formica rufa (Marikovsky, 1962) .

The foregoing report establishes the presence of both microsporidan and
fungal diseases in S. invicta. Although we can report only the involvement of
the Thelohania sp. and M. anasopliae at this time, there are strong indications
that we can expect to find other genera of microsporida as well as “virus-like”
pathogens of members of the S. saevissima complex in Brazil. Current studies
are being conducted to determine the interrelationship of S. invicta and its
Thelohania parasite and other pathogens of the S. saevissima complex.

Literature Cited

Allen, G. E., Buren, W. F., Williams, R. N., de Menezes, M., and Whitcomb, W. H.
1974. The red imported fire ant, Solenopsis invicta ; distribution and habitat in
Mato Grosso, Brazil. Ann. Entomol. Soc. Am., 67 : 43-46.

Benjamin, R. K. 1971. “Introduction and Supplement to Roland Thaxter’s Contribution
Towards a Monograph of the Laboulbeniaceae” (Bibliotheca Mycol., Vol. 30). Cramer,
Lehre.

Benjamin, R. K. 1973. Laboulbeniomyces. In “The Fungi” (G. C. Ainsworth, F. K.

Sparrow, and A. S. Sussman, eds.). Academic Press, New York. 4A: 223-246.

Buren, W. F. 1972. Revisionary studies of the taxonomy of the imported fire ants. J. Ga.
Entomol. Soc., 7: 1-26.

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New York Entomological Society

Buren, W. F., Allen, G. E., Whitcomb, W. H., Lennartz, F. E., and Williams, R. N.
1974. Zoogeography of the imported fire ants. J. N.Y. Entomol. Soc., 82: 113-124.

Chapman, H. C., Woodward, D. B., Kellen, W. R., and Clark, T. B. 1966. Host-
parasite relationships of Thelohania associated with mosquitoes in Louisiana (Nosemati-
dae: Microsporidia) . J. Invertebr. Pathol., 8: 452-456.

Charles, V. K. 1941. A preliminary check list of the entomogenous fungi of North
America. United States Dept, of Agric. Bur. Plant. Ind. Inst. Pest Survey Bull., 21:
707-785.

Cooke, M. C. 1889. New Australian fungi. Grevillea, 18: 1-8.

de Hoog, G. S. 1972. The genera Beauveria, Isaria, Tritirachium and Acrodontium Gen.
nov. Stud. Mycol., No. 1, pp. 1-41.

Dlusskii, G. M. 1967. “Ants of the genus Formica.''1 (K. V. Arnol’di ed.), Izdatel’stvo
“Nauka” (“Nauka” Press), Moscow. Pp. 232.

Kellen, W. R., Chapman, H. C., Clark, T. B., and Lindegren, J. E. 1966. Transovarian
transmission of some Thelohania (Nosematidae: Microsporidia) in mosquitoes of

California and Louisiana. J. Invertebr. Pathol., 8: 355-359.

Kudo, R. 1924. A biologic and taxonomic study of the microsporidia. 111. Biol. Monogr.,
9: 1-268.

Leatherdale, D. 1958. A host catalogue of British entomogenous fungi. Ent. Mon. Mag.,
94: 103-105.

Mains, E. B. 1948. Entomogenous fungi. Mycologia, 40: 402-415.

Marikovsky, P. I. 1962. On some features of behavior of the ants Formica rufa L.
infected with fungous disease. Insectes Sociaux, 9(2): 173-179.

McEwen, F. L. 1963. “Cordyceps Infections.” In Insect Pathology: An Advanced

Treatise (E. A. Steinhaus, ed.) Academic Press, New York. II. Pp. 273-290.

Petch, T. 1932. Notes on entomogenous fungi. Trans. Brit. Mycol. Soc., 16: 209-245.

Smith, M. R. 1946. Ant hosts of the fungus, Loboulbenia formicarum Thaxter. Entomol.
Soc. Wash., 48: 29-31.

Steiger, U., Lamparter, H. E., Sandri, C., and Akert, K. 1969. Virus-ahnliche Partikel im
Zytoplasma von Nerven — und Gliazellen der Waldameise. Arch. ges. Virusforsch., 26:
271-282.

Steinhaus, E. A. and Marsh, G. A. 1962. Report of diagnoses of diseased insects, 195 1—
1961. Hilgardia, 33 : 349-490.

Steinhaus, E. A. and Marsh, G. A. 1967. Previously unreported accessions for diagnosis
and new records. J. Invertebr. Pathol., 9: 436-438.

Van Pelt, A. 1958. The occurrence of a Cordyceps on the ant Camponotus penns ylvanicus
(De Geer) in the Highlands, N.C., region. J. Tenn. Acad. Sci., 33: 120-122.

Weber, N. A. 1972. “Gardening Ants, The Attines,” Mem. Amer. Phil. Soc. American
Philosophical Society, Philadelphia. 92: Pp. 146.

Weiser, J. 1961. Die Mikrosporidien als Parasiten der Insekten. Monogr. z. Ang. Ent.,
17: Pp. 149.

Wheeler, W. M. 1907. The fungus-growing ants of North America. Bull. Amer. Mus.
Nat. Hist., 23: 669-807.

Vol. LXXXII, June, 1974

131

A Supplement to the Revision of the Ant Genus Basiceros
(Hymenoptera: Formieidae)

William L. Brown, Jr.

Department of Entomology, Cornell University, Ithaca, New York 14850
Received for Publication January 7, 1974

Abstract: The genus Basiceros is expanded to include Aspididris due to the finding of a

new species, B. conjugans (Amazonian Ecuador and Colombia), which connects them.
Basiceros is redefined and the male caste formally described, and keys are provided for the
known forms of both sexes. The known d'stribution of B. singularis is extended to northern
Mato Grosso and of B. discigera to Espirito Santo State, Brazil, and to subandean Colombia.
B. singularis is confirmed as a termite predator.

INTRODUCTION

The genera Basiceros and Aspididris were treated by Brown and Kempf
(1960: 171-181) as part of a world revision of the myrmicine tribe Basicerotini.
At that time, we said of the status of Aspididris (op. cit., p. 179) :

This genus, known from workers and females, includes two neotropical species with the
basic characters of Basiceros , but in which the posterior half of the head has been transformed
into a disc-like structure, with the vertex convex, but the lateral and posterior occipital
borders drawn out into a sharp, upturned, saucer-like margin that is ornamented with a row
of clavate hairs. In A. militarise this margin is continuous around the back of the head,
from near one compound eye to the other, while in A. discigera, it is slightly interrupted
posteromedially. ... A. discigera has been placed in Basiceros by previous authors, and it is
clearly transitional in head shape between a species like B. convexiceps and the extreme
Aspididris militaris. Thus, while the generic split seems almost academic, the distinction
can still be drawn rather clearly on a practical basis, and there seems to be no good reason
to synonymize Aspididris unless further intergradient species are found.

The Wheelers have shown that the larva of A. militaris is very similar to that of Basiceros.
The two Aspididris species are known from Trinidad and southeastern Brazil, and both are
uncommon. We have no biological data on them beyond the fact that they are collected in
moist forested areas.

DISCUSSION

The “further intergradient species” has now been found, and it is described
below as Basiceros conjugans. This new species so clearly and completely links
Basiceros and Aspididris that there is no longer any excuse for recognizing the
latter as a genus apart, and the formal generic synonymy is recorded here.
The genus Cr eight onidris, with the sole species C. scambognatha, is closely
related to Basiceros but is separated on the basis of its extremely aberrant
mandibles.

The present paper, offered as a supplement to the revision of 1960, also

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describes the adult male caste of Basiceros in modern terms and presents some
new information on the distribution and biology of the species. A new key
to the Basiceros species is appended.

This article is dedicated to the memory of Dr. William S. Creighton, who
in 1950 breathed new life and reason into ant taxonomy with “The Ants of
North America.”

I should like to acknowledge the help of Dr. Henry Hermann, University of
Georgia, and Drs. Stuart and Jarmila Peck, Carleton University, Ottawa,
Canada, for furnishing material critical for this study. My own collecting
and other aspects of the research contributing toward this paper were supported
in large part by U.S. National Science Foundation, Grants GB-2175 and GB-
31662X.

BASICEROS

Basiceros Schulz, 1906, Spolia Hymenopt., p. 156, nom. pro Ceratobasis F. Smith. Type:
Ceratobasis singularis = Meranoplus singularis F. Smith.

Basiceros : Brown and Kempf, 1960: 171; see for complete synonymy; nomenclature and
history on pp. 168-169.

Aspididris Weber, 1950: 3. Type: Aspididris militaris Weber, by original designation.

New synonym.

Aspididris : Brown and Kempf, 1960: 179.

The diagnosis of the genus has to be modified in part to include the characters of the two
Aspididris species.

Worker. Head trapezoidal, oblong or disc-like, the posterior and lateral borders separate
and either rounded or crested, or else combined into a curving, continuous or near-continuous
crest around the back of the cranium. Mandibles sub-porrect, triangular, with straight,
opposable, multidenticulate masticatory borders; blade narrowed before insertion, the re-
sulting peduncle either partly exposed or entirely hidden beneath clypeus, so that an interspace
between basal mandibular and anterior clypeal borders is present or absent in varying degrees.
Propodeal teeth lamelliform, more or less acute.

Malpighian tubules 5.

Queen : Like worker, but more robust and with developed pterothorax bearing wings in
virgins; ocelli present.

Male. (Generic description based on B. discigera, B. conjugans, and B. singularis ): Size a
little smaller than the conspecific queens and workers, and more slender. Head broadest
across the large, bulging eyes (which are situated at or a little in front of midlength) rather
suddenly narrowed in front of eyes and tapering moderately anteriad; median vertex and
ocelli prominent. Clypeus broad, its postero-median lobe convex and truncate or rounded,
extending about to level of frontal lobes; its anterolateral lobes concave, free margin with a
thin, sharp, yellowish edge, transverse or concave in front and rounded-divergent on sides.
Frontal area variably distinct, semicircular or transverse, more or less impressed; rugose or
carinate in the middle, and more or less distinctly delimited behind by an arched carina or
rugulae that tend to connect the two frontal lobes. Frontal lobes prominent and projecting
forward, laterad and dorsad, their free margins rounded sharply in front and broadly laterad,
antennal insertions on their ventral faces. Lateral bases of lobes continued laterad as sharply

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133

raised arching carinae running nearly to the eye on each side, then curving forward to bound
deeply excavated, subreniform antennal scrobes, which are bounded in front by the
cariniform posterior borders of the lateral wings of the clypeus. (Similar arrangements
are found in many Attini, but in these the scrobes are usually not so deep or so sharply
bounded.) Posterior vertex bordered along the cervical limit by a lamelliform margin
bearing short longitudinal costulae; space between this and posterior ocelli either steep or
gradual, depending on whether the head is much drawn out behind or not. A continuous or
nearly continuous, sharp but irregular, ventrolateral carina extends from posterior corner
of head to mandibular insertions, bordering a subrectangular piece of the cheek extending
between eye and mandibular insertion, and bounded mesad by the carinate outer scrobe
margin.

Mandibles subtriangular, with curved outer borders converging rapidly in the apical half,
meeting along the masticatory borders, and the sharp apices crossing; gently downcurved
and the dorsal faces gently convex. Masticatory borders serially 8- 12 -dentate. Mandibles
petiolate or not, with or without anteclypeal space, and form of labrum in general as in
conspecific workers.

Antennae long and slender, 13-merous. Scape very short, only about twice as broad as
long, its base oblique, with the more acutely rounded angle on the outside, and the obtuse
angle inside (mesal), tapered toward the truncate apex; a little thicker than the remaining
segments. First funicular segment (pedicel) only about half as long as scape; succeeding
segments all much longer than broad; apical segment longest, third antennal (funiculus II,
counting from base) also very long.

Trunk robust; prescutum with a more or less distinct anteromedian carina; notauli deep
and complete, the arms of the Y forming rows of deep punctures separated by the inter-
calated costulae. Parapsidal furrows in the form of fine shining lines; parapsides more or
less impressed behind, but each with a sharp, raised posterolateral margin. Prescutellum
separated from scutellum by an impression or transverse row of punctures, or else the
middle part impressed and not distinct from scutellum; lateral wings of prescutellum with
a laterally marginate, posteriorly pointed process or blunt hook on each side. Scutellum
narrower than prescutellum, forming an elongate near-semicircle as seen from above, free
borders marginate, but posteromedian portion concave; posterior aspect broadly Y- or
U-shaped. Metanotum narrow, with a blunt median tumosity. Propodeum with dorsal face
flat, rectangular, steeply sloping toward the rear, separated from rectangular declivitous face
by a transverse carina. As seen from the side dorsal and declivitous faces of propodeum
meeting at an obtuse angle; declivity marginate on each side.

Petiole clavate, with anterior peduncle and long, low rounded node, usually bent slightly
downward near base of posterior peduncle; spiracles papillose, prominent. Postpetiole
broader than long and a little broader behind than in front and broader than petiole ;
rounded above, sternum shallow; attached its full width behind to gaster, which is slightly
concave in front to receive it. Gaster with first segment occupying most (70 percent or
more) of its length; four visible apical segments subequal in length. Genital capsule slender;
parameres slightly broadened, bluntly rounded and curved mesad at apices, but tapered to a
blunt end as seen from the side ; volsellae sock-shaped, as usual in Myrmicinae ; pygidium
and subgenital segment unremarkable, with moderately narrowly rounded apical margins.

Legs slender, tibiae of middle and hind pairs without apical spurs; tarsal claws slender
and simple. Wings brownish, with opalescent bluish reflections (both sexes) and dense brown
microtrichiation. Forewing veined as in queen of Cr eight onidris (Brown and Kempf, 1960:
173, fig. 8) except that m-cu is usually present as a spur from M, or as a complete crossvein.
Hind wing with only two longitudinal veins issuing from the median cell (apical abscissae
of R and Cu), with the tip of Sc branching off from fused Sc -(- R (Rfl lacking) as in the

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Tranopelta male (Kusnezov, 1962: 371, fig. 23). Anal loop (A + cu-a) short, without a
spur of A, but with a break or weak place at a longitudinal fold line. Hamuli 5-9, sub-
median.

Sculpture very finely and densely punctulate, opaque or nearly so, including legs, mandibles
and antennae. Vertex with overlying loose rugulae, especially behind compound eyes and in
and around ocellar triangle; loose rugulation also on trunk, especially posterior half of
mesonotum and sides of propodeum. In some species, varying parts of mesopleura smooth
and shining, or rugulose.

Pilosity of fine tapered hairs, golden brown in color, mostly erect or suberect on body
(some also appressed on gaster and clypeus in some species) ; mandibles, antennae and legs
with hairs becoming shorter, more abundant and decumbent passing from base toward
apices of these appendages. Mesal face of antennal scape with two or more long fine hairs
and some shorter ones.

Color black; legs and antennae brown.

Contains six species as known at present: conjugans n. sp., convexiceps, discigera, manni,
militaris and singularis. Basiceros militaris is a new combination.

DISTRIBUTION AND BIOLOGY

Basiceros has been found only in wet tropical and subtropical forests of
Central and South America and Trinidad at low and moderate altitudes. All
of the colonies for which data are available have been found in rotten logs, or
at least in pieces of rotting wood of fairly substantial size. The adults usually
move very slowly, and they feign death for long periods when disturbed,
rivaling the attine Apterostigma in their ability to escape detection by this
means in the forest gloom.

Weber (1950: 6) noted that he had found a worker of B. singularis near
midday carrying a dead termite in Guyana, and I found headless termites in a
nest of this species in Mato Grosso (see below under B. singularis). Food of
the other species is unknown, but they are almost certainly predatory, perhaps
on termites, judging by the hardened incrustations that many workers bear.

Basiceros conjugans, n. sp.

Holotype worker. TL 5.8, HL 1.24, HW 1.05 (Cl 85), ML 0.43, greatest diameter of eye
0.13, scape L 0.82, WL 1.51 mm.

Paratype worker from type locality. TL 5.9, HL 1.29, HW 1.09 (Cl 84), ML 0.42, greatest
diameter of eye 0.14, scape L 0.84, WL 1.56 mm.

Paratype workers (two) from near Leticia, Colombia. TL 6.2, 6.1; HL 1.32, 1.33; HW 1.14,
1.12 (Cl 86, 84) ; ML 0.44, 0.45; greatest diameter of eye 0.14, 0.16; scape L 0.85, 0.87, WL
1.63, 1.63 mm.

Form of head and body well shown by Figs. 1 and 2. Sides of head bordered by a distinct
raised margin that continues around the posterior corners and across the back of the head
as a less distinct margin with a shallow dip in the middle. Cephalic disc shallowly concave
inside the lateral margins, convex in the middle, but the convexity itself with a shallow median
impression running back from about the level of the eyes. Clypeus gently convex in both
directions, with a feebly concave free margin. Mandibles with concave external borders
in full-face view, eleven strong teeth on each, triangular except for the basalmost tooth,

Vol. LXXXII, June, 1974

135

which is broad and rounded. Mandibular peduncles very short, mostly hidden under
clypeus, leaving a short space between clypeal margin and basal borders. Extensor margin of
scape with a broadly rounded, translucent, crenulate lobe around basal angle. Labrum
elongate, tapered cuneiform, terminating in paired acute contiguous lobes separated by a
narrow cleft. Palpi each consisting of two recognizable segments, which, however, are
solidly fused in both maxillary and labial palpi to make one long crooked unit of each
maxillary palpus, and one long, curved, clavate unit of each labial palpus. Cervical border
of head with a strong raised margin.

Promesonotum forming a subglobular mass tapering sharply behind to the metanotum-
propodeum, which is only a little more than half as wide seen from above. Promesonotal
suture very faintly indicated above; metanotal groove broad and deeply impressed, longitu-
dinally costate, succeeded posteriorly by a sloping, shelf-like propodeal dorsum that has a
sharply downsloping declivitous face continuing into the final declivity of the propodeum.
The declivity is bounded by carinae above and on each side; the transverse upper carina
connects a pair of acute triangular teeth, hidden from side view by thick squamiform hairs.
Bullae of metapleural glands prominent, projecting.

Petiolar node with a distinct anterior peduncle having a longitudinal carina on each side
of dorsal surface; node distinct, with steep anterior face, mesally emerginate anterodorsal

v?'

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Vol. LXXXII, June, 1974

137

border and posteriorly sloping rounded dorsum, the latter obscured by squamiform hairs,
but its disc (without posterior peduncle) longer than broad. Subpetiolar processes 5 (4-6
in paratypes, sometimes adjacent processes partly fused). Postpetiole rounded in both
directions, nearly twice as broad as petiole (pilosity excluded) and slightly broader than
long, with a light-colored translucent anterior margin ; attached behind its full width
to concave anterior part of gaster as seen from above. Gaster with just the barest suggestion
of a broad median longitudinal sulcus, visible only in certain lights. Sting retracted in
holotype, but in a Colombian paratype worker, it is extended and has a shaft nearly 0.6
mm long.

Integument (where free of secretion and dense pilosity) prevailingly smooth and shining,
with coarse punctures, becoming smaller and spaced out on clypeus, and still smaller on
mandibles. Punctures large and contiguous or subcontiguous in those areas bearing the dense
heavy pilosity: posterior vertex, mesonotum, antero-dorsal shelf of propodeum, lateral edges
of propodeal coxae, and both nodes of the waist. Propodeal declivity punctate-rugose, smoother
ventrad. Pleural plates of meso- and metathorax and propodeum nearly free of punctures
except along edges and sutural lines. Gastric segment I densely sown with closely spaced but
separate medium punctures, less crowded along lateral curves of tergum; interspaces smooth
and shining. Normally exposed tergal surfaces of terminal segments (abdominal V, VI, VII)
finely and densely punctulate, opaque, but margins of these segments smooth and shining.
Antennal scapes and legs smooth or with very fine superficial roughening, and coarse punctures
for the hairs ; in general shining ; funiculi and distal halves of tarsi finely and densely punctu-
late, opaque to subopaque.

The shapes and location of the various kinds of pilosity are well shown in Figs. 1 and 2.
The thicker squamiform and clavate hairs have a complicated microstructure. Under high
magnification, the surfaces of these hairs appear fluffy, with ribs of free fibers running
longitudinally, represented sometimes as fine lines in Fig. 2. A pair of erect clavate hairs on
the vertex is not so easily distinguished in the figures, since they are close to the posterior
borders of similar hairs on the vertex, but this pair straddles the ocellar triangle in the
queen and corresponds to a similar pair in B. discigera and B. militaris. The pilosity is off-
white, contrasting with the deep brownish-red (approaching mahogany) of the integument;
appendages medium brownish-red.

Queen (alate), one of eight alates and dealates from type nest series. TL 6.3, HL 1.32, HW
1.09 (Cl 83), ML 0.46, greatest diameter of compound eye 0.24, scape L 0.86, WL 1.68,
forewing L 4.9 mm.

Male, one of three from type nest series. TL 4.9, HL 0.92, HW across eyes 0.88 (Cl 96),
HW behind compound eyes 0.76, ML 0.25, greatest diameter of eye 0.30, scape L 0.14, WL
1.46, forewing L 3.8 mm.

Head of the short type, not produced behind, and with a narrow flange on cervical border.
Rugulae behind eye shorter and weaker than in B. discigera. Most of mesanepisternum and
upper middle part of mesokatepisternum smooth and shining. Petiole claviform, with node
indistinctly set off from anterior peduncle ; front of node bordered by a dorsolateral ruga
on each side; subpetiolar processes: 1 large anterior tooth, plus 1-4 smaller teeth or lamellae,
very inconstant.

Fig. 2. Basiceros conjugans, new species. Lateral view of holotype worker. Drawing by
Susan Poulakis, X40.

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Holotype and a few other workers, queens and males were taken together
at Limoncocha, Ecuador, by Dr. Henry R. Hermann, Sept, through Nov. 1964;
a separate winged queen was taken at the same locality by Dr. Hermann. The
paratype series also includes two workers taken in a rain forest leaf litter
berlesate 7 km north of Leticia, Colombia, by S. and J. Peck (B-230) in
February 1972.

This species is intermediate between Aspididris discigera and Basiceros manni,
the latter representing the “typical” members of its genus. B. conjugans has
partially developed ridges framing the posterior vertex in a manner intermediate
between discigera and manni , and the erect clavate hairs on the back of the
head are concentrated along the posterior edge in an intermediate kind of pat-
tern. The exact shape of the head, the distribution of the peculiar broadened
hairs on the trunk, and particularly their thick clustering on the petiole and
postpetiole are sufficient characters to separate B. conjugans from all the other
species.

Basiceros discigera

This species is widespread in southeastern Brazil, and I can extend the range
northward into Espirito Santo State: Reserva Nova Lombardia, 4 km north
of Santa Teresa, 900 m, 24 Feb. 1967 (W. L. Brown, Jr.). The nest was in a
small fragment of a rotten log on the floor of wet upland forest, and contained
two winged males.

A much greater and more surprising extension of the range is provided by
a record from the eastern slope of the Andes in Colombia: Quebrada Susamuko,
23 km NW of Villavicencio, Dept. Meta, 1000 m, two workers in leaf litter
berlesate (B-234), S. and J. Peck leg. The male is characterized in the key
at the end of the paper.

Basiceros manni

I took a number of workers of this species in a large fragment of a rotten log
found in the middle of a rain forest trail west of the bridge at Rio Toro Amarillo,
near Guapiles, Costa Rica. The log fragment contained also workers of Pro-
ceratium goliath. The record represents only a northern “fill-in” of the range
on the Atlantic Plain of Costa Rica; the species is known from Honduras and
probably occurs through the forested lowlands of Nicaragua.

Basiceros singtdaris

In addition to the records of this species from the Guianas, Trinidad and
Amazonas, Brazil, I collected it in the forest at the Fazenda Junqueira Vilela,
Mun. Diamantino, northern Mato Grosso State, Brazil, on July 17, 1973. The
nest was in a thoroughly rotten log in deep shade, and the headless bodies of
three termites were found with the workers, winged queens and males in what
appeared to be rude chambers. The adults simulate death for long periods when

Vol. LXXXII, June, 1974

139

disturbed and are exceedingly hard to distinguish by eye. Many are heavily
encrusted with a light brownish or whitish material, apparently a hardened
secretion. I take it that the material represents the hardened defensive allomones
of prey termite species (nasutes?), although it is not altogether impossible that
the secretion is produced by the ants themselves. Callow and near-callow
workers and winged forms of both sexes in the nest lack the incrustation.

Some workers confined in a glass-topped plaster nest avoided or showed
no apparent interest in live larvae of Tribolium and workers of Zootermopsis
termites, though the latter are much larger than the ants and the termites
found as apparent prey in the original nest. The ants did feed on crushed
housefly pupae, and two eggs that must have been laid by workers in the queen-
less group developed to half-grown larvae in the six months I maintained the
ants alive.

I dissected ten workers to determine how many Malpighian tubules were
present. Of these, eight had five long tubules each, and two had four tubules.
Evidently the counts of four represent specimens that lost a tubule during
dissection, which is difficult because of the thick integument and the small size
of the opening at the apex of the first gastric segment. At least some of the
tubules are attached to the rectum.

The male of this species is characterized in the key to that sex below. In
addition to the characters cited, the sides of the metanotum-propodeum and
the dorsal surface of the scutellum are more heavily rugose than in the other
two species keyed, and the body size is larger.

Basiceros — Revised Key to Workers and Queens

1. Posterior half or more of head disc-like, subcircular in outline, the margins forming

a strong, continuous or nearly continuous raised crest 2

Posterior half of head trapezoidal or subrectangular, not disc-like, the lateral borders
of the vertex distinct from the posterior border, and not forming a continuous
semicircular crest 3

2. When head is viewed full-face, the arcuate crest or flange around the back of the

vertex is medially emarginate and confluent at this point with the median convexity

of the vertex (SE Brazil, subandean Colombia) discigera

Arcuate crest around back of vertex continuous and entire, and separated from the
median convexity of the vertex by a broad, uninterrupted sulcus that follows the
crest (Trinidad) militaris

3. Labrum a shield-shaped piece with rounded free margin, not divided medially, at

least on its dorsal (extensor) face 4

Labrum narrow, cuneiform, tapered apicad and with a distinct median division or
groove 5

4. Head narrow (Cl < 75) and nearly parallel-sided; clypeus and mandibles with

abundant and conspicuous appressed squamiform hairs; petiole with 1-3 ventral
processes, and usually at most 1 of these is well-developed and spiniform; base of
first gastric sternite with a short but sharp, angulate longitudinal carina (Trinidad
to N. Mato Grosso) singularis

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New York Entomological Society

Head wider (Cl > 75) and more triangular; clypeus and mandibles with punctures,
but no appressed hairs; petiole with 4-7 ventral processes, usually all or nearly all
slender spiniform ; base of first gastric sternite without a sharp longitudinal carina
(Central America) manni

5. Posterior dorsal half of head (vertex) continuously convex except for median sulcus;
head wide, worker Cl > 90; petiole and postpetiole with scanty pilosity, not hiding

sculpture; 1 subpetiolar process (SE Brazil) convexiceps

Vertex with raised lateral margins and a median sulcate tumosity; Cl < 90; petiole
and postpetiole covered densely with fat squamiform hairs that conceal the surface
beneath; 4-6 subpetiolar processes (W. Amazon Basin, Figs. 1, 2) conjugans

Basiceros — Key to Males of Three Species

1. Viewed full-face, head with a broad drawn-out neck longer than space occupied by

ocellar triangle ; mesokatepisternum opaque, covered by strong interlocking rugae ;

petiole about 3X as long as postpetiole (Trinidad to N. Mato Grosso) singularis

( B . manni from Central America would probably key out here, though the head and
petiole may be somewhat shorter than in B. singularis.)

Viewed full-face, head not produced behind, though with a flange along the cervical
margin that is much shorter than the ocellar triangle ; mesokatepisternum finely
punctate, sometimes with upper part smooth and more or less shining; petiole
about twice as long as postpetiole 2

2. Anterior border of clypeus concave in the middle; more than half of mesanepisternum

smooth, and even the punctate part strongly shining; upper middle part of meso-
katepisternum smooth and shining (W. Amazon Basin) conjugans

Anterior border of clypeus entire; only the anterior half of anepisternum smooth and
shining, remainder densely punctate and nearly opaque ; mesokatepisternum densely
punctate throughout, only weakly shining in upper middle part between punctures

(SE Brazil, subandean Colombia) discigera

( B . militaris from Trinidad, possibly occurring also on the mainland, and B. con-
vexiceps from SE Brazil probably key to couplet 2, but I have seen no male speci-
mens.)

Literature Cited

Brown, W. L., Jr. and Kempf, W. W. 1960. A world revision of the ant tribe Basicerotini.
Studia Entomol. (n.s.), 3: 161-250.

Kusnezov, N. 1962. El ala posterior de las hormigas. Acta Zool. Lilloana, 18: 367-378.
Weber, N. A. 1950. New Trinidad Myrmicinae, with a note on Basiceros Schulz (Hy-
menoptera, Formicidae). Amer. Mus. Novitates, 1465: 1-6.

Vol. LXXXII, June, 1974

141

Myrmicine Trail Pheromones: Specificity, Source and Significance

Murray S. Blum

Department of Entomology, University of Georgia, Athens, Georgia 30602

Received for Publication December 22, 1973

Abstract: The poison gland secretion is the source of the trail pheromones in the

myrmicine genera Myrmica, Manica, Pogonomyrmex, and Veromessor. Transposition studies
demonstrate that poison gland products of Myrmica, Manica, and Pogonomyrmex species
lack intra- and intergeneric specificity. The unpredictable lack of trail specificity identified
with myrmicine venoms is discussed in terms of common trace natural products which may
be utilized as trail pheromones by species in unrelated taxa. The persistence of chemical trails
is discussed as a function of the foraging strategies employed by myrmicine species.

INTRODUCTION

The sources of trail pheromones in the Formicidae are quite protean, especially
in the large subfamily Myrmicinae. Releasers of trail-following behavior have
been localized in the poison gland (Moser and Blum, 1963), Dufour’s gland
(Wilson, 1959) and metathoracic tibial glands (Fletcher and Brand, 1968) of
a wide range of myrmicine species, which clearly emphasizes the polyphyletic
origins of trail following in this subfamily. Indeed, with the exception of some
ponerine species which generate trails with poison gland secretions (Fletcher,
1971), species in formicid subfamilies other than the Myrmicinae are not known
to utilize the above named organs for producing these chemical releasers.

The variability in the glandular sources of trail following in the Myrmicinae
is exceeded by the variability in specificity of the products synthesized in these
social organs. In some cases constituents in the poison gland secretions release
trail following in different species in the same genus (Blum, 1966), but in other
cases these secretions may be completely species specific when evaluated among
members of one genus (Blum and Ross, 1965). Furthermore, the natural
product complex in the venoms of some myrmicines can release strong trail fol-
lowing in species in completely unrelated genera when assayed by an artificial
trail technique (Blum and Ross, 1965; Blum and Portocarrero, 1966). On the
other hand, it has not been ascertained whether these singular examples of non-
specificity reflect the utilization of the same pheromone by unrelated species, or
whether different poison gland secretions are enriched with common con-
stituents, some of which may serve as trail pheromones for unrelated species.

Acknowledgments: I am very grateful to P. B. Kannowski, G. L. Ayre, E. O. Wilson, G.

Scherba, H. Spangler, and R. R. Snelling for providing many of the species used in this
investigation. Special thanks go to P. B. Kannowski for providing facilities at the University
of North Dakota where some of this research was undertaken. The technical assistance of
G. N. Ross is gratefully acknowledged.

New York Entomological Society, LXXXII: 141-147. June, 1974.

142

New York Entomological Society

Table 1. Percentage of Myrmica workers responding to artificial trails prepared from

poison gland extracts

^mirrp

Test Species

UvUi CC

species

americana

brevinodis

brevispinosa emeryana

fracticornis

monticola

rubra

americana

brevinodis

brevispinosa

90 65

emeryana

80 95

fracticornis

—

monticola

rubra

The present investigation was undertaken in order to determine the source
and specificity of the trail pheromones of myrmicine species in a wide range
of genera. The results again clearly emphasize that these pheromonally-rich
secretions have a remarkable lack of intra- or intergeneric specificity.

METHODS

Workers of the following myrmicine species were utilized for studies of either
the source or specificity of trail pheromones: Aphaenogaster fulva Roger,

Myrmica americana Weber, M. brevinodis Emery, M. brevispinosa Wheeler, M.
emeryana Forel, M. monticola Wheeler, M. fracticornis Emery, M. rubra (L.),
Manica bradleyi (Wheeler), M. hunteri (Wheeler), M. mutica (Emery), Pogono-
myrmex badius (Latreille), P. barbatus (F. Smith), Novomessor cockerelli (E.
Andre), V eromessor pergandei (Mayr), Chelaner antarcticum (Wheeler),
Pheidole dentata Mayr, Crematogaster lineolata (Say), Monomorium minimum
(Buckley), Solenopsis invicta Buren, and Trachymyrmex septentrionalis
(McCook).

The presence of trail pheromones was examined by preparing methylene
chloride extracts of poison glands, Dufour’s glands, and hind guts. Four organs
were crushed in 2 ml of solvent and 0.2 ml of this extract was applied to a
circular trail 15 cm in diameter. Groups of ten workers were subsequently
introduced into the center of the circle and if a worker traveled around the entire
circumference after encountering it, a positive response was recorded (Moser
and Blum, 1963).

Specificity studies were undertaken by using the same technique. Six repli-
cates, consisting of ten workers each, were employed for each test species.

RESULTS

No evidence of trail following could be demonstrated when workers of
Aphaenogaster fulva and Novomessor cockerelli were exposed to circular trails
treated with extracts of their own sting-associated glands or hind guts. On the

Vol. LXXXII, June, 1974

143

Table 2. Percentage of myrmicine workers responding to artificial trails prepared from

poison gland extracts

Test Species

Source

species

Myrmica

americana

Myrmica

brevinodis

Manica

bradleyi

Manica

hunteri

Manica

mutica

Pogono-

myrmex

badius

A. fulva

—

Manica

bradleyi

M. hunteri

—

M . mutica

—

Myrmica

americana

M. brevinodis

—

M. brevispinosa

—

M. emeryana

—

M. fracticornis

—

M. monticola

—

M . rubra

N . cockerelli

—

P. badius

—

P. barbatus

—

V. pergandei

—

other hand, workers of Veromessor pergandei, Pogonomyrmex badius, Manica
spp., and six of the seven Myrmica spp. readily followed artificial trails gen-
erated with extracts of their own poison glands. None of the species responded
to extracts of either their Dufour’s glands or their hind guts.

The results of the specificity studies are presented in Tables 1 and 2. Table 1
presents the results of intrageneric studies utilizing the seven Myrmica spp.
Table 2 illustrates the responses of two Myrmica spp., three Manica spp., and
P. badius workers to artificial trails treated with poison gland extracts derived
from 15 myrmicine species in six genera.

With the exception of M. monticola, Myrmica spp. were almost equally
sensitive to trail extracts of each other’s poison glands (Table 1). M. monticola
workers showed almost no propensity to follow their own poison gland extracts,
whereas the other Myrmica spp. readily followed trails prepared from glandular
extracts of this species. Myrmica workers readily followed these artificial trails
and often circled them repeatedly. Trails prepared from M. americana glandular
extracts were strongly active 1 hr after their preparation but were only weakly
active after 3 hrs. Males and females of M. emeryana followed artificial trails
as faithfully as workers of this species.

Poison gland extracts of the three Manica spp. did not appear to possess any
specificity for the species in this genus (Table 2). The Manica spp. also
followed artificial trails prepared from Myrmica poison glands as effectively

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New York Entomological Society

as they did their own, and two Myrmica spp. were equally responsive to extracts
of Manica poison glands. As in the case of the Myrmica spp., Manica workers
exhibited sustained trail following when exposed to these artificial trails. Fur-
thermore, Myrmica americana and M. brevinodis readily followed poison gland
extracts prepared from Pogonomyrmex badius and P. barbatus workers. Simi-
larly, workers of Manica bradleyi exhibited trail following when exposed to trails
containing extracts of these two species of Pogonomyrmex (Table 2). On the
other hand, neither Myrmica nor Manica workers responded positively when
bioassayed with poison gland extracts prepared from Aphaenogaster julva,
N ovomessor cocker elli, and V eromessor pergandei.

Workers of V. pergandei were somewhat responsive (40 percent) to their own
poison gland extracts but they seldom adhered to the circular trail for more
than one complete traversal of its circumference. Similarly, workers of P. badius,
which showed moderate trail-following activity in the presence of their own
poison gland extracts as well as those derived from Manica bradleyi and
Myrmica brevinodis (Table 2), exhibited very ephemeral trail following.

Workers of T. septentrionalis, M. minimum, C. antarcticum, P. dentata, S.
invicta, and C. lineolata did not react to poison gland extracts of M. brevinodis .

DISCUSSION

At this juncture, the poison gland appears to be the primary source of odor
trail pheromones in the Myrmicinae. The presence of chemical releasers of trail
following in the venoms of Myrmica, Manica, and Pogonomyrmex species brings
to 12 the number of myrmicine genera in which the trail pheromone has been
localized in the poison gland secretion. Furthermore, since pheromonally-rich
venoms have been identified in both primitive ( Myrmica ) and highly advanced
( Atta ) myrmicine genera, it seems clear that the utilization of the poison gland
as a social organ is widespread in the Myrmicinae. On the other hand, Dufour’s
gland has been demonstrated to be the source of the trail pheromone in only two
myrmicine genera, Solenopsis and Pheidole (Wilson, 1959, 1963). The utiliza-
tion of the metathoracic tibial glands to generate chemical trails appears to be
limited to the genus Crematogaster (Fletcher and Brand, 1968; Leuthold,
1968) and probably reflects an evolutionary specialization correlated with the
apparent unsuitability of the gaster to function in trail laying.

Since the poison gland secretions of Myrmica and Manica species are almost
totally lacking in specificity, it would appear that they may be utilizing the same
or very similar trail pheromones, which would be consistent with the close rela-
tionship of these two myrmicine genera (Creighton, 1950). On the other hand,
the genus Pogonomyrmex is certainly not closely related to Myrmica and
Manica and the ability of workers in the latter two genera to follow artificial
trails prepared from Pogonomyrmex poison gland extracts, and vice versa, may
simply indicate that these three genera share common natural products in their

Vol. LXXXII, June, 1974

145

venoms. Indeed, other investigations demonstrate that phylgoenetic relation-
ships are of little value in predicting the trail specificity of myrmicine poison
gland secretions.

The poison gland secretion of the primitive ant Daceton armigerum (Latreille),
a non-trail-laying myrmicine, releases strong trail following in attine species
in three genera (Blum and Portocarrero, 1966). Similarly, the poison gland
secretion of Tetramorium guineense (F.) is active as a trail pheromone for
workers in two attine genera and vice versa. However, it is impossible to
generalize about the specificity of the poison gland secretions of Tetramorium
and the attines, since that of T. caespitum (L.) is not followed by workers of
T. guineense or attine workers (Blum and Ross, 1965). Significantly, in trans-
position studies with poison gland secretions of Monomorium species, it has
been established that the venom of one species releases strong trail following
in another species but not vice versa (Blum, 1966). Thus, M. minimum workers
will follow artificial trails prepared with M. pharaonis venom extracts as well
as their own, but workers of M. pharaonis will not follow artificial M. minimum
trails. Presumably, the venom of M. pharaonis contains the trail pheromone of
M. minimum , but the latter species does not produce the trail pheromone of
M. pharaonis in its poison gland secretion.

The trail pheromones derived from myrmicine venoms are certainly trace
constituents which are not identified with the proteinaceous compounds which
appear to dominate most of these secretions. The venoms of the Manica,
Myrmica, and Pogonomyrmex species examined in this investigation are rich
in proteins which are obviously not soluble in the solvent utilized to prepare
active trail extracts. Tumlinson et al. (1971) have identified the major trail
pheromone in the poison gland secretion of Atta texana (Buckley), methyl 4-
methylpyrrole-2-carboxylate, as a trace constituent of a proteinaceous venom.
Each ant is estimated to contain about 0.6 ng of this compound but four addi-
tional fractions are active in releasing trail following (Tumlinson et al., 1972).
Possibly, these other trail pheromones may confer a degree of specificity which
is unattainable with a single compound. It has also been suggested that the
products of the Dufour’s gland may be secreted in admixture with the poison
gland secretion in order to obtain a more specific trail pheromone.
Holldobler and Wilson (1970) have demonstrated that Pogonomyrmex badius
workers lay recruitment trails with the poison gland secretion, whereas the
Dufour’s gland products are utilized to set orientation marks. A combination
of these two glandular exudates may produce a trail of much greater species
specificity than could be obtained with either secretion alone.

Field studies with M. brevispinosa have demonstrated that trails are never
laid by a foraging worker that is capable of bringing a food find back to the
nest. Similarly, Eidmann (1927) noted that a worker of M. rubra did not lay
a trail unless it was unable to transport the food to the nest by itself. Sig-

146

New York Entomological Society

nificantly, Myrmica trails appear to be short lived both in the laboratory and
in the field. In nature, the persistence of trails laid by myrmicine species varies
greatly, which may be correlated with the recruitment strategies of the species
as well as the vapor pressures of the trail pheromones. Thus, Ayre (1969) has
demonstrated that workers of M. americana, after establishing a well-developed
trail to a honey solution, quickly switch to topographical landmarks as a means
of orienting between this food source and the nest. Since the trail is no longer
being reinforced, it rapidly dissipates. Therefore, the relatively volatile trail
pheromones which are characteristic of Myrmica species may be ideally suited
to the foraging strategies employed by species in this genus. On the other
hand, it would be selectively advantageous for those myrmicine species that do
utilize the same trails daily to secrete trail pheromones which possess a rela-
tively low vapor pressure. Indeed, artificial trails prepared from the poison
gland secretion of Atta texana are highly active one week after their prepara-
tion (Blum et al., 1964). Members of this genus lay some of the most durable
trails encountered in the Formicidae.

Among the species of Myrmica, M. monticola appears to be especially
aberrant since it does not readily follow trails prepared from its own poison
gland secretion or those of other Myrmica species. This species is also unusual
because its mandibular gland chemistry is radically different from that of other
Myrmica species which have been examined (Crewe and Blum, 1970). How-
ever, the singularity of M. monticola should serve to emphasize the variability
in either behavior or natural product chemistry that may be encountered within
the species in a genus. Ultimately, M. monticola may be demonstrated to be
typical of many species which, because they lack some significant generic
characters, are especially important as key indicators in the evolution of the
genus.

Literature Cited

Ayre, G. L. 1969. Comparative studies on the behavior of three species of ants (Hy-
menoptera: Formicidae). II. Trail formation and group foraging. Can. Ent., 101:
118-128.

Blum, M. S. 1966. The source and specificity of trail pheromones in Termitopone,
Monomorium and Huberia and their relation to those of some other ants. Proc. Roy.
Ent. Soc. Lond. (A), 41: 155-60.

Blum, M. S., Moser, J. C., and Cordero, A. D. 1964. Chemical releasers of social be-
havior. II. Source and specificity of the odor trail substances in four attine genera
(Hymenoptera: Formicidae). Psyche, 71: 1-7.

Blum, M. S. and Portocarrero, C. A. 1966. Chemical releasers of social behavior. X. An
attine trail substance in the venom of a non-trail laying myrmicine, Daceton armigerum
(Latreille). Psyche, 73: 150-155.

Blum, M. S. and Ross, G. N. 1965. Chemical releasers of social behaviour. V. Source,
specificity, and properties of the odour trail pheromone of Tetramorium guineense
(F.). J. Insect Physiol., 11: 857-868.

Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. Harvard,
104: 1-585.

Vol. LXXXII, June, 1974

147

Crewe, R. M. and Blum, M. S. 1970. Alarm pheromones in the genus Myrmica (Hy-
menoptera: Formicidae). Z. Vergl. Physiol., 70: 363-373.

Eidmann, H. 1927. Die Sprache der Ameisen. Rev. Zool. Russe, 7: 39-47.

Fletcher, D. J. C. 1971. The glandular source and social functions of trail pheromones in
two species of ants (Leptogenys) . J. Ent. (A), 46: 27-37.

Fletcher, D. J. C. and Brand, J. M. 1968. Source of the trail pheromone and method
of trail laying in the ant Crematogaster peringueyi. J. Insect Physiol., 14: 783-788.

Holldobler, B. and Wilson, E. O. 1970. Recruitment trails in the harvester ant
Pogonomyrmex badius. Psyche, 77 : 385-399.

Leuthold, R. H. 1968. A tibial gland scent-trail and trail-laying behavior in the ant
Crematogaster ashmeadi Mayr. Psyche, 75: 233-248.

Moser, J. C. and Blum, M. S. 1963. Trail marking substance of the Texas leaf-cutting
ant: source and potency. Science, 140: 1228.

Tumlinson, J. H., Moser, J. C., Silverstein, R. M., Brownlee, R. G., and Ruth, J. M.
1972. A volatile trail pheromone of the leaf-cutting ant Atta texana. J. Insect
Physiol., 18: 809-814.

Tumlinson, J. H., Silverstein, R. M., Moser, J. C., Brownlee, R. G., and Ruth, J. M.
1971. Identification of the trail pheromone of a leaf-cutting ant, Atta texana. Nature,
234: 348-349.

Wilson, E. O. 1959. Source and possible nature of the odor trail of fire ants. Science,
129: 643-644.

Wilson, E. O. 1963. The soil biology of ants. Ann. Rev. Ent., 8: 345-368.

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The New York Entomological Society

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Incorporating The Brooklyn Entomological Society

Incorporated May 21, 1968

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The New York Entomological Society

Organized June 29, 1892 — Incorporated February 25, 1893

Reincorporated February 17, 1943

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The Brooklyn Entomological Society

Founded in 1872 — Incorporated in 1885
Reincorporated February 10, 1936

The meetings of the Society are held on the first and third Tuesday of each month (except

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Officers for the Year 1974

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President, Dr. Daniel J. Sullivan, S.J.

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Vice-President, Dr. Peter Moller

Fordham’ University, New York 10458

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Secretary, Dr. Chables C. Porter

American Museum of Natural History, New York 10024

Fordham University, New York 10458

Assistant Secretary i Dr. Louis Trombetta

Pelham Manor, New York 10803

Treasurer, Dr. Winifred B. Trakimas

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State University of New York, Farmingdale, New York 11735

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Assistant Treasurer, Ms. Joan DeWind

American Museum of Natural History, New York 10024

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Trustees

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Class of 1974

Dr. Lee Herman
Class of 1975

Dr. Howard Topoff

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Mf. Edwin Way Teale

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Dr. Pedro Wygodzinsky

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Mailed November 29, 1974

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The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press
Inc., 1041 New Hampshire, Lawrence, Kansas 66044. Second class postage paid at New Brunswick, New
Jersey and at additional mailing office.

Known office of publication: Waksman Institute of Microbiology, New Brunswick, New Jersey 08903.

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Journal of the

New York Entomological Society

Volume LXXXII September 1974

No. 3

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Waksman Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lawrence E. Limpel
Helen McCarthy

Publication Committee

Dr. Kumar Krishna Dr. Ayodha P. Gupta

Dr. James Forbes, Chairman

CONTENTS

The Anthomyiidae and Muscidae of the Great Smoky Mountains and Mt.
Mitchell, North Carolina (Diptera) H. C. Huckett ISO

Notes on the Life Cycle and Natural History of Butterflies of El Salvador.

V. A. Pyrrhogyra hypsenor (Nymphalidae-Catonephelinae)

Alberto Muyshondt 163

Tenuicoris myrme forme : A New Genus and Species of Myodochini (Hemip-

tera: Lygaeidae) James A. Slater and Jane E. Harrington 173

A New Genus and Two New Species of Achipteriidae from New York State

(Acari: Cryptostigmata: Oribatei) F. Reese Nevin 177

Two New Tabanidae from Southeastern United States (Diptera)

L. L. Pechuman 183

The Distribution of Brood Ten of the Periodical Cicadas in New Jersey in

1970 John B. Schmitt 189

Terrestrial Mites of New York (Acarina: Prostigmata), I — Tarsocheylidae,

Paratydeidae, and Pseudocheylidae

Mercedes D. Delfinado and Edward W. Baker 202

Book Reviews 188, 201, 212, 213

Announcement 211

Proceedings of the New York Entomological Society 214

New York Entomological Society Guest Speakers, 1974/1975 219

XV International Congress of Entomology 220

150

New York Entomological Society

The Anthomyiidae and Muscidae of the Great Smoky Mountains
and Mt. Mitchell, North Carolina (Diptera)

H. C. Huckett

Long Island Vegetable Research Farm, Cornell University
Riverhead, New York 11901

Received for Publication September 25, 1973

Abstract: A preliminary listing is presented of the anthomyiid and muscid flies occurring
in the Great Smoky Mountains National Park and in the vicinity of Mt. Mitchell. The
data are based on collections made in the vicinities of Clingmans Dome and Mt. Mitchell in
1957 by members of the Entomology Research Institute at Ottawa, by the author when
visiting Mt. Le Conte in 1958 and 1959 and Mt. Mitchell in 1960 and 1961 and also on an
examination of additional material from these regions. Fifty-seven species of Anthomyiidae
and one hundred and five species of Muscidae were recognized, of which fourteen and twenty-
eight respectively were regarded as occurring chiefly within the eastern and midwestern areas
of the United States and transition zones of the Canadian provinces; five anthomyiid and
eight muscid species as boreal in habitat; two species of Anthomyiidae, namely, Chirosia
delicata (Huckett), Pegomya atlanis Huckett, and one species of Muscidae, Phaonia aberrans
Malloch, as restricted in their occurrence to the middle Atlantic states.

INTRODUCTION

The following work was undertaken as a means of bringing together the little
known records of anthomyiid and muscid flies captured within the boundaries
of the Great Smoky Mountains National Park and in the region of Mt.
Mitchell, and as a contribution in a series of studies toward a wider knowledge
of their presence and distribution on the higher slopes of the Appalachian
Range.1

The Great Smoky Mountains National Park lies within the southern limits
of the Appalachian Range, extending from slightly west of longitude 83 °W
to longitude 84°W, and varying irregularly in width between the lower parallels
of latitude 35°N. It is approximately 54 miles long, and at its widest point,
19 miles. The park extends over nearly 500,000 acres, or about 780 square
miles, and contains 16 forested peaks over 6,000 feet in altitude. To the
east, about 45 miles, lies Mt. Mitchell in the Black Mountains, where the
terrain was sadly denuded of timber during the earlier decades of the century.

PREVIOUS RECORDS

The present literature records of anthomyiid and muscid flies are few,
scattered, and sometimes of an ambiguous nature. Brimley (1938) in his list
of the insects of North Carolina refers to four species of Anthomyiidae from

huckett, H. C. 1972. The Anthomyiidae and Muscidae of Mt. Katahdin, Maine
(Diptera). Journal New York Entomological Society, 80 (4): 216-233, 1 map.

New York Entomological Society, LXXXII: 150-162. September, 1974.

Vol. LXXXII, September, 1974

151

the “mountain region/’ and four species of Muscidae as “Statewide.” Later,
under North Carolina, Wray (1967) in the third supplement to the above
work, added 19 nominal species to the list of Anthomyiidae and 22 to the
Muscidae, adopting the citations given for “N. C.” by Stone et al. (1965)
in “A Catalog of the Diptera of America North of Mexico.”

Whittaker (1952), in a study of the summer foliage insect communities
in the park, recorded the presence of 27 nominal species of Muscidae sens. lat.
The labels on these specimens bear the name Gatlinburg, a town outside the
boundaries of the park and contain much detailed information without men-
tioning the locality as given on maps of the park. Without actual reference
to his paper it would be difficult to determine where specimens were collected.
I have, therefore, with the cooperation of Dr. Whittaker, attempted to convert
the topographical data on these labels to names of localities and trails indicated
on official maps of the park.

Chillcott (1961) recorded 10 species of Fanniinae from locations in the
park, although unwittingly the name Gatlinburg is cited in several instances.
Three species are recorded from Mt. Mitchell.

MATERIALS

The survey is based on the following material that I have had the privilege
of examining: the collections of Iowa State University, containing specimens
taken by Dr. R. H. Whittaker in June and July 1947, and by the late Pro-
fessor J. L. Laffoon in the same months of 1958, to whom I am greatly
indebted for the extended loan of this and additional pertinent contributions
from neighboring counties in the Piedmont region; specimens from the col-
lections of the late Dr. R. R. Dreisbach, taken on a visit to the park in June
1946, and in August 1947, in company with Mr. D. S. Bullock; the small
collection of anthomyiid and muscid flies preserved in the museum at park head-
quarters, containing the only records on hand of specimens from Mt. Buckley,
taken by Dr. K. D. Snyder on July 21 and August 2, 1957, a loan made
available through the courtesy of Mr. Arthur Stupka, Park Naturalist. Also
there are my own collections during a brief visit to the park in May 1957
and to Mt. Le Conte during the latter half of May 1958 and 1959. I stayed
at the lodge, situated at an altitude of 6,200 feet, and thus was able to work
conveniently on the various trails traversing the extensive slopes of the
mountain mass (see Map 1). From mid May to early June 1960 and 1961,
I visited Mt. Mitchell, basing my operations from the inn at Stepps Gap,
at an altitude of about 6,000 feet (see Map 2) .

I have recently received a large assemblage of specimens from the Canada
National Collection collected in the park during the summer of 1957 by various
members of the Entomology Research Institute at Ottawa. The collections were
made on numerous occasions from May to August, the bulk of the specimens

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New York Entomological Society

being taken by Dr. J. R. Vockeroth and the late Dr. J. G. Chillcott. The latter
extended his visit to include a brief trip in mid August to Mt. Mitchell and
Roan Mountain, 6,200 ft., in North Carolina, and to a return to Clingmans
Dome on June 3, 1962, and May 22, 1965.

The collections in the park were chiefly made at the higher altitudes, between
Indian Gap at 5,100 feet to the summit of Clingmans Dome at 6,642 feet.
A few captures were made at lower levels from 2,000 to 2,100 feet at Greenbrier
Cove, Mingus Creek and Cherokee. To all these collectors I am deeply indebted,
and in particular to Dr. J. R. Vockeroth for his cooperation in having the speci-
mens assembled and thus made available for study. Their names may be
found among the list of collectors at the end of this article.

ABBREVIATIONS

In order to save space, the various locations in the Great Smoky Mountains
National Park from which specimens were obtained have been assigned a letter,
such as A, B, or C, and also for each species the numbers of specimens from
all localities have been combined. Similarly, for each species the numbers of
specimens captured on the various trails and logging roads in the Mt. Mitchell
region have been combined and the number of locations reduced to the naming
of three main collecting areas, as Mt. Mitchell, Clingman’s Peak, Mitchell
Falls, Roan Mountain in Mitchell County.

A. Park Headquarters, 1,500 ft.

B. Mt. Le Conte, upper trails, see map

C. Mt. Le Conte, lower trails2

D. Greenbrier Cove and Porters Creek, 2,000-3,000 ft.

E. Ramsey Cascade trail

F. Brushy Mountain, 4,911 ft. and Mountain trail

G. The Chimneys and camp

H. Newfound Gap, 5,048 ft.

I. Indian Gap to Clingmans Dome, 5,200 to 6,600 ft.

J. Indian Gap, 5,200 ft.

K. Mt. Collins, 5,900 ft. and Collins Gap, 5,700 ft.

L. Clingmans Dome, upper trail, 6,300 to 6,642 ft.

M. Forney Ridge trail to Andrews Bald

N. Mt. Buckley, summit, 6,100 ft.

O. Sailers Bald, summit, 5,620 ft.

P. Narrows, nr. Sailers Bald, 5,400 ft.

Q. Elkmont

R. Fighting Creek

S. Cades Cove

T. Spence Field trail camp, nr. Thunderhead Mtn.

U. Mingus Creek and Cherokee, 2,000 to 2,100 ft.

V. Smokemont

W. Mt. Sterling, summit, 5,842 ft.

Bullhead, Rainbow Falls, Cherokee Orchard, Trilium Gap, Alum Cave.

Vol. LXXXII, September, 1974

153

Sketch Map 1. The upper trails of Mt. Le Conte from the Lodge: a. Main Top, b. Cliff
Top, c. Myrtle Point, d. Boulevard, e. Alum Cave, f. Trilium Gap, g. Bullhead, h. Cherokee
Orchard, i. Lodge, stables, and campground. Distances from lodge to Cliff Top are one-third
mile; to Main Top, one-quarter mile; to Myrtle Point, two-thirds mile (by letter, Arthur
Stupka, Park Naturalist) .

Sketch Map 2. Trails and logging roads at Mt. Mitchell and Clingman’s Peak. Distances
by trail from summit of Mt. Mitchell to Stepps Gap are 2% miles; to Mt. Craig, 1% miles; to
Mitchell Falls, 2 miles; to Camp Alice, 2 miles (North Carolina Department of Natural and
Economic Resources) ; to Clingman’s Peak, 3% miles.

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New York Entomological Society

LIST OF SPECIES AND LOCALITY RECORDS
Family Anthomyiidae sens. str.

Chirosia delicata (Huckett)

94 $, 309 2. B, K, L. Mt. Mitchell, 73 $,31 2 ; Clingman’s Peak, 61 $, 8 2.

Chirosia hystrix (Brischke)

2 2. L, P.

Chirosia pusillans (Huckett) N. COMB.

2 $, 8 2. P.

Chirosia stratifrons (Huckett)

21 $, 13 2. Ba, d, g, I, J, K, L, M. Mt. Mitchell, 1 $.

Hylemya alcathoe (Walker)

38 $,28 2. A, Bb, d, g, C, D, F, G, I-L, T. Mt. Mitchell, 1 $ , 15 2 ; Roan Mtn. 1 2.
Hylemya latifrons (Schnabl)

1 $ , 12. Bg, L. Mt. Mitchell, 1 2 .

Hylemyza partita (Meigen)

1 $ . Clingman’s Peak.

Delia antiqua (Meigen)

1 2 . Mt. Mitchell.

Delia arnolitra (Huckett)

5 $,26 2. Ba-i. I, L. Mt. Mitchell, 20 $ , 34 2 ; Clingman’s Peak, 12 $ , 14 2 .

Delia echinata (Seguy)

1 $ , 2 2 . Ba, e, g. Mt. Mitchell, 10 $ , 3 2 ; Clingman’s Peak, 3 $ , 1 2 .

Delia inconspicua (Huckett)

2 $ . Mitchell Falls.

Delia laevis (Stein)

1 $.A.

Delia platura (Meigen)

532 $, 365 2. A, Ba-i, C, F, G, I-P, T. Mt. Mitchell, 116 $, 108 2 ; Clingman’s Peak,
44 $, 26 2 ; Mitchell Falls, 1 $ ; Roan Mtn. 2 2.

Delia winnemana (Malloch)

3 $ . Bg.

Botanophila inornata (Stein)

8 $ , 2 2 . A, Bd, g, J, O, P. Mt. Mitchell, 2 $ .

Paregle cinerella (Fallen)

1 $, 15 2. F, J, N, O. Mt. Mitchell, 1 2 ; Roan Mtn. 1 2.

Paregle radicum (Linnaeus)

2 2 . L. Roan Mtn. 1 $ .

Lasiomma abietis (Huckett)

1 $. Mitchell Falls.

Lasiomma anthracinum (Czerny)

8 $ , 1 2 . Be, g. Mt. Mitchell, 1 $ .

Lasiomma octoguttatum (Zetterstedt)

12 $, 29 2. Bb, c, f, g, h, I, L. Mt. Mitchell, 3 $ ; Roan Mtn. 1 $.

Pegohylemyia fugax (Meigen)

4 $ , 6 2 . Bb, d, f, g, L. Mt. Mitchell, 1 $ , 1 2 .

Pegohylemyia trivittata (Stein)

1 $, 1 2. C, J.

Acrostilpna atricauda (Zetterstedt)

5 $, 2 2. F, J, K.

Vol. LXXXII, September, 1974

155

Acrostilpna latipennis (Zetterstedt)

1 5- J.

Alliopsis species
3 2 . Mt. Mitchell.

Eremomyia pilimana (Ringdahl)

16 3, 11 2. A, B, C. Mt. Mitchell, 3 5 , 16 $ ; Clingman’s Peak, 15,5 $ ; Mitchell
Falls, 9 $ .

Nupedia infir ma (Meigen)

1 5 . J. Roan Mtn. 15,12.

Nupedia nigroscutellata (Stein)

1 $ . A. Mt. Mitchell, 1 5 , 3 2 ; Clingman’s Peak, 2 2 .

Pseudonupedia intersecta (Meigen)

1 5 , 3 2 . Mt. Mitchell.

Pegomya atlanis Huckett

1 $ . Clingman’s Peak.

Pegomya bicolor (Wiedemann)

2 $ . Bg, L.

Pegomya carduorum Huckett

1 2. Bb.

Pegomya connexa Stein

3 5 , 4 $ . A, Bg, D, L. Mt. Mitchell, 7 5, 12.

Pegomya finitima Stein

3 9. F.

Pegomya flavifrons (Walker)

1 $ . J. Roan Mtn. 1 5 .

Pegomya frigida (Zetterstedt)

1 $. P.

Pegomya geniculata (Bouche)

3^,5 2. A, C, D, J, L, U.

Pegomya hyoscyami (Panzer) var.

1 2. Bd.

Pegomya incisiva Stein
15,12. J, U.

Pegomya juvenilis (Stein)

2 5,3 2. A, Bg, D, K.

Pegomya lipsia (Walker)

6 5,6 2. A, C, D, G, Q. Mt. Mitchell, 3 2 ; Clingman’s Peak, 1 5 •

Pegomya mallochi Huckett
2 2. J.

Pegomya palposa (Stein)

2 5,4 2 . Mt. Mitchell.

Pegomya rubivora (Coquillett)

1 5 . Mt. Mitchell.

Pegomya tabida (Meigen) = Anthomyza gilva Zetterstedt (Hennig, 1973: 643)

15,12. C, P.

Pegomya univittata (von Roser)

4 5. F, P, R.

Pegomya winthemi (Meigen)

1 5,2 2. D, F.

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New York Entomological Society

Emmesomyia apicalis Malloch3

1 $ , 6 $ . A, Bb, d, g, i. Mt. Mitchell, 3 $ ; Clingman’s Peak, 1 $ .

Emmesomyia socialis (Stein)

2 9. A, K.

Hydrophoria divisa (Meigen)

1 $ . Clingman’s Peak.

Hydrophoria subpellucida Malloch
1 $. D.

Hydrophoria uniformis Malloch

27 S ■ A, Ba-c, g, i, J. Mt. Mitchell, 154 $ ; Clingman’s Peak, 109 $ ; Mitchell Falls,
13 $.

Anthomyia pluvialis (Linnaeus)

1 $. D.

Leucophora johnsoni (Stein)

13,2 9. Bf, F, J.

Leucophora marylandica (Malloch)

1 $ . Clingman’s Peak.

Paraprosalpia silvestris (Fallen)

4 $. J. Mt. Mitchell, 6 S, 2 $ ; Clingman’s Peak, 1 $.

Eustalomyia vittipes (Zetterstedt)

13,1$. C, F.

Family Muscidae

Schoenomyza chrysostoma Loew
13,1$. L.

Schoenomyza dorsalis Loew

36^,47$. B, F, I, J, L, N, O, P. Mt. Mitchell, 1 $.

Coenosia tigrina (Fabricius)

1 $ . Mt. Mitchell.

Limosia atrata (Walker)

217 3, 379 $. A-D, F, I, J-P, T, U. Mt. Mitchell, 40 $ , 46 $ ; Clingman’s Peak,

2 $ , 7 $ .

Limosia conforma Huckett
13,29. C, K.

Limosia errans (Malloch)

5 $ , 7 $ . I, J, L. Mt. Mitchell, 1 $ .

Limosia frisoni (Malloch)

4 $. J, L.

Limosia lata (Walker)

4 $ , 8 9. Ba-d, g, F, O, P. Mt. Mitchell, 4^,2$; Clingman’s Peak, 1 $.

Limosia nivea (Loew)

10 $, 24 $. A, Bb-g, C, I, J, K, L. Mt. Mitchell, 1 $, 11 9.

Hoplogaster intacta (Walker)

13,59. C, D, P.

3Steyskal (1973) has indicated that the two taxa Emmesomyia apicalis Malloch and
E. socialis (Stein) are but color variants of the same species. In apicalis the palpi are
brown, and in socialis yellow, and in the latter I find that the aristal hairs are slightly
longer. Tentatively I list the two forms separately.

Vol. LXXXII, September, 1974

157

Hoplogaster nigritarsis Stein

25 3, 56 $. A, C, D, F, I, J, K, L, O, P, T, U.

Neodexiopsis basalis (Stein)

1 <2, 2 9. Bb, C, L.

Neodexiopsis calopyga (Loew)

6 3, 29 9. A, Bb, d, g-i, C, D, F, I, J, L. Mt. Mitchell, 1 <5, 10 9 ; Roan Mtn. 1 9.
Neodexiopsis major (Malloch)

13,79. Bb, d, D, J, K, L, P. Mt. Mitchell, 1 9 .

Neodexiopsis occidentis (Stein ) = Coenosia rujitibia Stein (Huckett, 1972: 170)

2 3, 14 9. Bb, d, g, D, H, J, L. Mt. Mitchell, 13, 4 9.

Neodexiopsis ovata (Stein)

5 3, 16 9. A, Be, g, C, F, H, I, J, L, O. Mt. Mitchell, 2 3,12.

Macrorchis ausoba (Walker)

6 3,7 9 . Mt. Mitchell, 2 3 ■ Clingman’s Peak.

Lispocephala alma (Meigen)

2 3,7 9. Bf, g, T, J, K. Mt. Mitchell, 1 9 ; Roan Mtn. 1 9.

Lispocephala erythrocera (Robineau-Desvoidy)

1 3 • Mt. Mitchell.

Pentacricia aldrichii Stein

1 3 • Bi. Mt. Mitchell, 2 $ ; Clingman’s Peak, 1 9 .

Lispe albitarsis Stein

7 3, 10 9 . A, Bb-e, g, i, L. Mt. Mitchell, 11 3, 19 9 ; Clingman’s Peak, 4 3, 6 9;
Mitchell Falls, 13,29.

Lispe sociabilis Loew

1 9 . Ba. Mt. Mitchell, 3 3,2 9 .

Lispoides aequifrons (Stein)

2 3,1 2 . Mt. Mitchell.

Spilogona caroli (Malloch)

13,72. C, L. Mt. Mitchell, 2 3 .

Spilogona parvimaculata (Stein)

6 3,3 2. C, F, J.

Spilogona torreyae (Johannsen)

2 2. A.

Limnophora discreta Stein

2 2 . Mt. Mitchell.

Limnophora narona (Walker)

3 3,2 2 . Bb, g. Mt. Mitchell, 1 3,4 2 ; Clingman’s Peak, 2 3,1 2 .

Pseudolimnophora nigripes (Robineau-Desvoidy)

1 2. H.

Gymnodia arcuata (Stein)

1 3 , 29 2 . Ba, b, d, g, C, D, J, U. Clingman’s Peak, 1 9 ; Mitchell Falls, 1 $.

Helina johnsoni Malloch

2 9. A, L.

Helina obscurinervis (Stein)

2 3. F, J.

Helina rufitibia (Stein)

2 3,2 9. C, I, P. Mt. Mitchell, 2 3, 19.

Helina troene (Walker)

1 3. A.

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New York Entomological Society

Quadrularia laetifica (Robineau-Desvoidy)

22 2. I, J, K, L, N. Mt. Mitchell, 43,22.

Hebecnema affinis Malloch
62 3 , 35 $ . Ba, b, d, f, g. Mt. Mitchell, 1 $ .

Hebecnema umbratica (Meigen)

2 $ . C, J. Mt. Mitchell, 1 $ .

Hebecnema vespertina (Fallen)

8 $ . A, I, J, L, U. Mt. Mitchell, 2 2 ; Clingman’s Peak, 13,22.

Mydaea brevipilosa Malloch

13,19. Bg, C.

Mydaea discimana Malloch
1 2. L.

Mydaea flavicornis Coquillett
1 2. Bd.

Mydaea neglecta Malloch

15 2. Bb, g, D, F, J, L. Mt. Mitchell, 2 $.

Mydaea neobscura Snyder
6 $ . A, Bg, D.

Mydaea nubila Stein

9 3,3 2- A, I, J.

Mydaea obscurella Malloch

13, 5 $ . Bg, C, F, T. Clingman’s Peak, 1 2 .

Mydaea occidentalis Malloch
1 3. C.

Mydaea palpalis Stein
15 3,3 2 . Bb, d, f, g.

Mydaea urbana (Meigen)

1 2 . Mt. Mitchell.

Xenomydaea otiosa (Stein)

1 2 • J. Roan Mtn. 1 3 .

Myospila meditabunda (Fabricius)

46 3, 115 2. Ba-d, g, i, D, J, L. Mt. Mitchell, 25 3 , 25 2; Clingman’s Peak, 7 2 ;
Mitchell Falls, 2 2 .

Fannia americana Malloch

3 3, 2 2. A, Bg, D.

Fannia bifimbriata Collin

1 3, 12 2. Bb, d, f, i, C, P.

Fannia brevipalpis Chillcott
1 3 • A. Mitchell Falls, 3 3 •

Fannia brooksi Chillcott
8 2. C, D, J.

Fannia canicularis (Linnaeus)

13,12. Bb, D.

Fannia ceringogaster Chillcott

1 2. L.

Fannia depressa (Stein)

2 2. M, Q.

Fannia fuscula (Fallen)

8 3,8 2. C, D, E, F, J, Q.

Vol. LXXXII, September, 1974

159

Fannia immaculata Malloch
2 3, 1 9. I, J, L.

Fannia manicata (Meigen)

1 $ . Clingman’s Peak.

Fannia melanura Chillcott

1 2. J.

Fannia metallipennis (Zetterstedt)

2 9. M, O. Mt. Mitchell, 1 $ .

Fannia pellucida (Stein)

1 2 . Mt. Mitchell.

Fannia penepretiosa Chillcott
1 2. C.

Fannia postica (Stein)

4 2. D, L. Mt. Mitchell, 2 2 .

Fannia rondanii (Strobl)

2^,82. C, I, K, L, P. Mt. Mitchell, 2 2 ; Roan Mtn. 1 $.

Fannia scalaris (Fabricius)

1 2 . Mt. Mitchell, 1 2 . Clingman’s Peak.

Fannia serena (Fallen)

1 3, 5 2. Bd, g. Mt. Mitchell, 4 3, 3 2.

Fannia sociella (Zetterstedt)

2 3, 11 2. Bg, C, D, I, K, L, P. Mt. Mitchell, 1 3.

Fannia spathiophora Malloch

7 2. Bf, g, D, J, L.

Fannia ungulata Chillcott

2 2. A.

Coelomyia subpellucens (Zetterstedt)

8 3, 2 2. Ba, f, g, L, M. Mt. Mitchell, 3 3 ; Clingman’s Peak, 1 3.

Azelia cilipes (Haliday)

2 3, 11 2. Bb, C, D, F, I, L. Mt. Mitchell, 1 2.

Azelia gibbera (Meigen)

1 3. Q.

Hydrotaea houghi Malloch

15 2 . A, Ba, b, d, g, C. Mt. Mitchell, 2 2 ; Clingman’s Peak, 6 2 .

Hydrotaea militaris (Meigen)

4 2 • Bb, g. Mt. Mitchell, 6 2 ; Mitchell Falls, 13,29; Roan Mtn. 43, 12.
Hydrotaea occidta (Meigen)

1 2 . Bg.

Hydrotaea pilitibia Stein

9 2. A, F, J, K, L, T.

Hydroteae spinifemorata Huckett

2 2. K, P.

Lasiops innocuus (Zetterstedt)

1 3. L.

Lasiops rufisquama (Schnabl)

2 3,3 2. Bg; K, L.

Dialyta flavitibia Johannsen

8 3,7 2. Ba, g, J, K, L.

Dendrophaonia marguerita Snyder
2 9. N.

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Dendrophaonia querceti (Bouche)

1 2 . Mt. Mitchell.

Dendrophaonia scabra (Giglio-Tos)

u. c.

Phaonia aberrans Malloch

169 3 , 12 $ . Ba, b, d-g, i, L. Mt. Mitchell, 38 3,4 2; Clingman’s Peak, 14 3 .
Phaonia apicata Johannsen

16 3 , 46 2 . A, Bb-e, g, h, C, D, G, I-L. Mt. Mitchell, 7 2 ; Mitchell Falls, 4 <2 , 5 2 .
Phaonia apicata var. solitaria Stein
4 3 . Mitchell Falls.

Phaonia atlanis Malloch

1 2 . A. Mt. Mitchell, 2 $ , 1 2 .

Phaonia bysia (Walker)

4 3, 9 2 . A, Ba, g, C, D, J, K, L, P, U. Mitchell Falls, 1 2 .

Phaonia cauta Huckett

12 3 , S3 2 . Bb, d, f, g, i, C, D, K, L, P. Mt. Mitchell, 1 2 ; Clingman’s Peak, 1 2 .
Phaonia curvipes (Stein)

1 3 , 31 2 . Bb, d, f, g.

Phaonia deleta (Stein)

1 2. A.

Phaonia errans luteva (Walker)

6 2. I, J. Roan Mtn. 1 2 .

Phaonia fuscana Huckett

5 2 . C, E, I, K, L. Mt. Mitchell, 1 3 .

Phaonia laticornis Malloch

1 2. A.

Phaonia serva (Meigen)

5 3, 16 2. Bg, C, H, I, J, K. Mt. Mitchell, 1 2.

Muscina assimilis (Fallen)

13,62. Bb, g, D, L. Mt. Mitchell, 2 2 .

Muscina stabulans (Fallen)

1 3,7 2 . Bb, g, i.

Pararicia pascuorum (Meigen)

1 3. Ba.

Graphomya maculata (Scopoli)

1 2 . Clingman’s Peak.

Mesembrina latreillii Robineau-Desvoidy

1 3, 91 2. A-D, H-L, N, P, W. Mt. Mitchell, 2 2 ; Clingman’s Peak, 2 2.

Morellia micans (Macquart)

1 3, 16 2. A, Ba, d, g, K, S, T.

Pyrellia cyanicolor Zetterstedt

4 3, 17 2. A, Bf, g, D, I, L, N, U. Mt. Mitchell, 13,42; Roan Mtn. 2 3,3 2.
Orthellia caesarion (Meigen)

1 2. L. Mt. Mitchell, 13,22; Clingman’s Peak, 23,12.

Stomoxys calcitrans (Linnaeus)

3 3,9 2. A, Bb, c, f, g, i, G, J, V.

RESULTS OF THE SURVEY

A total of 57 species of Anthomyiidae sens. str. and 105 species of Muscidae were collected.
Fourteen anthomyiid and 28 muscid species were regarded as chiefly restricted in their

Vol. LXXXII, September, 1974

161

distribution to eastern and midwestern areas of the United States and the transition zones
of the Canadian provinces, namely, Family Anthomyiidae: Chirosia hystrix (Brischke), C.
pusillans (Huckett), C. stratifrons (Huckett), Delia arnolitra (Huckett), D. laevis (Stein),

D. winnemana (Malloch), Eremomyia pilimana (Ringdahl), Pegomya juvenilis (Stein),
P. lipsia (Walker), P. mallochi Huckett, P. palposa (Stein), Emmesomyia apicalis Malloch,

E. socialis (Stein), Leucophora johnsoni (Stein). Family Muscidae: Limosia errans

Malloch, L. nivea (Loew), Hoplogaster intacta (Walker), H. nigritarsis Stein, Neodexiopsis
basalis (Stein), N. calopyga (Loew), N. major (Malloch), N. occidentis (Stein), Lispe
albitarsis Stein, L. sociabilis Loew, Spilogona caroli (Malloch), S. parvimaculata (Stein),
S. torreyae (Johannsen), Helina johnsoni Malloch, H. obscurinervis (Stein), Mydaea
flavicornis Coquillett, M. neglecta Malloch, M. neobscura Snyder, Fannia americana
Malloch, F. brooksi Chillcott, F. ceringogaster Chillcott, Dialyta jlavitibia Johannsen, Den-
drophaonia marguerita Snyder, Phaonia apicata Johannsen, P. atlanis Malloch, P. cauta
Huckett, P. curvipes (Stein), P. laticornis Malloch.

Five anthomyiid and eight muscid species were regarded as boreal in habitat and as having
reached their southern limits of distribution, namely, Family Anthomyiidae: Acrostilpna

atricauda (Zetterstedt) , A. latipennis (Zetterstedt) , Alliopsis species, Pegomya jrigida
(Zetterstedt) , P. incisiva Stein. Family Muscidae: Mydaea obscurella Malloch, M. palpalis
Stein, Fannia melanura Chillcott, Hydrotaea pilitibia Stein, H. spinifemorata Huckett, Lasiops
rufisquama (Schnabl), L. innocuus (Zetterstedt), Mesembrina latreillii Robineau-Desvoidy.

Two species of Anthomyiidae, namely, Chirosia delicata (Huckett), Pegomya atlanis
Huckett, and one species of Muscidae, Phaonia aberrans Malloch, are known to be re-
stricted in their occurrence to the middle Atlantic states, from Long Island to north
Georgia.

NAMES OF COLLECTORS

Bullock, D. S.
Chillcott, J. G.
Dietrich, H.
Dreisbach, R. R.
Durden, C. J.
Hines, C. D.
Huckett, H. C.
Kelton, L. A.
Kukowitch, R. F.

Laffoon, J. L.
Mason, W. R. M.
Richards, W. R.
Shannon, R. C.
Sharp, A. V.
Snyder, K. D.
Stupka, A.
Vockeroth, J. R.
Whittaker, R. H.

Literature Cited

Brimley, C. S. 1938. “The Insects of North Carolina.” North Carolina Dept, of Agri-
culture. 560 pp.

Chillcott, J. G. 1961 A revision of the nearctic species of Fanniinae (Diptera:
Muscidae). Canad. Ent., (1960) 92 (Suppl. 14), 1-295; 289 figs.

Hennig, W. 1973. In Lindner, E., Die Fliegen der palaearktischen Region, Bd. 7 63a.
Anthomyiidae, Lief. 29 7, pp. 593-680, Taf. 78-85.

Huckett, H. C. 1972. Francis Walker’s little known North American specimens of the
families Anthomyiidae and Muscidae (Diptera) in the British Museum (Natural
History). Ent. News, 83: 169-172.

Steyskal, G. C. 1973. The genus Emmesomyia Malloch in North America (Diptera,
Anthomyiidae). U. S. Dept. Agr. Coop. Econ. Ins. Rpt., 23(22): 331-332, 2 figs.

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Stone, Alan, C. W. Sabrosky, W. W. Wirth, R. H. Foote, J. R. Coulson. 1965. “A
Catalog of the Diptera of America North of Mexico.” Handbook No. 276. U. S. Dept,
of Agriculture. 1696 pp.

Whittaker, R. H. 1952. A study of summer foliage insect communities in the Great
Smoky Mountains. Ecological Monographs, 22: 1-44, 13 figs., 9 tabs.

Wray, D. L. 1967. Insects of North Carolina. Third Supplement. North Carolina Dept,
of Agriculture. 181 pp.

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163

Notes on the Life Cycle and Natural History of Butterflies of
El Salvador. V.A. Pyrrhogyra hypsenor
( Nymphalidae-Catonephelinae )

Alberto Muyshondt

101 Ave. Norte #322, San Salvador, El Salvador

Received for Publication December 3, 1973

Abstract: For a period of three years eggs, larvae and pupae of Pyrrhogyra hypsenor
Godman & Salvin have been collected, reared, observed, and photographed in El Salvador.

In this paper the results of the observations are published for the first time, placing em-
phasis on the morphological and behavioral similarities existent with other Catonephelinae
and at least with some Callicorinae. Record is made of the larval food plants of the species
in Central America. The strong probability that the species is protected against predators
is inferred from the conspicuous coloration of the larvae and adults and from the known
poisonous properties of the food plants. Finally it is noted that apparently there is a prefer-
ence for parasitizing Diptera and Hymenoptera to deposit their eggs on the larvae of species
protected against predators by the unpalatable and/or poisonous substances sequestered
from their food plants, probably to ensure the safety of their own eggs and larvae.

INTRODUCTION

This is the fifth article of a series revealing our observations on the early
stages, behavior, and food plants of local butterflies belonging to the Catoneph-
elinae, group of the Nymphalidae. The series will include at least some of the
Callicorinae to emphasize the close relation between the two groups, as evi-
denced by the many morphological and behavioral similarities existent in their
respective early stages. This at the same time will establish the great differences
that exist with other groups of the Nymphalidae, so as to make one wonder
if the common characteristic of the adults having only two pairs of ambulatory
legs is a criterion strong enough on which to base a family. Not long ago Papili-
onidae and Pieridae were grouped together on the basis of the two groups hav-
ing three pairs of ambulatory legs.

We had observed and collected adults of Pyrrhogyra hypsenor Godman &
Salvin, since 1958, in ravines and creeks running through coffee plantations
in the neighborhood of San Salvador (600 to 900 m altitude), but owing to
our deficient knowledge of butterfles, we had always placed them among the

Acknowledgments: Once again we, my sons and I, express our obligation to Dr. Alexan-
der B. Klots, of the American Museum of Natural History, New York, for his generous
advice in our studies and for reading and constructively criticizing our manuscript. We
are thankful also to S. Stainhauser for kindly determining the butterfly species, and Drs.
C. W. Sabrowski, B. D. Burks, and J. L. Herring, of the United States Department of
Agriculture, who determined the parasites and predator mentioned in this article.

New York Entomological Society, LXXXII: 163-172. September, 1974.

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Figs. 1 to 7. Pyrrhogyra hypsenor Godman & Salvin

1. Egg showing prominent ribs. About 1 mm.

2. First instar larva. About 2 mm.

3. Second instar larva. About 3.8 mm.

4. Third instar larva. About 7.5 mm.

5. Fourth instar larva. About 1.6 cm.

6. Close up of head of a fifth instar larva.

7. Fifth instar larva. About 3 cm.

local Adelpha species, which they somewhat resemble, and with which they
share the habitat. It was not until late 1970, when we were searching a Paul-
linia pinnata L. vine for eggs and larvae of Morpho polyphemus polyphemus
Doubleday and Hewitson, that we found one larva, which unmistakably per-
tained to a Catonephelinae and which eventually produced our “pseudo-Adel-
pha,” that we realized our error. The butterfly was identified by S. Steinhauser.

Once the food plant was known, it was a matter of only a few weeks to find
a female in oviposition. A number of eggs were collected and put in clear plas-
tic bags. The larvae hatched from them were fed until pupation on leaves of
the same plant. Photos were made of the eggs, the different larval instars, and

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165

Figs. 8 to 10. Pyrrhogyra hypsenor Godman & Salvin

8. Pupa dorsal view. About 2 cm. long.

9. Pupa lateral view.

10. Pupa ventral view.

the pupae. Measures of the different stages and the time spent in each one were
recorded. The bags were kept at all times under ambient conditions of light
and temperature. Specimens of the early stages were preserved in alcohol and
sent to the American Museum of Natural History, New York, with specimens
of the adults.

This is one of the species reared the most, with similar results every time.

LIFE CYCLE STAGES

Egg. Bright light yellow, truncated-cone-shaped, with 11 prominent yellow ribs from
micropylar zone to base. About 1 mm long. Hatches in 5 days.

First instar larva. Head naked, roundish, light brown. Body greenish-yellow, cylindrical,
naked, with light brown legs. About 1.5 mm when recently hatched, growing to 2.2 mm
before moulting in 4-5 days.

Second instar larva. Head brown with short thick horns on epicrania. Body greenish-
brown, with scattered whitish tubercules and small forked spines, growing to 3.5 or 4 mm
in 5 days.

Third instar larva. Head cordiform, light brown, with dark brown, long horns on epi-
crania, bearing three rosettes of accessory spines, and slender spines on lateral margins.
Body light brown, except for orange caudal segments, with whitish tubercules and black

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Figs. 11 and 12. Pyrrhogyra hypsenor Godman & Salvin

11. Dorsal view of male and female adults.

12. Ventral view of male and female adults.

spines arranged as follows when seen laterally: on first thoracic segment (T-l), 2 sub-

dorsal simple spines, 1 subspiracular simple spine. On T-2, 1 prominent forked subdorsal
spine, 1 supraspiracular forked spine, 1 subspiracular simple spine. On T-3, 1 very promi-
nent forked subdorsal spine, 1 forked supraspiracular spine, 1 subspiracular simple spine.
On the first abdominal segment (A-l), 1 subdorsal simple spine, 1 supraspiracular simple
spine, and 1 subspiracular simple spine. From A-2 to A-6, 1 subdorsal forked spine, 1
supraspiracular simple spine, 1 subspiracular simple spine, and 1 supraventral simple spine.
On A-7 and A-8, 1 prominent forked spine at meson in addition. On A-9, 1 supraspiracular
forked spine only, directed posterad. The larva grows to .75 cm in 4-5 days.

Fourth instar larva. Head as in third instar but reddish with black horns and spines.
Body’s ground color brownish-orange with rings of yellow spots mostly dorsally, with black

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dorsal stripes across T-3, A-3, A-5, and A-7. Subdorsal spines on T-3, and median spines
on A-7 and A-8 very prominent, the latter orange-colored. Spines on A-9 orange with
black forks. Growing to 1.6 cm in 4-5 days.

Fifth instar larva. Head as in fourth instar, with longer horns. Body brownish-orange

ventrally, mostly yellow above, with thin black rings across segments and red wide bands
across T-2 and T-3 from base to base of subdorsal spines. Broad saddle-like zones on
dorsum of A-3, A-5, and A-7. Growing to 3 cm in 4-5 days.

Prepupa. All colors but black turn into green. Shortens and thickens considerably. One
day.

Pupa. All green except for dark brown cremaster and last abdominal segments. Body
thickens gradually to first abdominal segments which are the thickest part of the body
dorsoventrally and laterally. A projecting spur directed anterad on the dorsal part of the

thorax. Head slightly bifid. Spiracula very inconspicuous brown. Measures 1.5 to 2 cm

long, and lasts 7 days.

Adults. This species does not show a marked sexual dimorphism. Wingshape the same in
both sexes. Front wing with costal margin slightly convex with rounded apex, slightly
concave and sinuose outer margin, rounded tornus and almost straight inner margin. Hind-
wing with convex costal margin, rounded outer angle, sinuose outer margin, with a more
pronounced curve projecting on M3 vein, forming almost a “tail,” rounded anal angle and
convex inner margin, with a slight fold. Dominant dorsal color dark brown, darker in
males. Forewing with two squarish white bands aligned from midcostal margin to mid-
inner margin. Females with, in addition, a small round white spot subapically. Hindwing
with an elongated white stripe, continuing the white bands of the forewing, starting from
midcostal margin, ending in a point before reaching the anal angle. Both sexes with a small
red spot between the point of the white stripe and the anal angle.

Dominant color ventrally, white. Fore wing with a light brown band covering the distal
portion of the wing from % costal margin to % inner margin, covering the apex, outer
margin, and tornus. A thinner brown branch arising from the brown zone, about the discal
area, reaching the costal margin at a 60° angle. A secondary brown branch originating from
the primary ending at the base of the wing. Basal third portion of the costal margin and all
around the white island formed by primary branch, thinly lined with red. Hindwing mostly
white with a brown zone alongside the outer margin, covering about Vs of the wing surface.
On this brown area, alternately light and dark brown thin lines running parallel to the
sinuose outer margin, with a row of dark brown spots alongside the basal limit. A thin
red line running along the costal margin of the white area continuing along the basal limit
of the brown zone.

Average size of adults about 5 cm from tip to tip of spread front wings. Total develop-
mental time varying from 34 to 38 days.

NATURAL HISTORY

In El Salvador the eggs of Pyrrhogyra hypsenor are found exclusively on the
tender shoots of at least two species of Paullinia , either on the new leaves or on
the tendrils, and even if only one egg is deposited per location, several eggs
per shoot are laid by a single female. As many as 15 eggs have been found on
a single tender stem, probably resulting from several females visiting the same
terminal.

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The recently hatched larvae eat the upper surface of the eggshell and part
of the adjacent wall also. They move afterward to the edge or to the tip of a
tender leaf and feed thereon, using small pellets of excrement woven with silk
to construct a resting perch that protudes from the leaf. The larvae quit this
station only to feed, which they do early in the morning or late in the after-
noon. While resting, the larvae usually have the head pointing outward, at
times holding onto the perch with their prolegs and lifting the anterior part
of the body. Second and at times third instar larvae behave similarly. From
the third instar on, the larvae roam about the plant but always on the younger
leaves on which they feed exclusively, even during later instars.

When the larvae are ready to pupate they move to older parts of the plant
or even to neighboring shrubs, choose a place of their liking, weave a silk pad,
clean their digestive tract, and affix their anal prolegs to it. Very often the
chosen location is the upper surface of a leaf, and the larvae do not hang but
lie parallel to the surface.

The pupae, usually found standing at an angle on top of a mature leaf, react
when disturbed by wiggling vigorously from side to side, producing an audible
squeaking sound. Shortly before adult emergence, the pupae become dark
brown.

The adults rapidly abandon the shell and hang from a suitable place, while
ejecting a reddish meconium, until the wings are rigid. The process takes about
15 minutes.

Pyrrhogyra hypsenor adults do not visit flowers but feed greedily on a variety
of fallen and fermenting fruits, on juices flowing from tree wounds, on excre-
ments and mud. After their long feeding sessions the adults fly to a nearby
shrub and sit on top of a well-exposed leaf where they stay motionless except
for occasional flappings of the wings, which might be held open most of the
time. The adults of this species imitate not only the general color pattern of
local Adelpha spp. but even their peculiar jerky and sliding flight. We have
observed females of P. hypsenor alight repeatedly on leaves of a plant commonly
used by at least two species of Adelpha for oviposition and act as if depositing
eggs on it. Although we have observed the species for a number of years, we
have never been able to witness copulations.

The tender shoots of the food plants of P. hypsenor are currently invaded by
a species of aphid, which are tended by ants. The young parts of the plants
are also used as food by several species of Theclinae (T. marsyas, T. mykon ,
and others).

The larvae and pupae of P. hypsenor quite often bring forth tachinid and
chalcidid parasites. The tachinidae that were sent to the U.S. Department of
Agriculture were determined by C. W. Sabrowski as “ genus spA” [sic], the
chalcidoidea were determined by B. D. Burks of the same institution as Sphilo-
chalcis persimilis Ashmead. (Both parasites occur also in Pseudonica flavilla

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169

canthara Doubleday.) We found, once only, a Pentatomidae nymph (deter-
mined by J. L. Herring, USDA, as “ genus sp. ?”) with a third instar larva of
P. hypsenor impaled on its beak. A frequent cause of larval mortality, both
in our insectarium and in the fields, is a disease that produces diarrhea, which
is followed by softening of body tissues and death by bursting. Very often we
find dead larvae in the fields hanging limp from a leaf, still holding on with
their prolegs.

The food plants on which we have found eggs and larvae of P. hypsenor be-
long to the genus Paullinia (Sapindaceae) ; P. pinnata L. is by far the most
usual, and P. fuscescens H.B.K. less usual. On both plants only the tender
shoots are used for ovipositioning and feeding by this species.

Paullinia pinnata is a robust, semiscandent, tendril-bearing plant, with al-
ternate, persistent, pinnate leaves consisting of 5 to 7 large (up to 10 cm long)
lanceolate remotely dentate, slightly coriaceous leaflets on a broadly winged
rachis. The inflorescence is an axilar raceme of small, whitish, 4-petaled flow-
ers, which produce 3 -celled, septicidal, roughly pyriform, thick-walled capsular
fruits about the size of a coffeebean that are green when young, becoming red-
dish-orange when mature and containing up to three shiny-black seeds, covered
basally by a white arillum. This plant is widely used in the Neotropics for
stupefying fish in streams and lakes, and, according to several authors (H.
Baillon, 1874; L. Beille, 1909; P. Standley, 1923), it is reputed to be very
poisonous and to contain an alkaloid, timboine. Standley says: “Some of the
Indians are said to have used the juice to poison their arrows and it is reported
that in the Antilles the Negroes have made use of the seeds for criminal poison-
ings.”

P. pinnata and P. fuscescens , the alternate food plant, are very widely dis-
tributed in the country along wooded ravines and creeks from near sea level
to about 1500 m altitude.

DISCUSSION

Seitz (1914) states that very little is known about the larvae of Pyrrhogyra
hypsenor and although he does not describe the pupa in his work, he states
that “The pupa shows the same peculiar attachment as that of Myscelia, be-
cause it is attached to the upper surface of the leaf.” We are not aware of any
other publication describing the early stages of the species, so it appears that
this is the first mention of them.

Although this species is evidently in close relation to Myscelia and thence
to the rest of the Catonephelinae, as grouped by Ebert (1969), we feel that it
is a link between that group and the Callicorinae, a very closely related group,
if we are to judge by the many similarities existent between the early stages
and their behavior; the shape of the eggs of this species is more like the shape
of the eggs of the latter group than the shape of the eggs of the Catonephelinae,

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which are crowned by a series of protuberances around the micropylar area.
It is also the only case we have found among the Catonephelinae with eggs
colored other than white (the eggs of the Callicorinae are light green). As for
the shape and behavior of the larvae and pupae, these conform to the general
shape and behavior of the Catonephelinae we have studied.

It is true that the morphological and behavioral characteristics of the Calli-
corinae show abundant similarities to the other group, but in general the larvae
of the Callicorinae are not armed with the profusion of spines peculiar to the
Catonephelinae, so that it is an easy task to tell them apart under a superficial
examination from the third stadium on.

A factor which is exclusive to this species and seems to indicate an evolution-
ary trend that may eventually lead the species to adopt a gregarious behavior
is the discriminatory use of the young sprouts of the plant only for egg laying
and larval feeding. This trait causes a concentration of eggs and larvae, not
necessarily originating from one female only, but still usually several from one
female, which are forced to live and grow within a limited space. As a result,
larval encounters are common events. These do not result in larval fights nor
in larval mortality that would be caused by wounds inflicted on each other by
contenders. This peaceful coexistence is a total deviation from the individualis-
tic behavior displayed by the rest of the Catonephelinae we have studied, whose
larvae fight intransigent^ against other roaming larvae that accidentally come
in contact with them. The larvae of P. hypsenor have learned also to accept im-
passively the continuous traffic of ants tending the aphids which often dwell
in their domains, and the occasional disturbances caused by neighboring larvae
of Theclinae which share their food.

The preference acquired by this species of feeding on young leaves might
be an indication that the young leaves of the Paullinia vines are chemically
quite different from the older ones, as is the case on Prunus , Kalmia , Laurus ,
Quercus , etc., whose young foliage is much more toxic than the older (Klots,
personal communication). If this is the case the larvae feeding on young leaves
of Paullinia would thus build up rapidly an effective predator-deterrent concen-
tration of toxins.

For the rest, the behavior of the larvae of P. hypsenor is exactly like that
of the larvae of the other Catonephelinae we have observed: The eggshell is
disposed of in the same manner, the larvae build a similar resting perch with
frass pellets, they feed at about the same times, and, in the later instars, they
crawl about the upper surfaces of the leaves. The pupae stand on the leaves
like those of the other species. So they use the cryptical strategy during the
early larval stadia, and flaunt their presence in the late instars, even more
than the others, on account of their showier colors. This seems to indicate that
the protection they might derive from the noxious constituents sequestered from
the food plants takes some time to reach the necessary concentration to protect

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them effectively against predation. The adult behavior and coloration of P.
hypsenor tends to support our speculation in this respect: They are the most
conspicuous of all Catonephelinae.

In this species we again find that the alleged protection against predation
derived from the poisonous properties of the food plant does not protect the
larvae against parasitism. In fact the resultant immunity may be an advantage
to the parasitic Diptera and Hymenoptera, whose larvae logically would benefit
from the repellent properties of the host. The same phenomenon has been
noticed in other species of butterflies classically accepted as protected against
predation as a result of the food plant having poisonous and/or bitter com-
ponents. Among these we count several local Danaidae ( D . plexippus , D. eresi-
mus, D. gilippus), Heliconiidae (H. charitonius, H. petiveranus, H. talchinia,
Eueides aliphera, E. cleobaea ), Ithomiidae ( Dircenna klugii), Papilionidae
(. Battus polydamas, Parides photinus, P. areas), feeding on Asclepiadaceae,
Passifloraceae, Solanaceae, and Aristolochiaceae, respectively, plants known
or reputed to contain noxious substances, and other species feeding on Sapin-
daceae, such as Morpho polyphemus (Young and Muyshondt, 1972), Temenis
laothoe liberia (Muyshondt, 1973a), Pseudonica jlavilla canthara (Muyshondt,
1973&); others on Piperaceae such as Anaea ( Consul ) fabius (Muyshondt,
1973c) and Anaea (C.) electra (Muyshondt, in prep.). All of these species
produce, very often during the late larval instar or during pupation, quite a
variety of Tachinidae or Hymenoptera parasites.

It is evident that the characteristic of most of the predation-protected larvae
of crawling about exposed, displaying their gaudy colors, makes them an easy
target for the parasitizing female. That could very well be nature’s way of
keeping in check the population of a species chemically protected against pre-
dation. Perhaps, after more evidence is gathered, it could be deduced that when-
ever a species is found to be very prone to parasitism, it is to be suspected that
that species is chemically protected against predation by food-plant derivatives.

Literature Cited

Baillon, H. 1874. “Histoire des plantes.” Hachette et Cie, Paris.

Beille, L. 1909. “Precis de botanique pharmaceutique.” A. Malone, Paris.

Ebert, H. 1969. On the frequency of butterflies in Eastern Brazil, with a list of the
butterfly fauna of Pocos de Caldas, Minas Gerais. Jour. Lep. Soc., 23: Sup 3.

Ehrlich, A. H. and Ehrlich, P. R. 1961. “How to Know the Butterflies.” Wm. C.

Brown Co., Publishers, Dubuque, Iowa.

Klots, A. B. 1960. “A Field Guide to the Butterflies.” Riverside Press, Cambridge, Mass.
Muyshondt, A. 1973a. Notes on the life cycle and natural history of butterflies of El

Salvador. III. A. Temenis laothoe liberia (Nymphalidae-Catonephelinae) . Jour.

New York Entomol. Soc. In press.

. 19736. Notes on the life cycle and natural history of butterflies of El Salvador.

IV. A. Pseudonica jlavilla canthara. (Nymphalidae-Catonephelinae). Jour. New
York Entomol. Soc. In press.

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— . 1973c. Notes on the life cycle and natural history of butterflies of El Salvador.

III. Anaea ( Consul ) jabius (Nymphalidae) . Jour. Lep. Soc. In press.

Seitz, A. 1914. “Macrolepidoptera of the World.” Vol. 5. Stuttgart.

Standley, P. C. 1923. Trees and shrubs of Mexico. Contrib. from the U.S. Nat. Herb.,
Vol. 23, Part 3, 701-3.

Young, A. M., and Muyshondt, A. 1972. Biology of Morpho Polyphemus (Lepidoptera-
Morphidae) in El Salvador. Jour. New York Entomol. Soc., 80, 1: 18-42.

Vol. LXXXII, September 1974

173

Tenuicoris myrmeforme: A New Genus and Species of Myodochini
( Hemiptera : Lygaeidae ) 1

James A. Slater and Jane E. Harrington
Biological Sciences Group, University of Connecticut, Storrs, Connecticut

Received for Publication December 13, 1973

Abstract: Tenuicoris myrmeforme is described as a new genus and new species from

Bolivia, Peru, and Brazil. Ant-mimetic characteristics are noted. The relationships of the
genus are stated to be with such neotropical species as Heraeus cincticornis Stal. A dorsal
view of the holotype is presented.

The tribe Myodochini is one of the largest taxa in the lygaeid subfamily
Rhyparochrominae. It is the most diverse and dominant element in the neo-
tropical rhyparochromine fauna.

Several Western Hemisphere myodochines are striking ant-mimics. As field
information on lygaeid behavioral patterns increases, it is becoming evident
that many additional species are also ant-mimics, although the morphological
modifications are so limited that this is not readily evident in museum specimens.

In the present paper we describe a new ant-mimetic species from South
America which represents an undescribed genus.

Tenuicoris, new genus

Head elongate, acuminate, swollen and formicoid in lateral view; interocular area flattened,
post-ocular area prolonged and tapering markedly but gradually from eye to insertion of
head, lateral margins of juga forming a sharp ridge extending posterior to and above
insertion of antennal segment 1 ; anterior pronotal lobe shining, strongly convex, with a distinct
narrow ring-like anterior collar, lateral margins of anterior lobe evenly rounded, transverse
impression deep and distinct, lateral margin of posterior lobe obtusely rounded, narrowing
from humeri to transverse impression at a 45° angle, posterior margin straight, anterior lobe
(except collar) impunctate, strongly polished, posterior lobe completely pruinose with
anterior % thickly pale gray to silvery pruinose and posterior % strongly differentiated as
less densely pruinose yellowish ; scutellum convexly elevated across anterior % ; hemelytra
attaining abdominal apex, clavus coarsely punctate, forming four or more very irregular
intermixed rows of punctures, lateral corial margins strongly constricted at level of claval
commissure ; legs elongate and slender, fore femora only slightly incrassate, bearing one large

Acknowledgments: We wish to extend our appreciation to Mr. Johann Becker (Museu

Nacional Brazil); Dr. R. C. Froeschner (National Museum of Natural History), and Dr.
Peter Wygodzinsky (American Museum of Natural History) for the loan of material; to
Drs. C. W. and L. B. O’Brien (Florida State University) for the gift of specimens; to Mrs.
Kathleen Schmidt (University of Connecticut) for the execution of the illustration; and
to Mrs. Darken Wilcox and Mrs. Elizabeth Slater (University of Connecticut) for aid in the
preparation of the manuscript.

1This work was supported by a grant-in-aid from the National Science Foundation.

New York Entomological Society, LXXXII: 173-176. September, 1974.

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and several small spines subdistally below; 1st metatarsal segment extremely elongate, three
times as long as length of segments 2 and 3 combined; antennae very long, slender, terete;
metapleural evaporative area extending well beyond scent gland orifice to occupy inner
% to % of metapleuron, laterally truncate; abdominal spiracles 2, 3, and 4 located dorsally;
scent gland scars present between abdominal tergites 3 to 4, 4 to S, and 5 to 6; inner latero-
tergites absent; abdomen moderately constricted basally.

This ant-mimetic genus belongs to the rhyparochromine tribe Myodochini. It has all of
the typical tribal characters such as abdominal spiracles placed dorsally on segments two,
three, and four, inner latero-tergites absent, conventional, generalized trichobothrial pattern
and laterally rounded pronotum.

Tenuicoris is most closely related to Heraeus Stal and appears to be derived directly from
species of the latter. Heraeus at present contains a rather diverse assemblage of species held
together chiefly by the tendency of the head behind the eyes to be narrowed to form a
short “neck.” Tenuicoris myrmeforme is probably derived from a Heraeus stock rather
similar to that at present represented by Heraeus cincticornis Stal. Like Tenuicoris
myrmeforme, cincticornis is a large, slender species with very elongate legs and antennae.
Its lateral jugal margins are noticeably carinate and the head, when viewed laterally, is
rather myrmecoid in appearance. There is a conspicuous elongate white macula distally on
the corium and another at the mesal apex of the membrane. The connexival area on sterna
4 and 5 is also pale. There are other large neotropical species of Heraeus that also have
carinate lateral jugal margins so that T. myrmeforme seems to represent a highly derived
taxon that has evolved from a Heraeus cincticornis -like ancestor.

Tenuicoris myrmeforme is readily distinguishable from any species of Heraeus by the
striking condition of the pronotum. The anterior lobe is smooth and polished while the
posterior lobe appears banded with its anterior third very heavily pruinose and its posterior
two-thirds sharply demarked as less pruinose. Heraeus species generally have both pronotal
lobes dull or subshining and never present the contrasting highly polished anterior lobe of
Tenuicoris. The longitudinally oval eyes of T. myrmeforme are quite unlike the condition
found in Heraeus species. The basally constricted abdomen and strongly mesally depressed
head in the area of the eyes and juga are also distinctive features, and we have not examined
any species of Heraeus which has such reduced armature of the fore femora as is found in
T. myrmeforme.

Type species: Tenuicoris myrmeforme, n. sp., monobasic.

Tenuicoris myrmeforme, n. sp.

Please see Fig. 1. Head, antennal segment 1 and scutellum chestnut to tawny; antennal
segments 2, 3, and 4 sordid buffy yellow with distal portions of 2 and 3 red, proximal and
distal ends of segment 4 shaded with buffy brown; pronotum, thorax laterally and ventrally,
femora and tibiae tawny, lightly suffused with brownish red, proximal % of femora pale,
tarsi tawny to buffy yellow; clavus and corium cinnamon, membrane becoming buffy brown
and fuscous along apical margins of corium, central area of clavus and on corium adjacent to
claval suture and over most of membrane; hemelytra marked with strongly contrasting
white coloration as follows: a narrow stripe along entire lateral claval margin adjacent

to claval suture widened at distal end, a short narrow macula on corium adjacent to claval
suture at level of anterior % of claval commissure, an oblique mesally tapering vitta near
posterior end of corium running antero-mesad from lateral corial margin almost to middle

Fig. 1. Tenuicoris myrmeforme, n. sp. Dorsal view, holotype.

Vol. LXXXII, September, 1974

175

H. Schmidt

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New York Entomological Society

of apical corial margin, a broad white median stripe on distal Vs of membrane extending
anteriorly almost to level of apex of corium ; abdomen fuscous, becoming tawny on sternum
2, connexivum ventrally with sordid white macula covering most of segment 5 and central
V2 of segment 6; head granulose and obscurely transversely rugulose, anterior pronotal lobe
impunctate, shining, posterior lobe with distinct discrete punctures; scutellum shining with
a few large scattered punctures on elevated area and coarsely punctate laterally; entire body
surface except membrane clothed with fine short ( circa 0.08) 2 decumbent hairs, hairs most
dense on and lending a subshining appearance to antennae, tibiae, abdomen and laterally in
transverse impression of pronotum, anterior pronotal lobe nearly glabrous; anterior pronotal
collar pruinose and punctate ventrally.

Head slightly declivent anteriorly, eyes longitudinally oval, sessile, length head 1.86, width

I. 23, interocular distance 0.61; length anterior pronotal lobe 1.14; length posterior lobe 0.61,
maximum width anterior lobe 1.14; width across humeri 1.54; length scutellum 0.99, width
0.84; length corium 3.31, distance apex corium to apex membrane 1.38; labium extending
beyond posterior margin of prosternum, length labial segments I 0.84, II 1.03, III 0.76,
IV 0.38; bucculae very short, scarcely reaching over proximal end of 1st labial segment,
extending beyond apex of tylus; length antennal segments I 0.95, II 1.80, III 1.48, IV 1.75;
total length 8.24.

Holotype. Bolivia: $ Rurrenabaque Beni Oct. 1921 (W. M. Mann), Mulford Bio. Expl.

1921-22. In National Museum of Natural History No. 71224.

Paratypes. Bolivia: 1 $ Reyes-beni XII-12-1956 — 1 $ Prov. Sara (Steinbach). Peru: 1 $
Tingo Maria VII-10-1968 night (C. E. & L. B. O’Brien). Brazil: 1 $ Barbacena, M. Gerais,
Feb. 1962 (M. Alvarenga) on Urticaceae — 1 $ Caceres M. Gerais, Dec. 1955 (Alvarenga).
In Museu Nacional Rio de Janeiro, American Museum of Natural History, J. A. Slater and

J. E. Harrington collections.

The two Brazilian and the Peruvian paratypes are darker than the holotype and Bolivian
paratypes. Their general coloration is between dark chestnut and dusky brown or fuscous
rather than the light chestnut to tawny of the holotype. The number of small spines
present ventrally on the fore femora also seems to be a variable condition. However, the
body form and proportions, distinctive white color pattern of the hemelytra, and unique
pruinose banding on the posterior pronotal lobe and all other significant morphological
features are constant.

The Villalobos color chart (Palmer, 1962) has been used as a standard in the above
description.

Literature Cited

Palmer, R. S. 1962. “Handbook of North American Birds.” Vol. I, Loons through
Flamingos. New Haven and London, Yale University Press.

2 All measurements are in millimeters.

Vol. LXXXII, September 1974

177

A New Genus and Two New Species of Achipteriidae from New York
State (Acari: Cryptostigmata: Oribatei)

F. Reese Nevin

Department of Biological Sciences, State University of New York,

College of Arts and Science, Plattsburg, New York 12901

Received for Publication March 5, 1974

Abstract: Dentachipteria, a new genus, and two species of oribatid mites are described. Spe-
cies are recognized by the large downward pointing lamellae the broad rostrum, and presence
of denticles on the distal margins of the pteromorphs. In D. ringwoodensis there are many
pteromorphic denticles, in D. highlandensis there is a single large denticle. In D. ring-
woodensis leg 1 is held beneath the lamellae, in D. highlandensis leg 1 is free.

INTRODUCTION

In the late thirties and early forties a number of collections of mites were made
and preliminary studies started. Balsam mounts, specimens preserved in a
killing-fixing and preserving fluid, and specimens preserved in beechwood
creosote serve as the basis for the following study.

Dentachipteria , n. gen.

Generic characteristics. The body is truncated anteriorly giving a squared appearance.
The prodorsum is strongly bent ventrally so that the tips of the lamellae point downward.
The lamellae are broad and flat covering most of the prodorsum. The lamellar setae
(le) are rough and pressed against the lamellae. The interlamellar setae are long. The
translamella is present or may be incomplete. The sensilli are rodlike. The notogaster
is cup-shaped to more elongate, rounded posteriorly; the pteromorphs are bent strongly
ventrally with the distal margin dentate and the anterior margin without a pronounced
curve to the lateral tip. The notogaster bears 10 pairs of setae which decrease in size
posteriorly. There are one to three pairs of sacculi. The ventral setae are long and
geniculate. There are 6 pairs of genital setae, 2 pairs of anals, and 3 pairs of adanals.
The fissure iad is near the anterior end of the anal field. The tarsi are tridactylous. Tarsus
II bears a large branched seta on its ventral surface. The mandibles are chelate and the
maxillary palps are five-jointed.

Dentachipteria ringwoodensis, n. sp.

Color , Mahogany red. Size , Mean for 19 adult specimens. Length: 0.674 mm; range:

0.62 mm-0.74 mm. Width: 0.516 mm; range: 0.47 mm-0.53 mm (Figs. 1-4).

Mean depth for three specimens 0.42 mm. The greatest width was in the posterior
region of the hysterosoma through the anal plates.

Acknowledgments: I wish to thank Dr. Marie Hammer of Roland, Fredenborg, Denmark
for the examination of some of the drawings and for her suggestion about generic status.
I also wish to thank Dr. E. Piffl of the Zool. Inst, of University of Wien, Vienna, Austria for
his interpretation of the position of Parahypozetes into its correct family.

New York Entomological Society, LXXXII: 177-182. September, 1974.

178 New York Entomological Society

ringwoodensis, n. sp. Ventral view. 3. Dentachipteria ringwoodensis, n. sp. Lamella
with lamellar and interlamellar setae. 4. Dentachipteria ringwoodensis, n. sp. Sketch
of leg I held in position by the lamella and the rostrum and showing the rostral seta.

Shape. Oval, truncated anteriorly, the gnathosoma sharply bent ventrally.

Prodorsum. The broad lamellae with their broad thin cusps cover most of the prodorsum.
Because of the bending of the gnathosoma the tips of the lamellae point downward and
are not readily seen in dorsal view. Each lamellar cusp bears a pointed tip in an anterio-
lateral position and a rounded median tip. A fleshy appearing lamellar hair arises as an
extension from a thickened area along the median margin of the lamellar cusp and extends
beneath the cusp or along its apical margin. It does not extend straight forward. The
free ends of the lamellar cusps are wrinkled with a few small nodules among the wrinkles.
The interlamellar setae are long and straight with a few short spines. Their origin is
at the midpoint of the lateral margins of a clear area between the slatlike parts of the
lamellae. The rostral bristle is not visible from the dorsum. It is almost smooth, is
straight and directed anteriorly.

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179

The rodlike sensillus originates from the mesial margin of the bothridium, curves
laterally and anteriorly to a point opposite the genus of leg II, then bends inward toward
the lamellae extending to a point opposite the middle of tibia II. The sensillus enlarges
slightly from the bend to the tip and bears many short barbs especially toward the tip.
The mandibles are well developed and are chelate in type.

Notogaster. The notogaster extends broadly anteriorly along the midline slightly beyond
the anterior margins of the pteromorphs. The integument of the notogaster as well as of
the pteromorphs is punctate and of a finer granular nature in the middorsal region
which is rounded and in some specimens is marked off by its fine granularity from the
rest of the notogaster. There are ten pairs of notogastral setae, the shorter setae being
near the posterior end. What appears to be openings of the sacculi, Sa, are found anterior
to seta ti in some specimens but could not be seen in all. Si is anterior to seta r3; S2 is
near the base of r2. Crescent shaped bodies appearing much like the openings of sacculi
are found in the hypodermis near the posterior end of the notogaster and interfere with
the detection of possible saccular openings. It is necessary to remove some of the pigment
before the setae and sacculi can be studied. There is no lenticulus.

Ventral Region. The genital plates bear 6 pairs of setae arranged as follows: gi and

g2 are along the anterior border of the plate; g3, g4, gs and g6 are arranged in a row
extending in a line posterior to a point midway between the bases of gi and g2. In a few
specimens g3 forms a line directly posterior to g2. The space between g4 and g5 is slightly
greater than the space between g3 and g4 or g5 and g6 which are evenly spaced from one
another.

There are 2 setae on each anal plate, one near the center at the anterior end of the
plate and one near the center at the posterior end of the plate. Seta ad3 is located midway
between the anterior and posterior ends of the anal plates, posterior and lateral to the
slit-pores iad. Each hair of the ventral surface is inserted beside a hair pore giving the
appearance of the hair encircling the hair pore and described by Hammer (1967) in
Parahypozetes as geniculate. The anal plates are larger than the genital plates. The
distance between the genital and anal plates is greater than the length of the anal plates.

In the epimeral region the following setae are found: la and lb, 2a and 2b, 3a and 3b,

and 4a.

Clear less densely pigmented areas are found between the camerostome and the genital
plates, anterior to the apodeme of leg 1 and also betweeen the apodemes of leg I and leg II.
Small circular, less deeply pigmented areas appear in the integument between the genital
and anal plates.

Legs. All tarsi are tridactylous, the middle claw being much heavier than the other two.
Leg I differs from the other legs in being held between the lamellae and the rostrum.
The femur of leg I fits into a pocket of the propodosoma. The genus of leg I bends at
near a 90° angle. The tarsus of leg II only bears one large branched ventral seta. Both
the genus and tarsus of leg II bears a heavy blunt smooth spine.

Materials. The holotype and 30 paratypes (some dissected or crushed to study the struc-
tures) are adult specimens mounted individually in balsam on slides. All slides will be
deposited in the collection of the New York State Museum in Albany, New York.

Type locality. Ringwood Preserve near Ithaca, New York. Specimens were collected
by me on November 19, 1939 from among liverworts and mosses in a wooded area at
the margin of a small muck pond just off the highway.

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Figs. 5-6. 5. Dentachipteria highlandensis, n. sp. Dorsal view. Enlargement of
lamella and lamellar seta. 6. Dentachipteria highlandensis, n. sp. Ventral surface of
pteromorphs to show single denticle.

Dentachipteria highlandensis, n. sp.

Color , Mahogany red. Size, length: 0.60 mm; width: 0.46 mm (Figs. 5-6).

Shape. Oval, truncated anteriorly.

Prodorsum. The lamellae are large, covering most of the prodorsum. Outer ends of the
lamellar cusps are distinctly separated. Each cusp ends in a lateral point which may not
be seen except in a semi-face view. The median margin of the cusp bears a lamellar seta
visible through the cusp. It arises along the mesial margin of the lamella and may appear
entirely ventral to the lamellae. It extends outward to a point near the lateral tip of the
lamella. Rostral setae are attached far back on the rostrum and curve inward beneath
the lamellae and extend slightly beyond the tip of the rostrum. The rostral setae are heavily
plumose especially along the lateral margin. The long interlamellar setae are spinose from
base to tip.

Vol. LXXXII, September, 1974

181

The sensilli bear several rows of short bristles . They are more abundant apically and
disappear near the bend of the stalk of the sensillus.

Notogaster. The notogaster is longer than broad. It bears a distinct lenticulus. The noto-
gastral setae are similar to those of D. ringwoodensis. Only one sacculus, Si, was found.
The integument over the mid- notogaster is rounded up and is finely punctate. The entire
dorsal surface including the upper surface of the pteromorphs is coarsely punctate. The
large pteromorphs bear a single large denticle on the distal margin.

Ventral surface. The numbers and arrangement of the ventral setae are similar to those

of D. ringwoodensis.

Legs. Leg I, free from the lamellae and the rostrum, is readily visible. Legs I and II
each bear a heavy blunt spinous seta on the genus and on the tibia. The genus of leg IV
also bears a similar heavy spinous seta. The tarsus of leg II bears a branched seta which
is glovelike in appearance. The distal hairs of the tarsi are sharply pointed.

Materials examined. One female specimen designated as holotype. Collected by me from
a sample of grasses and soil at 623 Highland Road, Ithaca, New York, on August 29,
1940. The type specimen will be deposited in the collection of the New York State Museum
in Albany, New York.

REMARKS

The two new species of Dentachipteria may be separated readily by the
concealed first pair of legs in D. ringwoodensis and by the presence of 1
to 14 or more denticles on the distal margin of the pteromorphs to only 1
denticle in D. higlilandensis. A lenticulus is present in D. highlandensis.

Dentachipteria species closely resemble species of Parahypozetes Hammer
1967 and will key in Balogh (1972) to both Parahypozetes and to Austra-
chipteria Balogh and Mahunka 1966. I sent drawings of Dentachipteria
species to Dr. Hammer and she has indicated her belief that the specimens
described do represent a new genus. The following are some of the reasons for
considering Dentachipteria a new genus. In Parahypozetes both the lamellae
and the lamellar hairs point forward. The tips of the lamellae are always seen
in dorsal view. The anterior end of Dentachipteria has a blunt squared off
appearance due to the ventral bending of the gnathosome. In Parahypozetes
the pteromorphs flair outward, but bend sharply ventrally in Dentachipteria.
The distal margins of the pteromorphs are not dentate in known species of
Parahypozetes. The ventral setae in particular are much longer in the new
genus.

On the question of the position of the genus in its correct family I am
indebted to Dr. Hammer, and to Dr. E. Piffl of the Zoologisches Institut of
the University of Vienna, Austria who was referred to me by Dr. Hammer.
Dr. Hammer had sent a specimen of Parahypozetes bidentatus to Dr. Piffl
for comparative study. Dr. Piffl considers Parahypozetes distinct from Austra-
chipteria Balogh and Mahunka but none the less considers Parahypozetes a

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member of the Achipteriidae rather than of the Ceratozetidae as originally
described. According to Dr. Piffl a study of the immature forms is necessary
for a definitive classification of the groups. Since the specimens which I have
described appear to be closely related to the genus Parahypozetes considered
by Dr. Piffl to be a genus of the Achipteriidae I have placed the specimens
described in the Achipteriidae and have named them accordingly.

Literature Cited

Balogh, J. and S. Mahunka. 1966. New Oribatids (Acari) from Australian Soils.
Folia Ent. Hungarica, 19 (33): 553-568.

Balogh, J. 1972. “The Oribatid Genera of the World.” Akad. Kaido, Budapest: 1-188,

71 plates.

Hammer, M. 1967. Investigations of the Oribatid Fauna of New Zealand, Part II.
Biol. Skr. Dan. Vid. Selsk., 15(4): 1-64, 40 plates.

Vol. LXXXII, September 1974

183

Two New Tabanidae from Southeastern United States (Diptera)

L. L. Pechuman

Department of Entomology, Cornell University, Ithaca, New York 14853
Received for Publication April 8, 1974

Abstract: Asaphomyia floridensis from Highlands County, Florida, is described as new.
Asaphomyia includes only one other described species, A. texensis Stone, known from three
counties in Texas. Chrysops dixianus, a species related to Chrysops pudicus Osten Sacken,
is also described as new; specimens were seen from Virginia, North Carolina, South Carolina,
Florida, Alabama, Mississippi, and Louisiana.

Introduction

With really distinctive species, there is always a temptation when making
determinations to spend little time looking at them. In the Tabanidae, I know
of no Nearctic species more distinctive than Asaphomyia texensis Stone (1953).
When specimens of what seemed to be this species, known only from three
counties in Texas, were collected in Florida, I considered it only an interesting
extension of range. It was only when I began a comparative study of the an-
tennae of the Florida specimens that I found they represented a species quite
distinct from the one in Texas.

For some years I had noted in routine identifications and had found in some
collections under Chrysops pudicus Osten Sacken a southeastern Chrysops that
was undescribed. I had hoped for the male of the species before describing it
as new but now a name is needed for some manuscripts in preparation covering
both a faunal study and the immature stages.

The types of both are retained in my collection for the present.

Asaphomyia floridensis, n. sp.

Holotype $. Length, 11 mm. Wing, 10 mm.

Head. First 2 antennal segments brown, each about as long as wide, with short black hairs;
third antennal segment with basal annulus almost round, as wide as first segment, brown,
slightly paler at base with a few black hairs and many very short silver hairs ; remainder
of annuli brown, in form of a style % width of first annulus at base and tapering to % of

Acknowledgments: The loan of a $ paratype of Asaphomyia texensis by Pedro Wygodzin-
sky of the American Museum of Natural History and a comparison of the holotype $ of
A. texensis with the holotype $ of 4. floridensis by George Steyskal of the Systematic
Entomology Laboratory, U.S.D.A., is greatly appreciated. Specimens which made this
study possible were received from the following: T. R. Adkins, R. G. Beard, W. B. Ezell,
G. B. Fairchild, S. W. Frost, J. T. Goodwin, H. M. Henry, J. E. Lloyd, D. C. Sheppard,
R. E. Silberglied, M. A. Tidwell, and R. L. Watson.

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width near apex; the style of the right antenna has six apparent segments and that of
the left five such segments and, in both, the last segment is equal in length to the total of
the preceding style segments; the last style segment has a tuft of stiff black hairs at the
tip. Frons brown pollinose with no trace of calli, 1.8 times as high as width below and
slightly widened below; the ocelli are on a low, brown pollinose tubercle; vertex behind
ocelli with a clump of stiff black hairs and a row of shorter black hairs which rim upper
occipital margin. Clypeus and genae brown with black hairs. Beard black. Palpi dark
brown, second segment stout at base tapering to a truncate apex, both segments with
long black hairs. Proboscis shorter than palpi, brown with black and brown hairs.

Thorax. Dorsum brown, unstriped, with a few black hairs and many recumbent golden
hairs. Pleurae uniformly brown. Legs brown, mostly brown and black haired with some
scattered golden hairs; hind tibial spurs short. Wings uniformly brown; bifurcation of
third longitudinal vein with a long appendix.

Abdomen. Rather uniformly brown dorsally and ventrally except seventh segment a
darker shade of brown and incisures of second and third tergites slightly paler ; with many
dark brown and golden brown hairs.

Allotype {$). Length, 10.5 mm. Wing, 10 mm.

Head. Antennae similar to $ except basal portion of third segment a bit narrower and
slightly paler in color; 5 apparent segments in the style of each antenna. Frontal triangle,
cheeks and genae dark brown pollinose, the latter and beard with long black hairs. Ocelli
on a slightly raised, grayish brown pollinose tubercle, which posteriorly has a tuft of long
stiff black and golden brown hairs. Palpi dark brown, second segment stouter and more
acutely tapered than in $ , with long black hairs. Proboscis subequal to palpi, dark brown
with dark hairs.

Thorax. Dorsum, pleurae, wings, legs, and halteres as in $ except fewer golden hairs on
dorsum and legs.

Abdomen. Incisures of second, third, and fourth tergites a little paler than in $ ; fifth
and following tergites darker brown than anterior tergites. The porportion of golden to
dark hairs is greater than in $ .

Holotype and Allotype. Archbold Biological Station, Lake Placid, Highlands County,
Florida, 7 June 1966, 15 w. UV blacklight (R. Silberglied) .

Paratypes. AS S Archbold Biological Station, Highlands Co., Florida: 20 May 1968

(S. W. Frost), 8 June 1966 (Robert G. Beard), 20 June 1966, 15 w. UV blacklight (R.
Silberglied); 2$ S 2 mi. NE of intersection of rte 70 and Fla. 27, Highlands Co., Florida,
oak palmetto scrub, 9 July 1969 and 8-9 July 1969; the latter specimen carries notation
“ ‘asleep’ on twig of shrub 1 m. high.”

Paratypes will be deposited in the collections of Cornell University, U.S. Museum of
Natural History, and G. B. Fairchild.

Variations. The paratypes range in length from 9 to 12 mm with an average of 10.4 mm.
The apparent segments of the antennal style range from 3 to 6 and these vary in the same
specimen in number and distinctness; in all cases the terminal annulus is longer than any
of the others.

A. floridensis is a more slender-appearing insect than A. texensis and differs in a number
of characters: The tubercle on which the ocelli rest is less raised and is pollinose, including

Vol. LXXXII, September, 1974

185

Fig. 1. Wing of Chrysops dixianus, n. sp.

the area between the ocelli, in considerable contrast to the high, shining tubercle in
texensis ; the antennal style is less slender, darker, and more likely to be subdivided. When
viewed laterally the occiput is practically invisible, whereas in texensis it is wide and
conspicuous; many of the recumbent hairs on the dorsum of the thorax are golden rather
than black; the wings are uniformly brown rather than darker anteriorly. The wings of
floridensis are narrower, the ratio of greatest width to greatest length in the $ holotype
being 1:3.30 and in the $ $ ranging from 1:3.14 to 1:3.53 with an average of 1:3.34,
whereas in texensis the ratio is 1:2.41 in the $ holotype, 1:2.77 in a $ paratype and
1:2.64 in a $ paratype.

The $ holotype of A. texensis is from Columbus (Colorado County), Texas, and
carries no collection date. The type series included a $ from Victoria (Victoria County),
Texas, 3 May 1913 and 2$ $ and 2$ $ from Weser (Goliad County), Texas, 11 May 1952.
No specimens from Texas or elsewhere have been reported subsequent to Stone’s description
and it is somewhat of a surprise to encounter a second species in Florida.

Chrysops dixianus, n. sp.

Holotype ($). Length-. 8.25 mm (Fig. 1).

Head. First antennal segment yellow, second a deeper shade of yellow, basal portion of
flagellum yellow-brown, annuli dark brown, nearly black; first two segments with black
hairs. Frons grayish yellow pollinose with a scattering of fine yellow hairs, most dense near
the vertex; frontal callus yellow, % as high as wide, somewhat pointed above. Fronto-
clypeus shining yellow with no dark spots; cheeks shining yellow below, yellow pollinose
above. Palpi dark yellow with short black hairs and a few longer yellow hairs. Proboscis
dark brown.

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Thorax. Dorsum dark brown in ground color with a median brown pollinose stripe which
is flanked by grayish yellow pollinose stripes ; 2 sublateral subshining brown stripes
merge with the median stripe near the scutellum ; the stripes immediately above the wing
bases are grayish yellow pollinose; scutellum dark brown with apical half orange-brown.
Pleurae dark brown with 2 broad yellow pollinose stripes. Hairs of thorax pale yellow.
Halteres dark brown. Forelegs with coxae and femora yellow, the latter somewhat darker
at apex, tibiae with basal half yellow, apical half and tarsi dark brown ; middle coxae
dark brown, femora, tibiae, and metatarsi yellow, balance of tarsi dark brown; hind
coxae and basal 4/5 of femora brown, apical % of femora and basal % of tibiae yellow-
brown gradually shading in the latter to dark brown, metatarsi and adjoining segment
yellow-brown, balance of tarsi dark brown. Hairs on legs match the ground color of the
integument. Wing as figured; hyaline triangle not quite reaching second longitudinal
vein, fifth posterior cell mostly hyaline, apical spot crossing slightly more than half of
the upper branch of the third longitudinal vein.

Abdomen. First tergite yellow shading to yellow-brown beneath scutellum; second tergite
yellow, the anterior half entirely so, the posterior half with a dark brown marking in
the shape of a flattened inverted “V” which continues as a dark shadow to the posterior
margins of the segment where the color is intensified to form a small brown spot ; third
tergite with a dark brown band, shading to chestnut brown laterally, covering the anterior
half of the segment, shallowly indented by the yellow posterior border of the segment;
fourth tergite similar to third but indentation even shallower and with chestnut brown
portion more extensive ; fifth and following tergites dark brown with a grayish yellow
posterior border. Venter pale yellow with a vague indication of a dark median spot on
third and fourth sternites, such a spot distinct on fifth sternite, sixth sternite dark brown.

I have seen two damaged specimens of what may be the male of this species but it
seems advisable to withhold a description until specimens in better condition are available.

The name dixianus is derived from the area in which the species is found, known in
the vernacular as “Dixie.”

Holotype. Wedge Plantation, McClellanville, South Carolina, 28 May 1970 (LLP).

Paratypes. Virginia: Sussex Co, 8 June 1973 and Greensville Co., 20 June 1973 (Steve
Jones). North Carolina: Williamston, 8 July (G. Fairchild). South Carolina: Wedge
Plantation, McClellanville, 28 May 1970 (LLP) ; Hobcaw (Baruch) Plant., Georgetown, 29
May 1970 (Pechuman & Burton) ; Sumter, 24, 25 June 1970 (T. R. Adkins, Jr.) ; Boykin,
Sumter Co., 27 June 1968 (W. B. Ezell, Jr.) ; Sumter Co., 27 July 1971 (D. C. Sheppard) ;
Sweden, Orangeburg Co., 2 July 1968 (Adkins, Ezell, Krebs) ; Marlboro County, 5 June 1970
(Sheppard) ; Berkeley Co., 3 July 1970 (T. R. Adkins, Jr.) ; Berkeley Co., 1 July 1960, 6, 27
July 1971 (D. C. Sheppard). Florida: Cody, 18 May 1935; Wacissa, 5 June 1935; Green-
ville, 12 June 1935; Highlands Hammock St. Pk., Highlands Co., 11 May 1965 (LLP) ; Welaka,
11, 26 May 1961 (A. & H. Dietrich) ; Levy Co., 2 June 1960 (F. S. Blanton) ; Wakulla Springs,
5 July 1950 (A. G. B. Fairchild) ; Gainesville, Alachua Co., 8 May 1965, 21 May 1964 (J. E.
Lloyd) ; Jackson Co., 31 May 1965 (F. J. Moore) ; 3 mi. SW of Cantonment, Escambia Co.,
22 May 1965 (Ray Tidwell). Alabama: Blue Girth Creek, Dallas Co., 19 June, 18 August
1964 (R. L. Watson) ; Bear Creek, Autauga Co., 3 August 1966 (Hays and Watson). Missis-
sippi: Logtown, Hancock Co., 23 June 1966 (Diamond and Bradford). Louisiana: Approx.
1 mi. S. of Pearl River, St. Tamany Par., 15 June 1969 (Mac Tidwell).

Paratypes will be deposited in the collections of: American Museum of Natural History,
Auburn University, British Museum (Natural History), Canadian National Collection,
Clemson University, Cornell University, Florida State Collection of Arthropods, Museum

Vol. LXXXII, September, 1974

187

of Comparative Zoology, Ohio State University, Pennsylvania State University, SUNY
College of Environmental Science and Forestry, U.S. Museum of Natural History, John
F. Burger, W. B. Ezell, G. B. Fairchild, J. T. Goodwin, C. B. Philip, R. H. Roberts, D. C.
Sheppard, and Mac A. Tidwell.

Variations. The series of specimens is quite uniform. Length varies from 7.5 to 9 mm
with an average of 8.25 mm. The wing pattern is uniform and the characters of the
head and thorax show only slight variations ; the proportion of brown and yellow-
brown on the hind legs differs to some extent and the pale thoracic stripes in a few
specimens have a greenish tinge. The dark marking on the second abdominal tergite
in a few specimens is composed of two dashes connected by a brownish shadow; in others
it is more extensive than in the type, nearly reaching the lateral margins of the segment.
The size and intensity of the dark median markings on the venter differ but in all cases
the first and second sternites are completely yellow.

As mentioned above, Chrysops dixianus was found in several collections under C.
pudicus. From that species, dixianus may be separated by the brown thorax, pale tipped
scutellum, broader apical spot, hyaline triangle not reaching the second longitudinal vein,
completely yellow frontal callus, no distinct dark spot on abdomen under the scutellum
and pale median markings of second and third abdominal tergites broader and less distinct.
C. pudicus is a variable species and occasional specimens have a broader than usual apical
spot and/or hyaline triangle not reaching the second longitudinal vein; the other characters
mentioned above were found to distinguish dixianus from these specimens.

Chrysops dixianus will run to couplet 46 in my recent (1973) key to the species of
Chrysops found in Virginia. A modification of this portion of the key to include dixianus
follows:

46. Abdominal markings black and median marking of second segment usually reaches

anterior margin; frontal callus normally black but sometimes yellow; usually

at least basal portion of hind femora black dimmocki Hine

Abdominal markings pale to dark brown, sometimes evanescent; median marking
of second abdominal segment rarely attains anterior margin ; frontal callus
yellow; hind femora yellow to brown 47

47. Thorax greenish-gray with fuscous stripes; outer margin of crossband usually

sinuous celatus Pechuman

Thorax brown or yellowish in ground color with brown stripes; outer margin of
crossband concave, straight, bowed, or sinuous 48

48. Dark median marking of second abdominal segment reaching about % across

segment ; outer margin of crossband usually straight or somewhat concave ;

hind femora yellow jlavidus Wiedemann

Dark median marking of second abdominal segment reaching only about half-way
across segment; outer margin of crossband frequently bowed or sinuous; hind
femora partly or all brown - 49

49. Apical spot occupying upper half of second submarginal cell and sharply outlined;

fifth posterior cell largely hyaline; smaller species averaging 8.25 mm dixianus , n. sp.
Apical spot indefinite in outline, extending into lower half of second submarginal
cell as a paler infuscation which may continue into apical portions of first,
second, and third posterior cells; fifth posterior cell largely infuscated; larger
species averaging 9.5 mm reicherti Fairchild

That Chrysops dixianus can be a common pest is indicated by 124 specimens
collected on 6 July 1971 in Berkeley County, South Carolina, by D. C. Sheppard.

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Literature Cited

Pechuman, L. L. 1973. The insects of Virginia No. 6. The horse flies and deer flies
of Virginia (Diptera: Tabanidae). V. P. I. and State Univ. Research Div. Bull.,
81: 1-92.

Stone, Alan. 1953. New tabanid flies of the tribe Merycomyiini. Wash. Acad. Sci. J.,
43(8): 255-258.

BOOK REVIEW

The South Asiatic Olethreutini (Lepidoptera, Tortricidae) . A. Diakonoff. Zool.
Mon. Rijksmuseum van Nat. Hist. No. 1. Brill, Leiden. 1973. XXI + 699 pp., 15 pis. (1
col.), 732 figs. 208 guilders.

This is a highly important taxonomic monograph of the southern Asiatic members of a
large, worldwide group treated by various authors as a tribe, a subfamily, or even a family.
It is based on all known material in the collections of the world. The author is a recognized
authority on these and related moths, on which he has published voluminously. He himself
lived and collected in Java for many years. The present work is especially valuable since
the Palaearctic and Australian faunas are being intensively studied by other authors. The
Nearctic fauna, long overdue for revision, is also being studied. The author’s opinions of
the taxonomy of the larger taxa will therefore be especially important. The present work
covers 12 subtribes, 94 genera, 17 subgenera, 430 species, 14 subspecies, and 2 “formae.” Of
these, 11 tribes, 39 genera, 3 subgenera, 176 species, and 7 subspecies are described as new.
Many new combinations are also made. Keys to these taxa are given, based on all usable
characters, and very thorough descriptions of all taxa are included. Dates, localities, and
institutional locations of specimens are given, including, of course, types when these are
known. Both male and female genitalia are figured whenever possible, as well as many
heads and venations. Food-plant records are also given for many species.

A preliminary section contains, among other things, a discussion of the general classifica-
tion of the Tortricidae, past and present, and of the morphology of certain genitalic and
scent organs. A discussion of the Palaearctic genera is given for comparison. A new term,
“apallotype,” is proposed for a supplemental type of the opposite sex from the type, a cate-
gory sometimes confusingly, called “neallotype.” It is hardly necessary to state that this
is a taxonomic work of the highest quality, one that will be essential for all students of
this and related groups anywhere in the world.

Alexander B. Klots

The American Museum of Natural History

Vol. LXXXII, September 1974

189

The Distribution of Brood Ten of the Periodical
Cicadas in New Jersey in 1970* 1

John B. Schmitt

Rutgers — The State University, New Brunswick, N. J.

Received for Publication April 26, 1974

Abstract: The last thorough study of the distribution of Brood X of the periodical cicadas
in New Jersey ( Magicicada spp.) was made in 1902. Data collected on the distribution
of the 1970 emergence indicates a disappearance since 1902 from the following localities:
Mercer County except Princeton; eastern Somerset County; Prospertown-Colliers Mills,
Ocean County; Jacobstown-Ellisdale, Burlington County; Cherry Hill Township, Camden
County; Salem and Woodstown, Salem County; and Shiloh in Cumberland County.
Hitherto unreported populations were found on Lower Powhatcong Mountain, Warren
County; near Middletown, Monmouth County; and Quinton and Alloway in Salem
County. Forty populations were found in Hunterdon County, west and south of the
South Branch of the Raritan River. The chief factors in the disappearance of the insect
since 1902 appear to have been the destruction of woodlands, forest fires, and urbanization.
The possibility that forest losses caused by the gypsy moth may play a part in the loss
of periodical cicada populations is suggested.

INTRODUCTION

The periodical cicadas are well known for the fact that adults of the six species
emerge from the soil after either 13 or 17 years of nymphal existence. Alexander
and Moore (1962) provide a table showing past emergence dates since 1621 and
predicting the future emergence dates until the year 2028, of all known broods
of both the 13-year and the 17-year species. A “brood” may be defined as con-
sisting of all the populations of the species complex (either 13 -year or 17-year)
emerging in any year. Since the years of emergence follow a well-defined cycle,
the various generations of a brood may thus be recognized and identified by a
Roman numeral. In New Jersey, six broods of the 17-year species were known
to exist in the early decades of this century (Weiss, 1916; Davis, 1926). As
predicted, adults of Brood X appeared in 1970. The preceding years of emer-
gence of Brood X in this century were 1902, 1919, 1936 and 1953.

Acknowledgments: At various points in this paper, the writer has endeavored to recognize
the individuals whose generous participation made possible a more complete or more
satisfying solution of some questions regarding Brood X in 1970. The contributions of
two individuals, however, should be especially acknowledged. I am particularly indebted
to Dr. Lyle E. Hagmann and Mr. Joseph D. Stewart of the Department of Entomology
at Rutgers for their very considerable help.

1 Paper of the Journal Series, New Jersey Agricultural Experiment Station, Cook College,
Rutgers University — The State University of New Jersey, Department of Entomology and
Economic Zoology, New Brunswick, New Jersey 08903.

New York Entomological Society, LXXXII: 189-201. September, 1974.

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New York Entomological Society

In 1969 the writer reviewed the literature which had been published on the
distribution of Brood X in New Jersey since Smith (1903) described its distri-
bution in the emergence of 1902. Weiss (1916) added nothing to Smith’s data.
Marlatt (1907) also described the 1902 emergence, and lists several counties
and localities not cited by Smith. However, Smith was aware of these additions
through correspondence with Marlatt, and in his 1903 report comments on them
substantially as follows:

Middlesex County. Marlatt cited a report from Deans which stated that the
cicadas occurred “by the millions.” Smith says he searched the area and found
no trace of them.

Morris County. Marlatt reported them from Boonton. Smith searched through
Morris County, and especially Boonton, without being able to verify the record.

Gloucester County. Marlatt’s record, according to Smith, was based on a news-
paper report of an occurrence in the Swedesboro-Harrisonville area. Smith found
no trace of the insect in Gloucester County. Although Smith failed to confirm
Marlatt’s records, the writer made a special effort to find these populations in
1970, without success.

Davis (1926) considerably extended the list of counties in New Jersey over
that provided by Smith for Brood X. However, a careful reading of Davis’
paper shows that he based his additions solely on the annual report of the
Department of Entomology of the New Jersey Agricultural Experiment Station
for 1919 (Headlee, 1919). Unfortunately, these records did not mean that
actual specimens were received or identified, but only that correspondence con-
cerning the cicada was received from residents in the various counties. Much
of this correspondence was dated in months of the year when the insects were
underground, and it seems very probable that such correspondence was prompted
by newspaper accounts predicting the forthcoming emergence of the cicadas.
However, as in the case of Marlatt’s records, an intensive effort was made in
1970 to determine whether Brood X exists in the disputed counties. No litera-
ture could be found regarding the emergences of 1936 and 1953 which extended
the distribution described by Smith in the 1902 emergence.

The writer decided to undertake a thorough study of the 1970 emergence to
learn what, if any, changes in distribution had taken place since 1902, a period
of time representing four cicada generations. It was obvious that some measure
of public assistance in finding local populations would be valuable. Accordingly,
in the spring of 1970 news stories alerting the public to the coming of the cicadas
were distributed to the newspapers of the State through the courtesy of the
Communications Center of our State Cooperative Extension Service. The writer
also sent a personal letter to each county agricultural agent asking for records
of cicada emergence and explaining the purpose of the study. A similar appeal
was sent to the superintendent of each county mosquito control agency. Col-

Vol. LXXXII, September, 1974

191

leagues in the Department of Entomology and Economic Zoology at Rutgers
were also reminded of the predicted emergence and their cooperation sought.
During and after the emergence period, the writer made a number of field trips
to check on distribution, and a record of adult cicada distribution in 1970 was
thus obtained which the writer believes is fairly complete. This paper will
compare that record with the observations of Smith (1903). It should perhaps
be noted that in the study of these cicadas, it is the existence of large local
populations that is significant, not the occurrence of individuals separated from
a large population. Such large local populations typically contain many thou-
sands of individuals of both sexes, and their presence is advertised by the daytime
din of their song and by oviposition injury to deciduous trees.

OBSERVATIONS

Figure 1 summarizes the distribution of Brood X in New Jersey in 1902
(Smith’s data) and in 1970. Isolated localities known to Smith are marked by
circles. If cicadas appeared in a given locality in 1970 also, the circle is solid;
if cicadas could not be found in 1970, the circle is open. The squares represent
populations seen in 1970 in localities apparently not known to Smith. The
numeral accompanying each locality marker serves to identify the locality in
the text.

The large, lightly-shaded area in the central-western part of the map indicates
the general distribution of Brood X in 1902 in that area. The smaller, heavily-
shaded area represents the general distribution of Brood X in the area in 1970.
The distribution of individual populations in 1970 in most of that area (Hunter-
don County and adjoining areas) is shown in Figure 2 as numbered circles. These
localities are also geographically identified by number in the text.

The isolated populations indicated in Figure 1 will be identified first. In
1902 Smith reported, from correspondence, a population at Roxbury, in Warren
County. His map shows it extending inland from the Delaware, a few miles
south of Belvidere. In 1970, the writer did find a small population at the
western end of the area indicated by Smith, near Harmony Station (Fig. 1, 1).
Roxbury itself is at the northern end of Scott’s Mountain. No cicadas were found
at Roxbury, but near the village of Montana, a few miles south, they were very
abundant (Fig. 1, 2).

Other Warren County populations were observed in 1970 at Stewartsville and
New Village. These localities are part of the Lower Powhatcong Mountain
forested area (Fig. 1, 3). Apparently Smith did not know of this locality;
neither his text nor his map indicates it. Smith did list Carpentersville, Warren
County, and a few cicadas were found there. However, the Delaware River at
this point represents a gap in the woodland of no more than fifty yards. On the
Pennsylvania side there was a large population (Fig. 2, 50); probably the
cicadas seen at Carpentersville were only strayed individuals from the Pennsyl-

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New York Entomological Society

Fig. 1. Map of New Jersey showing distribution of Brood X in 1902 and 1970. See
text for details.

Vol. LXXXII, September, 1974

193

vania population. Finesville, also in Warren County, is on the border of the
great Hunterdon distribution (Fig. 2, 15).

Sussex County, just north of Warren, is one of the counties cited by Davis
(1926) on the basis of the Headlee report (1919). Since it is heavily forested
with oak, particular attention was given to the area. County agent John W.
Raab, at the request of the writer, spent considerable effort in inquiries and
travel, without finding a single population. The writer toured the localities cited
by Headlee without uncovering any evidence of cicadas, and no correspondent
provided knowledge of a single population. Neither could any resident be found
who remembered ever hearing or seeing the insects within the county. Similar
efforts in both Morris and Passaic Counties were equally unsuccessful. Brood II
is well known to residents of these two counties, but Brood X is not. The County
agents of both of these counties, and of Bergen, Union and Essex also could not
find a single instance of Brood X, nor could the writer.

As Figure 1 shows, the locality of Princeton (Fig. 1, 4) was included in the
general distribution of cicadas in Mercer County in 1902. Smith found the in-
sects to occur abundantly as far south as the Pennsylvania Railroad main line,
and westward along that line to Lawrence Station. He concluded that the
cicadas were “pretty generally distributed” in Mercer County “except in the
extreme south.” In 1970, Mr. Charles M. Holmes, senior county agent, supple-
mented the writer’s observations, and was unable to find any cicadas in Mercer
County other than the large population on the western edge of Princeton Borough,
and along the Mercer-Hunterdon line (Fig. 2, 38, 42). Also, no populations
could be found in either Somerset or Middlesex County representing the former
eastward extension of the great central area shown in Fig. 1 . The only Somerset
populations found were on the Sourland Mountain ridges, to be described later.

In Monmouth County, a population in the Navesink Highlands (Fig. 1, 5)
had been described by Smith. In 1970, its decendents were very abundant in
the same locality, and as the area has apparently changed very little during this
century, the insects were probably nearly as abundant as they were in 1902.
Individual cicadas were found in Fair Haven, separated from the Navesink
Highlands population by about one-half mile of open water, but as no evidence
of emergence could be found at Fair Haven, the writer assumes that these speci-
mens were strays from the Highlands population. A second population was found
in Monmouth County near Middletown, in the low wooded hills known as the
Telegraph Hill formation (Fig. 1, 6). Smith apparently did not know of this
colony, located about eight miles from the Navesink Highlands population.

Smith recorded a colony in 1902 in the northwestern corner of Ocean County,
between Collier’s Mills and Prospertown (Fig. 1,7). In 1970 no trace could be
found of that colony. This area is heavily wooded, with few access roads or
human inhabitants. Searches and inquiries in both this area and in the nearby
town of New Egypt failed to provide any evidence of the cicadas. It therefore is

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probable that this colony has perished. One possible explanation is the fact
that the area between Collier’s Mills and Prospertown has been devastated by
a number of very severe forest fires since 1902. The resulting tree loss might
have destroyed the cicadas. However, the apparent disappearance of another
population reported by Smith in nearby Burlington County, of which no descen-
dents could be found in 1970, could not be accounted for. Smith reported (from
correspondence) a population between Jacobstown and Ellisdale (Fig. 1, 8).
Unable to find the cicadas himself, the writer enlisted the aid of Mr. Daniel
Kensler, who had been the county agricultural agent in Burlington County for
almost 40 years. Despite strenuous efforts on his part, no trace of this colony,
either in 1970 or in the past, could be found; either it has become extinct, or
Smith’s correspondent was in error.

A second locality in Burlington County was described by Smith from the
vicinty of Indian Mills. The writer was unable to find any trace of the insects or
reports of them, and he is indebted to Dr. Lyle E. Hagmann for finally dis-
covering them. They were found about halfway between Indian Mills and
Tabernacle, about one mile east of Route 206 (Fig. 1, 9). The precise location
was scaled from a Geological Survey map as 39°48'45" N and 74°42'30" W.
It is a large colony, as more than 100 acres of trees showed evidence of ovi-
position.

The only colony of Brood X in Camden County in 1902 was recorded by
Smith from Delaware Township, since renamed “Cherry Hill” Township. This
area is now highly urbanized. No trace of the colony could be found in 1970,
either by the county agent or by the writer. It is probable that the destruction
of woodlands since 1902 has destroyed this colony.

As regards Gloucester County, Smith mentions correspondence with Marlatt,
who, he says, sent him newspaper reports of the insects near Swedesboro and in
the woods between Harrison ville and Swedesboro. In 1970, however, no trace of
either of these colonies could be found.

Smith reported two colonies from Salem County in 1902. One of these was
described by a correspondent from the town of Salem as “occupying a large tract
of timber land which is, unfortunately, gradually becoming exterminated”
(Fig. 1, 11). A second colony was described as being near Yorktown (Fig. 1,
12). Mr. Robert Gardner, the county agent in Salem County, became keenly
interested in the matter of cicada distribution, and expended a great deal of
effort in trying to find populations in 1970. Only two were found. One of these
inhabited a woodland near the “Happy Hill” Nursery, Alloway (Fig. 1, 13),
and the other was discovered near Quinton (Fig. 1, 14). No trace of the York-
town colony could be found (Yorktown is about five miles from Alloway).
Whether either of the two populations that were found represents the Salem
colonies described by Smith is uncertain because of the vagueness of Smith’s
record, but if the locality was near either Alloway or Quinton, it is perhaps odd

Vol. LXXXII, September, 1974

195

that his correspondent did not use those names, as both communities are old
and well-known localities.

Smith reported a single record in 1902 from Cumberland County, in the
vicinity of the village of Shiloh (Fig. 1, 15). No trace could be found of this
colony in 1970. Here again the writer was very fortunate in the fact that Mr.
Kenneth E. Pickett, the county agent of Cumberland County, has lived most
of his life in Shiloh and took a keen interest in the matter. Despite all his efforts,
he could not find any trace of the Shiloh colony. No other evidence of Brood X
in Cumberland could be found by Mr. Pickett or the writer.

Turning to the northwestern area of the state, Smith in 1902 found Brood X
existed from just north of Trenton along the Delaware River upstream to a
point just south of Phillipsburg in Warren County (Fig. 1). Eastward, the
cicadas were found by Smith as far east as Bound Brook. The extent of Brood
X distribution in Warren and Mercer Counties has already been considered.
The disappearance of the cicadas from their eastern range in Somerset County
is indicated in Figure 1.

In Hunterdon County, Smith described the insect in 1902 as “generally
present from the Delaware River east to the line of the Central Railroad of
New Jersey, and from the Mercer County line north to the Warren County line.”
The 1970 emergence in Hunterdon was found to be concentrated in three well-
defined physiographic areas:

1. The Musconetcong Mountain and ridge area: populations numbered 1

through 21 (Fig. 2). This is a mountain of granitoid gneiss of Precambrian
age and is heavily forested. Just south of the mountain in Union and
Alexandra townships, there is a forested ridge area of rubbly, glaciated soils
derived from the gneiss. These forests were also heavily populated by the
cicadas.

2. The Hunterdon Plateau; populations numbered 24 through 30. This is an
area of hard sandstone and argillite west of Flemington, about 8 miles wide
at Baptistown. The southern portion of the plateau is heavily wooded,
primarily because poor soil drainage tends to discourage agriculture, and it is
this area which supports the cicadas.

3. The Sourland Mountain; populations numbered 33 through 42. This is a
ridge of crystalline rocks, mostly diabase, which extends from the Delaware
River to the vicinity of Belle Meade in Somerset County, a distance of about
16 miles. Its western part is a series of forested hills, but its eastern area,
which extends into Somerset County, is a continuous plateau. Much of the
land is wooded because of steep slopes and stoniness. Populations of the
cicada extend a few miles east of the distribution shown in Fig. 2, into
Somerset County as far as Belle Meade.

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Fig. 2. Map of Hunterdon County and adjacent areas showing distribution of Brood X
in 1970. See text for details.

The western boundary of Hunterdon County, along the Delaware River, is in
most places a steep escarpment, usually wooded. Smith described the 1902 emer-
gence along the river in these words: “Running south along the Delaware, the
Warren County area of infestation extends into Hunterdon County and for its
full length. It is broken, of course, at several points, notably at towns and settled
areas, but practically the ridge back of the river is all cut by the Cicada.”
(His expression “cut by the cicada” refers to oviposition injury to trees.)

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197

In 1970, very little cicada activity could be found along this same route.
South of Milford, only three populations were found. One of these, No. 25, was
found one mile north of Byram, and another, No. 31, was at Raven Rock but
no other evidence could be found of the extensive emergence described by Smith
as occurring between Raven Rock and Tumble Falls. The third population,
No. 33, was found at Goat Hill, on the Hunterdon County line, at the end of
the Sourland outcrop.

The total list of Hunterdon county localities is as follows (Fig. 2):

Hunterdon County Localities
Figure 2

No. Locality

1. Musconetcong Mountain, north of Polktown

2. Musconetcong Mountain, route 41, 1 mile south of Bloomsbury

3. Ridge, 3 miles south of Bloomsbury, on the Pattenberg-Bloomsbury road

4. Musconetcong Mountain, portion known as “Bloomsbury Mountain”

5. Ridge, west of route 579

6. 7. Along township road, south of Hickory Corner

8. West of route 579 at Mechlin’s Corner

9. One mile west of Pittstown

10. Little York

11. Musconetcong Mountain, north of Riegel Ridge

12. Musconetcong Mountain, on the south side of route 519

13. East of route 519, between Riegel Ridge and Spring Mills

14. One mile west of Spring Mills, off the Amsterdam road

15. Musconetcong Mountain, Finesville (Warren County)

16. Along Delaware River, Musconetcong Mountain (Mt. Joy)

17. Along Delaware River, Musconetcong Mountain (Riegelsville Curve)

18. Along Delaware River, Milford-Holland road, continuous for 1 mile

19. Hickory Corner, east side of route 579

20. East side, route 579, north of Mechlin’s Corner

21. East side, route 579, Mt. Salem

22. Everittstown, south side of route 513

23. Kingwood Township, route 519, 2 miles south of Baptistown

24. Kingwood Township, east of route 519, along Kingwood-Locktown Road

25. Along Delaware River, 1 mile north of Byram

26. Croton, north of state highway 12

27. Hardscrabble Hill, 2 miles west of Flemington

28. Route 579, 2 miles south of Croton

29. One mile south of Locktown

30. East of route 519, 1 mile north of Rosemont

31. Along Delaware River at Raven Rock

32. Route 523 at Sand Brook

33. Along Delaware River at Goat Hill

34. West of Hunterdon Hills Regional High School

35. West of Rocktown

36. West of Lin vale, 1 mile

37. West of route 518, Snydertown road

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New York Entomological Society

38. On the Mercer County line, west of the Wertsville-Hopewell road

39. One mile west of Buttonwood Corners

40. On the Mercer County line, Wertsville-Zion road

41. Zion (Somerset County)

42. On the Mercer County line, one mile west of Harbourton

Although Smith’s map of the 1902 emergence bears some discrepancies with
his text, it is clear that the extent of Brood X distribution in the Warren-Hunter-
don-Mercer-Somerset region has been greatly reduced in the intervening 68 years.
Whether the remaining populations have much prospect for continued existence is
an interesting question. Since the Delaware River is no more than one hundred
yards wide at Frenchtown, and becomes much narrower upstream, there is some
possibility that future reestablishment from Pennsylvania might occur if
ecological conditions permit, in the event of the loss of the Hunterdon popula-
tions. Accordingly, several field trips were made to scout for cicada populations
within five miles of the river in Pennsylvania. In a single day of field work,
16 populations were found, shown on Figure 2 as follows:

Pennsylvania Localities:

50. Raubsville

51. Riegelsville (Pa.)

52. Durham Furnace (ruins)

53. Kintnersville

54. Ferndale

55. Opposite Holland, N. J.

56. Upper Black Eddy

57. Tohickon Park

58. Ralph Stover State Park

59. Erwinna-Ottsville road

60. Tinicum Park

61. Stover Mills

62. Lumberville

63. Opposite By ram, N. J.

64. Opposite Raven Rock, N. J.

65. Solebury

The extension of Musconetcong Mountain into Pennsylvania, sometimes
called the “Reading Prong,” was reported by various correspondents to be
heavily populated by the cicadas, but no effort was made to determine their
distribution at points more than five miles from the river.

DISCUSSION

In 1902, Smith gathered his information on the distribution of periodical
cicada populations in three ways: (1) by general and professional correspon-

dence, (2) from the reports of 127 “official crop correspondents” scattered
throughout the state, and (3) by his own travels, chiefly by railroad. In 1970,

Vol. LXXXII, September, 1974

199

the relative ease of gathering information by automobile and the generous help
of colleagues and county extension agents, as well as many letters from the
general public, made the writer’s undertaking both a much easier task and,
presumably, a more thorough one. From the compared data it is at least clear
that a very marked reduction in the number of Brood X populations has taken
place between 1902 and 1970. This reduction involves both the loss of isolated
populations and a considerable reduction in the extent of the regional distribu-
tion now centered in Hunterdon County.

With respect to these losses, it is of some interest to consider a statement made
by Marlatt (1898): “The greatest check on the species has been the advent of
European man on this continent and the accompanying clearing of woodland
and the increase of settlement. The vast areas in the more-densely populated
East which were once thickly inhabited by one or the other of the broods of the
periodical cicadas, are rapidly losing this characteristic and the Cicada will
doubtlessly appear in fewer and fewer numbers in all settled districts.” Marlatt
illustrated this prophecy with an account of the fate of Brood XI in the Connect-
icut Valley, which appeared in great abundance in 1869, but seemed doomed to
virtual extinction by 1903 “as a result of the steady reduction of woodlands.”

Some of the loss of Brood X in New Jersey since 1902 was very probably due
to man’s direct interference, especially in Mercer and Somerset. The loss of
woodlands for agricultural use, however, has probably not been significant.
Rather, the development of homesites, especially as large-scale undertakings,
would appear to have been a more likely cause. Smith believed that both domes-
tic fowl and the English sparrow played a significant part in the extermination
of some populations, especially in the case of isolated counties.

The situation in Salem County, where two 1902 populations appeared to have
been lost, and two “new” populations were found in 1970, may involve nothing
more than a relocation of the 1902 populations in the four-generation interval.
Lloyd and Dybas (1966) point out that female cicadas are very prone to ovi-
posit in the young trees of an advancing forest edge. Whether such a mechanism
could result in such extensive relocations is not easily decided.

Perhaps future study of the Indian Mills population (Fig. 1,9) may provide
some information on this question. This population was found in woodlands
about one-half mile from the boundary of the State-owned Wharton Estate
and coextensive with the State lands. It is very unlikely that there will be
any future human activity deleterious to the cicadas.

The future of the Hunterdon County populations can be speculated upon only
with considerable uncertainty. The three populated regions described in this
paper (the Musconetcong Mountain and ridge area, the Hunterdon plateau, and
the Sourlands outcrops) offer so little advantage to agriculture that further
destruction of woodlands to that end seems unlikely. While there has been some
home-building in all three regions, the pressure for home sites does not appear

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to be very threatening. Also, a large part of the Sourlands area in Somerset
County has been purchased and set aside as preserved land, and may permit the
cicadas to endure in at least that much of the Sourlands.

Perhaps a more immediate threat exists in the destructiveness of another
insect, the gypsy moth. Mr. John Kegg of the N. J. State Department of Agri-
culture has kindly supplied the writer with detailed maps of defoliation caused
by the gypsy moth from 1971 through 1974. Past experience suggests that
considerable loss of oak and other deciduous trees is likely to result from such
defoliation after three successive years. The areas of heaviest defoliation in
Hunterdon County correspond very closely with the greatest concentrations
of periodical cicada populations (Fig. 2), but whether the cicada nymphs can
survive such tree losses is uncertain.

Perhaps a more immediate evaluation of the possible impact of tree loss
caused by gypsy moth defoliation on the periodical cicadas will be possible in
1979, when Brood II, the other major brood in New Jersey, would mature.
Smith (1912) compiled fairly detailed records on the distribution of Brood II
in 1911. The Wanaque Reservoir forests in Passaic County, an area heavily
populated by Brood II of the cicadas, have been studied intensively by Mr. Kegg
in an evaluation of gypsy moth activity, and have also, by virtue of being on a
protected watershed, been fairly free of human activities detrimental to the
cicadas. The fate of Brood II in this area in 1979 may therefore be of special
interest in determining the impact of tree loss caused by the gypsy moth on
the periodical cicada.

Apart from the possible effect of the gypsy moth on the Hunterdon County
cicada distribution, it should be of interest to determine in 1987, 2004, 2021,
et seq. what happens to the Hunterdon populations. It was primarily with the
hope of providing a basis for such determinations that the writer sought to locate
each Hunterdon population, although it must be admitted as quite possible that
some populations were missed. At any rate, the behavior and ecology of an insect
species with a 17-year life cycle offers an interesting challenge to the human
species.

Literature Cited

Alexander, Richard D. and Thomas E. Moore. 1962. The evolutionary relationships
of 17-year and 13 -year cicadas, and three new species (Homoptera, Cicadidae, Magi-
cicada) . Misc. Pub. Museum Zoology. Univ. Mich. No. 121; 59 pp., frontis (color)
and 10 text f.

Davis, William T. 1926. The cicadas or harvest flies of New Jersey. Circular No. 97.
N. J. State Dept, of Agriculture. 2 7 pp.

Headlee, Thomas J. 1919. Report of the Department of Entomology of the New Jersey
Agricultural College Experiment Station for the year ending June 30, 1919. 375-519.
Lloyd, Monte and Dybas, Henry S. 1966. The periodical cicada problem. I. Population
ecology. Evolution, 20: 133-149.

Vol. LXXXII, September, 1974

201

Marlatt, Charles L. 1898. The periodical cicada. Bull. No. 14, New Series. USDA
Division of Entomology. 147 pp.

. 1907. The periodical cicada. Bull. No. 71, USDA, Bureau of Entomology.

181 pp., frontis (color).

Smith, John B. 1903. The periodical cicada. Twenty-third annual report of the New
Jersey State Agricultural Experiment Station, pp. 474-489.

. 1912. The periodical cicada. Thirty-second annual report of the New Jersey

State Agricultural Experiment Station, pp. 454-466.

Weiss, Harry B. 1916. The distribution of the periodical cicada in New Jersey. Entomo-
logical News, 27: 337-340. 1 plate.

BOOK REVIEW

Tissue Culture: Methods and Applications. Paul F. Kruse, Jr., and M. K. Patterson,
Jr., eds. Academic Press, New York. 868 pp. $22.00.

This book describes the uses of tissue culture in a wide variety of disciplines. Entomolo-
gists will be particularly interested in Imogene Schneider’s chapters, “Dipteran embryos
and larvae (Diploid lines)” and “Characteristics of insect cells,” E. P Marks’ “Cockroach
and grasshopper embryo tissue,” and Arthur E. Greene and Jesse Charney’s “Invertebrate
cell cultures.” In addition, such chapters as Leonard Hayflick’s “Screening tissue cultures
for mycoplasma infections” and Michael F. Barile’s “Mycoplasma contamination of cell
cultures: Incidence, source, prevention, and problems of elimination” are of pertinent in-
terest to all engaged in attempting to grow insect cells and tissues in vitro. More than 100
authors contributed to this volume ; it should serve as a reference source for both experts
and beginners using tissue culture for years to come. Its usefulness as a guide is enhanced
by a detailed author and subject index, totaling 39 pages. Excellent illustrations of cul-
tured cells and karyotypes, as well as of specialized equipment, add to the value of this
book.

Karl Maramorosch

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New York Entomological Society

Terrestrial Mites of New York (Acarina: Prostigmata), I —
Tarsocheylidae , Paratydeidae , and Pseudocheylidae

Mercedes D. Delfinado* 1

New York State Museum and Science Service, Albany, New York 12224
Edward W. Baker

Systematic Entomology Laboratory IIBIII, Agricultural Research Service,
USDA, Beltsville, Maryland, 20705

Received for Publication June 21, 1974

Abstract: The mites here described were collected from Long Island, Lake Champlain region,
and the Mohawk Valley area, New York, in June-October of 1973. The new species described
are: Tarsocheylidae, Hoplocheylus similis, H. amerieanus; Paratydeidae, Scolotydaeus

simplex; Pseudocheylidae, Anoplocheylus transiens. Twenty-six figures are presented. The
genus Neotydeus Baker is synonymized with Scolotydaeus Berlese.

For many years New York has been a favorite collecting ground for various
arthropods, and extensive collections have been accumulated in different state
institutions (Leonard, 1928). This has not been the case with mites, however,
and our knowledge of the mite fauna of this area is almost nonexistent. A survey
of terrestrial mites was started in New York in the summer of 1973 by M. D.
Delfinado. This collection forms the basis of a proposed series of papers on the
mites of New York and neighboring areas.

The present paper deals only with the free-living or primary-feeding and
predaceous mites of the families Tarsocheylidae, Paratydeidae and Pseudo-
cheylidae. Members of these families are rather uncommon and only rarely
collected. They occur in soil, forest litter and debris, under tree bark and rotten
wood, and in moss. One species of Tarsocheylidae, however, was found under
the elytra of a passalid beetle in the Congo (Cooreman, 1951). Other Prostig-
mata collected will be dealt with in later papers.

The mites reported here were collected by the authors and M. Abbatiello from
Long Island, the Lake Champlain region and the Mohawk Valley area in June-
October 1973, by use of Tullgren-Berlese funnels from forest soil, litter and
debris, tree holes and hollow tree trunk debris.

Acknowledgments: Sincere thanks are due Michael Abbatiello and the administration of
the Biology Department at New York State University at Farmingdale, Long Island, who
generously provided laboratory facilities and space.

1 Published by permission of the Director, New York State Science Service, Journal
Series No. 159.

New York Entomological Society, LXXXII: 202-211. September, 1974.

Vol. LXXXII, September, 1974

203

Family Tarsocheylidae
Genus Hoplocheylus Atyeo and Baker, 1964

Hoplocheylus Atyeo and Baker, 1964, Bull. Univ. Nebraska St. Mus. 4: 247. Type-species,

Tarsocheylus atomarius Berlese, by original designation.

The genus Hoplocheylus has the following general characteristics of the family: presence
of dorsal hysterosomal plates and a pair of pseudostigmatic organs on propodosomal plate;
reduced palpal tarsus and absence of femoral division and pretarsal pedicels on all legs;
and presence of simple peritremes with stigmata located on the shoulders of propodosoma
as in the Tarsonemini. Atyeo and Baker (1964: 246) in a key to the genera used prin-
cipally the presence or absence of empodia on legs I (absent in Hoplocheylus , present in
Tarsocheylus ) and the structure of palpal tarsus (papilliform in Tarsocheylus , indistinguish-
able or missing in Hoplocheylus) . Seven species were known in Hoplocheylus. Two new
species are present in the collection from New York.

Hoplocheylus similis, n. sp.

(Figures 1-11)

H. similis may be distinguished from the closely related species: H. discalis Atyeo and
Baker, H. pickardi Smiley and Moser and H. americanus, n. sp. by having the distal solen-
idion short and not reaching beyond tarsal claws I ; by the forked distal setae on tarsi II-IV ;
by the narrow first medial dorsal plate with sides bulging at the level of the setae, and
by the very long posterior dorsal setae on the third hysterosomal plate surpassing the
posterior margin of the fourth plate.

Female. Length of body including gnathosoma, 574 microns. Palpus with genu and femur
completely fused and with a small inner protuberance ; tibiotarsus with 5 simple, long setae,
one rodlike solenidion and 2 small unequal subterminal spines as in figures 4 and 5.
Chelicerae fused into a single unit and truncate at apex, with 2 pairs of dorsal setae;
gnathosoma with two pairs of ventral setae, posterior pair about 3 times as long as anterior
pair. Dorsal propodosomal plate bearing a pair of clavate pseudostigmatic organs near
lateral margin and 3 pairs of dorsal setae. Peritremes as in figure 3, with stigmata on shoul-
ders of propodosoma, distal ends of tracheae converging medially between propodosomal setae.
Hysterosoma (figure 1) dorsally with 4 medial plates and a pair of lateral or humeral
plates; first dorsal plate narrow, about twice as long as wide, with sides bulging at level
of setae; second plate squarish, about as wide as long, with a pair of short setae; third plate
; slightly wider than long, with 2 pairs of unequal setae, posterior pair reaching beyond
i posterior margin of fourth plate ; fourth plate with 2 pairs of posterior setae, median pair
| about 3 times as long as lateral pairs. Dorsal anal region with a pair of terminal setae.

I Venter as in figure 2, with large rectangular hysterosomal plate, 2 elongate plates between
coxae IV each with 2 setae, and 2 large paragenital plates each bearing pair of setae.
Leg chaetotaxy as follows; the numbers represent coxa, trochanter, femur, genu, tibia and
S tarsus:

Leg I. 4-1-4-5-6 + 2-13 + 2
Leg II. 3 - 1 - 3 - 4 - 5 + 1 - 8 + 1
Leg III. 3-2-2-4-5+1-8
Leg IV. 2 - 1 - 2 - 5 - 5 - 7

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New York Entomological Society

Hoplocheylus similis , n. sp. 1, dorsal surface of female; 2, ventral surface of female;
3, peritremes; 4, dorsum of palpal tibia-tarsus; S, venter of palpal tibia-tarsus; 6, tibia
and tarsus I; 7, tibia and tarsus II; 8, tibia and tarsus III; 9, tibia and tarsus IV; 10, claws
of leg I; 11, claws and empodium of leg II.

Vol. LXXXII, September, 1974

205

Tarsus I lacks empodium ; empodia present on tarsi II-IV ; claws present on all legs ;
solenidion present on tibia I— III, absent on IV ; anterior distal seta on tarsi II-IV forked
apically; tarsi I and II with 2 and 1 short solenidia respectively; coxae III not entirely
separated from coxae IV.

Male. Not known.

Holotype. Female, collected from tree hole debris, Sunken Meadow, North Shore, Long
Island, New York, June 26, 1973, by M. D. Delfinado and M. Abbatiello, deposited in the
New York State Museum and Science Service, Albany.

Paratypes. Four females, same data as holotype, in the U.S. National Museum and New
York State Museum and Science Service collections.

Hoplocheylus americanus, n. sp.

(Figures 12-16)

This new species resembles H . longispinus Atyeo and Baker and H. canadensis Marshall in
most respects, and the 3 species are evidently closely related morphologically. The most dis-
tinctive characters of H. americanus are the long solenidia on tarsus and tibia of leg I, and
the very small subterminal spines on the palpal tibia and the shape of the ventral hysterosomal
plate.

Female. Length of body including gnathosoma, 466 microns. Palpus with genu fused with
femur; tibiotarsus with 5 simple setae, one rodlike solenidion and 2 very small, equal in
size subterminal spines as in figure 15. Chelicerae fused into a single unit and truncate
apically, with 2 pairs of dorsal setae ; gnathosoma with 2 pairs of ventral setae, posterior
pair only slightly longer than anterior pair. Dorsal propodosomal plate with a pair of
pseudostigmatic organs near lateral margin and 3 pairs of dorsal setae. Peritremes situated
on shoulders of propodosoma. Hysterosoma (figure 12) with 4 medial dorsal plates and
a pair of lateral or humeral plates; first medial dorsal plate longer than wide, with 2 setae;
second plate squarish, with 2 setae; third plate large, about as broad as long, with 2 pairs
of setae, the posterior pair longer than anterior pair but not reaching posterior margin
of fourth plate; fourth plate with 2 pairs of posterior setae nearly equal in length. Dorsal
anal region with a pair of terminal setae. Venter as in figure 13 ; hysterosomal plate large
with rounded posterior margin. Leg chaetotaxy as follows; the numbers represent coxa,
trochanter, femur, genu, tibia and tarsus:

Leg I. 4-1-5-5-6 + 2-14 + 2
Leg II. 3- 1 -3-4-5 + 1- 7 + 1
Leg III. 3-2-2-4-5+1-8
Leg IV. 2 - 1 - 2 - 5 - 5 + 1 - 7

Tarsus I lacking empodium; empodia present on tarsi II-IV; claws present on all legs;
tibia I-IV each with a solenidion; tarsi I and II with 2 and 1 long solenidia respectively;
the distal solenidion on tarsus I very long, reaching apices of claws; coxae III fused with
coxae IV.

Male. Not known.

Holotype. Female, collected from soil and pine debris, Lake Champlain region, New York,
October 15, 1973, by M. D. Delfinado and E. W. Baker, and deposited in the New York
State Museum and Science Service, Albany.

Paratypes. Eight females, same data as holotype, in the U.S. National Museum and New
York State Museum and Science Service collections.

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New York Entomological Society

Hoplocheylus americanus, n. sp. 12, dorsal surface of female; 13, ventral surface of female;
14, leg I; 15, dorsal view of palpus; 16, dorsal view of leg II.

Vol. LXXXII, September, 1974

207

Family Paratydeidae
Genus Scolotydaeus Berlese, 1910

Scolotydaeus Berlese, 1910, Redia 6: 214. Type-species, Scolotydaeus bacillus Berlese,

by monotypy.

Neotydeus Baker, 1950, Jour. Wash. Acad. Sci. 40 (6): 289. Type-species, Neotydeus

ardisannae Baker, by original designation. New synonymy.

The monotypic genus Scolotydaeus was previously known only from a brief description,
figure and notes by Berlese (1910), Thor (1933), and Baker (1949) who placed it in the
family Tydeidae. Baker (1950) later placed it in the Paratydeidae with Paratydeus Baker,
1949, and Neotydeus Baker, 1950. Neotydeus has proved to be a synonym of Scolotydaeus.

The genus Scolotydaeus primarily possesses the characters of the family (Baker, 1949,
1950) in that the hysterosoma is divided transversely at the third pair of legs; the palpus
is simple; tarsal claws are present on all legs, with small, clawlike empodia, and the proximal
venter of femora has a tiny, broadened dark ‘seta.’ The peritremes are simple, arising
from the bases of the chelicerae. The propodosoma lacks the lenselike eyes of Paratydeus.
The genus now includes 3 species; the one from New York is being described as new.

Scolotydaeus simplex, n. sp.

(Figures 17-22)

Scolotydaeus simplex is similar to S. ardisanneae (Baker) in several respects. It is dis-
tinguished by its very long solenidia on tarsus and tibia of legs I and much longer posterior
(third) propodosomal, humeral and posterior dorsal hysterosomal setae.

Male. Length of body including gnathosoma, 466 microns. Palpus 4-segmented, femur-
genu and tibia each with 2 long setae, tarsus with 3 rodlike and 4 short slender setae and
one small lateral solenidion. Chelicerae coalesced, suture obvious, movable chela curved
and strong, fixed chela not developed (fixed and movable chelae not opposed) ; gnathosoma
with 2 pair of setae, anterior pair shorter. Peritremes simple, short, lightly hooked distally
and arising from cheliceral bases. Propodosoma with anterior lateral peglike solenidia;
integument striate anteriorly, with 3 pair of slender setae, the anterior median pair long
and slender, the posterior pair slightly longer than anterior pair; eyes lacking. Hysterosoma
elongate, divided transversely at third pair of legs; humeral setae long, dorsal setae short
and slender; areas posterior to third pair of legs with first 2 pairs of setae in longitudinal
row; posterior setae in transverse rows, posterior lateral setae shorter. Venter as in
figure 18, ventral hysterosomal setae longer than setae at genital region with 4 pairs of
genital and 6 pairs of paragenital setae, transverse row of posterior ventral setae and 2
pairs of anal setae. Internally, genitalia with 5 pairs of short setae, and 4-5 pairs of short
spines on internal “spermatophore” apparatus (not figured). Leg chaetotaxy as follows; the
numbers represent coxa, trochanter, femur, genu, tibia and tarsus:

Leg I. 4-0-3 + 5 (*) -6-8 (7 + 1) -12 + 1

Leg II. 3- 1-2 -2 -4-7

Leg III. 2-1-3-2-3-5

Leg IV. 2-0-3 + 1(*) -1-3-5

Claws large, uncinate; empodia of all legs small and uncinate; femora I and IV divided
into basi- and telofemur with setal count as above (*). All solenidia on tarsus and tibia
of legs I very long and nearly equal in length.

Female. Similar to male, except genitalia much longer. Length same.

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New York Entomological Society

Scolotydaeus simplex, n. sp. 17, dorsal surface of male; 18, ventral surface of male;
19, details of gnathosoma; 20, dorsal view of leg I; 21, ventral view of leg I with detail of
tarsal claw; 22, dorsal view of tibia II.

Holotype. Male, collected from pine debris, bark and roots, Hague, Lake George, Adirondack
Park, New York, October 11, 1973, by M. D. Delfinado and E. W. Baker, deposited in the
New York State Museum and Science Service, Albany, N.Y.

Paratypes. Two females, with the above data, in the U.S. National Museum and New
York State Museum and Science Service collections.

209

Vol. LXXXII, September, 1974

Family Pseudocheylidae
Genus Anoplocheylus Berlese, 1910

Pseudocheylus, subg. Anoplocheylus Berlese, 1910, Redia 6: 210. Type-species, Pseudocheylus
( Anoplocheylus ) europaeus Berlese, by original designation.

Rhagina Womersley, 1935, Rec. So. Australian Mus. 5 (3): 336. Type-species, Rhagina

protea Womersley, by original designation.

This genus is characterized by the absence of claws on all legs which terminate with
a stalked membranous empodia. The peritremes are simple, chambered and located in the
membrane connecting the gnathosoma and propodosoma ; the palpal tarsus complex is
lacking; the chelicerae are attached basally and are movable laterally; a pair of lenslike
eyes is located on the anterior outer margins of the propodosomal plate. Five species were
previously known in the genus Anoplocheylus. The new species here described from New
York is the first records of the genus in North America.

Anoplocheylus transiens, n. sp.

(Figures 23-26)

Anoplocheylus transiens is similar to A. aegypticus Baker and Atyeo and A. tauricus
Livshitz and Mitrofanov in having the subcuticular reticulate bands on the propodosoma.
It differs in having all dorsal hysterosomal setae of approximately equal length. We have
examined adult and immature specimens of aegypticus and confirmed the presence of 3
coxal setae as figured. The text (Baker and Atyeo, 1964: 268) is in error stating that

coxa II has 4 setae.

Female (?). Length of body including gnathosoma, 530 microns. Palpus without thumb-
claw complex, with 4 distinct segments and a terminal claw. Peritremes chambered through-
out, arising at bases of chelicerae and situated on membrane separating gnathosoma from
propodosoma. Chelicerae hinged at bases and capable of lateral movement. Propodosomal
plate with fine, longitudinal striae; a single pair of lenslike eyes; 4 pairs of short setae,
the median pair located between anterior trichobothria ; the posterior marginal pair quite
long and slender; subcuticular reticulate bands on propodosoma posterior to trichobothria.
Hysterosoma with transverse striae anteriorly and posteriorly, longitudinal in region of
coxae III and IV; humeral setae long, slender; dorsal body setae all short except for
posterior setae of varying lengths. Genitalia longitudinal, usually with 4 pairs of short genital
setae and 3 pairs of paragenital plates. Sternal area with 2, 3 or 4 short setae. All legs
ending in stalked membranous empodia, claws lacking. Leg chaetotaxy as follows, the
numbers refer to coxa, trochanter, basifemur, telofemur, genu, tibia and tarsus:

Leg I. 5-1-8-6-7-8+1-19 + 4
Leg II. 3-1-2-4-5-5-9 + 1
Leg III. 3-2-2-3-4-5-9
Leg IV. 2 - 1 - 1 - 2 - 4 - 6 - 9

Male. Not known.

Holotype. Female (?), collected from leaf litter, Rensselaerville, New York, October, 1973
(no exact date), by M. D. Delfinado, deposited in the New York State Museum and
Science Service, Albany.

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New York Entomological Society

Pseudocheylus transiens, n. sp. 23, dorsal surface of female; 24, distal portion of venter
of palpus; 25, palpus I and distal portion of tibia I; 26, genitalia.

Vol. LXXXII, September, 1974

211

Paratypes. Seven females (?), 4 with the above data; 1 from litter, Taconic Parkway,

New York, June 16, 1973; 1 from litter, Heckscher Park, Long Island, New York, June 14,

1973; and 1 from debris, Rt. 87, 36 miles from New York City, July 22, 1973, all collected

by M. D. Delfinado, deposited in the U.S. National Museum and New York State Museum

and Science Service collections.

Literature Cited

Atyeo, W. T., and Baker, E. W. 1964. Tarsocheylidae, a new family of Prostigmatic
mites (Acarina). Bull. Univ. Nebraska St. Mus. 4(11): 243-256, figs. 1-16.

Baker, E. W. 1949. Paratydeidae, a new family of mites. Proc. Ent. Soc. Washington
51(3): 119-122, figs. 1-8.

. 1950. Further notes on the family Paratydeidae (Acarina) with a description

of another new genus and species. Jour. Washington Acad. Sci. 40(9): 289-291,

figs. 1-7.

and Atyeo, W. T. 1964. A review of the mites of the family Pseudocheylidae

Oudemans, 1909 (Acarina, Prostigmata). Bull. Univ. Nebraska St. Mus. 4(12):
257-272, figs. 1-30.

Berlese, A. 1910. Acari nuovi. Redia 6: 200-234, pis. 17-21.

Cooreman, J. 1951. Notes et observations sur les Acariens (IV). Bull. Inst. roy. Sci.
nat. Belg. 27(1): 4-7.

Leonard, M. D. 1928. A list of the insects of New York. Mem. Cornell Univ. Agric.
Expt. Sta. 101: 5-1093.

Livshitz, I. Z., and Mitrofanov, V. I. 1973. A new species of the genus Anoplocheylus
(Trombidiformes, Pseudocheylidae) from Crimea. Zool. Zh. Akad. Nauk SSSR
52(5): 770-771. (In Russian).

Marshall, V.G. 1966. Une nouvelle espece d’acarien (Tarsocheylidae: Prostigmata)

du sud-est du Canada. Acarologia 8(1): 45-48.

Smiley, R. L., and Moser, J. C. 1968. New species of mites from pine. Proc. Ent.
Soc. Washington 70(4): 307-317, figs. 1-14.

Thor, S. 1933. Fam. Tydeidae, Ereynetidae. Das Tierreich, lfg.60, pp. 1-45.

Centennial of Entomology at Cornell

The faculty of the Department of Entomology at Cornell University will
celebrate 100 years of entomology at Cornell on October 14 and 15 with a special
symposium. John Henry Comstock graduated from Cornell in 1874 and we take
this opportunity to honor the man who founded our department.

The symposium will bring many invited guests to the University and affords
an opportunity for persons to discuss the dynamic aspects of entomology. The
complete program will be carried in the September issue of the Bulletin of the
Entomological Society of America.

212

New York Entomological Society

BOOK REVIEW

The Gunong Benom Expedition, 1967: Parts 11-13. R. Traub. Bulletin of the British
Museum ( Natural History) Zoology , Vol. 23, No. 9-11. London, 1972. Notes on zoogeogra-
phy, convergent evolution and taxonomy of fleas (Siphonaptera) , based on collections from
Gunong Benom and elsewhere in Southeast Asia. I. New taxa (Pygiopsyllidae, Pygiopsyllinae) ,
pp. 201-305, 58 plates. II. Convergent evolution, pp. 307-387, 20 plates. III. Zoogeography,
pp. 389-450.

In the first paper of this series a new genus for the 5. robinsoni group is erected, the
hosts and distribution of the Malayan peninsular species of the group are discussed, a new
genus for the S. ferinus group is described, and keys to the new and old forms are provided.
Heretofore unknown males and females of various species are described for the first time.
The molding influence of the environment on these fleas and the principles involved in their
evolution and adaptation are described in the second article. In the third paper the author
presents evidence that fleas in the family Pygiopsyllidae originated in the Australian region
and moved to the mainland of Asia. He gives convincing data concerning the Australian
roots of the genus Medwayella, which probably originated in Borneo, thereafter moving to
the Asian mainland and Indochina, as well as to the Philippines. The speculation and dis-
cussion concerning the transport by rats of Palearctic fleas from the west and northwest,
with at least one species, Sigmacteus, reaching New Guinea, are most interesting. Malaya,
Sumatra, Java, and Borneo share many faunal features, but there are significant differences
between the mammals and fleas of Sarawak and those of Sabah, with those of the former
resembling Malaya more than the latter.

The descriptions of methods of collecting fleas in the tropics will be of special interest
to field workers. The major collecting areas were in forests, at elevations between 800 and
2000 feet, usually in primary jungle but also in secondary forests and bamboo areas. Rats,
tree shrews, and tree and ground squirrels were trapped and examined by the author while
he served as Commanding Officer of the U.S. Army Medical Research Unit in Malaya from
1948 to 1959. In addition, collections were made by others throughout the Southeast Asian
region. There were inherent disadvantages so far as collecting fleas on trapped animals
was concerned. Fleas tend to leave their hosts soon after feeding, or leave the animals when
the animals become excited and agitated. Heavy rain, a daily occurrence in the tropical
rain forest, also depletes the flea population on trapped animals. Even more disastrous is
the situation when killed animals are examined, because, within minutes, dead rats or
squirrels invariably attract swarms of ants, rarely leaving fleas on the carcasses. Therefore,
trapping was supplemented by shooting, usually at night, when the eyes of mammals would
glow in the light from powerful flashlights.

Entomologists will find the descriptions of the new taxa, the discussion of the convergent
evolution, and the zoogeography of fleas a useful guide and reference source for every as-
pect of flea research.

The definitive descriptions of fleas of Southeast Asia and the Indo-Australian Archipelago
contained in these three superbly illustrated issues of the Bulletin of the British Museum
will be of interest not only to taxonomists but also to medical officers and students of evo-
lution. Altogether this monumental work, containing 244 pages and 78 plates, is truly out-
standing. It constitutes exciting reading for everyone interested in the intriguing aspects
of collecting and handling fleas and the formidable difficulties that may be encountered.
Throughout the vast area of collection, Stivalius sensu lat. is a potential vector of plague,
and, in fact, it has been found infected with plague in India and Java.

Karl Maramorosch

New York Entomological Society, LXXXII: 212-213. September, 1974.

Vol. LXXXII, September, 1974

213

BOOK REVIEW

The Common Insects of North America. Lester A. Swann and Charles S. Papp. 1972.
Harper & Row, New York, xiii + 750 pp., 8 color plates, 2,450 drawings. $15.00

All but four very minor orders and most suborders, superfamilies, and families (275) are
covered and characterized in some detail. A total of 1,422 species are illustrated and treated
in some detail, and a great many others are mentioned and described briefly. Not only the
adults but the early stages (when known) are described and often are figured. The range
and chief environment of each species are given, as well as much information about habits,
economic importance, foods, and chief natural controlling agents such as predators, parasites,
and virus and bacterial diseases. An introductory section covers such important features
as the general characteristics of insects, the chief phyla of animals and classes of arthropods,
insect development and metamorphosis, predators and parasites, structures and some physi-
ology, insect defense mechanisms (unfortunately, too short) and “the value of insects.”
There is a very usable pictorial key to the orders and an excellent geologic time chart cover-
ing the main groups of plants and animals, as well as the insects. There is also a very good
glossary and a bibliography (perhaps too long) that includes many small papers and refer-
ences in economic entomology. Throughout, the authors have used as simple and nontechni-
cal language as possible. The black and white illustrations are mostly excellent, although
some Lepidoptera do not show the patterns very well and there is some distortion of wing
shapes. The scales of magnification or reduction are quite erratic. This can be a bit con-
fusing, even though the size measurements are given. In the copy at hand the color repro-
duction is not very good.

It is always a problem to a ye viewer to decide how much he is justified in listing errors,
a good many of which are liable to creep into a book of this magnitude. For example, is
such notice useful for corrections in subsequent editions? A couple of slips in the Lepidop-
tera, with which I have some acquaintance, are: the anal prolegs are not lacking in noto-
dontid larvae, although reduced or greatly modified in many; the tympana of “most moths”
are not in the mesothorax but in the metathorax (Noctuoidea) . No mention is made of the
abdominal tympana of the very large superfamily Pyraloidea. The enormous family Noc-
tuidae has been short-changed; more of the abundant and biologically interesting members
should have been included. And why was a highly aberrant specimen used to illustrate
the American Copper?

A very large proportion of the insects included are of economic importance. It is hard
to fault this, especially since such species are often abundant and likely to be noticed. But
as a result many more ecologically significant and interesting species have been left out. I
feel, too, that much more should have been included about the ecologic status of insects in
their communities and their great importance in energy cycles, subjects in which, it is good
to note, very large numbers of people are becoming interested. There is much information
about the control of many species by natural means but very little about insecticides, on
which we are still dependent in a great many cases (many “instant ecologists” would bene-
fit by some hard facts here).

The geographic coverage is extremely good and is a welcome change from books that
give undue importance to Eastern species. Canada and the West are justly represented.
The classification and nomenclature are up-to-date, although there will always be subjective
differences of opinion in these fields. Undoubtedly this book will be valuable to anybody
with an interest in natural history and environmental studies as well as to many entomolo-
gists, especially students and those engaged in economic work.

Alexander B. Klots

The American Museum of Natural History

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New York Entomological Society

Proceedings of the New York Entomological Society

(Meetings held in Room 129 of the American Museum of Natural History unless otherwise
indicated.)

Meeting of October 2, 1973

The meeting was called to order by Dr. Howard Topoff, President, at 8:10 p.m. 24 mem-
bers and 19 guests were present.

The minutes of the meeting of Tuesday, May 15, 1973, were approved as read.

Dr. Lawrence Limpel of Yonkers, N.Y., was proposed for Active Membership. His ento-
mological interests are insect control and insect physiology. Mr. Lamar Holsheimer of
Portland State College was proposed for Student Membership. His interests are Lepidop-
tera and Hymenoptera. Ms. Rosa Montes of Pace College was proposed for Student Mem-
bership. Her interests are myrmecology and general entomology. Ms. Mercedes Delfinado
was proposed for Active Membership; her interests are Diptera and free-living terrestrial
mites.

PROGRAM.

After a couple of short announcements by members of the Society Dr. Topoff introduced
Dr. Norman Lin who talked about social insects. The title of his paper was “Evolution
of Sociality in Insects.”

Father Sullivan announced that the speaker for the meeting on October 16, 1973, will be
Dr. Louis D. Trombetta, Isaac Albert Research Institute, Kingsbrook Jewish Medical Cen-
ter. His topic will be: “Abnormal development in Tenebrio caused by a juvenile hormone
analogy.”

Peter Moller, Sec.

Meeting of October 16, 1973

The meeting was called to order by Dr. Howard Topoff, President, at 8:10 p.m. 7 mem-
bers and 9 guests were present.

The minutes of the meeting of Tuesday, October 2, 1973, were approved as read.

Ms. Mercedes D. Delfinado, of Albany, N.Y., was elected to Active Membership; her in-
terests are in taxonomy of Diptera and free-living mites. Dr. Lawrence Limpel, of Yonkers,
N.Y., was elected to Active Membership; his interests are in insect control and insect physi-
ology. Ms. Rosa M. Montes, of Pace College, was elected to Student Membership. Her
interests are in myrmecology and general entomology. Mr. Lamar Holsheimer, of Portland
State College, was elected to Student Membership; he is interested in Lepidoptera and
Hymenoptera.

Ms. Betty Lane Faber, of New Brunswick, N.J., was proposed for Active Membership.
She is interested in insect behavior. Mr. S. M. Ulagaraj, of the University of Florida, was
proposed for Student Membership; he is interested in behavior and bionomics of Orthop-
tera. Mr. Henry M. Knizeski, Jr., of Fordham University, N.Y., was proposed for Student
Membership; he is interested in systematics and ecology in Hymenoptera. Mr. Charles
William Calmbacher, of Fordham University, N.Y., was proposed for Student Membership.
His interests are in Hymenoptera, systematics, and ethology of Sphecidae. Mr. James
Wangberg, of University of Idaho, was proposed for Student Membership.

New York Entomological Society, LXXXII: 214-218. September, 1974.

Vol. LXXXII, September, 1974

215

PROGRAM.

Father Sullivan introduced Dr. Louis D. Trombetta, of the Isaac Albert Research Institute
of Kingsbrook Jewish Medical Center. Dr. Trombetta presented a fascinating paper on
“Abnormal Development in Tenebrio caused by a juvenile hormone analog.”

The first meeting in November was cancelled because of Election Day.

Father Sullivan announced that the speaker for the meeting on November 20, 1973, will
be the Society’s own secretary, Dr. Peter Moller, of the Department of Psychology, Hunter
College, and the Department of Animal Behavior, American Museum of Natural History.
His topic will be: “How does a spider find its way home?”

The meeting was adjourned at 9:20 p.m.

Peter Moller, Sec.

THE EFFECTS OF A JUVENILE HORMONE ANALOG ON THE
DEVELOPMENT OF THE ANTENNA OF TENEBRIO MOLITOR

The developmental morphology and histology of the antenna of Tenebrio molitor as well
as its musculature were described and compared with antennae of insects treated with a
juvenile hormone analog. The juvenile hormone analog was code labeled JM-1-46 (4-
Ethylphenyl 2-(2-Etoxy Etoxy) Ethyl Acetal). It was topically applied with a microliter
syringe to the frontoclypeal suture of the pupa at a dose concentration of 3/ig//d of acetone.

Three extrinsic antennal muscles and three intrinsic antennal muscles were described. The
intrinsic antennal muscles all originated on the same surface of the scape.

The histology of the adult antenna revealed that the cuticle of the newly emerged insect
was composed of two layers, the exocuticle and endocuticle, separated in some regions by a
mesocuticle that alters considerably during the first week of development. The cuticle of
the intersegmental membrane consisted of two layers that were continuous with the endo-
cuticle. The epidermis was a simple epithelium that varied from cuboidal- to columnar-
shaped cells, depending on the location and density of the underlying nervous tissue. The
changes in the epidermis from the newly emerged to the one-week-old insect were described.
Johnston’s organ and an antennal blood vessel were also described. The antennal nervous
and respiratory systems were similar to those in other insects, and the changes that occurred
in these systems from the newly emerged to the one-week-old insect were described.

The morphogenesis of the antenna was divided into three stages, each of which was
marked by specific characteristics in the developmental sequence. The early stage extended
for the first four days after the larval-pupal molt, the intermediate stage continued through
days 5,6,7 postpupation, and the late stage was days 8 through 12. The cuticle, epidermis,
nervous system, tracheae, and blood vessel were described histologically at the larval-pupal
molt, 24 and 48 hours postpupation, and 7 and 12 days postpupation. All the above struc-
tures were shown to undergo significant alterations during development.

Contrasted to the above findings were insects treated with the juvenile hormone analog.
The development of the antenna of a treated insect determined the degree of muscle devel-
opment. The antennal pupal case was very delicate and much thinner than the normal one
and had the same sensory structures that were present on the normal pupal cuticle. The
antennae of the treated insects showed four different morphological conditions. First, the
antenna of the adult resembled the antenna at the larval-pupal molt, but was highly convo-
luted and had a very irregular cuticle. The cuticle was said to be juvenilized. Second, the
antenna had both juvenilized regions and regions that appeared adultlike. Third, the distal

216

New York Entomological Society

segments of the antenna were rounded in a manner not characteristic of the adults. Juvenil-
ized patches of cuticle were scattered over the antennal surface. Fourth, two pupal cases
were covering the antenna. The outer case resembled the original pupal case but the inner
one was not as well defined.

Two general histological patterns were described for the treated antennae. The first
pattern was for antennae where the cuticle appeared morphologically pupal and the second
pattern was for antennae of insects where the cuticle appeared morphologically adultlike. A
few aberrant antennal forms unlike the above were described.

Reversal of metamorphosis by juvenile hormone as stated by some previous investigators
was shown to be unlikely. Rather, it seems more probable that the juvenile hormone analog
acts upon the cell nucleus to produce abnormal characteristics.

Louis D. Trombetta
Kingsbrook Jewish Medical Center

Meeting of November 20, 1973

The meeting was called to order by Dr. Howard Topoff, President, at 8:10 p.m. IS members
and 15 guests were present.

The minutes of the meeting of October 16, 1973, were approved as read.

The following membership elections were held:

Mr. Henry M. Knizeski of Fordham University was elected to Student Membership. He is
interested in the systematics and ecology of Hymenoptera.

Mr. Charles W. Calmbacher of Fordham University was elected to Student Membership.
Mr. Calmbacher works on the systematics and ethology of the hymenopterous family
Sphecidae.

Mr. S. M. Ulagaraj of the University of Florida at Gainesville, who specializes in the be-
havior and bionomics of Orthoptera, and Mr. James Wangberg of the University of Idaho,
were elected to Student Membership.

Ms. Betty L. Faber of New Brunswick, New Jersey, whose main interest is behavior, was
proposed for Active Membership.

program. Dr. Topoff introduced Dr. Peter Moller of the Department of Animal Behavior
of the American Museum of Natural History. Dr. Moller, our Secretary, was greeted with
vigorous and sustained applause. His talk provided us with brilliant and provocative answers
to the question “How does a spider find its way home?” Making effective use of slides and
other illustrative material, he considered various aspects of aranean orientation behavior.
The lecture was followed by prolonged and heated debate.

Our next meeting is scheduled for December 4, 1973, at which time Dr. David C. Rentz of
the Department of Entomology of the Academy of Natural Sciences of Philadelphia will
consider the question of mechanical reproductive isolating mechanisms in a talk entitled
“The lock and key as an isolating mechanism in katydids.”

The meeting was adjourned at 9:30 p.m.

Charles C. Porter, Asst. Sec.

Vol. LXXXII, September, 1974

217

Meeting of December 4, 1973

The meeting was called to order by Dr. Howard Topoff, President, at 8:15 p.m. 10 mem-
bers and 5 guests were present.

The minutes of the meeting of November 20, 1973, were approved as read.

program. Father Sullivan introduced Dr. David C. Rentz, Department of Entomology,
Philadelphia Academy of Natural Sciences, who told us briefly about his experiences as
president of the Philadelphia Entomological Society before he started his interesting talk
about “The lock and key as an isolating mechanism in katydids.” His talk was followed
by a lengthy discussion.

Father Sullivan announced that the speaker for the next meeting on December 18 will be
Dr. Ross H. Arnett, Department of Biology, Siena College. He will talk about “Pollen-
feeding beetles.”

The meeting was adjourned at 9:10 p.m.

Peter Moller, Sec.

Meeting of December 18, 1973

The meeting was called to order by Dr. Howard Topoff, President at 8:20 p.m. 12 members
and 7 guests were present.

The minutes of the meeting of December 4, 1973, were approved as read.

Mr. Alberto Muyshondt of San Salvador was proposed for Active Membership. His ento-
mological interests are Rhopalocera of Tropical America.

program. Father Sullivan introduced Dr. Ross H. Arnett, Department of Biology, Siena
College. In a fascinating talk illustrated with color slides Dr. Arnett introduced his audience
to the phenomenon of “pollen-feeding beetles.” A very interesting and heated discussion
followed.

It was announced that the next meeting will be held on January 15, 1974. The speaker
will be Mr. Frederick H. Miller, Jr., Nassau County Medical Center, who will talk about
“The scanning electron microscope — A tool for entomologists.”

The meeting was adjourned at 9:45 p.m.

Secretary’s note: For the records it should be mentioned that this meeting was the last
one chaired by President Howard Topoff, who, after two years of office, leaves the ranks
of officers in the New York Entomological Society. The Society is grateful for his many
innovations and hard work.

Peter Moller, Sec.

THE ROLE OF POLLEN FEEDING IN COLONIZATION OF SMALL
POPULATIONS WITH PARTICULAR REFERENCE TO COLEOPTERA

An understanding of some of the problems of small populations of colonizing species
has been gained through a study of the role of pollen feeding by quantitative experimental
ecological population studies. Selected species of oedemerid beetles, all obligate pollen

218

New York Entomological Society

feeders, and all well-known taxonomically, have been studied in particular, along with other
pollen-feeding beetles in general, by field experimentation involving population sampling,
feeding experiments, and karyotype determination.

Many plants suffer very heavy predispersal pollen predation by a large variety of animals.
In spite of the generally held view that insects are the responsible and required pollinators
for many plants, it is certain that most of the produced pollen in these and other, noninsect
pollinated plants goes as insect food without any self- or cross-pollination. In fact, some
plants develop feeding anthers that produce a degenerate pollen used solely for food con-
sumption. The development of these special pollen-feeding, nonpollinating structures, the
chemical secretions used as attractants, and the ethological patterns in beetles that make
this a mutual relationship are very poorly understood.

Oedemerid beetles are ideally suited for such studies because: 1) They are now relatively
well known taxonomically through the previous research on the group by this investigator;
2) the breeding populations of almost all the species are very small, and there is abundant
evidence to show that they are actively colonizing; and 3) they are all obligate pollen
feeders with a specialized pollen rumen used when pollen foraging.

Two things are clear from the sketchy studies made of the pollen-feeding phenomenon: It
is 1) a highly evolved chemical, structural, and ethological process, and 2) a large and
important but as yet not fully exploited field of study. This, coupled with the need to
know more about the factors operating during colonization attempts, has resulted in the
accumulation of a wealth of data, but many questions remain unanswered.

Ross H. Arnett, Jr.

Siena College

Vol. LXXXII, September, 1974

219

New York Entomological Society

PROGRAM SCHEDULE 1974/75
Guest speakers at forthcoming regular meetings:

October

1974

Robert R. Granados

Boyce Thompson Institute, Yonkers, N.Y.

October

15,

1974

Bert Holldobler

Harvard University, Cambridge, Mass.

November

1974

Thomas Eisner

Cornell University, Ithaca, N.Y.

November 19,

1974

Vincent G. Dethier

Princeton University, Princeton, N.J.

December

1974

Kenneth D. Roeder

Tufts University, Medford, Mass.

December

17,

1974

Jerome S. Rovner

Ohio University, Athens, Ohio

January

1975

Arthur H. McIntosh

Rutgers University, New Brunswick, N.J.

January

21,

1975

Karl Maramorosch

Rutgers University, New Brunswick, N.J.

February

1975

Gary D. Bernard

Yale University, New Haven, Conn.

February

18,

1975

Rudolf Jander

University of Kansas, Lawrence, Kan.

March

1975

Peter N. Witt

N.C. Dept. Human Resources, Raleigh, N.C.

March

18,

1975

open

April

1975

Bertrand Krafft
University of Nancy, France

April

15,

1975

Walter C. Rothenbuhler

Ohio State University, Columbus, Ohio

May

1975

Neal A. Weber

Florida State University, Tallahassee, Florida

May

20,

1975

open

All meetings

will be held at the American Museum of Natural History, Central

Park West at 79th Street, at 8:00 P.M. For further information (dinner reser-
vation etc.) call Dr. Peter Moller (212-873-1300 ext. 385).

220

New York Entomological Society

Washington DC
USA

August 19—27
1976

XV International Congress of Entomology
First Announcement

The 15th International Congress of Entomology will be held in the beautiful
capital city, Washington, D.C., U.S.A., August 19-27, 1976, under the sponsor-
ship of the National Academy of Sciences and the Entomological Society of
America. Sessions will be held in the excellent meeting facilities of the Washing-
ton Hilton Hotel. Special events are being planned at national scientific and
cultural centers. Two international airports near Washington give direct access
from abroad. University housing will be available in addition to hotel facilities.

The Organizing Committee for the Congress is composed of Curtis W.
Sabrosky (Chairman and President of the Congress), Ernest C. Bay (Secretary-
General), Wallace P. Murdoch (Treasurer), William G. Eden, Gordon E. Guyer,
E. F. Knipling, Robert L. Metcalf, John V. Osmun, Ray F. Smith and Edward
O. Wilson.

The program will emphasize plenary symposia, invitational speakers, special-
ized symposia/work groups/panel discussions, and special interest groups or
informal conferences. Thirteen program sections cover Systematics, Genetics,
Physiology and Biochemistry, Toxicology, Ecology, Behavior, Social Insects
and Apiculture, Biological Control, Medical and Veterinary Entomology, Agri-
cultural Entomology and Pest Management, Forest Entomology, Stored Products
and Structural Insects, and Pesticide Development, Management and Regulation.

A Congress Brochure and application forms will be mailed in May, 1975. The
Brochure will contain information on highlights of the scientific program,
receptions, tours, ladies program, scientific, historical and other features of the
Washington area, and useful data for visitors.

PLEASE NOTE: Announcements of this Congress are not being sent to
individuals, but are being publicized in journals and circulated to museums,
departments, and other institutions. If you are interested in receiving future
information, including registration forms, please send a postcard to the under-
signed with your name and address, typed or in block letters, and also the
section of your major interest.

Dr. ERNEST C. BAY, Secretary General
XV International Congress of Entomology
P.O. Box 151
College Park, Md.

USA 20740

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JOURNAL of the

NEW YORK ENTOMOLOGICAL SOCIETY

The JOURNAL of the NEW YORK ENTOMOLOGICAL SOCIETY is de-
voted to the advancement and dissemination of knowledge pertaining to insects
and their related forms.

Please make all checks, money-orders, or drafts payable to the NEW YORK
ENTOMOLOGICAL SOCIETY.

ORDERS and inquiries for back issues and complete sets should be sent to
our agent, S-H Service Agency, Inc., 866 Third Ave. New York, N. Y. 10022.
Complete files of back issues are in stock.

INFORMATION FOR AUTHORS

Submit manuscript in duplicate (original and one carbon) to the Editor, New
York Entomological Society, Boyce Thompson Institute, Yonkers, N.Y. 10701.

i '' ' C _ / I / L. 3

1. GENERAL POLICY. Manuscript submitted must be a report of unpub-
lished research which is not being considered for publication elsewhere. A
manuscript accepted and published in the JOURNAL must not be published
ftgain in any form without the consent of the New York Entomological Society.

A page charge of $15 per printed page is assessed except that the charge may
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IT;

The New York Entomological Society
Incorporating The Brooklyn Entomological Society
Incorporated May 21, 1968

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The New York Entomological Society

Organized June 29, 1892 — Incorporated February 25, 1893

Reincorporated February 17, 1943

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The Brooklyn Entomological Society

Founded in 1872 — Incorporated in 1885
Reincorporated February 10, 1936

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Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00.

Members of the Society will please remit their annual dues, payable in January, to the
Treasurer.

Officers for the Year 1974

President, Dr. Daniel J. Sullivan, S.J.

Fordham University, New York 10458

Vice-President, Dr. Peter Moller

American Museum of Natural History, New York 10024
Secretary, Dr. Charles C. Porter

Fordham University, New York 10458
Assistant Secretary, Dr. Louis Trombetta

Pelham Manor, New York 10803

Treasurer, Dr. Winifred B. Trakimas

State University of New York, Farmingdale, New York 11735
Assistant Treasurer, Ms. Joan DeWind

American Museum of Natural History, New York 10024

Class of 1974

Trustees

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Dr. Lee Herman
Class of 1975

Dr. Howard Topoff

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Mr. Edwin Way Teale

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Dr. Pedro Wygodzinsky

Mailed February 24, 1975

71:

The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press
Inc., 1041 New Hampshire, Laurence, Kansas 66044. Second class postage paid at New Brunswick, New

Jersey and at additional mailing office.

Knbwn/ office of publication: Waksman Institute of Microbiology, New Brunswick, New Jersey 08903.

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life

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, '

Journal of the

New York Entomological Society

Volume LXXXII December, 1974

No. 4

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Waksman Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lois Keller, RSM
Dr. Herbert T. Streu

Publication Committee

Dr. Kumar Krishna Dr. Ayodha P. Gupta

Dr. James Forbes, Chairman

CONTENTS

William Couper, Taxonomist-Entomologist F. Martin Brown 222

Nest Biology of the Eucerine Bee Thygater analis (Hymenoptera, Antho-
phoridae) Jerome G. Rozen, Jr. 230

Notes on the Natural History of a Rare Adelpha Butterfly (Lepidoptera :
Nymphalidae) in Costa Riean High Country Allen M. Young 235

Revision of the Genus Holcostethus in North America (Hemiptera: Penta-
tomidae) F. J. D. McDonald 245

Digger Wasps as Colonizers of New Habitat (Hymenoptera: Aculeata)

Howard E. Evans 259

Seasonal Variation in Tachysphex terminatus (Smith) (Hymenoptera: Sphec-
idae, Larrinae) Nancy B. Elliott and Frank E. Kurczewski 268

Two New Genera and Two New Species of Acantliosomatidae (Hemiptera)
from South America, with a Key to the Genera of the Western Hemisphere

L. H. Rolston and R. Kumar 271

New or Little-Known Crane Flies from Iran. I (Diptera: Tipulidae)

Charles P. Alexander 279

Index of Scientific Names of Animals and Plants for Volume LXXXII 285

Index of Authors for Volume LXXXII iii

222

New York Entomological Society

William Couper, Taxidermist-Entomologist

F. Martin Brown

6715 So. Marksheffel Road, Colorado Springs, Colorado 80909

Received for Publication December 14, 1973

Abstract: Records are made of what little is known about William Couper, a Canadian field
naturalist of the latter half of the 19th century. This is based upon the literature and
Couper’s letters to Herman Strecker. Couper’s collecting trips to Labrador and Anticosti
Island are recounted. Considerable information is given about the controversy involving
Papilio brevicauda Saunders and Papilio anticostiensis Strecker, and that involving Lycaena
pembina Edwards and Lycaena couperi Grote.

Some years ago while carrying out the N.S.F. mission of cataloging and
preserving the thousands of letters that Herman Strecker accumulated, we found
a small bundle of them written in the early 1870’s by William Couper of Mon-
treal. Since none of the usual sources of biographical material about entomolo-
gists contains information about Couper I thought it worth-while to present to
you something about the man gained from his letters. Couper is memorialized
by Glauco psyche lygdamus couperi Grote, originally described from Anticosti
Island.

Couper’s activity in the province of Quebec was outlined by Comeau (1965) in
an address, “ A Glance at the History of Entomology and Entomological Collec-
tions in Quebec” presented at “The Lyman Entomological Semicentennial
Symposium.” This was delivered on December 30, 1964, as part of the A.A.A.S.
meetings held at Montreal. Comeau noted that Couper built the third entomo-
logical collection for the province. The first, that of Pierre Boucher, Governor
of the city of Three Rivers, was made around 1664 and lost. Pierre Chasseur’s
collection was bought by the government of Lower Canada in 1827 and lost
by fire in 1832. Couper’s was given to Morin’s College in the city of Quebec
in 1871 and was destroyed by pests.

While living in Quebec Couper helped found the first entomological society
in the province. This occurred in June 1862 in league with Provancher and
Leclerc. It affiliated with the Entomological Society of Ontario in 1868 but
died in 1871 when Couper left Quebec. In 1873 Couper was instrumental in
organizing what became the Montreal branch of the Entomological Society of
Quebec. Couper had a falling out with William Saunders, the editor of CANA-
DIAN ENTOMOLOGIST. This caused him to bring together a group of
entomologists, who had accidentally met on a mountain side, to form the
Montreal Entomological Society.

With his letter of December 8, 1873, to Strecker, he included a clipping from
a newspaper — name and date unknown to me — that opens as follows:

New York Entomological Society, LXXXII: 222-229. December, 1974.

Vol. LXXXII, December, 1974

223

“ENTOMOLOGICAL SOCIETY.— The monthly meeting of this Society
was held on Wednesday night at the residence of the President, Mr. William
Couper, No. 67 Bonaventure Street. The following members were present — -
The President, Messers Kolmar, Kuetzing, Caulfield, C. W. Pearson and
G. B. Pearson. Mr. Alexander Gibbs was proposed for membership, and
Mr. Andrew Johnson was elected a member . . .”

The “Commemorative Programme” for the 85th Annual Meeting of the
Entomological Society of Ontario, 3-5 November 1948, celebrated the 75th
anniversary year of the Montreal Branch of the society, the result of Couper ’s
original Montreal Entomological Society. In the program there is “Short History
of the Montreal Branch . . .” and on p. 15 a photograph and the signature of
Couper are reproduced. On p. 6 of the program it states “. . .he left Montreal
for New York in 1884. ... it is thought that he died at his son’s residence at
Troy in 1890.”

Couper made an early collecting trip in 1867 to Labrador and a second in 1872.
Also in that year, 1872, and upon two other occasions, he visited and collected
insects upon the Island of Anticosti. I have only found references to an earlier
trip (in 1865?) to Labrador but there appears to be first-hand information in
the contemporary newspapers of Montreal to which I do not have access. His
letters to Strecker at least give us an outline of his second trip (1872) and of
his trips to Anticosti.

In the late 1860’s and at least to March of 1871, Couper served as Assistant
Curator and Taxidermist for “The Literary and Historical Society of Quebec.”
He apparently lived at 38 Bonaventure Street, Montreal. In 1873 he decided to
set up as an independent taxidermist and established himself at 67 Bonaventure
Street in Montreal. This was done after his second trip to Anticosti Island.
On that trip he had some official position with the Anticosti Company, the only
way he could reside upon the island. He sailed from Montreal on May 15th on a
vessel chartered for the trip by the Anticosti Company. A letter dated August
6th, 1873 opens “I have just returned from Anticosti.” Thus he spent about
9 or 10 weeks there on this trip. His subscribers, at $12 a head, were Grote,
W. H. Edwards, a Mr. Chase, and a Mr. Wassemann of England. Strecker,
always parting with money only under duress, finally contributed his $12 to
Mrs. Couper after the vessel had sailed. Couper hoped to gain permanent em-
ployment with the Anticosti Company but this seems never to have materialized.

In addition to bringing to you something about one of the early Canadian
collectors of Lepidoptera, these notes from Couper’s letters touch upon two
interesting taxonomic problems : the relationships of Papilio brevicauda Saunders
and antic os tiensis Strecker, and the identity of Lycaena pembina Edwards.
I will let Couper tell you of his travels and collecting experiences by means of
direct quotations from his letters.

224

New York Entomological Society

“September 30, 1872”

“I have returned home after an absence of over four months. I left Quebec
on May 18 last on board the government schooner “Stella Maris”, for Anticosti.
I remained on the island two weeks, when I took passage for the coast of Labra-
dor, arriving at Natashiquan, and collected between latitude 50 and 51 success-
fully, obtaining a good number of diurnal lepidoptera. Everything went on well
until I turned my face homeward by the western North Shore route, traversed
by me about 7 years ago, terminating at a place called Mingan. At the
latter place, I wished to obtain Argynnis Boisduvalli and Colias interior , 2
species occurring there about the 22 July. While thus engaged, and during
my absence from camp (at Mingan), the whole of my former collection, the
result of 5 weeks work on the lower portion of that coast, was destroyed by
the mountain Indians. These miserable beings not only destroyed my collec-
tions, but robbed me of provisions, etc. 1 appealed to the priest then in charge
of the Mission, who told me that he had no control in the matter of this nature,
but that he would make inquiry, and help me all in his power. The only relief
obtained from him was that he wished me to leave their territory or the juris-
diction of their chief as soon as possible, as he would not be responsible for
their actions regarding my life or property. The tribe indicated their determina-
tion to punish me, in fact, to shoot me down. They looked on me as a government
spy, and I am since informed that some English person told them who I was,
and that I wrote in the Quebec papers about 7 years ago, that they speared
salmon on the spawning grounds. This statement is true, for I did describe
their disgraceful modus operandi in destroying salmon, but never anticipated
that it would end so unfortunately for me. As I am now situated, I cannot,
this year, fulfill my agreement with you. The species collected on Anticosti are,
however, safe as they were not in camp at the time. The Anticosti collection
is not large in species, but will be honestly divided between the 5 subscribers
who advanced money for the object. These are Mr. Edwards of San Francisco,
Mr. Mead of N. York, the Ent. Soc. of Ontario, Mr. Morrison of Boston and
yourself. In order to carry out my contract with you, I propose going to Labra-
dor (but not to Mingan) next season at my own expense, and if God spares
me, you will be furnished with the missing species. I will send you the box
containing the Anticosti species in a few days. I remain,

Yours truly,

/s/ William Couper”

“February 3, 1873
“Montreal

“Your favor of 7th ult. came duly to hand, and I have delayed answering
for the reason that on its receipt I communicated with the Rev. Cannon [sic]
Innes and Mr. W. Saunders of London, Ontario, asking for information regarding

225

Vol. LXXXII, December, 1974

Papilio brevicauda. Up to this instant, no answer from either. It appears to me
that both wish to evade my queries, and I enclose Mr. S.’s last to me in order to
show that he makes no illusion to it. Now, I wish to inform you that I know
something regarding how Mr. S. obtained his specimen of the insect which he has
named as above. The Rev. Mr. Innes, who has a cabinet of Lepidoptera, lived at
Quebec during my residence there. After my return from my northern tour,
about 6 years ago, I presented him with 2 or 3 specimens of a Papilio taken in
Labrador, which I then supposed was P. asterias. He had, at the time, a few
similar, but smaller, specimens of the same insect from Newfoundland. Mr.
Innes removed afterwards to London, Ontario, where Mr. Saunders resides and,
of course, the Papilio attracted the attention of the latter. I have not seen
Mr. S.’s description, and I wish you to inform me what locality is given. Thanks
for your information regarding the Lycaena, I am so pleased that it is new, and
long to see it described and figured by you.”

W. H. Edwards in volume 2 of his superb “Butterflies of North America”
figured the imagoes of brevicauda on Plate Papilio VIII. The models for female
figures definitely are from specimens taken in Newfoundland and supplied to
Edwards by Saunders. The source of the male figure is in question. It may
have been based upon an Anticosti specimen from Couper. It conforms in color
of the band on the upper sides of the wings better with Anticosti males than
with Newfoundland males. The mature larva figured on this plate is erroneously
colored. Edwards corrected this with Plate Papilio VII IB and noted the
error in the text that accompanies that plate. The figures of the immature
stages presented by Edwards as of brevicauda actually apply to antic ostiensis
Strecker. At the end of the text for Plate VIII B is a letter from T. L. Mead in
which he compared the larvae and pupae of brevicauda with the figures on the
plate. Males bred by Mead from Newfoundland larvae varied from some that
were as free of fulvous suffusion as shown on Plate VIII, to some with a little
fulvous suffusion. So the true source of the model for Edwards’ figure of the
male still is in doubt.

Edwards’ Plate VIII was issued in December 1875. His notebooks for the
period preceding this have been searched for information about brevicauda. He
sent the pickled mature larva and egg shown on Plate VIII to Mrs. Peart on
August 25, 1873. (ms Ent. Journal “1872” p. 235). In March of 1875 he
paid Mrs. Peart for drawing the figures for Plate VIII (ms Ent. Journal “1872”
p. 227). This notebook also contains quotations from Couper’s letters to Edwards
about brevicauda, but nowhere have I found any notation of the source of the
male figured on Plate VIII.

The Lycaena mentioned in this letter of Couper later was named couperi
by Grote. How Grote, not Strecker, came to name the taxon is explained in a
letter that will be quoted further on.

226

New York Entomological Society

The following are quotations from letter of April 15, 1873:

“Pap. Anticostiensis is what I may term an uncommon butterfly in every
locality on the shores of the lower St. Lawrence visited by me. When I arrived
at Fox Bay, Anticosti, last June, it was extremely rare, and I captured only
4 specimens in the course of 15 days. The specimens were apparently fresh
on 20 June — they generally flew low, frequenting the flowers of a species of
wild pea which occurs abundantly on the banks of the river on Anticosti and
Labrador. I experienced great difficulty in approaching them with the net.
On first appearance its flight is rapid and low extending along the margin
of the rocky cliffs and grassy portions of the Bay, near tidemark. I never
noticed them in the woods. They appeared to me to keep within the circuit
of the Bay, and I remarked the same fact on the Labrador coast, where I also
found them on the flowers of the wild pea. Indeed, they hovered about it so
much that I expect to find its larva feeding on it this season. If I do, I will
take descriptions of them, and then the difference between it and Asterias
will be so far settled. I noticed toward the end of July, that their strength
gives way and if the weather is cool, added specimens may be taken by hand
from the flowers of the pea. It is the only species of Papilio so far noticed by me,
either on Anticosti or Labrador.”

“August 6, 1873

“Montreal

“I have just returned from Anticosti. The west end of the island so far
visited by me, produced only 12 species of butterflies, viz. P. anticostiensis
Strecker, which is found throughout the whole island; P. Turnus of which I took
only 2, and I have not yet compared them with sp. taken here; an Argynnis
not yet determined, and of which 1 have only 28 specimens. The only butterfly
new to me is a Colias which was exceedingly rare at Ellis Bay, and I have
taken notes on its habits. I have only 12 specimens of the latter, but you shall
have a pair of the best. This Colias is evidently different from Interior. I fear
that the last named does not occur south of the Labrador coast. The form which
I have may be Labrador ensis . I have also 9 specimens of Hesperia paniscus
which was also rare. I took the same Lycaenidae which were met with last year:
M. Batesii, P. frigida — the common nettle butterfly and a Grapta — the latter
rare. However, I will do my best to distribute the material fairly. I am sorry
that I have not met with a greater number of species, but that cannot be helped,
perhaps I may have little better luck next time in a new locality. I will send
such things as I believe will please you, and the rest of the material will be
fairly distributed to my subscribers.

“I have discovered the food plant of P. Anticostiensis and have both eggs
and larvae. The caterpillar feeds on two plants — viz. — Archangelica antho-
purpurea or G. A. of Hoffmann, and Heracleum lunatum Michx. I have 4

Vol. LXXXII, December, 1974

227

notes regarding it. I have also discovered the food plant of P. jrigida , and have
eggs and larvae. The plant is the L. M. — Larritis striata Graham. I consider
these investigations worth the time and trouble taken. I had a narrow escape
from shipwreck on my way home.”

That is almost the entire letter. The rest of it goes into personalities.

I am recording a good deal of the next letter, of August 26, 1873.

“Ellis Bay is calculated to be 117 miles west from Fox Bay, as you can
see on the map of Anticosti. I sent [send?] you a few extracts from my field
book regarding the species remarked in your letter. P. Antic ostiensis was
noticed and a specimen taken at Ellis Bay on 14 June, the day of my arrival.
From the latter date, as the weather became warmer, 40 specimens were taken
up to 26 June. On 25 June I noticed a female depositing an egg on the food
plant, Archangelica antho purpurea, which occurs common throughout the sec-
tion of the island. The egg, (one in bottle sent), is laid singly on the upper
surface of the leaf, where it is exposed to the full force of the sun’s rays. The
egg is spherical, pale yellow. On fine days, between 10 and 3 o’clock is the
best time to capture them, but should the weather become cold or windy, not
one would be seen. About half the specimens taken have short tails. I cannot
say that the larvae sent to you is a full grown specimen. I took the largest
which I could find only a day or two prior to leaving the Bay at the end of July.
They are the only mature ones I could obtain, and have evidently the skin of
the last moult. You can see that it differs from Asterias larva, by having
oblique black lines on the sides of the body, besides other minute points. I
have also sent you a young larva of about 10 days old, which shows the light
yellow band in the center of the body. The regularity of the markings of the
perfect insect (males and females) are to my observations of sufficient value
to make a good species. The question remains what is the difference between it
and Brevicauda. Saunders said that there is a vast difference, but time will settle
that matter, and I will sift it next season. Every entomologist who has seen
P. Anticostiensis since my return, believes that there is little or no connection
between it and Asterias.

“I took 12 specimens of Colias at Ellis Bay — all in one locality — at different
times. I extract the following from my notebook: — ”

And then in columns he gives these dates each followed by “one specimen.”
The dates are: June 3, 30, July 3, 5, 8, 14, 16, 19, 23, 24. “It is evidently a
rare insect at Ellis Bay, another old resident informed me that it has never been
common in that locality. Its flight differs from other species known to me.
It is extremely restless, very zigzag and quick, and it goes over a great extent
of ground in a short time; indeed, I had much difficulty to capture the few
I obtained. It was only about 5 June that I noticed one light on a flower. It
has a peculiarity when at rest, of lying half on its side, as if enjoying the heat

228

New York Entomological Society

of the sun. I am sorry that the two examples sent you are both males, as it was
not my intention to have done so; true, I did not think, as you have since
informed me, they might be 2 species, and I have still doubts as to the likelihood
of their being so. However, you will soon determine that when they are spread.
It may be that the bright one is Interior. I have sent 2 to Edwards, 2 to Chase
and 2 to Grote; the latter by the way called on me last week on his way to the
Science Association at Portland. I do not think that the smaller one is rubbed —
you better examine it closely.

Couper was a fairminded man. It irked Couper to read Strecker’s intemperate
remarks about other entomologists printed in his “Lepidoptera: Rhopaloceres —
Heteroceres.” The matter came to a head over Grote describing Glaucopsyche
couperi. The first inkling of this trouble appears in the letter dated December 8,
1873, in which Couper wrote “Grote described the Glaucopsyche as I spoke to
him about it when he visited Montreal this summer and I told him that you
intended to name it Couperi. I did so because I deemed it necessary to present
duplicate descriptions. I only wish that you will figure it in order to make
the species bona fide Strecker did figure the taxon (Lep. R.-H., pi. 10, f. 10,
11, 1874) but insisted that it was nothing more than pembina Edwards and
that Grote was in error thinking it otherwise. This was said by Strecker in a
vituperative manner. On March 28, 1874, Couper wrote Strecker “I note your
decision that Glaucopsyche couperi Grote is identical with Pembina Edw.
Still I cannot overlook the fact that you stated “It makes little difference who
names a species, so long as it is well done.” Of course, I am not prepared to
say that Grote is wrong, as Mr. Edwards’ description of Pembina is now before
me and I cannot make it agree with the Ellis Bay Lycaena. There is also my
knowledge of the species as a nondescript as far back as 1867, on my first
visit to Labrador. You also informed me that Grote did what you intended to
do. This was the cause of writing and quoting your statements and I trust
you will excuse my being candid in saying that your answer to the above is
comparatively vague. Moreover, Canadian entomologists of my acquaintance,
who read your remarks on Mr. Grote, do not appreciate the style of epiplionema,
considering in a scientific light, it would be well if they were omitted.”

Strecker’s reply to this was to terminate Couper’s subscription to his book.
Couper insisted upon reinstating the subscription a year later and there followed
a few brief letters before the correspondence closed. This is an excerpt from a
letter 17 April 1874, and what I quote is a quotation from a letter sent to
Couper by W. H. Edwards.

“With regard to the Lycaena from Anticosti I presume Mr. Scudder is correct.
The original Pembina came from Lake Winnipeg, a single specimen or a single
pair, several years ago. These types were afterwards lost in a box of insects
sent by me to California. I had forgotten them, and somehow another species

Vol. LXXXII, December, 1974

229

had been assumed to be Pembina by Scudder and others, and I had fallen into
the error myself of thinking with them, that Pembina was allied to Lygdamus.
I discovered the fact last year, and called Mr. Scudder’s attention to it, while
he was here on a visit. I think this Couperi was what had been thought to be
Pembina and Grote was correct in naming of CouperiP

The problem of the identity of pembina Edwards is still unsettled. Currently,
and for some time past, it has been accepted as a subspecies of Plebejus icarioides
Boisduval. Dr. John Downey, who has been studying this complex species for
many years has not been able to find any specimens anywhere in collections
that hail from the reputed type locality. In the original description Edwards
stated that the types were collected by Scudder at “Lake Winnipeg.” Scudder
himself stated that he took the specimens in a glade on the banks of the
Saskatchewan River northwest of Lake Winnipeg. Dr. Downey has proven
that the food plants of the larvae of the icarioides complex all are confined to
the genus Lupinus. Dr. David Dunn, the ranking authority upon this genus for
North America, tells me by letter that there are no known records of Lupines in
the region published as the type locality of pembina. This blank area in the
distribution of the genus Lupinus is a real one, not the result of insufficient
collecting. All of this poses a problem, or three of them: first, is pembina

an icarioides that has changed its food plant?; second, did Scudder really
collect the types of pembina where he said he did or did Kennicott?; third, is
pembina now applied to a taxon that is not the same species as that to which
Edwards first applied the name?

Edwards’ original description of pembina credited Kennicott with the capture
of the type. This he had been told by Baird. However, research has given much
greater support to John Pearsall being the captor and the locality being in
western Montana, not “Lake Winnipeg.” Apparently McDunnough had come
to a similar conclusion about the type locality. (See Brown, 1970, pp. 397-402.)
The name pembina Edwards now is used for Icaricia icarioides from Montana,
Alberta and British Columbia in the foothills and mountains.

Literature Cited

Anonymous. 1948. Commemorative Programme, 85th Annual Meeting, Entomological
Society of Ontario.

Brown, F. Martin. 1970. The types of lycaenid butterflies named by William Henry
Edwards, Part III. Plebejinae. Transaction of the American Entomological Society,
96: 353-433.

Comeau, Noel-M. 1965. A glance at the history of entomology and entomological
collections in Quebec. Annals of the Entomological Society of Quebec, 1965:
85-90.

Couper, William. Manuscript letters to Herman Strecker preserved in the Dept, of
Entomology, Field Museum of Natural History, Chicago, Illinois.

Edwards, William H. Manuscript entomological journals. State Archives, Charleston, West
Virginia.

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Nest Biology of the Eucerine Bee Thygater analis
(Hymenoptera, Anthoplioriclae)

Jerome G. Rozen, Jr.* 1

Received for Publication December 20, 1973

Abstract: Details of the nest of Thygater analis are described and diagramed. Information on
cell provisions, larval feeding activity, defecation and cocoons is also given.

I had an opportunity to excavate a nest of an unidentified bee at Antonina,
Parana, Brazil on November 7, 1971. Because of the fine texture and cohesive
nature of the soil, the excavation was made with considerable accuracy and a
rather clear understanding of the nest structure resulted. No adult female
was associated with the nest but I was able to determine that the bee belonged to
the Eucerini because of larval anatomy, cocoon structure and fecal pattern.
Emergence of a female and a male from cocoons on November 7 and 12, 1972
respectively permitted Padre J. S. Moure of the Universidade Federal do Parana
to identify the species as Thygater analis (Lepeletier). Although a number of
workers have described various aspects of the biology of this species (for refer-
ences see Urban, 1967) none has described the nest in detail. For that reason
and because of the unusual structure of the nest, I offer the following diagram
and account to which I have added other information on nesting biology.

Description of Nesting Site. The nest was located on a moderately sloping barren
stretch of ground (fig. 1) next to the roadway leading to Antonina, Parana,
Brazil. Only a single nest was located although a search was made for others.
The nest entrance was unshaded by the tropical vegetation which surrounded
the area. The soil was extremely fine, with almost no rock inclusions, and moist
except on the surface.

Description of Nest. The nest entrance was located in a depression, presumably
caused by a rock having been removed. The main burrow (fig. 2) entered the
ground nearly horizontally in the side of the depression. The tumulus, which
was abundant and moderately coarse, filled the lower part of the depression and
partly obscured the nest itself. Circular in cross section and with a diameter of

Acknowledgments: I would like to thank Padre Moure and his staff, both for the

species identification and for their hospitality and courtesies while I was in Brazil. The
research was supported by National Science Foundation Grant GB32193. Specimens of
cells, cocoons, larvae, as well as reared adults are in the collection of The American Museum
of Natural History.

1 Deputy Director for Research and Curator of Hymenoptera, The American Museum
of Natural History, Central Park West at 79th Street, New York, New York 10024.

New York Entomological Society, LXXXII: 230-234. December, 1974.

Vol. LXXXII, December, 1974

231

Fig. 1. Site of nest of Thygater analis near Antonina, Parana, Brazil. Padre Moure
is standing by the nest.

7.0 mm, the main burrow descended for the first several centimeters hori-
zontally but then turned downward and descended vertically. The wall of the
main burrow was extremely smooth and when examined under the microscope,
faint markings, obviously created by the pygidial plate of the female, could be
observed on all sides. Descending open and vertically to about the depth of 20
cm, the burrow then turned horizontally, rose somewhat, descended again, and
ended blindly. A single branch, 3.5 cm long and filled with soil, entered the
burrow near where it first started to curve horizontally. Connected to the branch
was a single, nearly vertical cell (fig. 2, cell 13) containing a small feeding larva
and provisions. Attached at the lower end of the cell was a tunnel that descended
more or less vertically and then curved horizontally, rose over a distance of
about 6 cm before bending downward, at which point it was lost. Three branches,
again all filled and about 3.5 cm long, connected to this tunnel and each ended
in a single nearly vertical cell. This tunnel was open except near where it
attached to the cell 13. Cell 12, farther along in the series, contained a moder-
ately small feeding larva, the next one (cell 11) held a somewhat larger one,
and cell 10, closest to cell 13, contained the largest larva. Descending obliquely
from the lower end of cell 10 was another more or less horizontal tunnel, filled
near the cell with coarse soil but open for the rest of the way. This tunnel led
to five filled branches and cells, very much as described for the above tier. Cell

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Fig. 2. Diagram of the nest of Thygater analis.

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233

9, farthest along, contained a large feeding larva which, however, was the
youngest in this series. Cell 5 at the other end held a larva that had finished
feeding and was starting to defecate. Connected to this cell was yet another
lower, more or less horizontal tunnel filled at the beginning but open elsewhere.
This lowest tunnel gave rise to four cells with their filled branches, the one
(cell 4) farthest from this connection holding the youngest larva, a form that
was defecating but not yet cocoon spinning. The other three cells housed larvae
that were either spinning or had spun their cocoons. This account seems to
amplify the general description of the nest provided by Michener and Lang
(1958).

On the assumption that a cell is constructed, provisioned, oviposited in and
closed before the next cell is started, the order of cell construction and provision-
ing was from cell 1 to cell 13 because of the ages of the larvae. There is no
evidence to determine whether the entire tunnel system was constructed before
the first branch and the cell were excavated or whether the female dug a vertical
tunnel and a lateral to form the tunnel of the lowest tier and then constructed,
provisioned and oviposited in the first four cells all before excavating a lateral to
form the tunnel of the next higher tier. In each tier the cell closest to the
connection to the tier above was constructed and provisioned first; furthermore,
the cell in the tier above that received the tier below also contained the oldest
individual of the tier.

All tunnels and branches were approximately 7.0 mm in diameter; the fill in
branches was loose, coarse soil.

The cells, similar to those of Svastra obliqua (Say) (Rozen, 1964), were
elongate, 17-18 mm long and had a maximum diameter of 8 mm but little
wider than the tunnel. Vertical or nearly so, they possessed an extremely smooth,
shiny wall and there was no obvious indication of a built-in lining. The wall
was faintly embossed, presumably with the pygidial plate of the female, and
was coated with a special semitransparent lining which was obviously water-
proof as evidenced by the nature of the provisions. The closures were a somewhat
concave spiral on the inside with four to five rows to the radius. The deepest cell
was about 44 cm from the surface, the highest, approximately 24 cm.

Provisioning. The source of the pollen was not known. Provisions occupied the
lower part of the cell, gave off a faintly aromatic odor, and were approximately
7 mm in depth, at least in one instance. They were apparently stratified into an
upper clear layer containing almost no pollen and having a slightly sweet taste,
and a lower more opaque yellowish layer, only slightly less liquid and containing
pollen.

Development. Young larvae rested on their side while feeding in the soupy
provisions. Feeding actively, they curled so that their dorsum often adhered
to the cell while they submerged their head and anterior part of their body into

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the food. The orientation of older larvae was not ascertained except in one
instance in which the larva was found curled around a small quantity of the
semiliquid food. Upon finishing feeding, a larva defecates and applies the feces
to the area of the cell cap, as was described for Svastra obliqua (Rozen, 1964).
After the feces are attached to the cell cap, the larva begins to spin its cocoon.

The fecal material and the cocoon are similar to those of Svastra obliqua
(Rozen, 1964, fig. 2). As in Svastra three layers of the cocoon can be detected,
the outermost being actually the cell lining that adheres to the cocoon itself.
The apparent middle layer is thin, brownish, semitransparent and nonfibrous.
The inner layer, closely applied to the middle one, is thin, greyish brown, semi-
opaque, fibrous and moderately thin. Also as in Svastra the top of the cocoon
is domed by a moderately thick roof and there are a number of silken partitions
separating air spaces between the roof and the feces.

The fact that adults emerged a year after they were collected as larvae seems
to indicate that there is a single generation a year.

Parasitism. No parasitic bees were found in the vicinity of this nest and larvae
of none were recovered from cells.

Literature Cited

Michener, Charles D., and Rudolf B. Lange. 1958. Observations on the ethology of
neotropical Anthophorine bees (Hymneoptera: Apoidea). Univ. Kansas Sci. Bull.,

vol. XXXIX, 3: 69-96, figs. 1-24.

Rozen, Jerome G. Jr. 1964. The biology of Svastra obliqua obliqua (Say), with a
taxonomic description of its larvae (Apoidea, Anthophoridae) . Amer. Mus. Novitates,
2170: 1-13, figs. 1-15.

Urban, Danuncia. 1967. As especies do genero Thygater Holmberg, 1884. (Hymenoptera,
Apoidea). Bol. Univ. Federal do Parana, II, 12: 177-309, figs. 1-22.

Vol. LXXXII, December, 1974

235

Notes on the Natural History of a Rare Adelpha Butterfly
(Lepidoptera: Nymphalidae) in Costa Rican High Country

Allen M. Young

Department of Biology, Lawrence University, Appleton, Wisconsin 54911
Received eor Publication January 16, 1974

Abstract: The nymphalid butterfly Adelpha leucophthalma tegeata Fruhstorfer is a rare

member of the macrolepidopterous fauna in the central high (montane) country of

Costa Rica. The life stages and developmental time are described for the first time, along

with observations on the behavior of the larva and adult. The egg-to-adult developmental

time is 51 days on Pentagonia wendlandia Hook (Rubiaceae). The egg is laid singly

on the dorsal surface of older leaves of the food plant and both the larva and pupa
(chrysalis) are very cryptic in morphology, color, and behavior. As with most species
of this genus, the adults of A. leucophthalma are very skittish. This is one of the few
reports on a Central American Adelpha from montane environments.

INTRODUCTION

Many species of Adelpha butterflies (Lepidoptera: Nymphalidae) are well
known by entomologists working in lowland tropical wet forests of Central and
South America. It was Godman and Salvin (1870-1901) who originally pictured
several of these species, followed by the descriptions of Fruhstorfer (1915).
Most of the lowland tropical species of Adelpha are medium-sized butterflies
with rich chocolate-brown wings bearing a single, bold white or combination
orange and white band on the forewing (fractionated into an anterior orange
section and posterior white section), and they are frequently encountered along
sunny forest paths and edges where they rest on low vegetation. Although these
butterflies are well-known in tropical lowlands, Miller and Miller (1970) have
emphasized that very little is known about the more elusive species of this
interesting genus in montane regions of the Neotropics. In fact, the apparent
paucity of information even on the geographical distribution and taxonomy
of montane species of Adelpha led Miller and Miller (1970) to their discovery
of two rare species in the high country of Hidalgo, Mexico.

Acknowledgments: This research was funded by COSIP (College Science Improvement
Program) Grant GY-4711 (National Science Foundation) through Lawrence University.
The author is very grateful for this support. Dr. Lee D. Miller of Allyn Museum of
Entomology (Sarasota, Florida) kindly identified the butterfly and provided background
information of the species. The larval food plant was identified by Drs. Luis Diego Gomez
P. (Museo Nacional de Costa Rica) and Dieter C. Wasshausen (National Museum of
Natural History). Color transparencies (35 mm.) on the early stages of this butterfly
may be borrowed from the author by interested researchers and collectors.

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Fig. 1. Adelpha leucophthalma tegeata Fruhstorfer,
Scale in mm.

Wild-caught female, dorsal view.

Adelpha leucophthalma tegeata Fruhstorfer (which is taxonomically near
A. diocles Godman & Salvin, Lee D. Miller, pers. comm.) belongs to that
constellation of rare montane species of Adelpha in Central America. One of
the few adults in my possession is shown in Figure 1. The color pattern of the
wings of this little known species represents a significant departure from the
more typical appearance of other members of the genus: the dorsal surface of
the forewing has a thick, bright orange band and the round spot on each hind-
wing is vivid white. This species is said to be “rare” in the sense that there are
very likely very few specimens in museum collections anywhere, and nothing was
known about its natural history. The purpose of the present paper is to describe,
for the first time, the early stages of the little known Adelpha , along with a
larval plant food record, developmental time, and other aspects of natural history.

HABITAT AND METHODS

The high country to the northeast of San Jose, the capitol city of Costa Rica,
includes a series of mountain valleys of virgin rain forest that lead into the foot-
hills of the Caribbean drainage of these mountains (Central Cordillera). One

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of the most prominent of these montane moist valleys is one that occurs east of
Volcan Poas. The road that connects San Jose with Puerto Viejo runs along
the western ridge of this valley. The altitude of the valley where this study was
done is about 1000 m and the depth of the valley itself is about 150 m. The
bottom of the valley is the Rio Sarapiqui, and at the study area this river is
filled with very large boulders and it is several m in width with very swift cur-
rent. On the western edge of the river, there is a small plains area that includes
virgin forest and heavily-disturbed areas. The habitat (“study area”) where
oviposition by A. leucophthalma was observed consisted of a rectangular patch
of very recently cut (1-2 weeks) old secondary forest. This site is just to the
right before the girder bridge on the trail from the Puerto Viejo road, down the
valley, across the river, and up the other side to a penal colony. Much of the
forest had been cleared (machete) by a “squatter” farmer for bananas and cattle.

It was in this patch of freshly-cut forest trees that I saw oviposition by a
single female of A. leucophthalma at 12:30 P.M. on July 2, 1971. The sky was
very overcast with the threat of rain and this butterfly flew low and swiftly
among the still fresh leaves of the various felled trees lying in the area. Two eggs
were laid on two different old leaves of a sapling-size (4 m tall) individual of
Pentagonia wendlandi Hook (Rubiaceae).

The two eggs were collected, confined to a clear plastic bag and allowed to
hatch in San Jose. The two larvae from these eggs were reared in this manner
and supplied with fresh leaves of the food plant. The big, papery, food plant
leaves were always retrieved from this individual of Pentagonia at the study
site and perhaps owing to the heavy rainfall at this time of year, several fresh
leaves remained available on the cut tree. Photographs and color transparencies
were made of the life stages. Although one larva died in the fourth instar, the
other larva survived to adulthood. Measurements (in mm) were made on life
stages and gross external features of morphology were noted. Searches were
made on subsequent visits for other ovipositing adults and adults of both sexes
in general, but to no avail. I was able, however, to observe some adult behavior
from the girder bridge high over the Rio Sarapiqui: on two different dates
during July 1971, I watched adults flying and resting among leaves of trees
overhanging the river near the bridge and about ten m above the water. Other
than these observations, adults were very seldom seen. The single adult obtained
from rearing was eventually sent to Lee D. Miller of the Allyn Museum of
Entomology for identification. Despite many other visits to the area ( 1972—
1973), it was not until July 1973, that I was able to capture another specimen
of this very elusive butterfly, and this specimen is shown in Figure 1. It was
collected about 60 m from the study site.

Observations were also made on the feeding behavior of the larvae in the
plastic bags. Interest here concerned perch construction for resting periods, and
the pattern of leaf damage resulting from feeding.

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Fig. 2. Life stages and behavior of A. leucophthalma. (A) egg, (B) first instar (lateral)
resting on perch constructed from the bared apical leaf midrib and silk, (C) second instar
in similar position, (D) third instar, (E) fourth instar on perch, and (F) fifth instar
(lateral), emphasizing the raised position of the spiracles and first thoracic segment and
eighth abdominal segment.

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RESULTS

Life stages. The light bluish-green spherical egg (Fig. 2-A) is about 1.0 mm in diameter
and the chorionic surface is highly sculptured with ridges forming distinct facets. Tiny
hair-like projections arise from the facets, giving the egg a fuzzy appearance (Fig. 2-A).
One day prior to hatching, the egg turns light tan in color.

The first instar larva (Fig. 2-B) is 3.0 mm long at the time of hatching, and it possesses
a large, orange-yellow head and light green trunk region. Both the head and trunk are
covered with rows of very small, tubercle-like scoli. Since I was unable to preserve larval
specimens (due to small sample size), the precise distribution and structure of scoli for
all instars could not be studied at this time. By the first molt, the larva is about 9.0 mm
long. The second instar (Fig. 2-C) has a dark reddish-brown head and dark green body;
sets of prominent tubercles become noticeable on the third thoracic segment, and first,
third, and eighth abdominal segments (Fig. 2-C, and also Fig. 2-D, for third instar as
well). This pattern of prominent, dorsal tubercles, and the arched condition of the third
thoracic segment, is very reminiscent of the larva of the North American Limenitis
(Nymphalidae) . In fact, Lee Miller has told me (pers. comm.) that some authors place
Adelpha as a subgenus of Limenitis. By the second molt, the active larva is about 13.0
mm long.

The third instar (Fig. 2-D) is characterized by a noticeable change in coloration:
the head has become very dark brown bearing white tubercles, and the body is mottled
in shades of dark brown and gray. This very cryptic instar attains a body length of
about 17.0 mm by the third molt. The fourth instar (Fig. 2-E) retains the studded
condition of the integument of the previous instar, the body being covered with many
tiny whitish-gray tubercles, and it is also very cryptically colored in shades of brown
and gray. A prominent light gray saddle-like area develops dorsally (but with lateral
extensions) in the posterior abdominal region (visible but slightly out of focus in Fig. 2-E).
A pronounced transformation has taken place in the size and shape of the prominent
tubercles of thoracic and abdominal sectors: all tubercles have undergone considerable

elongation, and the set of the metathoracic segment is curved anteriorly. The set of
the third abdominal segment is strongly curved towards the posterior end of the body,
as is the set on the eighth segment (Fig. 2-E). All of these tubercles bear many tiny
stiff hairs. The rich brown color of the tubercles is continuous with the brown coloration
of the body. Figure 2-E purposely emphasizes the morphology of the head capsule of
this instar. The reasons for this are two-fold: (1) the general shape of the head capsule

is now very different in that it is strongly forked dorsally whereas before it was round,
and (2) the head capsule color pattern is very different since it now consists of a pair
of prominent, vertical cream bands on an otherwise brown background, whereas before
it was entirely brown. The head capsule retains the highly studded surface texture of
previous instars and as before, all studs are white, cream, or very pale green. By the
fourth molt, this instar attains a body length of about 21.0 mm.

As with the transformation to the fourth instar, the advent of the fifth instar is marked
by new pronounced changes in gross external morphology. So profound are these changes
in appearance that it is worthwhile to emphasize both head and body structure as shown
in the series of Figures 2-F and 3-A through C. The system of prominent tubercles of
thoracic and abdominal regions has become even more pronounced in this instar: the

tubercles are greatly elongated and scoli appear on all of them as prominent projections
(Fig. 2-F; Fig. 3-A). The set of lateral, sub-spiracular tubercles relatively reduced in
previous instars is now elongated with similar spiny projections. All of these tubercles

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Fig. 3. Life stages and behavior of A. leucophthalma. (A) fifth instar, emphasizing
the structure and distribution of trunk tubercles, (B) fifth instar (dorsal), emphasizing
cryptic resemblance to a moss-covered twig, (C) fifth instar, emphasizing the morphology
of the head capsule, (D, E) lateral and ventral of the chrysalis, respectively, and (F)
leaf damage pattern by fifth instar larva.

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are dull green and there are also green areas on lateral portions of the body. The fifth
instar is very spiny in general appearance (Fig. 3-B).

There are also some changes in morphology of the head capsule of the fifth instar,
as emphasized in Figure 3-C. Although the coloration remains the same as in the previous
instar, there has been considerable expansion on the lateral row of studs that surround the
head posteriorly. The two forks remain anterior to this pronounced row of studs on
the dorsum, appearing as twin darkened cones. Just below these head capsule forks,
there is a pair of large but stubby white tubercles, followed ventrally by a second, smaller
pair.

Another conspicuous feature of the fifth instar is the raised condition of the spiracles
on the first (prothoracic) thoracic segment and on the ninth abdominal segment (Fig. 3-A).
The spiracles of these two segments are positioned more dorsally than those on other
segments.

The position of tubercles and scoli deserves comment as they are very noticeable on
the fifth instar. There are two rows of spiny tubercles: the uppermost row is dorso-

lateral and the second row is sub-spiracular. The first pair of upper tubercles on the
metathoracic segment is almost vertical to the body, while those of the first abdominal
segment are smaller or more oblique to the body. The next pair is about the same length
(7.5 mm) as those of the metathoracic segment, vertical, and positioned on the third
abdominal segment. The several segments between this segment and the eighth bear
short tubercles, all about the same length as those of the first abdominal segment (4.5 mm) .
The pair on the eighth abdominal segment is oblique and about the same length as the
larger anterior ones; the set on the ninth segment is slightly shorter and more vertical.
All of the sub-spiracular tubercles are about the same length (5.0 mm). Other details
of tubercle position and structure are omitted since they were not studied due to a lack
of preserved material.

Together, the coloration, studded integument of head and body, along with the struc-
ture and distribution of spiny tubercles, endows the fifth instar larva of A. leucophthalma
with a very cryptic resemblance to a short section of moss-covered twig (Fig. 3-B).
As to be outlined below, the behavior of the larva suggests further that crypsis in this
insect is employed during both resting and feeding. The active fifth instar form attains
a length of 26.0 mm prior to pupation.

There is no distinct prepupa, unlike several other neotropical Nymphalidae. The angular
pupa (Fig. 3-D, E) is very dark brown but lustrous, with some irregular silver flecks
on the ventral thoracic area. It is about 20.0 mm long and resembles a shriveled up dry
leaf. The conspicuous lateral and ventral aspects of the pupa are self-explanatory in
Figure 3-D, E.

The total egg-to-adult developmental time is 51 days for the single individual reared
to the imago. The egg stage lasts eight days, the total larval period 31 days, and the
pupa 12 days. This individual was a male.

Larval Behavior. Upon hatching, the first instar larva immediately devours the
empty egg shell; how consistent this behavior pattern is among the species and
subspecies cannot be determined from this study. As the egg is usually affixed
to the leaf either at the edge or near a hole, the larva then moves to the very tip
of the leaf. Throughout the first three or four instars, the larva when not feeding
rests on a perch made from the bared midrib of the leaf. If the larva is disturbed
experimentally while it is feeding, its immediate response is to crawl rapidly back
to the perch and stay there for several minutes. This behavior pattern is very

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consistent. Very interestingly, the larva during the first two instars weaves a
thick, silvery mat of silk around the midrib section forming the perch, and the
mat extends partially onto the intact leaf surface. This structure is very much
reminiscent of the woven construction of the hibernating tube of first or second
instar Limenitis at northern latitudes. But here there is no enclosure (tube)
formed after the mat is built. The resting perch of younger instars is shown in
Figure 2-B, C.

The fifth instar rests on both sides of food plant leaves under laboratory con-
ditions. It generally rests near large brown spots on the edges of these leaves, and
it feeds along the edges rather than from the tip (Fig. 3-F). During all instars
the larva is a diurnal feeder. When disturbed, the fifth instar ceases to feed and
folds the head down beneath the thoracic region. The spines on the prominent
tubercles do not produce a rash on the back of the hand when they are rubbed
against them, suggesting that the spines are functional as part of a general
morphological adaptation for passive defense (crypsis) rather than for offensive
chemical defense against attackers.

Adult behavior. What little can be stated concerning adult behavior in this
butterfly has to do with oviposition and play. Oviposition is very fast: there is
a rapid flight movement over the foliage with sudden stops to lay a single egg.
It is so fast that I was not able to determine the stance assumed during oviposi-
tion. The egg is laid on the dorsal leaf surface: in the two instances observed,
one egg was laid near the edge, and another (on a different leaf) was laid near a
hole in the interior region of an old leaf (Fig. 2-A).

As typical with many species of Adelpha , adults of leucophthalma exhibit
flying play behavior amidst sun-flecked leaves overhanging the Rio Sarapiqui.
I define play behavior as the flitting among different leaves by adults, including
momentary perching on leaves in sunny places. It is likely that both sexes are
involved. Based on these observations, and the fact that oviposition occurred
on the leaves of a felled Pentagonia that is four m tall, it is likely that adults are
active primarily in forest strata that occur about four to six m above the ground.
On the morning of July 4, 1973, a sunny day at the study site, I observed one
fresh adult of A. leucophthalma resting on the broad leaf of an epiphytic palm
about five m from the ground; this was in the uncut forest about 100 m from the
cut (now regenerating) forest site. I was unable to net this individual. This
individual was resting with the wings outstretched, possibly thermogulating,
as noted for other montane species of Adelpha (Miller and Miller, 1970). I
have never seen adults resting with their wings closed in this species.

DISCUSSION

Adelpha leucophthalma is probably absent from the tropical lowlands of Costa
Rica. I have never seen it either in Guanacaste or in Sarapiqui (“tierre

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243

caliente”). As for high country, I have seen it both at Cuesta Angel, the site of
the present study, and also southwest of here at Bajo la Hondura. At both of
these localities, the insect is solitary, and I have seldom seen more than three
adults on a given day. The factors contributing to the apparent “rarity” of this
butterfly in Costa Rican high country are not known at this time. Based on these
field observations, the scarcity of the species in museum collections is indicative
of it being rare in the wild. In the tropical lowlands, many other species of
Adelpha appear to be locally abundant.

Miller and Miller (1970) suggest that some Adelpha are Q^erc^-feeders as
larvae; Quercus is a member of the Fagaceae (beech family). The present
record of A. leucophthalma on a rubiaceous food plant may be a first record for
a member of this genus on something other than oaks.

In terms of adjusting to the local community of plants and animals, the
natural history of this Adelpha , as in other species of this genus, entails an
adaptive response to some spectrum of potential predators in the form of crypsis.
As perceived by humans, this crypsis is best expressed in this insect during the
larval and pupal (chrysalis) periods of ontogeny. Such adaptations are very
likely most effective against attacks by visual-hunting predators such as foliage-
foraging insectivorous birds, lizards, and perhaps larger predatory arthropods
possessing compound eyes capable of color discrimination. The behavior of the
larva in returning to a thin, isolated perch upon disturbance is interpreted here
as a means of positioning itself in a place where (1) crypsis is enhanced, and
(2) it is less accessible to predators. It is doubtful, however, that the latter
aspect is an adaptation to large vertebrate predators that glean leaves for insects,
but rather it may be most effective against ants and other smaller predatory
arthropods that hunt by odor and tactile means in addition to vision. A bird or
lizard would gobble the larva very quickly should the crypsis be penetrated by
the searching behavior of these forms, but there may be more time for escape
when attack is by a single ant or beetle. The predator-defense adaptations of
the egg and adult stages are obscure. The hair-like projections from the egg may
be functional in discouraging attack by predatory insects. Depending on the
size and diversity of the guild of leaf-chewing insects that attack the older leaves
of Pentagonia at the study site, the eggs of this butterfly and those of other
insects may be subject to varying degrees of predation through passive uptake
as leaves are consumed. The adults are very swift, agile fliers, as are all members
of the genus. The general habits of the adults of several insular species of
Adelpha have been given in Barcant (1970), and those of adult A. leucophthalma
conform closely to A. naxia (Fldr.).

Literature Cited

Barcant, M. 1970. “Butterflies of Trinidad and Tobago.” London: Collins, 314 pp.

Frtjhstorfer, H. 1915. Adelpha. In A. Seitz, Die Gross-schmetterlinge der Erde,
5: 510-533.

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Godman, F. D., and O. Salvin. 1870-1901. “Biologia Centrali-Americana. Insecta.
Lepidoptera-Rhopalocera.” London, 2 vols.

Miller, L. D., and J. Y. Miller. 1970. Notes on two rare Mexican Adelpha and
related Central American species (Nymphalidae) . J. Lepid. Soc., 24: 292-297.

Vol. LXXXII, December, 1974

245

Revision of the Genus Holcostethus in North America
(Hemiptera: Pentatomidae)

F. J. D. McDonald

Department of Plant Pathology and Agricultural Entomology,

University of Sydney, Sydney, Australia

Received for Publication February 21, 1974

Abstract: A diagnosis is given for the genus Holcostethus and descriptions are provided for
six species. A new species is described from Arizona and a key is provided for the identifica-
tion of all North American species.

Holcostethus is a widespread genus with representatives in North America,
Europe, North Africa and Asia. The holarctic distribution of the genus is con-
firmed by the similarity of the seven North American species to the two European
species examined, H . sphaecelatus (the type species) and H. vernalis (Figs.
4-17).

All the species examined have a pair of structures lying dorsally in the base
of the pygophore that are unusual but not unique to the genus. These structures
have been observed by many authors and given many names (Tuxen, 1970).
Here they are termed pseudoclaspers.

Holcostethus Fieber, 1860

Holcostethus Fieber, 1860, Europ. Hem., p. 79; Kirkaldy, 1909, Cat. Hem., p. 47.

Peribalus Mulsant and Rey, 1866, Ann. Soc. Linn. Lyon (2) 14, p. 185; Stal, 1872, Ofv.
Svenska Vet-Ak. Forh. 29 (3), p. 34; Distant, 1880, Biol. Cent. Amer. Het. 1, p. 65;
Jakovlev, 1902, Ent. Obozr. 2, p. 158; Van Duzee 1904, Trans. Amer. Ent. Soc. 30,
p. 32; Zimmer, 1912, Univ. Nebraska Studies 11, p. 221; Van Duzee, 1917, Cat. Hem.,
p. 32; Blatchley, 1926, Het. E. N. Amer., p. 105; Froeschner, 1941, Amer. Mid. Nat.
26, p. 127.

Dryocoris Mulsant and Rey, 1866, Ann. Soc. Linn. Lyon (2), 14 p. 190.

Type species : Cimex sphacelatus Fabricius, 1794.

Diagnosis. Oval brown pentatomids ranging in size from 4-6 mm in width (across lateral
angles) and 8-10 mm long (tip of head to membrane apex). Head. Jugae slightly longer

Acknowledgments: I should like to thank the following: Dr. P. Wygodzinsky of the

American Museum of Natural History, Dr. Paul Arnaud, Jr. of the California Academy
of Science, Dr. C. Triplehorn, Ohio State University, and Dr. R. C. Froeschner, United
States National Museum for the loan of type and other material used in this study;
Dr. Per. Inge Person, Swedish Museum of Natural History, and Dr. W. R. Dolling,
British Museum (Natural History) for the loan of type material; Dr. David C. Renz,
Academy of Natural Sciences of Philadelphia for the loan of original Uhler material.

I am especially grateful to Professor L. H. Rolston, Entomology Department, Louisiana
State University, for suggesting this study and for the use of material and notes he had
made on this genus. I am also indebted to him for reading and correcting the manuscript.

New York Entomological Society, LXXXII: 245-258. December, 1974.

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Figs. 1-10. H. sphacelutus . 1. Prosternum. 2. Prosternum, lateral view. 3. Metasternum.
4. Pygophore, dorsal view. 5. Ventral margin of pygophore. 6. Right pseduoclasper.
7. Right clasper. 8. Aedeagus, lateral view. 9. Aedeagus, ventral view. 10. Spermathecal
bulb and pump, stink gland opening and sulcus (A. p.), apical tubercle (A. t.), basal
tubercle (B. t.), conjunctival appendage (C. a.), clasper (Cl.), coxal cavity (Cox.), distal
flange of pump (D. f.), dorsal margin (D. m.), endophallic duct (E.), evaporative area
(Ev.), flange (F.), keel (K.), median penal lobe (M. p.), proctiger (P.), pseudo clasper
(P. c.), process (Pr.), tubercle (T.), theca (Th.).

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247

than tylus, generally convergent apically, in some species meeting completely around tylus.
Apex of rostrum generally reaching hind coxae. Thorax. Pronotum trapezoidal, deflexed
between anterolateral margins. Scutellum extending two-thirds way down abdominal
terga, apex narrowing beyond frena and bluntly rounded. Prosternum with two distinct
keels forming a V between the coxal cavities (Figs. 1, 2). Metasternum with large granulose
evaporative areas (Fig. 3) ; odoriferous gland openings with a long sulcus. Abdomen.
Females generally with connexiva visible beyond folded hemelytra; males with connexiva
covered by folded hemelytra.

Male genitalia. Pygophore with a pair of palmate pseudoclaspers, one on either side
(Figs. 4, 6). Claspers L-shaped with a large basal tubercle (Fig. 7). Theca well-sclerotized
with a pair of basal and apical tubercles, latter absent in one species (Fig. 8). One pair
of conjunctival appendages present. Median penal lobes paired, plate-like, apically acute,
united at base on either side of S-shaped endophallic duct (Fig. 9).

Female genitalia. External genitalia plate-like ; eighth paratergites without spiracles ; ninth
paratergites oblong, well-separated (Fig. 26). Spermatheca typically pentatomoid; sper-
mathecal bulb with 2 or 3 fingerlike processes (Fig. 10).

KEY TO THE SPECIES OF HOLCOSTETHUS IN N. AMERICA

1. Ventral surface of abdomen dark chocolate brown, margins may be outlined

in yellow 2

Ventral surface of abdomen either yellow or buff or reddish brown, with or
without black markings 5

2. Long gray setae found on dorsal surface especially on pronotum

hirtus (Van Duzee)

Dorsal surface without conspicuous setae 3

3. Scutellum with distinct yellow tip; anterolateral margins of prothorax straight

or slightly concave; ventral margin of pygophore as in Fig. 38; theca lacking

apical tubercles piceus (Dallas)

Scutellum concolorous or with very faint white tip; anterolateral margins of
prothorax convex; theca with apical tubercles 4

4. Small species, not more than 7.5 mm long (apex of head to tip of membrane)

and 4.0 mm wide (between lateral angles of pronotum) ; ventral border of

pygophore with a small protuberance below median notch (Fig. 53)

ruckesi McDonald

Larger species, over 7.5 mm long and 4.0 mm wide; ventral border of pygo-
phore without protuberance (Fig. 46) tristis (Van Duzee)

5. Reddish brown species with distinctive zig-zag black markings on ventral

surface of abdomen; jugae not meeting in front of tylus; restricted to

E. States fulvipes (Ruckes)

Brown colored species without distinctive markings on abdomen 6

6. Anterolateral margins of pronotum convex, submarginally impressed; connexiva

with distinctly alternating pattern of yellow and black along margins (Fig. 19).
Ventral border of pygophore as in Fig. 21; dorsal border bearing a large

pair of spines (Fig. 20) abbreviatus Uhler

Anterolateral margins of pronotum straight or slightly concave (Fig. 29) ;
connexival margin entirely yellow (Fig. 30) ; ventral border of pygophore
as in Fig. 31; dorsal border without spines limbolarius (Stal)

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Figs. 11-17. H. vernalis. 11. Pygophore, dorsal view. 12. Ventral margin of pygophore.
13. Left pseudoclasper. 14. Left clasper. 15. Aedeagus, ventral view. 16. Aedeagus,
lateral view. 17. spermathecal bulb and pump, basal tubercle (B. t.) conjunctival ap-
pendage (C. a), clasper (Cl.), dorsal margin (D. m.), endophallic duct (E.), median penal
lobe (M. p.), proctiger (P.), pseudoclasper (P. c.), tubercle (T.), Theca (Th.)

Holcostethus abbreviatus Uhler, 1872

Holcostethus abbreviatus Uhler, 1872, Prelim. Rep. U.S. Geol. Surv. Mont., p. 397;
Uhler, 1876, Bull. U.S. Geol. Surv. 1, p. 289; Uhler, 1877, Bull. U.S. Geol. Surv. 3, p. 403;
Uhler, 1895, Proc. Calif. Acad. Sci. (Ser. 2) 4, p. 230; Gillette and Baker, 1895, Hem.
Colo., p. 16; Kirkaldy, 1909, Cat. Hem., p. 47.

Peribalus abbreviatus Van Duzee, 1904, Trans. Amer. Ent. Soc. 30, p. 33 ; Snow, 1906,
Trans. Kan. Acad. Sci. 20(1), p. 177; Zimmer, 1912, Univ. Nebr. Stud. 11, p. 224;
Van Duzee, 1917, Cat. Hem., p. 32.

Peribalus eatoni Bliven, 1960, Occ. Ent. 1, p. 36.

Jugae meeting in front of tylus. Anterolateral margins of pronotum distinctly bowed
out and submarginally impressed (Fig. 18). Anterior face of pronotum very slightly sloping.

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249

Figs. 18-28. H. abbreviatus. 18. Pronotum. 19. Lateral margin of abdomen, dorsal
view. 20. Pygophore. 21. Ventral margin of pygophore. 22. Left clasper. 23. Aedeagus,
lateral view. 24. Right median penal lobe, lateral view. 25. Median penal lobes, ventral
view. 26. Female genitalia. 2 7. Spermatheca. 28. Spermathecal bulb and pump, apical
tubercle (A. t.), spermathecal bulb (B.) , basal tubercle (B. t.), conjunctival appendage
(C. a.), clasper (Cl.), distal flange (D. f.), dorsal margin (D. m.), dilation of spermathecal
duct (Dl.), endophallic duct (E.), first gonocoxae (1 Gx.), second gonocoxae (2 Gx.),
median penal lobe (M. p.), proctiger (P.), pseudoclasper (P. c.), proximal flange (P. f.),
paratergite 8 (Pt. 8), paratergite 9 (Pt. 9), sclerotized rod (R.), spine (S.), sternum 10
(S. 10), sclerite (Sc.), tubercle (T.), thecal shield (T. s.), theca (Th.).

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Conspicuous pattern of alternating black and yellow squares along connexival margins
visible beyond hemelytra in females (Fig. 19).

Male genitalia (Figs. 21-25). Ventral margin of pygophore broadly V-shaped with small
emarginations at lateral extremities (Fig. 21); dorsal margin arched and bearing two
large spines, one on each side of mid-line (Fig. 20). Proctiger box-like with a posterior
lip. Claspers L-shaped, apically spatulate and serrate on inner surface, basally bearing
a large rounded tubercle (Fig. 22). Theca well sclerotized, bearing laterally on each side
a pair of small heavily sclerotized basal tubercles and a larger pair of anterior tubercles
(Fig. 23) ; apical margin bearing a large thecal shield (Fig. 23). One pair of membraneous
conjunctival appendages present. Median penal lobes narrow leaf-like structures, apically
acute, lying one on either side of S-shaped endophallic duct, basally fused by a cross bar
around duct (Figs. 24, 25).

Female genitalia (Figs. 26-28). External genitalia plate-like (Fig. 26); paratergites 8
without spiracles; paratergites 9 oblong, lying widely separated on either side of second
gonocoxae. Spermatheca typically consisting of a balloon-like dilation and apical sperma-
thecal bulb (Fig. 27) ; latter bearing two finger-like processes, one short, the other longer
and reaching proximal flange of pump (Fig. 28). Opening of spermathecal duct sur-
rounded by a ring sclerite and a small U-shaped cap sclerite.

Type. Three specimens from P. R. Uhler’s collection in the United States National
Museum are identifiable as part of the syntype series. From these the following specimen
is designated as the lectotype: $ S. Diego, Cal (segment 5 of left-antenna missing),

United States National Museum, Washington. Paralectotypes: $ N.E. Col; $ Ks, 5, 80.

Distribution. Nebraska, Kansas, Colorado, Utah, Montana, Arizona, California, Nevada,
Oregon, British Columbia.

Synonymy. The type of H. eatoni Bliven was not available for study. This name is placed
in synonymy on the basis of the description and illustration of the pygophore.

Holcostethus limbolarius (Stal, 1872)

Peribalus limbolarius Stal, 1872, Svenska Vet.-Akad. Handl. 10(4), p. 34; Uhler, 1877,
Bull. U.S. Geol. Survey 3, p. 403; Distant, 1880, Biologia Cent. Amer. Het. 1, p. 65,
pi. 6, fig. 19; Van Duzee, 1894, Bui. Buf. Soc. Nat. Sci. 5, p. 171; Gillette and Baker,
1895, Hem. Colo., p. 16; Uhler, 1904, Proc. U.S. Nat. Mus. 27, p. 351; Van Duzee,
1904, Trans. Amer. Ent. Soc. 30, p. 32; Snow, 1906, Trans. Kan. Acad. Sci. 20(1),
p. 177; Van Duzee, 1917, Cat. Hem., p. 33; Blatchley, 1926, Het. of E. N. Amer., p. 105;
Froeschner, 1941, Amer. Mid. Nat. 26, p. 135; McDonald, 1966, Quaest. Ent. 2, pp. 18,
51 figs. 106-110, 469-470 (genitalia).

Holcostethus limbolarius Kirkaldy, 1909, Cat. Hem., p. 48.

Peribalus modestus Uhler, 1872, U.S. Geol. Surv. Mont., p. 396; Uhler, 1876, Bull. U.S.
Geol. Surv. 1, p. 289.

Jugae meeting in front of tylus. Anterolateral margins of pronotum straight, sometimes
slightly concave (Fig. 29). Anterior face of pronotum declivous. Connexiva with a continuous
narrow yellow margin (Fig. 30).

Male genitalia (Figs. 31-33). Ventral margin of pygophore deeply incised on lateral
extremities, less deeply incised medianly, forming two distinct oblong plates (Fig. 31).
A pair of palmate pseudoclaspers present, these not observed in McDonald’s (1966)
description.

Remainder of male genitalia described by McDonald (1966) and Baker (1931).

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251

Figs. 29-35. H. limbolarius. 29. Pronotum. 30. Lateral margin of abdomen, dorsal
view. 31. Ventral margin of pygophore. 32. Aedeagus, ventral view. 33. Right median
penal lobe, lateral view. 34. Female genitalia. 35. Spermathecal bulb and pump, apical
tubercle (A. t.), basal tubercle (B. t.), conjunctival appendage (C. a.), endophallic duct
(E.), first gonocoxa (1 Gx.), second gonocoxa (2 Gx.), median penal lobe (M. p.),
process (Pr.), paratergite 8 (Pt. 8), paratergite 9 (Pt. 9), sternum 10 (S. 10), thecal
shield (T. s.), theca (Th.).

Female genitalia (Figs. 34, 35). Described by McDonald (1966).

Types. From the syntype series the following specimen is designated as the lectotype:
No. 202 $ Texas, Belfrage, Naturhistoriska Riksmuseet, Stockholm. 13 remaining specimens
are designated paralectotypes: 192 $ N. York, Belfrage; 193-4 $ Illinois, Belfrage;

195 $ N. York, Belfrage; 196-199 $ Texas, Belfrage; 200-1, 3 $ Texas, Belfrage;

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204 $ Mexico, Doucard; 205 $ Mexico, Salle. Specimen 192 has a label Typus; 192,
193 have been labelled Paratypus and 195 Allotypus. None of these can be considered
valid designations.

Distribution. Throughout U.S.A., southern Canada and northern Mexico.

Note. This species can be distinguished from H. abbreviatus by the following features:
Anterolateral pronotal margins straight (convex in H. abbreviatus and impressed behind) ;
connexiva with a continuous yellow margin (alternating black and yellow in H. abbreviatus ) ;
pygophore with no spines on dorsal margin (two present in H. abbreviatus ) ; spermathecal
bulb with one process not much longer than the other (Fig. 35) (in H. abbreviatus one
process much longer, reaching proximal flange of pump (Fig. 28).

Holcostethus piceus (Dallas, 1851)

Pentatoma piceus Dallas, 1851, List. Hem. 1, p. 236.

Holcostethus piceus ; Kirkaldy, 1909, Cat. Hem., p. 48.

Peribalus piceus ; Gillette and Baker, 1895, Hem. Colo., p. 16; Van Duzee, 1904, Trans.
Amer. Ent. Soc. 30, p. 34; Van Duzee, 1917, Cat. Hem., p. 33; Blatchley, 1926, Het.
E. N. Amer., p. 106.

Dark brown species. Anterolateral margins of pronotum straight as in P. limbolarius,
margined with yellow (Fig. 36). Narrow continuous pale yellow margin on connexiva.

Male genitalia. (Figs. 37-41). Ventral margin of pygophore deeply cleft centrally and
with a shallow emargination laterally on either side (Fig. 38) ; dorsal margin smoothly
arched (Fig. 37). Proctiger resembling that of H. abbreviatus , differing slightly in shape.
Claspers similar to H. tristis but bearing a distinct keel on the outer arm (Fig. 39).
Aedeagus similar to H. tristis ; no apical tubercles on theca (Figs. 40, 41).

Female genitalia. (Figs. 42, 43). Similar to H. tristis; two processes on spermathecal
bulb approximately same length (Fig. 43).

Type. Holotype. British Museum, Type No. HEM 970. $ Hudsons Bay. Type examined.

Distribution. Iowa, S. Dakota, Colorado, Montana, Illinois, Alberta, Ontario.

Note. This species can be distinguished from H. tristis by the distinct yellow tip of the
scutellum, straight anterolateral pronotal margins (convex in H. tristis), lack of distinct
flanges on the ventral margin of the pygophore (in H. tristis the flanges are impressed)
and absence of anterior tubercles on the theca (present in H. tristis).

Holcostethus fulvipes (Ruckes, 1957)

Peribalus fulvipes Ruckes, 1957, Bull. Brook. Ent. Soc. 52, p. 39.

Holcostethus fulvipes. New Combination.

Reddish brown species. Jugae not meeting in front of tylus. Femora red or rosy.
Abdominal sterna with distinct black, zig-zag markings in two parallel lines on each side
of mid-line.

Male and Female genitalia. Identical to those of H. abbreviatus .

Type. Holotype. The American Museum of Natural History, $ Lake George, N. Y.,
J. L. Zabriskie, 22 Aug. 1893; Paratype, $ (same data as holotype). Type examined.

Distribution. New York, New Hampshire.

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253

Figs. 36-43. H. piceus. 36. Dorsal view. 37. Pygophore, dorsal view. 38. Ventral margin
of pygophore. 39. Left clasper. 40. Aedeagus, lateral view. 41. Right median penal
lobe, lateral view. 42. Female genitalia. 43. Spermathecal bulb and pump, basal tubercle
(B. t.), conjunctival appendage (C. a.), clasper (Cl.), distal flange (D. f.), dorsal margin
(D. m.), endophallic duct (E.), first gonocoxa (1 Gx.), second gonocoxa (2 Gx.), median
penal lobe (M. p.), proctiger (P.), paratergite 8 (Pt. 8), paratergite 9 (Pt. 9), sternum
10 (S. 10), tubercle (T.), thecal shield (T. s.), theca (Th.).

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Holcostethus tristis (Van Duzee, 1904)

Peribalus tristis Van Duzee, 1904, Trans. Amer. Ent. Soc. 30, p. 33; Van Duzee, 1917, Cat.
Hem., p. 33.

Holcostethus tristis. New Combination.

Nearly unicolorous dark brown species with convex anterolateral margins of pronotum
margined in lighter brown (Fig. 44). Alternating pattern of light brown or yellow and
black squares along margins of connexiva.

Male genitalia (Figs. 45-48). Ventral margin of pygophore with a deep median notch
and two smaller emarginations at lateral extremities (Fig. 46) ; dorsal margins evenly
arched (Fig. 45). Claspers L-shaped; apex flattened on inner margin and bearing rows
of minute serrations; base with a large prominent tubercle (Fig. 47). Theca heavily
sclerotized; basal and apical tubercles present; apical margin produced into a thecal
shield (Fig. 48). Conjunctival appendages membraneous, bifid. Median penal lobes
similar to H. abbreviatus but somewhat broader (Fig. 49).

Female genitalia. (Figs. 50, 51). Similar to H. abbreviatus. Usually two and sometimes
three processes on spermathecal bulb (Figs. 50, 51).

Type. Lectotype. California Academy of Sciences, $ Vancouver Is., B.C., G. Taylor,
20 Aug. 1897. Type examined.

Distribution. California, Oregon, Washington, Idaho, Montana, British Columbia.

Holcostethus hirtus (Van Duzee, 1937)

Peribalus hirtus Van Duzee, 1937, Pan. Pacific Ent. 13, p. 25.

Holcostethus hirtus. New Combination.

Unicolorous dark brown species with long grey setae on dorsal and ventral surfaces.
Anterolateral margins of pronotum convex. Connexiva with a continuous narrow lighter
brown margin.

Male genitalia. No males available for examination.

Female genitalia. Similar to H. abbreviatus.

Note. This species resembles H. tristis but can be distinguished by the presence of long
grey setae on the dorsal surface of the body and the narrow light brown margin of the
abdominal connexiva (H. tristis has an alternating pattern of light brown and black squares).

Type. Holotype. California Academy of Sciences, $ Sequoia Nat. Pk., Calif. Alt. 3-
5000 ft., Collector E. C. Van Dyke, 20 June 1929. Four paratypes, all $ (same data
as holotype). Type examined.

Distribution. California.

Holcostethus ruckesi n. sp.

Dark brown species; male 7 mm long (apex of head to tip of membrane) 4 mm wide
(between lateral angles of pronotum); female 6. 5-7.3 mm long; 3. 8-4.0 mm wide. Entire
dorsal surface heavily punctate. Jugae longer than tylus, broadly rounded and sometimes
meeting apically. Antennae dark brown. Rostrum light to dark brown, apex reaching

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255

Figs. 44-51. H. tristis. 44. Dorsal view. 45. Pygophore. 46. Ventral margin of pygo-
phore. 47. Left clasper. 48. Aedeagus, lateral view. 49. Left median penal lobe, lateral
view. 50. Spermatheca. 51. Spermathecal bulb and pump, anterior tubercle (A. t.),
spermathecal bulb (B.), basal tubercle (B. t.), conjunctival appendage (C. a.), clasper
(Cl.), dorsal margin (D. m.), endophallic duct (E.), median penal lobe (M. p.), proctiger
(P.), pseudoclasper (P. c.), proximal flange (P. f.), process (Pr.), sclerotized rod (R.),
tubercle (T.).

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Ap.

Figs. 52-58. H. ruckesi n. sp. 52. Metasternum. 53. Pygophore, dorsal view. 54.
Ventral margin of pygophore. 55. Right pseudoclasper. 56. Right clasper. 57. Aedeagus,
lateral view. 58. Aedeagus, ventral view, stink gland opening and sulcus (Ap.), apical
tubercle (A. t.), basal tubercle (B. t.), conjunctival appendage (C. a.), clasper (Cl.),
dorsal margin (D. m.), endophallic duct (E.), evaporative area (Ev.), median penal lobe
(M. p.), proctiger (P.), pseudoclasper (P. c.), protuberance (Pe.), tubercle (T.), thecal
shield (T. s.), theca (Th.).

hind coxae. Ventrolateral margins of pronotum brown or yellow, bowed out and sub-
marginally impressed; pronotal disk rugose. Scutellum bluntly rounded apically, narrowing
only slightly beyond frena. Coxae and trochanters amber, femora and tibiae dark brown,
tarsi light brown. Metapleural stink gland openings with a long apically acute sulcus
(Fig. 52). Evaporative area extending onto mesopleuron. Connexiva with alternating
markings on sternal margins of abdomen, a median light brown streak on sterna 3-6.

Male genitalia (Figs. 53-58). Ventral margin of pygophore sinuate with median and
lateral notches forming two plates (Fig. 54) ; a small protuberance present below median

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257

notch; dorsal margin smoothly arched (Fig. 53). Pseudoclaspers small, elongate and
brush like (Fig. 55). Claspers L-shaped; upper surface with a keel bearing a number
of setae (Fig. 56); apex flattened and inner surface finely serrate; a large tubercle present
on base. Theca bearing large apical and small basal tubercles (Fig. 57) ; apical margin
produced into a thecal shield. Conjunctiva with one pair of appendages, these broadly
bifid (Fig. 57). Median penal lobes leaf-like apically acute and basally fused by a cross
bar (Fig. 58) ; endophallic duct lying between margins of median penal lobes and not
extending beyond them.

Female genitalia. External genitalia and spermatheca similar to H. limbolarius.

Type. Holotype, £, Rustlers Park, Chiricahua Mts. Ariz. VII-30-55; P. D. Hurd Collector.
Deposited in The American Museum of Natural History. Left antenna minus segment
4 and 5, right antenna minus segment 5 ; forelegs minus tarsomeres 2 and 3. Paratypes ,
3 9 s: Rustlers Park, Chiricahua Mts. Ariz. VII-1-55, P. D. Hurd Collector; Chirc.
Mts. Ariz. 9-11-35, E. D. Ball; McMillan Camp, 13 miles, N. Silver City, Grant Co.,
New Mexico, July 18, 1961, 6800 ft., F., P. and J. Rindge (Deposited in The American
Museum of Natural History) ; 1 9 Rustlers Pk., Ariz., Chiricahua Mts., Cochise Co.,
July 2 7, 1955 (Deposited with L. H. Holston, Louisiana State University).

Remarks. This species is named after the late Dr. Herbert Ruckes who originally noted
it in The American Museum of Natural History Collections.

Literature Cited

Baker, A. D. 1931. A study of the male genitalia of Canadian species of Pentatomidae.
Canadian J. Research, 4: 148-220.

Blatchley, W. S. 1926. “Heteroptera of Eastern North American.” Nature Publishing
Co., Indianapolis, 1116 pp.

Bliven, B. P. 1960. Studies on insects of the Redwood Empire III New Hemiptera
with notes on others. Occidental Entomologist, I : 34-42.

Dallas, W. S. 1851. List of the specimens of hemipterous insects in the collection of
the British Museum 1. London, 368 pp.

Distant, W. L. 1880. “Biologia Centrali-Americana Insecta, Rhynchota 1.” London
XX -f- 462 pp., 39 pis.

Fieber, F. X. 1860. “Die europaischen Hemiptera.” Halbfluger (Rhynchota: Heteroptera).
C. Gerold, Wien, 444 pp.

Froeschner, R. C. 1941. Contributions to a synopsis of the Hemiptera of Missouri,
Pt. 1. American Midland Naturalist, 26: 122-146.

Gillette, C. P. and Baker, C. F. 1895. A preliminary list of the Hemiptera of Colorado.

Bull. Colorado Agr. Exp. Sta. 31(1): 137 pp.

Jakovlev, B. 1902. Les Peribalus (Hemiptera-Heteroptera, Pentatomidae) de la faune
palearctique. Rev. Russe Ent. 2: 157-159.

Kirkaldy, G. W. 1909. “Catalogue of the Hemiptera (Heteroptera) Vol. 1, Cimicidae.”
Felix Dahms, Berlin, XL + 392 pp.

McDonald, F. J. D. 1966. The genitalia of North American Pentatomoidea (Hemiptera:
Heteroptera). Quaest. Ent. 2: 7-150.

Mulsant, E. and Rey, C. 1866. Histoire Naturelle des Punaises de France. Ann. Soc.
Linn. Lyon (2) 14: 1-288.

Ruckes, H. 1957. New species of Pentatomidae from North and South America
(Heteroptera). Bull. Brook. Ent. Soc. 52: 39-41.

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Snow, F. H. 1906. Some results of the University of Kansas entomological expeditions
to Arizona in 1904 and 1905. Trans. Kansas Acad. Sci. 20(1): 155-181.

Stal, C. 1872. Enumeratio Hemipterorum. Svenska Vet. -Akad. Handlingar 10(4) : 1-159.

. 1872. Genera Pentatomidarum Europae disposuit. Ofversigt Svenska Vet.-Akad.

Forhandlingar. 29(3): 31-40.

Tuxen, S. L. 1970. “Taxonomists Glossary of Genitalia in Insects.” Munksgaard,
Copenhagen. 359 pp.

Uhler, P. R. 1872. Notices of the Hemiptera of the Western Territories of the United
States. Preliminary Rept. U.S. Geol. Surv. Montana, 392-423.

— -. 1876. List of Hemiptera of the region west of the Mississippi River including

those collected during the Hayden explorations of 1873. Bull. U.S. Geol. Geog.
Surv. 1: 269-361.

. 1877. Report upon the insects collected by P. R. Uhler during the explorations of

1875. Bull. U.S. Geol. Geog. Surv. 3: 355-475.

. 1904. List of Hemiptera-Heteroptera of Las Vegas, Hot Springs, New Mexico,

collected by Messrs. E. A. Shwarz and Herbert S. Barber. Proc. U.S. Nat. Mus.
27: 349-364.

Van Duzee, E. P. 1904. Annotated list of the Pentatomidae recorded from America
North of Mexico with descriptions of some new species. Trans Amer. Ent. Soc.
30: 1-80.

. 1917. “Catalogue of the Hemiptera of America North of Mexico.” Univ. Cal.

Publ. 2, Berkley, 902 pp.

. 1937. A few new Hemiptera. Pan Pacific Ent. 13: 25-31.

Zimmer, J. T. 1912. The Pentatomidae of Nebraska. Univ. Nebraska Studies. 11:
219-251.

Vol. LXXXII, December, 1974

259

Digger Wasps as Colonizers of New Habitat (Hymenoptera: Aculeata)

Howard E. Evans

Department of Zoology and Entomology, Colorado State University,

Fort Collins, Colorado 80521

Received for Publication February 22, 1974

Abstract: Twenty-nine species of solitary wasps occupied a newly bulldozed area of sandy
soil at Bedford, Mass., during the summers of 1972 and 1973. Of these, 5 species built
up large populations in only two years, having moved into the area from adjacent study
plots. Another 17 species occupied the newly bulldozed area in smaller numbers, while
the remaining 7 species, present in adjacent plots, failed to occupy the new substrate or
did so with no increase in numbers.

INTRODUCTION

Students of solitary wasps often seek out the nests of these insects in plots of
soil made bare by man, either in excavations or fresh fills. In the more wooded
parts of the country these wasps presumably once inhabited eroded slopes and
banks along watercourses, but man’s propensity for moving soil about has created
many new areas of suitable substrate. On the whole these areas are probably no
more or less permanent than the original nesting sites, for an eroded slope, no
less than a man-made gravel pit, undergoes its own cycle of development. At
first the soil is loose and bare; gradually mosses, grasses, and small herbs take
root, reducing the bare spaces and bringing about consolidation of the soil;
gradually a new topsoil is built up, and larger plants fill or shade the remaining
bare spots. Thus ground-nesting insects must be able to colonize new exposures
rapidly, to build up large populations, and to send out new colonizers. These
statements are less true of beaches and dunes, but even such areas have their
patterns of change to which ground-nesters must adjust.

Set against these facts is the common observation that aggregations of ground-
nesting Hymenoptera sometimes persist in the same site for many years. Females
tend to nest near the place they emerged, perhaps by some form of locality
imprinting, or perhaps simply because the soil near their emergence site is the
most suitable in the area. There is evidence that females of some species make
a series of nests in the site where they emerged, but later make one or more nests
some distance away (Evans, 1966). It is also evident that individual species
vary in their tendency to adhere to one site year after year and in their ability to
colonize new areas. Evidence on these points is, however, fragmentary.

Acknowledgments: This research was conducted at the Concord Field Station of the

Museum of Comparative Zoology, Harvard University. For much assistance in the field
work, I am indebted to Victoria Rowntree and to Fred Atwood. The flies were identified
by Lloyd Knutson, the weevils by Janice White, the spiders by Herbert Levi.

New York Entomological Society, LXXXII: 259-267. December, 1974.

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New York Entomological Society

DESCRIPTION OF STUDIES

An opportunity to study the relative motility of ground nesters was presented
to me in the spring of 1972, when an area was bulldozed at the Concord Field
Station of the Museum of Comparative Zoology at Harvard (Bedford, Mass.,
site). I had been studying an area immediately adjacent for several years. The
major occupant of this area was Philanthus gibbosus (Fabricius) (as reported by
Evans, 1973), but I also recorded all nests of other species within the plot. The
plot measured 5 X 9 m and had evidently been bulldozed many years earlier; it
had since become much overgrown with moss and herbs. It is here referred to as
plot X.

The area newly bulldozed in May, 1972, was much larger and the freshly
exposed, sandy soil was at first wholly devoid of vegetation, although parts had
been filled in by grasses and herbs by late summer of 1973. This area had previ-
ously been occupied by a grass-covered bank about 3 m high by 8 m wide, the
soil from the bank having been moved elsewhere. Hence the newly exposed soil
was at first wholly devoid of ground-nesting insects. The most suitable nesting
substrate, to which my studies were confined, was in a strip 8 m wide by 50 m
long. For convenience this strip was arbitrarily divided into 3 plots, A, B, and
C, A and B being separated by a narrow strip of less suitable substrate (Fig. 1).
Plot A was studied intensively, B only slightly less so; C was visited for brief
periods several times each day. Observations were recessed during inclement
weather, but at other times (mid-June to mid- Aug.) at least one and usually
two observers were on duty nearly full time during daylight hours.

Nests were marked with numbered stakes and followed from day to day. It
was impossible to record all the activity at even a few nests, since so many
were involved. However, there was little difficulty in identifying active nests
by the appearance of the burrow and the soil at the entrance. A few females of
each of several species were marked with paint of various colors to determine
whether they made more than one nest, and if so where the additional nests were
dug. Observations were also made on parasites, and a few selected nests were
excavated to determine the number of cells and the incidence of successful
parasitism.

During the summers of 1972 and 1973, 25 species of Sphecidae and 4 species
of Pompilidae were found nesting in these plots. Of these 29 species, only 13 had
been found nesting on plot X during the preceding several years. Of the 13, the
majority merely moved into plot A without increasing notably in numbers,
although 2 species increased greatly and extended over all three plots. The
remaining 16 species presumably migrated in from other sandy areas nearby,
although I am not aware of any major nesting sites within 1.5 km. Some of
these wasps appeared in small numbers, while others became common in only two

seasons.

Vol. LXXXII, December, 1974

261

OPEN

WOODLAND

Fig. 1. Relative position of study plots, Bedford, Massachusetts.

In the following list, the species are grouped as rapid colonizers, slow colo-
nizers, and noncolonizers. In each case a few notes are presented on the nature
and abundance of the nests. Only a few biological notes are presented, since most
of these are well-studied species and biological references can be found in the
Synoptic Catalog of Hymenoptera North of Mexico and its supplements. The
genera Ageniella and Anoplius are Pompilidae, all others Sphecidae.

RAPID COLONIZERS

Crabro monticola (Packard). From 1968 to 1971, the number of nests in plot
X varied from 0 to 2, although others were noted in paths in nearby woods. In
1972, 28 nests were counted in plots A and B (none in C). In 1973, the number
had increased to 123, of which a maximum of 88 were active at one time (22
June) (females commonly make a second nest after closing the first). Since the
nests of this species are surmounted by a prominent ring of soil, they could be

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New York Entomological Society

Fig. 2. Nests of Crabro monticola (circles) and Aphilanthops frigidus (triangles) in
plots B and C during summer of 1973 (for correct relative position of plots, see Fig. 1).

easily marked and followed. Prey consisted of flies, mainly Thereva jrontalis Say
(Therevidae), which the Crabro found in abundance in the adjacent woods.

Aphilanthops frigidus (Smith). This species nested in plot X each year, 1968—
71, the number of nests varying from 1 to 3. Only 4 were noted in the newly bull-
dozed plots in 1972, but in 1973 the number increased to 58. All of these were
dug between 14 July and 2 August, the nesting cycle of the wasp being closely
synchronized with mating flights of the prey, queen Formica ants. Numerous
colonies of Formica became established in the newly bulldozed area during 1972
and 1973, and abundance of Aphilanthops was undoubtedly partly a consequence
of the abundance of prey.

Bicyrtes quadrijasciata (Say). This is a common wasp in Massachusetts, but
none had been seen at the Bedford site until 1973. On 23-25 July a number of
males were seen flying over the ground in plot C, and over the next two weeks
an estimated 12 females nested in plots A-C. It was not possible to keep an

Vol. LXXXII, December, 1974

263

accurate count of nests, since there is no distinctive pattern of soil at the entrance
as in the preceding two species.

Oxybelus bipunctatus Olivier and O. subulatus Robertson. These two species
made their first appearance in 1972, in plot A, but only a few were noted. In
1973, there were at least about 30 nests of each species in plots A and B. It was
again impossible to make an accurate count, since these are very small wasps
and the nests of short duration. O. subulatus preyed exclusively upon therevid
flies, but used consistently a smaller species than Crabro ( Psilocephala frontalis
Cole), and only males, as reported by Peckham, Kurczewski, and Peckham
(1973) in their excellent paper on members of this genus.

SLOW COLONIZERS

Anoplius marginatus (Say) and A. semirufus (Cresson). Females of the first
species were observed 10 times and females of the second 3 times, in each case
carrying spiders from the woodland into areas A and B during 1973. A. mar-
ginatus sometimes nested from the walls of inactive Crabro burrows. Both
species had been seen in the area only rarely during previous seasons.

Astata unicolor Say. One nest was found in plot X and 1 in plot A during 1973.
The species had not been observed during previous years.

Tacky sphex similis Rohwer and T. tarsatus (Say). Both of these grasshopper-
predators appeared in small numbers in plot A in 1972 and showed no increase
in 1973.

Chlorion aerarium Patton. During 1973, 2 females constructed multicellular
nests from pre-existing holes, provisioning them with Gryllus crickets. The
species had not previously been observed in the area.

Sphex ichneumoneus (Linnaeus). This large wasp has nested in a gravel strip
at the Bedford site each year, the number of nests varying from 2—10 each year.
Nysson plagiatus Cresson was seen entering nests on several occasions, and in
1970 we reared a female N. plagiatus from cells of S. ichneumoneus. This species
had been reported as a possible parasite by Ristich (1953), but the relationship
had not been confirmed.

This nesting site was about 40 m from plots X and A. None were found
nesting in these plots, but 2 females nested in plot B in 1973.

Trionyx parkeri Bohart and Menke. One nest of this wasp was noted in 1972,
2 in 1973; the species had not been seen during previous years. The nests are
of short duration, and there were undoubtedly others that were not discovered.

Gorytes canaliculatus Packard and Hoplisoides nebulosus (Packard). These
two related species, both predators on Homoptera, were seen in small numbers
in 1972, but not previously. During 1973 we noted several nests of both species,
but it is doubtful if there were more than 3-5 active females of each.

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New York Entomological Society

Bembix americana spinolae (Lepeletier). Two nests were noted in plot B in
1972, 8 in plots A and B in 1973. During 4 previous years, the species was
sighted each summer, but no nests were ever found.

Lindenius columbianus errans (Fox). This is a minute wasp, but the nests are
distinctive and are maintained for several days. We noted none until July, 1973,
when 4 appeared within 2 m2 in plot A.

Ammophila procera Dahlbom and A. urnaria Dahlbom. These wasps were
seen only occasionally prior to 1973. During that season, we estimated 3-5
females of each species. They were often seen proceeding from the woodland
into the newly bulldozed area carrying caterpillars. Most nests were in or near
plot A.

Philanthus politus Say. One nest was found in plot B in 1972. None were found
in 1973, although males were seen on several occasions on the flowers of Achillea
millefolium and Chrysanthemum leucanthemum.

Philanthus gibbosus (Fabricius). This species maintained a nearly steady
population of 32-40 nests in plot X over a period of 4 years (one female usually
maintains one nest for the season) (Evans, 1973). In 1972, there were only 12
nests in plot X (now well overgrown with vegetation), but 11 females had
established themselves in adjacent parts of plot A. In 1973, only one nest was
dug in plot X, 20 in plot A (Fig. 3). Thus the number of females declined
slightly over a three-year period, during which time there was a gradual shift
into newly available bare soil, although over only a few meters.

Cerceris prominens Banks. This species was not recorded until 1973, when 3
nests appeared 8-14 July in area B, all within 1 m2. One of these nests was
excavated on 14 July, at which time it contained 7 cells at depths of from 12 to
17 cm. Freshly provisioned cells each contained 18-19 weevils, and there were
also 8 weevils at the bottom of the burrow, 8 cm deep, in compact soil. All
weevils were Baridinae, the 38 specimens preserved belonging to 4 species:
Baris sp. (17 3 3,1 9, 10 of unknown sex), Limnobaris sp. (3 3 3,3 9 9),
Pachygeraeus sp. (2 3 3 ), and Odontocorynus sp. (1 3,1 9).

It should be noted that this is quite a different complex of weevils than those
employed by any of the three following species. The four species of weevil-
hunting Cerceris occurring at this site showed no overlap in prey whatever,
providing an excellent example of competitive exclusion.

NONCOLONIZERS

Cerceris atramontensis Banks, C. halone Banks, and C. nigrescens Smith. These
three species all nested in small numbers in plot X during the summers of 1969—
71 (Evans, 1971). However, they appeared to be absent during 1972 and 1973

Vol. LXXXII, December, 1974

265

Fig. 3. Nests recorded in plot A during summer of 1973. Plot is marked off into
squares 2 m each side. Solid circles: Crabro monticola; solid triangles: Aphilanthops

frigidus ; squares: Lindenius columbianus errans ; hollow circles: Philanthus gibbosus ;

hollow triangle: Bembix americana spinolae ; A: Astata unicolor ; B: Bicyrtes quadri-

fasciata ; H: Hoplisoides nebulosus ; X: Anacrabro ocellatus.

except for one nest of C. halone each year, again in plot X. The original nesting
sites were well covered by moss in 1973, but all 3 species failed to establish them-
selves in plots A-C.

Lyroda subita (Say). This wasp constructed its cells from the walls of the
burrows of both Philanthus gibbosus and Sphex ichneumoneus . Only one female
was noted during the summer of 1973, although the species had been fairly
plentiful during previous summers.

Ageniella conflicta Banks. During the summers of 1968-71, this species was
seen in some numbers within the Philanthus gibbosus nesting area. Females
were seen carrying spiders into inactive Philanthus burrows on several occasions.
These burrows were later closed by picking up small pebbles, bits of leaves,
and grass blades and placing them in the burrow and in a small pile over the top.
One cell was located at a depth of only 7 cm. Two prey spiders taken from wasps
both proved to be female Schizocosa bilineata (Emerton) (Lycosidae). Both
had all the legs amputated. This species was not observed in 1972, and only one
female was seen in 1973.

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New York Entomological Society

Fig. 4. Number of active females of three species over a 3 year period.

Ageniella partita Banks. This species was observed only in 1969, and like the
preceding was closely associated with inactive nests of P. gibbosus. One spider
taken as prey proved to be Zelotes sp., juvenile female (Gnaphosidae). The
wasp ran over the ground with her prey and made several short, hopping flights,
straddling the prey and holding it by the spinnerets. Only one leg had been
amputated. This is evidently the first record of this species from Massachusetts.

Anacrabro ocellatus Packard. One or two females nested in plot X each year,
1968-71. In 1972 one female nested in plot X, one in plot A, only 3 m away.
In 1973 one female nested in plot A. Thus the population remained essentially
stable despite the large amount of new substrate available.

DISCUSSION

It is evident that in this limited area and over a limited time period, some
wasps spread rapidly over newly available bare, sandy soil and increased their
numbers greatly. Others increased in numbers only slight^ or even showed a

Vol. LXXXII, December, 1974

267

decline. It is unlikely that exactly this same pattern would have been followed
under different circumstances. The proximity of the study area to a woodland
having a plentiful supply of therevid flies undoubtedly permitted Crabro monti-
cola and Oxybelus subulatus to flourish. The sudden abundance of Aphilanthops
jrigidus was also very probably related to the fact that its host, Formica jusca,
had also rapidly occupied the newly available substrate.

On the other hand, there were many blowflies and muscids around the nearby
animal pens, and solitary bees abounded in and around plots A-C. Thus there
appeared to be ample prey for species of Bembix and Philanthus. Members of
these genera are, in fact, known to remain attached to their nesting sites for
many years, and in this instance they showed little tendency to avail themselves
promptly of new potential nesting sites.

The differential effect of parasites also undoubtedly plays a role in controlling
the numbers of these insects. By colonizing new sites rapidly, species such as
Crabro monticola and Aphilanthops jrigidus may in some measure evade the
attacks of miltogrammine flies. The latter species is known to be especially sus-
ceptible to attacks by these flies (Ristich, 1956). We did not excavate any
Aphilanthops nests, but of the 7 Crabro nests excavated, all but 1 had at least
one cell containing maggots of miltogrammine flies. In all, 15 of 37 cells were
parasitized (40%). Metopia argyrocephala Meigen was reared from 3 nests,
Senotainia trilineata Wulp from one. Neither species of wasp is known to be
attacked by mutillid wasps, the behavior patterns of which seem especially
adapted for more gregarious species which persist in one site from year to year.
I believe the decline of Philanthus gibbosus to be related to the abundance of
its parasite, Dasymutilla nigripes (Fabricius).

Literature Cited

Evans, H. E. 1966. “The Comparative Ethology and Evolution of the Sand Wasps.”
Cambridge, Mass.: Harvard Univ. Press.

— . 1971. Observations on the nesting behavior of wasps of the tribe Cercerini.

Jour. Kansas Ent. Soc., 44: 500-523.

. 1973. Burrow sharing and nest transfer in the digger wasp Philanthus gibbosus

(Fabricius). Anim. Behav., 21: 302-308.

Peckham, D. J., Kurczewski, F. E., and Peckham, D. B. 1973. Nesting behavior of
nearctic species of Oxybelus (Hymenoptera: Sphecidae). Ann. Ent. Soc. Amer.,

66: 647-661.

Ristich, S. S. 1953. A study of the prey, enemies, and habits of the great golden
digger wasp Chlonon ichneumoneum (L.). Canad. Ent., 85: 374-386.

Ristich, S. S. 1956. The host relationship of a miltogrammid fly, Senotainia trilineata
(VDW). Ohio Jour. Sci., 56: 271-274.

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New York Entomological Society

Seasonal Variation in Tachysphex terminatus (Smith)

( Hymenoptera : Spliecidae, Larrinae)

Nancy B. Elliott, Frank E. Kurczewski

Department of Entomology, State University of New York
College of Environmental Science and Forestry, Syracuse, New York 13210

Received for Publication March 11, 1974

Abstract: Comparisons of morphological characters between successive generations of males
and females of Tachysphex terminatus (Smith) in upstate New York were made. Significant
differences were found for the samples of females but not for those of males. However,
the males showed greater variability within a generation than did the females.

Introduction

Tachysphex terminatus (Smith), a common North American digger wasp,
is bivoltine in the Northeast with generations emerging in June and August.
As part of a study of morphological variation in the species throughout its
range (Elliott, 1971), it was deemed desirable to determine whether there
was significant morphological variation between individuals of successive
generations.

MATERIALS AND METHODS

Samples of males and females were collected near Chittenango, New York,
in June and August, 1969. The following size-related characters were measured
for each specimen: head width, interocular distance across vertex, clypeal

width, forewing vein length along costal margin of the wing to the distal end
of the marginal cell, and length of flagellomere 2. We tested for seasonal dif-
ferences in these characters by using an F test for comparing two means.

RESULTS AND DISCUSSION

Results of comparisons for each sex are given in Table 1. Of the characters
measured, only the length of flagellomere 2 showed significant differences
between samples of males. Comparisons of the same characters in females
demonstrated differences in interocular distance, clypeal width, forewing vein
length, and length of flagellomere 2. This analysis did not show a significant
difference for head width between samples of females. However, a one-way
analysis of variance comparing several samples of females from various U.S.
localities revealed significant differences for this character for June and August
samples from Chittenango.

Acknowledgments: We thank G. C. Gaumer, Department of Entomology, Texas A. & M.
University, for collecting many of the specimens used in this study.

New York Entomological Society, LXXXII: 268-270. December, 1974.

Vol. LXXXII, December, 1974

269

Table 1. Seasonal Variation in Morphological Characters of Tachysphex terminatus

Character

$ $

Head width

Groups =

21.13

Groups =

31.04

Ind. =

18.16

Ind. =

8.70

F =

1.16

F =

3.56

Interocular

Groups =

0.94

Groups =

9.04

Distance

Ind. =

1.91

Ind. =

0.30

F =:

0.49

F =

6.93*

Clypeal width

Groups =

4.23

Groups =

14.08

Ind. =

5.29

Ind. =

3.20

F =

0.80

4.40*

Forewing

Groups =

75.72

Groups =

190.48

Vein length

Ind. =

19.58

Ind. =

14.39

F —

3.87

F =

13.24*

Flagellomere 2

Groups nr

4.16

Groups =

6.05

Ind. r=

0.39

Ind. =

0.72

F=r

10.67*

F =

8.40*

* = significant at a = 0.05.

The fact that females showed greater morphological variation between genera-
tions than males leads one to ask whether or not females really are more variable.
Lewontin (1966) suggested comparing relative variability of two samples by
comparing variances of the logarithms of the measurements using F. Table 2
shows comparisons of this kind for our samples. Only in the case of forewing
vein length for the June samples did females show greater variability than
males. Conversely, males from the August sample were more variable than
females in every character compared. A similar comparison for a sample from
Lakin, Kansas, also showed more morphological variability among the males
(Elliott, 1971). Eickwort (1969) compared variability of males and females of
Polistes exclamans Viereck (Vespidae) and found the males to be more variable,

Table 2. Relative Variability of Morphological Characters in Males and Females of

Tachysphex terminatus

Head Width

F. V. L.

Femur 1

June, 1968
S2

0.00267

0.00713

0.001

log $
S2

0.00130

0.00291

0.00175

log 2

2.054

2.45*

1.75

August, 1968
S2

0.0004

0.0002

0.000533

log $
S2

0.00141

0.00264

0.00438

log $
F

3.500*

13.00*

8.218*

* =r significant at a = 0.05.

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New York Entomological Society

as expected, because they are haploid. She conceded that reports of other male
Hymenoptera which are less variable than conspecific females might reflect
decreased responsiveness to environmental selective pressures. In such species,
the females are involved in food gathering and nesting while the primary func-
tion of the males is copulation.

Females of T. terminatus are continually subjected to the selective pressures
of the environment, especially during nesting, hunting, and prey transport.
Apparently selective pressures act upon them, resulting in greater seasonal
variation than in males. The males, on the other hand, perhaps because they
are haploid, show greater intrapopulational variation.

In T. terminatus females, August samples had larger mean values for size-
related characters than June samples. Kurczewski (1964) observed that females
of this species stored not only more prey per cell in June than in August but
also a greater biomass; hence the individuals emerging in August have been
reared on more food than those emerging in June, probably accounting for
their larger size. Dow (1942) reported that larger cocoons of the cicada-killer
were found in cells stocked with two cicadas and smaller cocoons in cells stocked
with only one.

Literature Cited

Dow, R. 1942. The relation of the prey of Sphecius speciosus to the size and sex of
the adult wasp. Ann. Entomol. Soc. Amer., 35: 310-317.

Eickwort, K. R. 1969. Differential variation of males and females in Polistes exclamans.
Evolution, 23: 391-405.

Elliott, N. B. 1971. Morphological variation in Tachysphex terminatus (Smith) and
Tachysphex similis Rohwer (Hymenoptera: Sphecidae, Larrinae). Syracuse, N.Y.,

S. U. N. Y. College of Environmental Science and Forestry. Ph.D. thesis.

Kurczewski, F. E. 1964. A comparative ethological study of some Nearctic digger
wasps of the genus Tachysphex Kohl. (Hymenoptera: Sphecidae, Larrinae). Ithaca,
N.Y., Cornell Univ. Ph.D. thesis.

Lewontin, R. C. 1966. On the measurement of relative variability. Syst. Zool., 15:
141-142.

Vol. LXXXII, December, 1974

271

Two New Genera and Two New Species of Acanthosomatidae
(Hemiptera) from South America, with a Key to the
Genera of the Western Hemisphere

L. H. Rolston1 and R. Kumar2

Received for Publication June 8, 1974

Abstract: Two new acanthosomatid genera, each with one new species, are described:

Mazanoma, new genus, M. variada, new species and type species, type locality Guardia
Vieja, Los Andes, Aconcagua, Chile; and Tolono, new genus, T. decoratus, new species and
type species, type locality Loja Province, Ecuador. A key to the acanthosomatid genera of
the Western Hemisphere is given. Sinopla bicallosus Stal is transferred to Acrophyma.

Two new genera of acanthosomatids from South America, each with one new
species, are described. A key to the acanthosomatid genera of the Western
Hemisphere, the first since that of Stal (1867), relates the new genera to those
previously known in this region.

Synonymy and generic diagnoses will appear in a revision of the world genera
of acanthosomatids now being completed by R. Kumar. The format of the
generic descriptions given here and the terminology conform to that used in the
revision of world genera.

Tunaria Piran, 1957, (not Link, 1807; not Steinmann and Hoek, 1912),
represented by T. andicola Piran, 1957, was not available for study, and the
description does not permit placing this genus in the key. From the description
and figure Tunaria Piran cannot be distinguished from Blaudus Stal.

One generically misplaced species was noted during the preparation of the key:

Acrophyma bicallosa (Stal) New Combination

Sinopla bicallosus Stal, 1872, Sv. Vet. Ak. Handl. 10(4): 62.

KEY TO GENERA OF ACANTHOSOMATIDAE OF THE WESTERN HEMISPHERE

1. Median tubercle or spine present at base of abdominal venter 9

1' Base of abdominal venter smoothly convex 2

2(1). Distal end of first antennal segment clearly surpassing apex of head 3

2’ Distal end of first antennal segment reaching little if any beyond apex of

head 6

3(2). Longitudinal sulcus on prosternum before coxae as deep as diameter of rostrum,
little wider; distal diameter of first antennal segment usually about twice

basal diameter Cylindro enema Mayr

3' Longitudinal sulcus on prosternum absent or much broader than diameter of

rostrum ; first antennal segment subcylindrical 4

1 Department of Entomology, Louisiana State University, Baton Rouge, Louisiana 70803.

2 Department of Zoology, University of Ghana, P. O. Box 67, Legon, Ghana.

New York Entomological Society, LXXXII: 271-278. December, 1974.

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4(3). Length of first antennal segment more than .8 length of head measured

dorsally Planois Signoret

4' Length of first antennal segment less than .6 length of head measured dorsally S

5(4). Prosternum shallowly depressed lengthwise Nopalis Signoret

5' Prosternum transversely convex Ditomotarsus Spinola

6(2). Paraclypei far surpassing anteclypeus, usually contiguous before anteclypeus

Mazonoma n. gen.

6' Paraclypei not or scarcely surpassing anteclypeus 7

7(6). Scent gland spout reaching more than halfway from inner margin of ostiole

to lateral margin of metapleuron Hyperbius Stal

7' Scent gland spout short 8

8. Mesosternum weakly carinate; female with one pair of Pendergrast’s organs

Tolono n. gen.

8' Mesosternum without carina ; female with two pairs of Pendergrast’s

organs Praesus Stal

9(1). Mesosternal carina greatly produced, extending anteriorly beyond procoxae;

abdominal spine appressed to right side of posterior portion of mesosternal

carina 10

9' Mesosternal carina weakly developed or absent 11

10(9). Scent gland spout reaching a little more than halfway from inner margin of ostiole

to lateral margin of metapleuron Elasmucha Stal

10' Scent gland spout reaching about three-fourths distance from inner margin

of ostiole to lateral margin of metapleuron Elasmostethus Fieber

11(9). Abdominal spine surpassing mesocoxae 12

11' Abdominal spine not reaching mesocoxae 13

12(11). Scent gland spout reaching more than halfway from inner margin of ostiole
to lateral margin of metapleuron ; abdominal spine attaining procoxae

Blaudus Stal

12' Scent gland spout reaching about one-third distance from inner margin of
ostiole to lateral margin of metapleuron ; abdominal spine attaining head

Bebaeus Dallas

13(11). Paraclypei contiguous before anteclypeus Sniploa Signoret

13' Paraclypei not surpassing anteclypeus or if longer than anteclypeus neither

markedly convergent nor contiguous 14

14(13). Scent gland spout extending much farther than halfway from inner margin

of ostiole to lateral margin of metapleuron 15

14' Scent gland spout extending halfway or less from inner margin of ostiole to

lateral margin of metapleuron 16

15(14). Anterolateral pronotal margins serrate; first antennal segment not surpassing

apex of head; mesosternum slightly depressed lengthwise ____ Pseudobebaeus Distant
15' Anterolateral pronotal margins entire, somewhat rugose; first antennal seg-
ment slightly surpassing apex of head ; mesosternum weakly carinate

Phorbanta Stal

16(14). Scent gland spout reaching halfway from inner margin of ostiole to lateral

margin of metapleuron Lanopis Signoret

16' Scent gland spout reaching not more than one-third distance from inner

margin of ostiole to lateral margin of metapleuron 17

17(16). Spine at base of abdominal venter clearly extending onto metasternum 19

17' Base of abdominal venter tuberculate, tubercle not or scarcely surpassing

posterior margin of metasternum 18

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273

18(17). Apex of head broad, anteclypeus and each paraclypeus individually rounded,

sides scarcely concave before eyes Ea Distant

18' Apex of head a narrrow smooth parabola, sides distinctly concave before

eyes Acrophyma Bergroth

19(17). Paraclypei clearly surpassing anteclypeus, dehiscent; mesosternum weakly

carinate ; first antennal segment slightly surpassing apex of head Sinopla Signoret

19' Anteclypei slightly longer than paraclypei; mesosternum without carina;

first antennal segment not reaching apex of head Hellica Stal

Mazanoma, n. g.

Type species: Mazanoma variada, n. sp.

Head. Antenniferous tubercles unarmed. Basal segment of antennae reaching almost to
apex of head. Maxillary tubercle absent. Bucculae moderately elevated, covering about
three-fourths of distance from their anterior limit to base of head, extending a little
beyond distal end of first rostral segment, arcuately truncate at their posterior limit.
Apex of rostrum resting on metasternum.

Thorax and wings. Prosternum somewhat produced on each side of broad median sulcus;
rostrum lying in sharply defined sulcus of mesosternum ; metasternum concave. Scent
gland spout drop-shaped, expanding from ostiole, covering about one-third of distance
from ostiole to lateral margin of metapleuron; evaporative area not defined (Fig. 4).
Costal margin of coria arcuate with slight expansion above posterior limit of metapleura,
radial vein plicately elevated ; membranes nearly reaching or slightly surpassing apex
of abdomen (Fig. 1).

Abdomen and general body features. Body obovate. Abdomen spatulate due to dorsad
inclination of connexiva toward margins, broadest at fourth and fifth segments, appre-
ciably wider here than pronotum. Connexiva broadly exposed. Pendergrast’s organ
covering sixth and basal half of seventh abdominal sterna. Abdomen lacking median
spine or tubercle. One discernible trichobothrium caudad and mesad of each spiracle
on sterna 3-7.

Male genitalia. Maximum diameter of phallotheca near distal limit, no great ventral
enlargement basad of conjunctiva (Fig. 7). Conjunctiva with partially sclerotized pair
of dorsolateral processes. Seminal conducting canal tubular, not expanded into conducting
chamber.

Female genitalia. 8th paratergites truncate apically, continuing contour of connexiva,
bearing exposed spiracles; remaining genital plates together forming obovate area in deep
emargination of seventh sternum (Fig. 3).

Mazanoma variada, n. sp.

Paraclypei rounded distally, contiguous or dehiscent before anteclypeus, lateral margins
slightly sinuous. Elongate depression with black confluent punctures located on each
side of disk between eye and anteclypeus and running from ocellus to level of distal end
of antenniferous tubercle. Anterior to these depressions paraclypei slope upward from
anteclypeus to lateral margins, forming disk on anterior half of head into trough. Punc-
tation other than in depressions moderately dense with punctures mostly discrete, black
or castaneous. Antennae dark brown becoming fuscous on third segment or near base of
fourth; second segment slightly bowed; length of segments 0.4-0. 5 ; 1.2— 1.3 ; 0.8; 0.8-
1.0 mm.

New York Entomological Society

2 74

Figs. 1-7. Mazanoma variada n. sp. Fig. 1. General dorsal apsect. Fig. 2. Right
paramere. Fig. 3. Apex of female abdomen, ventral aspect; Pendergrast’s organ (o) ;
tenth sternite (s) . Fig. 4. Right metapleuron; scent gland spout (sg) ; Fig. S. Apex
of male abdomen, ventral aspect. Fig. 6. Genital cup. Fig. 7. Aedeagus; conjunctival
process (cp) ; phallotheca (t).

Pronotum subtriangular, truncate apically (Fig. 1). Anterior emargination evenly
concave behind head; anterolateral margins nearly straight, obtusely carinate, lacking
denticle at anterolateral angles ; humeri broadly rounded, protruding little beyond costal
margin of hemelytra at base. Transverse tumescence on anterior pronotal disk includes

Vol. LXXXII, December, 1974

275

indistinct cicatrices and area between them, separated from anterior pronotal margin
by narrow sulcus. Punctation rather evenly distributed excepting scattered patches of
dense black punctures, especially in and near anterior submarginal sulcus, in submarginal
impression before each humerus and on humeri. Color predominately ivory, relieved by
discrete or aggregated black punctures and caudad of tumescence by light castaneous
punctures and blotches. Width at humeri 4.0-4.5, mesal length 1. 7-1.9 mm.

Basal two-thirds of scutellar disk tumescent, elevated well above surface of coria, with
large central impression. Color ivory excepting large light castaneous to brown basal patches
on each side of impunctate median fascia dividing entire scutellum. Punctation mostly
black, usually aggregated along lateral borders and frequently in dark basal patches. Sides
converging somewhat arcuately along frena, parallel beyond frena; apex subangulate.
Width at base 2. 4-2. 6, length 2.3-2 .6 mm.

Punctations of coria rather fine, brown or rufous or black, black and aggregated in
broad irregular ivory band along membrane. Membrane vitreous, venation reticulate.
Broadly exposed connexiva immaculate brownish yellow in females, broadly banded
with fuscous along both sides of intersegmental sutures in males.

Head and thorax beneath brownish yellow, usually with some irregular rufous or
castaneous infusion. Punctation moderately strong and dense, concolorous to black.

Lateral half of mesopleura with conspicuous broad transverse depression. Deep marginal
depression on metapleura extending mesad about length of lateral lobe on posterior
margin of mesopleura and located almost equidistant from anterior and posterior meta-
pleural margins. Legs stout, predominately brownish to castaneous usually, with a broad
pale band of varying width beginning near proximal end of tibiae ; superior surface of tibiae
flattened. Abdomen brownish yellow with dense concolorous punctation ; maximum

width 4.2— 5.2 mm.

Length of body 8. 7-9. 6 mm.

Posterior pygophoral margin sinuously truncate from ventral aspect (Fig. 5), sinuously
rounded from dorsal aspect (Fig. 6). Floor of genital cup transversely rugose, with a low
broad median elevation. Apical half of parameres arcuate beyond sublinear stem, forming
setose cup proximally, bearing subapical truncate tooth and terminating in acute tooth,
both teeth along mesal edge (Fig. 2). Distal margin of phallotheca unpigmented, unclearly
differentiated from conjunctiva, appearing deeply emarginated ventrally. Sclerotized

rami of dorsolateral conjunctival processes ascending from W-shaped common base.
Gonopore apparently located near dorsolateral conjunctival processes, encircled by multi-
lobate ragged-appearing distal portion of conjunctiva.

Tenth sternum of females depressed mesially, subtriangular ; tenth tergum visible,
subvertical.

Types. Holotype. Male, labeled Guardia Vieja, Cord. Aconcagua, 12 Dic-1958, Leg.
G. Barria. Polyphore dissected; right antennae missing. Deposited in University of Chile,
Santiago. (Facultad de Agronomia Museo)

Paratypes. 2 $ $, 4 2 2. Same data as holotype. Deposited as follows: $ Louisiana

State Univ.; 2, U.S. Nat. Museum; $, 2 2 2 Luis Pena collection; 2 Univ. Nac.
La Plata, Arg. (Museo La Plata)

Tolono, n. g.

Type species: Tolono decoratus, n. sp.

Head. Antenniferous tubercle unarmed. Basal segment of antennae nearly reaching to
slightly surpassing apex of head. Maxillary tubercle absent. Bucculae moderately elevated,
joined posteriorly, covering about three-fourths of distance from their anterior limit to

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base of head, extending beyond distal end of first rostral segment, arcuately truncate at
their posterior limit. Basal segment of rostrum and bucculae prominent, inclined about
45° from longitudinal axis of head (Fig. 9) ; apex of rostrum reaching onto metasternum.
Eyes not contiguous with pronotum.

Thorax and wings. Prosternum with shallow impression on each side of weak median
carina; impressions and carina narrowing toward procoxae; mesosternum and meta-
sternum nearly flat, meson weakly carinate in former, shallowly sulcate in latter. Ostiole
auriculate ; auricle short, protruding, extending about one-sixth of distance from inner
margin of ostiole to lateral margin of metapleuron ; evaporative area well defined, matte,
on metapleuron covering about half the distance from ostiole to lateral margin of meta-
pleuron (Fig. 10). Costal margin of coria smoothly sigmoid; disk without hump or
fold; membrane extending a little beyond apex of abdomen (Fig. 8).

Abdomen and general body features. Body ovoid, broadest across third abdominal segment,
slightly wider here than across humeri. Connexiva not exposed. Pendergrast’s organ
small, one on each side near anterior margin of seventh abdominal sternum. Abdomen
lacking median spine or tubercle. Paired trichobothria on each side of abdominal sterna
3-7 paralleling posterior margin of sterna and on posterior margin of narrow shallow
transverse impression; outer trichobothrium of each pair on spiracular line (Fig. 13).

Male genitalia. Conjunctiva incompletely eversible, a dorsal and ventral fold remaining
when sides completely extended, bearing dorsomedian membranous lobe and on each
side one dorsolateral process, latter with pigmented ramus apparently arising within
dorsal conjunctival fold. Seminal vesical except near distal end enclosed by median penal
lobes, these curving abruptly ventrad at termination within phallotheca; portion of
seminal vesical within phallotheca obscured.

Female genitalia. First gonocoxae large, convex, together forming approximately half
of hemisphere; 8th paratergites cultriform, each bearing a spiracle; remaining genital
plates little exposed (Fig. 12).

Tolono decoratus, n. sp.

Black, shiny, marked with shades of yellow. Dorsum broadly marked with ivory as
follows: on pronotum a crescent curving from postero-lateral margins to anterior sub-
margin ; on scutellum a transverse basal band ; on each corium a band along costal margin
and along membrane, this marginal band connected by a diagonal band running from near
base of costa to inner angle or corium. Ventrally, edge of coria ivory; broad subbasal
band on tibiae, sometimes obscurely displayed on posterior tibiae only, and usually basal
band on second rostral segment ivory or sordid yellow; broad band along lateral margins
of abdomen and rectangular area on disk of abdominal sterna 4-6 pale orange, this area
a little longer than wide, usually enclosing dark semicircular spot or band at base of one
or more segments.

Anteclypeus longer than paraclypei, cuneiform, broadly rounded at apex. Lateral
margins of paraclypei sigmoid, before eyes reflexed and concave, largely exposing an-
tenniferous tubercles from above ; distal portion of paraclypei sloping upward from
anteclypeus to outer margins. Disk rugosely punctate excepting rather smooth basal area
which includes ocelli and extends on each side to eye. Antennal segments 0.6-0. 7 ; 0.6-0. 7 ;
0.6-0. 7 ; 0.9-1. 1; 1. 1-1.4 mm in length. Width of head across eyes 1.9-2. 2; length 1.7-
2.0 mm.

Vol. LXXXII, December, 1974

277

Figs. 8-15. Tolono decoratus n. sp. Fig. 8. General dorsal aspect. Fig. 9. Head,
lateral aspect. Fig. 10. Right metapleuron. Fig. 11. Right paramere. Fig. 12. Apex of
female abdomen, ventral aspect; Pendergrast’s organ (o) ; trichobothria (tr). Fig. 13. Apex
of male abdomen, ventral aspect. Fig. 14. Genital cup. Fig. 15. Aedeagus; conjunctival
process (cp).

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Pronotum subtriangular, apically truncate and moderately emarginate ; anterolateral
margins slightly sinuous, narrowly reflexed, lacking denticle at anterolateral angles;
humeri narrowly rounded, scarcely produced (Fig. 8). Shallow arcuate impression
traversing disk about midway between anterior and posterior margins, paralleling basal
margin of pronotum; no indication of cicatrices; punctation fine, sparce, excepting a line
of strong close punctures along anterior margin which continues with diminishing strength
along anterolateral margins and a similar line in impression of disk between arc of ivory
band. Width at humeri 3.6-4.3, length at meson 1.5-1. 9 mm.

Lateral margins of scutellum faintly convex along frena, curving sigmoidly from distal
end of frena to subacute apex; disk convex basally; punctation fine, sparce. Width at
base 2. 1-2 .5, length 2. 0-2 .4 mm. Hemelytra covering connexiva; costal margin of coria
noticeably reflexed along basal half ; punctation moderately strong and rather dense along
clavical suture, elsewhere on coria fine, sparce; membrane dark, translucent, veins few,
simple, inconspicuous.

Ventral surfaces of head and thorax with moderately strong sparce punctation ; abdomen
without obvious punctation. Legs of moderate size; superior surface of tibiae flattened
toward apex; posterior tibiae bowed dorsoventrally. Seventh abdominal sternum of
female slightly protruding mesally on posterior margin, a constriction extending laterad
on each side from this point to Pendergrast’s organ.

Length of body with membrane 7.4-9. 2 mm.

Posterior margin of pygophore sinuately truncate from ventral aspect (Fig. 13), rounded
from dorsal aspect (Fig. 14) ; dense patch of setae located along anterolateral borders
within genital cup. Parameres small, subcylindrical, with rather flat production at apex
extending cephalad and bearing a few transverse ridges opposite apical face (Fig. 11).
Phallotheca weakly sclerotized and little pigmented, indistinctly differentiated from
conjunctiva. Seminal vesical terminating distally as hyaline flagellate penisfilum (Fig. 15).

In female, 9th paratergites narrowly exposed along posterior border of basal plates.
Second gonocoxae projecting obscurely as carinate triangle. Tenth sternite small, sub-
rectangular, transverse (Fig. 12).

Types. Holotype. Male, labeled E. Loja, Ecuador, 2800 m, 21-Nov. 1970, Coll. L. E.
Pena. Pygophore dissected. Deposited in Univ. Chile, Santiago.

Paratypes. 26 $ $ and 36 2 2 . Same data as holotype, 2 2 2 deposited in Univ. of
Chile, $ in Luis Pena coll.; Colombia, Narino, Laguna La Cocha, IX-26-71, G. E. Bohart,
2 $ $ , 7 2 $ Utah State Univ., $ , 2 Univ. Nac. La Plata, $ , 2 Univ. Fed. Rio Grande
do Sul, 2 Naturhistoriska Riksmuseum, Stockholm, 2 Univ. Zool. Mus. Copenhagen;
Colombia, Narino, Lago de La Cocha, 2600 m, Dec. 1-3, 1970, B. Malkin, 17 $ $,
20 $ 2 Amer. Mus. Nat. Hist., 2 $ $ , 2 La. State Univ., $ , 2 Brit. Mus. (Nat. Hist.) ;
(a) La Sierra, Jan. 29, 1931, W. A. Archer (b) Colombia, S. A., W. A. Archer, 2 $ $
U.S. Nat. Mus.

Literature Cited

Piran, A. A. 1957. Tunaria andicola, especie y genero nuevos de la fauna de Bolivia
(Hemiptera:Pentatomidae) . Rev. Chilena. Entomol. 5: 19-21.

Stal, C. 1872. Enumeratio Hemipterorum II. Svenska Vet.-Ak. Handl. 10(4) : 1—159.

Vol. LXXXII, December, 1974

279

New or Little-Known Crane Flies from Iran. I (Diptera: Tipulidae)1

Charles P. Alexander
Amherst, Massachusetts 01002

Received for Publication June 24, 1974

Abstract: A short series of papers covering the crane flies of Iran, based on materials col-
lected by Dr. Fernand Schmid in 1955 and 1956, is begun with Part I discussing certain species
of the tribe Pediciini. The new species are Pedicia ( Tricyphona ) persica, P. ( T .) iranensis,
P. (T.) elburzensis, and P. ( T .) acuspica, from the Elburz Mountains in northern Iran, and
P. (T.) luteicolor from Jugoslavia, included here for completeness.

During 1955 and 1956 the distinguished entomologist, Dr. Fernand Schmid, of
Ottawa, collected extensively in northern Iran, his materials including numerous
Tipulidae that were acquired by the writer. Several undescribed species were
represented as well as a surprisingly large number of described species that
presently are known only from Europe. Most of the Schmid materials were
from various stations in the Elburz Mountains, in the Province of Mazanderan,
along the south shore of the Caspian Sea, taken at altitudes between 1700 and
2300 meters. In this initial paper I am describing five new species in the genus
Pedicia and expect to discuss further materials in later papers under this general
title. One species from this series was described earlier as Erioptera ( Psilo -
conopa) iranica Alexander (Journal N. Y. Ent. Soc., 81: 83-84; 1973). I wish
to extend my sincere thanks to Dr. Schmid for his interest in collecting these
flies throughout the Himalayas and adjoining regions to the west. All materials
in this series of papers are preserved in the Alexander Collection.

Pedicia ( Tricyphona ) persica, n. sp.

Size medium (wing about 13 mm) ; general coloration of thorax yellow, very restrictedly
patterned with darker; legs yellow; wings yellow, restrictedly patterned with brown,
venation of outer radial field very variable; male hypopygium with dististyle very large,
placed at apex of basistyle, outer surface with abundant erect black spinoid setae.

Male. Length about 13-14 mm; wing 12-13 mm; antenna about 1.8 mm.

Female. Length about 14-17 mm; wing 13-15 mm.

Rostrum yellow; palpi with basal segment yellow, remainder medium brown. Antennae
yellow throughout; flagellar segments short and crowded, outer ones shorter than their
verticils. Head pale gray, yellowed behind; anterior vertex broad, more than three times
the diameter of scape.

Prothorax yellow. Mesonotum yellow, centers of scutal lobes pale brown. Pleura
uniformly yellow. Halteres very pale yellow. Legs with coxae and trochanters clear
light yellow; remainder of legs slightly darker yellow, outer tarsal segments pale brown.

1 Contribution from the Entomological Laboratory, University of Massachusetts.

New York Entomological Society, LXXXII: 279-284. December, 1974.

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Wings (Figs. 1-3) yellow, costal border slightly darker yellow, stigmal region, cell Sc and
base of cell C very faintly darkened; narrow brown seams at origin and fork of Rs and
adjoining veins, with less evident darkenings at R2 and m. Venation: Radial field very

variable, as discussed later; r-m before fork of Rs in all available material; in holotype
(Fig. 1) forking into a long R2+2 and a short Ri+5; in allotype (Fig. 3) vein R2+ s+i preserved
as a short suberect element, with Ri on the upper fork; in paratype (Fig. 2) veins R2+s,
Ri and R- , all at fork of Rs; in holotype cell Mi short, subequal to its petiole, in other
specimens cell Mi deep, its petiole subequal to or slightly shorter than m. Variation in
this relatively small series is shown ; a second paratype not figured has the fork of Rs
shortly beyond r-m and with two and three adventitious crossveins in cell R\.

Abdominal tergites with proximal segments brown medially, yellow laterally, in holotype
more uniformly yellow; sternites clear light yellow; hypopygium darkened. Male hypo-
pygium (Fig. 7) with tergite, t, broadly transverse, posterior border slightly produced.
Basistyle, b , with interbase, z, a flattened pale blade with sparse setae. Dististyle, d,
very large, flattened, placed at apex of basistyle; outer angle with two short spines, apical
margin with abundant erect black spinoid setae, those of the inner group shorter and
more abundant.

Holotype. $, Haradan, Iran, September 11, 1956 (Schmid). Allotype : $, Zanus, Iran,

Elburz Mts., 2,000 meters, September 21, 1955 (Schmid). Paratypes , one $, one 9,
pinned with allotype.

In its hypopygial structure the present fly is generally similar to Pedicia {Tricy phono)
riedeli (Lackschewitz) , P. ( T .) straminea (Meigen) and some other European species,
differing most evidently in details of this structure, especially the very large dististyle.
Edwards (1938) referred these species to the subgenus Crunobia Kolenati but I prefer
to retain them in Tricy phona. The venation of the radial field of the wing is very variable as
shown by the few figures here provided. Such conditions of variation in the subgenus are not
rare and have been discussed and figured by several students, such as the Nearctic P. ( T .)
inconstans (Osten Sacken) by Johnson (Psyche, 34: 216-217, figs.; 1927) and the European
P. ( T .) claripennis (Verrall) and P. ( T .) immaculata (Meigen) by Edwards (Trans. Soc.
British Ent., 5: 56-57; 1938).

Pedicia {Tricy phona) iranensis, n. sp.

General coloration of head and thorax gray, praescutum with four polished black stripes;
halteres obscure yellow; legs with bases of femora yellow, passing into brownish yellow;
wings pale brown, stigma slightly darker, prearcular field more yellowed; abdomen dark
brown; male hypopygium with arms of tergal lobes slender, subapical in position.

Male. Length about 10 mm; wing 10 mm; antenna about 1.6 mm.

Rostrum and palpi black. Antennae of male 17-segmented, black; flagellar segments
subcylindrical, proximal ones longer than their verticils; terminal segment one-half longer

(Symbols: Male hypopygium — b , basistyle; <7, dististyle; g, gonapophysis ; i, interbase;
/>, phallosome ; t, 9th tergite.)

Fig. 1. Pedicia {Tricy phona) persica, n. sp.; venation, holotype.

Fig. 2. The same; venation, paratype, showing variation.

Fig. 3. The same; venation, allotype,

Vol. LXXXII, December, 1974

Fig. 4. Pedicia ( Tricyphona ) iranensis, n. sp.; venation.

Fig. 5. Pedicia ( Tricyphona ) luteicolor , n. sp.; venation.

Fig. 6. Pedicia ( Tricyphona ) acuspica , n. sp.; venation.

Fig. 7. Pedicia ( Tricyphona ) persica, n. sp.; male hypopygium.

Fig. 8. Pedicia ( Tricyphona ) iranensis , n. sp.; male hypopygium.
Fig. 9. Pedicia ( Tricyphona ) luteicolor , n. sp.; male hypopygium.
Fig. 10. Pedicia ( Tricyphona ) elburzensis, n. sp.; male hypopygium.
Fig. 11. Pedicia ( Tricyphona ) acuspica, n. sp.; male hypopygium.

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than the penultimate. Head gray, posterior vertex behind with a central darkening (perhaps
artificially produced) .

Pronotum gray. Mesonotal praescutum gray with four polished black stripes, intermediate
pair nearly contiguous, ending some distance before suture ; scutum gray, centers of lobes
vaguely paler; scutellum and postnotum light gray, parascutella obscure yellow, deeply
excavated. Pleura gray, dorsopleural membrane obscure yellow. Halteres yellow. Legs
with coxae and trochanters yellow, fore coxae slightly more darkened basally; femora yellow
basally, outwardly brownish yellow; tibiae light brown, darker distally; tarsi black;
claws slender, yellow. Wings (Fig. 4) pale brown, stigma slightly darker, prearcular

field more yellowed; veins dark brown. Longitudinal veins from slightly beyond the
arculus with trichia. Venation: Branches of Rs consisting of the long i?2+3 and very short

Ri+ 5, R* and Re subequal in length; cell 1st M2 small; m-cu at near one-third Ms+i.

Abdomen dark brown. Male hypopygium (Fig. 8) with arms of the tergal lobes, t,
basal in position, the lobes extended beyond their insertion. Basistyle, b, with the inter-
base a very small curved club, its outer end slightly dilated.

Holotype. Javardi, Iran, October 7, 1956 (Schmid).

The most nearly related regional species include Pedicia (Tricy phono) sakkya Alexander, of
Sikkim and Assam, and three European species, P. ( T .) claripennis (Verrall), P. ( T .) lucidi-
pennis Edwards, and the Corsican, P. ( T .) trifurcata (Edwards, 1928), all differing in details
of coloration and in hypopygial structure, especially the tergite and interbase. P. ( T .) lutei-
color, n. sp., is similar in venation and in the general structure of the hypopygium, differing
evidently in the yellow body coloration.

Pedicia ( Tricyphona ) luteicolor, n. sp.

Generally similar to lucidipennis, differing in the light yellow thoracic coloration and
hypopygial structure; legs yellow, tarsi darker; wings entirely light yellow, veins darker
yellow; male hypopygium with tergal arms slender; basistyle with two terminal lobes,
both with conspicouus black setae; dististyle a flattened yellow blade, the relatively few
setae restricted to the lower margin.

Male. Length about 8 mm; wing 9.5 mm; antenna about 1.4 mm.

Female. Length about 9 mm; wing 9 mm.

Rostrum yellow; palpi dark brown. Antennae 16-segmented ; scape and pedicel yellow,
flagellum black; flagellar segments oval, gradually decreasing in size outwardly, terminal
segment larger than the penultimate. Anterior vertex yellowish gray, posterior vertex light
gray.

Thoracic dorsum light yellow, praescutum with four scarcely differentiated more orange
stripes ; pleura clear light yellow. Halteres yellow, knob very slightly more darkened
apically. Legs with coxae and trochanters light yellow; femora darker yellow; tibiae
and tarsi brown, the latter darker. Wings (Fig. 5) entirely light yellow, veins darker
yellow. Venation: Rs about as long as cell 1st M2; r-m connecting with Re shortly before its
base; cell 1st M2 closed; cell Mi about one-third longer than its petiole; m-cu shortly
before midlength of M&+i.

Abdomen yellow. Male hypopygium (Fig. 9) with tergite, t , transverse, outer lateral
angles not produced, tergal arms long and slender. Basistyle, b, with two terminal lobes,
both with conspicuous blackened setae, those of the longer ventral lobe more abundant,
arranged in a double row; interbase, i, a small curved rod, as in subfigure. Dististyle, d,
a conspicuous flattened yellow blade, with very sparse small setae on the lower margin.

Vol. LXXXII, December, 1974

283

Holotype. $, Cipari, Jugoslavia, 1.400 meters, August 11, 1955 (Schmid). Allotype , $,
pinned with type.

The species is most nearly allied to certain other European species including besides
Pedicia ( Tricyphona ) lucidipennis Edwards, also P. ( T .) claripennis (Verrall) and P. ( T .)
trifur cata (Edwards), all of which have the thoracic coloration dark brown or gray, with
conspicuous brown or blackened stripes and all species differ among themselves in hypo-
pygial details.

Pedicia ( Tricyphona ) elburzensis, n. sp.

Size medium (wing 7.5-9 mm) ; general coloration of thorax orange yellow, abdominal
tergites yellowish brown, subterminal segments slightly darker; wings broad, nearly hyaline,
cell 1st M2 closed, cell Mi subequal to its petiole; male hypopygium with median region
of tergal border produced, with very long pale setae, lateral arms erect, inner angle of
apex produced into a slender acute spine; dististyle with rostrum broad, with conspicuous
setae, apex obtuse; gonapophyses with apices slightly extended into hyaline subtriangular
blades.

Male. Length about 6.5-7 mm; wing 7.5-9 mm; antenna about 1-1.2 mm.

Female. Length about 7-8 mm; wing 8-9 mm.

Rostrum light brown; palpi black. Antennae 15-segmented; scape and pedicel dark
brown, flagellum yellowed; proximal flagellar segments short and crowded, transverse,
outer segments more elongate. Head dark brown, heavily gray pruinose.

Thoracic dorsum orange yellow, including three vaguely indicated praescutal stripes
and the scutal lobes; pleura clear yellow. Halteres light yellow. Legs with coxae and

trochanters yellow; remainder of legs light yellow, tips of femora and tibiae and the
outer tarsal segments light brown, claws long, appressed. Wings broadest opposite end

of vein 2nd A; nearly hyaline, prearcular and costal fields of light yellow, no evident stigma;
veins light brown. Longitudinal veins of about the outer four-fifths of wing with abun-
dant short trichia. Venation: Ri+ 5 relatively short, forking about opposite midlength
of cell 1st Mo ; cell M 1 subequal to its petiole; m-cu at or shortly beyond the fork of M,
in cases about to one-fifth Ms+i.

Abdominal tergites yellowish brown, subterminal two segments slightly darker, sternites
and hypopygium clearer yellow. Male hypopygium (Fig. 10) with posterior border of
tergite, t, conspicuously produced medially, provided with very long pale setae; lateral
arms erect, very slightly enlarged at apex, the apical inner angle produced into a slender
acute spine. Dististyle, d, with base moderately enlarged, with relatively short setae;

rostrum broad, apex obtuse, surface with long conspicuous setae. Gonapophysis, g,
with outer half more slender, apex slightly expanded into a hyaline subtriangular blade.

Holotype. $, Quattekas, Elburz Mts., Iran, 1,800 meters, September 19, 1955 (Schmid).
Allotype : $, Zanus, 2,000 meters, September 21, 1955, pinned with one paratype. Paratypes:
3 $$, with the allotype; $ , Barajan, 2,000 meters, September 15, 1955 (Schmid).

The present fly is most nearly related to Pedicia ( Tricyphona ) acuspica, n. sp., which dif-
fers chiefly in hypopygial characters, including the tergal arms, dististyle, and apex of
the gonapophysis. As has been indicated by Savtshenko (in Mendl, Bull. Soc. Ent. Suisse,
46: 292; 1973) in Transcaucasia and also in Iran there are various generally similar
species or races in this genus that are closely interrelated and whose exact relationships
remain uncertain.

284

New York Entomological Society

Pedicia ( Tricyphona ) acuspica, n. sp.

Size medium (wing about 9 mm) ; general coloration of thorax orange yellow, head and
abdomen brown ; halteres yellow ; legs yellow ; wings almost uniformly subhyaline, costal
border light yellow, stigma lacking; vein Ri+ 5 relatively short with r-m at near midlength,
cell 1st M 2 closed; male hypopygium with beak slender, its lower margin with very long
pale setae, each lateral tergal arm gradually narrowed into a curved acute point.

Male. Length about 9 mm; wing 9 mm; antenna about 1.3 mm.

Rostrum and palpi brown. Antennae 15-segmented ; dark brown; proximal four or

five flagellar segments short and crowded, outer segments more elongate with verticils
that are subequal to the segments, the terminal one-third longer than the penultimate
segment. Head dark brown.

Thorax almost uniformly orange yellow with a poorly indicated slightly darker central
stripe, scutal lobes similarly patterned. Halteres yellow. Legs with coxae and trochanters
orange yellow; remainder of legs yellow, outer tarsal segments very slightly darker;
claws needlelike. Wings (Fig. 6) almost uniformly subhyaline, costal border light yellow,
stigma not indicated; veins pale brown, yellow in the costal field. Longitudinal veins

of outer three-fourths of wing with small inconspicuous trichia. Venation: i?1+5 relatively

short, with r-m at near midlength; cell 1st M» closed; cell Mi subequal in length to its petiole;
vein Cu-2 very faint to scarcely evident, ending about opposite one-third Cui.

Abdomen light brown, hypopygium slightly more yellowed. Male hypopygium (Fig. 11)
with dististyle, d , relatively small, body suboval, dorsal half relatively low, with abundant
blackened spinose setae ; rostrum slender, its lower margin with a row of very long pale
setae. Ninth tergite, t, with lateral arms distinctive, appearing as erect rods that narrow
gradually into an acute curved point, median region of posterior border low convex.
Phallosome, p, including a pair of slender apophyses, their apical third outcurved and
slightly enlarged, roughened.

Holotype. $ , Mughan, Iran, June 20, 1956 (Schmid).

The present fly is generally similar to the smaller Pedicia ( Tricyphona ) zwicki Mendl (Mit-
teil. Schweitz. Ent. Gesell., 46: 291-293, figs. 1-3, 1973), described from the Akiyama Pass,
Turkey, and from P. ( T .) elburzensis, n. sp. All three species are yellow flies having cell
1st M2 of the wings present, differing from one another in relative size, details of venation,
and in the male hypopygium, especially the dististyle and lateral tergal arms. P. ( T .)
livida (Madarassy) likewise agrees in its general yellow coloration, differing in the open
cell M2 of the wings and in hypopygial characters.

Vol. LXXXII, December, 1974

285

INDEX TO SCIENTIFIC NAMES OF ANIMALS AND PLANTS
VOLUME LXXXII

Generic names begin with capital letters. New genera, species, subspecies, and varieties
are printed in italics. The following articles are not indexed: “The Sphingidae of Turrialba,
Costa Rica,” by Richard P. Seifert, pp. 45-56; “The Anthomyiidae and Muscidae of the
Great Smoky Mountains and Mt. Mitchell, North Carolina (Diptera),” by H. C. Huckett,
150-162.

Acrophyma bicallosa, 271
Adelpha, 164, 168
diodes, 236

leucophthalma tegeata, 235
Ageniella conflicta, 265
parfita, 266
Agraulis, 39, 63
Agraulis vanillae, 33, 38, 62
lucina, 61
maculosa, 63
moneta, 62
Alternaria tenuis, 125
Ammophila procera, 264
urnaria, 264
Anacrabro ocellatus, 265
Anaea electra, 171
fabius, 171
Anasimyia, 16
Anoplius marginatus, 263
semirufus, 263

Anoplocheylus aegypticus, 209
tauricus, 209
transiens, 202

Apan teles melanoscelus, 2, 3, 4, 5
Aphaenogaster, 95
Aphaenogaster megommatus, 94
pythia, 94
treatae, 94, 96
ashmeadi and floridana, 96
fulva, 96, 142
splendida, 96
longiceps, 96
megommatus, 96
pallida, 96
rudis picea, 96
tennesseensis, 96
Aphilanthrops frigidus, 262
Apis melifera, 6, 126
Archangelica anthopurpurea, 226
Argynnis Boisduvalli, 224

Asaphomyia floridensis , 183
texensis, 183
Asiodidea, 17
Aspergillus flavus, 12 7
Aspididris, 131
militaris, 131
discigera, 131
Astata unicolor, 263
Asterias, 227

Atopomyrmex mocquerysi, 94
Atta sexdens rubropilosa, 125
texana, 127
Austrachipteria, 181

Baris, 264

Basiceros conjugans, 134
singularis, 131
discigera, 131
convexiceps, 131
manni, 138

Battus polydamas, 171
Beauveria densa, 127
Bebaeus, 272

Bembix americana spinolae, 264
Bicyrtes quadrifasciata, 262
Blaudus, 272
Bombus, 24
Brevicaude, 22 7

Calliopsis, 12

Camponotus pennsylvanicus, 127
castaneus, 127

Campanotus herculeanus, 122
Caonabo casicus , 57
Capicola, 10

braunsiana, 6, 11
Cardiocondyla, 73
ectopia , 76, 82

286

New York Entomological Society

emeryi, 81, 82
nuda, 81, 82
nuda minutior, 86
venustula, 81, 82
wroughtoni, 81
nuda mauritanica, 90
Carebara junodi, 94
jauzei, 94

Castanea dentata, 22
Ceanotheus, 22
Cerapachys, 103
Cerceris atramontensis, 264
halone, 264
nigrescens, 264
prominens, 264
Chelaner antarticum, 142
Chlorion aerarium, 263
Chrysops celatus, 187
dimmocki, 187
dixianus, 183
flavidus, 187
pudicus, 183
reicherti, 187
Cimex sphacelatus, 245
Circulifer tenellus, 42
Colias interior, 224
Conomyrma bicolor, 121
Colobopsis, 73
Cordyceps

Australis, 12 7
bicephala, 127
necator, 127
proliferans, 12 7
unilateralis, 12 7
Crabro

monticola, 261
Creightonidris, 131
Crematogaster larreae, 121
opuntiae, 122
navajoa, 122
lineolata, 142
Crunobia, 280
Cylindrocnema, 271

Danaus eresimus, 171
gilippus, 17 1
plexippus, 171
Dasymutilla nigripes, 267
Dasypoda, 9

Dentachipteria highlandensis , 177
ringwoodensis, 177
Dialictus, 8
Didea, 17
Dideomimia, 17
Dione, 37, 39, 62

juno, 33, 61, 62, 63
andicola, 61, 62
huascama, 61, 62
Dircenna klugii, 1 7 1
Discothyrea testacea, 122
Ditomotarsus, 272
Drosophila, 111
Dryadula, 39
phaetusa, 33
Dryas julia, 33
Dryocoris, 245

Ea, 273

Elasmostethus, 272
Elasmucha, 272
Eueides aliphera, 171
cleobaea, 171
Euphorbia, 88
serpens, 87
Euschistus, 59
Eusphinctus, 103

Forelius, 104
Formica, 262
fusca, 267
rufa, 125
lugubris, 126
Froggattella, 104

Glaucopsyche lygdamus couperi, 222
pembina, 228
Gorytes canaliculatus, 263
Grapta, 226
Gryllus, 263

Habrobracon, 111
Haplomelitta, 13

ogilviei, 6, 9, 10, 11, 13

Vol. LXXXII, December, 1974

Heliconius charitonius, 171
dorus, 33
erato, 37
hecale, 30
isabella, 37
melpomene, 37
petiveranus, 171
ricini, 37
hermatheria, 40
talchinia, 171
Hellica, 273
Helophilus, 16
Heracleum lunatum, 226
Heraeus cincticornis, 173
Hesperapis, 9
Hesperia paniscus, 226
Holcostethus abbreviatus, 246
fulvipes, 246
hirtus, 246
limbolarius, 246
piceus, 246
ruckesi, 246
sphaecelatus, 245
tristis, 246
vernalis, 245

Hoplisoicles nebulosus, 263
Hoplocheylus americanus , 202
canadensis, 205
discalis, 203
longispinus, 205
pickardi, 203
similis, 202

Hymenostilbi australienses, 127
Hyperbius, 272

Icaricia icarioides, 229
Iridomyrmex, 89, 92, 104
humilis, 88, 115

Kalmia, 170
Korinchia, 17, 18

Laboulbenia, 125
Lanopis, 272
Larritis striata, 22 7
Lasius neoniger, 118

Laurus, 170
Leptothorax, 106

curvispinosus, 106, 107, 108, 109, 110
duloticus, 107, 108
Limenitis, 239
Limnobaris, 264

Lindenius columbianus errans, 264
Lioperna, 103
Lupinus, 229
Lycaena couperi, 222
pembina, 222
Lyroda subita, 265

Macropsis, 9
Magicicada, 189
Malotta, 16, 27

unicolor, 16, 24
Manica bradleyi, 96, 142
hunteri, 96, 142
mutica, 96, 142
rubida, 96

M azonoma variada, 273
Megaponera foetens, 127
Melitta, 9

capensis, 6

Mesembryanthemum sensu lato, 8
Messor aegyptiacus, 96
alexandri, 96
arenarius, 96
barbarus, 96
capitatus, 94
caviceps, 96
orientalis, 96
semirufus concolor, 94
structor, 94

Metarrhizum anasopliae, 127
Metopia argyrocephala, 267
Milesia, 17

bacuntius, 15
Monomorium, 92, 95
chobauti, 96
niloticoides, 96
venustum, 96
salomonis, 96
minimum, 142
Monopsis simplex, 10, 12
Morpho polyphemus polypbemus, 164
Myolepta simulans, 27
Myrmica, 141

americana, 142

288

New York Entomological Society

brevinodis, 142
brevispinosa, 142
emeryana, 142
fracticornis, 142
laevinodis, 94
lobicornis fracticornis, 94
monticola, 142
rubra, 96, 142
ruginodis, 94
sabuleti americana, 94
schenki emeryana, 94
Myscelia, 169

Neotydeus, 202, 207
Nopalis, 272
Nosema apis, 126
Novomessor albisetosus, 97
cockerelli, 97, 142
manni, 97

Nysson plagiatus, 263

Odontocorynus, 264
Ornithoptera, 37
Orthroprosopa, 17
Oxybelus bipunctatus, 263
subulatus, 263, 267

Pachygeraeus, 264
Palumbia, 17, 18
Pangonia lasiophthalma, 28
Papilio anticostiensis, 222
asterias, 225
brevicauda, 222
turnus, 226
Parahypozetes, 177
bidentatus, 181
Parides areas, 171
areas mylotes, 31
photinus, 17 1
Parrhyngia, 17
Passiflora, 31, 62
vitifloria, 30
laurifloria, 38
cyanea, 38
mucronata, 38

alata, 38
speciosa, 38
violacea, 38
jileka, 38

Paullinia pinnata, 164
fuscescens, 169
Pedicia acuspica , 279
claripennis, 280
elburzensis, 279
immaculata, 280
inconstans, 280
iranensis, 279
livida, 284
lucidipennis, 282
luteicolor, 279
persica, 279
riedeli, 280
sakkya, 282
straminea, 280
trifurcata, 282
zwicki, 284

Pentagonia wendlandia, 235
Pentatoma piceus, 252
Perdita, 12
Peribalus, 245

abbreviatus, 248
eatoni, 248
fulvipes, 252
hirtus, 254
limbolarius, 250
modestus, 250
piceus, 252
tristis, 254
Philaethria, 30
dido dido, 30
wernickei, 30
dido pygmaleon, 30
dido wernickei, 30
Pheidole, 92, 121
bicarinata, 94
creightoni, 94
crassinoda, 97
dentata, 86, 142
megacephala, 94, 121
morrisi, 97
nari, 94

pilifera pacifica, 93
ridicula, 97
saxicola, 97
sculpturata, 97

Vol. LXXXII, December, 1974

289

sitarches, 94
xerophila, 97

Philanthus gibbosus, 260, 264
Phorbanta, 272
Phyracaces, 103
Planois, 272
Plebejus icarioides, 229
Podotricha, 39
Pogonomyrmex, 141
badius, 94, 142
barbatus, 94, 142
californicus, 94, 121
desertorum, 97
imberbiculus, 94
magnacanthus, 97
maricopa, 94
occidentalis, 94
owyheci, 97
rugosus, 94

Polistes exclamans, 269
Porthetria dispar, 2
Praesus, 272
Prionyx parked, 263
Prunus, 170
Pseudobebaeus, 272
Pseudocheylus europaeus, 209
transiens, 210
Pseudodichroa, 13

Pseudonica fla villa canthara, 168, 171
Pseudozetterstedtia, IS
unicolor, 15, 24
thoracicus, 16
Psilocephala frontalis, 263
Pteralastes, IS

bomboides, 18
borealis, 15
colei, 16
literatus, 27
nubeculosus, IS
perfidious, IS
thoracicus, 15
unicolor, 18

Pyrrhogyra hypsenor, 163

Quercus, 170, 243
Quichuana, 27

Rhagina protea, 209
Rhinotropidia, 17

Salpingogaster, 17
Schizocosa bilineata, 265
Scolotydaeus ardissannae, 207
bacillus, 207
simplex , 202
Scrapter, 10

Senofainia trilineata, 267
Simopone, 103
Sinopla bicallosus, 271
Sniploa, 272
Solenopsis, 92, 115
angulatus, 94
aurea, 117
blumi, 122
germinata, 86, 116
interrupta, 120
invicta, 94, 113, 125, 142
molesta, 94
quinquecuspis, 118
richteri, 94, 113
saevissima, 94, 114
saevissima richteri, 114
xyloni, 93, 116
Solidago, 22

Spartina alterniflora, 14
Sphecodes, 13
Sphecodopsis, 13
Sphenophorus pertinax, 14
pertinax peninsularis, 14
pertinax ludovicianus, 14
Sphex ichneumoneus, 263
Stenamma brevicorne, 94
Stilbum burmense, 127
Svastra obliqua, 233
Syrittosyrphus, 17

Tachysphex similis, 263
tarsatus, 263
terminatus, 268
Tarsocheylus atomarius, 203
Temenis laothoe liberia, 17 1
Temnostoma, 17
Tenuicoris myrmeforme, 173
Tetramorium caespitum, 94
Tetrastylis ovalis, 38
Teuchocnemis lituratis, 15
Thelohania, 125
There va frontalis, 262
Thygater analis, 230
Tolono decoratus, 275

290

New York Entomological Society

Trachymyrmex septentrionalis, 142
Tribolium, 139
Tricyphona, 280
Tunaria andicola, 271
Turneria, 103

Viburnum nudum, 22
Veromessor, 141
andrei, 94, 97
pergandei, 94, 142
j uliana, 97

lariversi, 97
lobognathus, 97
smithi, 97

Victorina (Metamorpha) stelenes, 39
Wasmannia, 92

Zammara sp., 31
Zelotes, 266
Zootermopsis, 139

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Journal

of the

New York

ENTOMOLOGICAL SOCIETY

Devoted to Entomology in General

VOLUME LXXXIII

Published by the Society
New York, N. Y.

INDEX OF AUTHORS

ADAMS, ROGER G„ JR., JOHN H. LILLY, and ADRIAN G. GENTILE. Increased

Gladiolus Spike Growth with use of Certain Systematic Insecticides 251

AHMAD, SAMI and ANDREW J. FORGASH. Changes in Tolerance of Porthetria
dispar (L.) (Lepidoptera: Lymantriidae) to Insecticides in Relation to Larval

Growth and Mixed-Function Oxidase Activity 271

ALEXANDER, CHARLES P. New or Little-Known Crane Flies from Iran. II (Diptera:

Tipulidae) 2

ALEXANDER, CHARLES P. New or Little-Known Crane Flies from Iran. Ill

(Diptera: Tipulidae) 121

ALEXANDER, CHARLES P. New or Little-Known Crane Flies from Iran. IV

(Diptera: Tipulidae) 129

ALLEN, WILLIAM A. Results of an Insect Scouting Program in Virginia Soybeans 256

ASCERNO, M. E. Effects of the Insect Growth Regulator Altozar on the Parasitoid,

Microctonus aethiops, and Its Host, Hypera postica 243

ASH, NOREEN and BERNARD GREENBERG. Differential Cold Survival of Two

Sibling Species of Blow Flies, Phaenicia sericata and Phaenicia pallescens 33

BARROWS, EDWARD M. Territoriality in Male Bees (Hymenoptera: Apoidea) 280

BAUGHER, DOUGLAS G. and WILLIAM G. YENDOL. Intra-instar Respirometric
and Phase Distribution Differences in Trichoplusia ni (Hiibner) (Lepidoptera:

Noctuidae) Larvae 2 73

BECKWITH, ROY C. Douglas-Fir Tussock Moth, Orgyia pseudotsugata (McD.)

(Lepidoptera: Lasciocampidae) : Influence of Host Foliage 282

BENTON, ALLEN H. and ANDREW J. CRUMP. The Relationship of Coleomegilla
maculata (DeGeer) (Coleoptera:Coccinellidae) to the Cocoon of its Parasite Perilitus

coccinellae (Schrank) ( Hymenoptera :Braconidae) 60

BENTON, ALLEN H. and DANNY L. KELLY. An Annotated List of New York

Siphonaptera 142

BODE, WILLIAM M. Bionomics of the Tufted Apple Budmoth, Platynota idaeusalis

(Walker) (Lepidoptera: Tortricidae) , in Pennsylvania Apple Orchards 268

BOWMAN, JAMES S. An Improved Insect Pest Management Program on Sweet Corn

in New Hampshire 249

BRYANT, D. G. The Importance, Biology and Control of the Birch Casebearer,
Coleophora fuscedinella Zeller (Lepidoptera: Coleophoridae) , an Imported Pest in

Insular Newfoundland 285

BUSH, G. L. Genetic Changes occurring in Flight Muscle Enzymes of the Screwworm

Fly During Mass Rearing 275

CAMPBELL, ROBERT W. The Bimodality of Gypsy Moth, Porthetria dispar (L.)

(Lepidoptera: Lymantriidae) Populations 287

CAPINERA, JOHN L. and PEDRO BARBOSA. Dispersal of First-Instar Gypsy Moth

Larvae in Relation to Population Quality 258

CURL, G. D., P. P. BURBUTIS, and C. P. DAVIS. Rearing the European Corn Borer,

Ostrinia nubilalis (Hiibner) (Lepidoptera: Pyralidae) on a Lima Bean Medium 265

DELFINADO, MERCEDES D. and EDWARD W. BAKER. Mites (Acarina) asso-
ciated with Popilius disjunctus (Illiger) (Coleoptera: Passalidae) in Eastern United

States _ 49

DENNO, ROBERT F. Wing Polymorphism in Salt Marsh Inhabiting Fulgoroidea 253

DROOZ, A. T. Current Research with Telenomus alsophilae Viereck, an Egg Parasite of
the Fall Cankerworm, Alsophila pometaria (Harris) (Lepidoptera: Geometridae) .... 283

ELDRIDGE, B. F„ R. R. PINGER, JR., J. F. BURGER and D. E. HAYES. Environ-
mental Control of Diapause in Three Species of North American Aedine Mosquitoes

(Diptera: Culicidae) 249

FAIN, A. and K. E. HYLAND. Speleognathinae Collected From Birds in North America

(Acarina: Ereynetidae) 203

FISHER, G. T. and J. TURMEL. Control of the Apple Leaf Curling Midge, Dasyneura

mali (Kieff) (Diptera: Cecidomyiidae) in New Hampshire 244

FITZPATRICK, GEORGE and DONALD J. SUTHERLAND. Temefos Residues in the

Salt Marsh Snail Melampus bidentatus Say (Bassommatophora: Ellobiidae) 267

GITTELMAN, STEVEN H. Depth Selection in Buenoa (Heteroptera: Notonectidae) .. 265
GORDH, GORDON. Some Evolutionary Trends in the Chalcidoidea (Hymenoptera)

with Particular Reference to Host Preference 279

GRANETT, J., R. M. WESELOH and D. M. DUNBAR. Dimilin Toxicity to Apanteles
melanoscelus (Ratzeburg) (Hymenoptera: Braconidae) and Effects on Field Popu-
lations 242

HARRIGAN, W. R. and J. L. SAUNDERS. Honeylocust Pod Gall Midge, Dasyneura

gleditschae Osten Sachen (Diptera: Cecidomyiidae), Control with Dacamox® 259

HENDRICKSON, R. M., JR. Mass Rearing of Diglyphus isaea (Walker) (Hymenoptera:

Eulophidae) on Liriomyza trifoliearum Spencer (Diptera: Agromyzidae) 243

HOWER, A. A., JR. and J. E. LUKE. Response of the Alfalfa Weevil Parasitoid,
Microctonus colesi (Drea) (Hymenoptera: Braconidae), to a Recommended Insecticide

Treatment in Pennsylvania 263

HOY, M. A. Improving the Quality of Laboratory-Reared Insects - 276

HUETTEL, M. D. Monitoring the Quality of Laboratory-Reared Insects 276

HULL, L. A., D. ASQUITH and P. D. MOWERY. Mite Consuming Capacity of

Stethorus punctum (Leconte) (Coleoptera: Coccinellidae) 262

HUNG, A. C. F. and S. B. VINSON. Notes on the Male Reproductive System in Ants
(Hymenoptera: Formicidae) 192

HUSSAIN, MANZOOR. Predators of the Alfalfa Weevil, Hypera postica in Western
Nevada — a Greenhouse Study. (Coleoptera: Curculionidae) 226

JONES, JACK COLVARD and DOROTHY HOELZER. The Oenocytes of Tenebrio

molitor Linnaeus (Coleoptera: Tenebrionidae) 274

JOWYK, EUGENE A. and ZANE SMILOWITZ. Growth and Development of
Hyposoter exiguae (Viereck) (Hymenoptera: Ichneumonidae) on Two Instars of

Trichoplusia ni (Hiibner) (Lepidoptera: Noctuidae) 261

KOK, L. T. Survival of Aestivating Adult Rhinocyllus conicus Froelich (Coleoptera:

Curculionidae) at Different Temperatures and Photophases 251

KOSTELC, J. G., L. B. HENDRY and R. J. SNETSINGER. A Sex Pheromone Com-
plex of the Mushroom-Infesting Sciarid Fly, Lycoriella mali Fitch 255

LEONARD, DAVID E. Parasitization of the Spruce Budworm, Choristoneura fumiferana
(Clemens) (Lepidoptera: Tortricidae) by Brachymeria intermedia (Nees) (Hyme-

noptera: Chalcididae) 269

McDANIEL, IVAN N. Is a Black Fly Survey Worthwhile? 245

MCDONALD, JOHN L. Mosquito Control in Unusual Breeding Sites in Southern Italy
(Diptera: Culicidae) 267

MILIO, JOHN and ELTON J. HANSENS. Evaluation and Control of a Nuisance Fly
Problem (Diptera: Muscidae) at Monmouth Park Jockey Club, Oceanport, New

Jersey 252

MILLER, RICHARD C. and FRANK E. KURCZEWSKI. Comparative Behavior of
Wasps in the Genus Lindenius (Hymenoptera: Sphecidae, Crabroninae) 82

MINOT, MILDRED C. and DAVID E. LEONARD. Influence of Physical Factors on
the Behavior and Development of Brachymeria intermedia (Nees) (Hymenoptera:
Chalcididae) 269

MOORE, IAN and E. F. LEGNER. Revision of the Genus Endeodes LeConte with a
Tabular Key to the Species (Coleoptera: Melyridae) 70

MOORE, RICHARD C. Determination of Seasonal Activity of Four Fruit Pests Using
Pheromone and Other Traps 264

MORRIS, GLENN K., RON B. AIKEN and GORDON E. KERR. Calling Songs of
Neduba macneilli and N. sierranus (Orthoptera: Tettigonidae: Decticinae) 229

MUYSHONDT, ALBERTO. Notes on the Life Cycle and Natural History of Butterflies
of El Salvador. VI A. — Diaethria astala Guerin. (Nymphalidae-Callicorinae) 10

MUYSHONDT, ALBERTO and ALBERTO MUYSHONDT, JR. Notes on the Life
Cycle and Natural History of Butterflies of El Salvador. I B. — Hamadryas februa
(Nymphalidae-Hamadryadinae) 157

MUYSHONDT, ALBERTO and ALBERTO MUYSHONDT, JR. Notes on the Life
Cycle and Natural History of Butterflies of El Salvador. II B. — Hamadryas guate-
malena Bates (Nymphalidae-Hamadryadinae) 170

MUYSHONDT, ALBERTO and ALBERTO MUYSHONDT, JR. Notes on the Life
Cycle and Natural History of Butterflies of El Salvador. Ill B. — Hamadryas amphi-
nome L. (Nymphalidae-Hamadryadinae) 181

NORTON, ROY A. Elliptochthoniidae, A New Mite Family (Acarina: Oribatei) From
Mineral Soil in California 209

PADHI, SALLY B. and ARTHUR H. McINTOSH. The Use of Autoradiography to
Detect RNA in Polyhedral Inclusion Bodies of Insect Nuclear Polyhedrosis Viruses ____ 270

PEAIRS, F. B. and J. H. LILLY. Parasites Reared from Larvae of the European Corn
Borer , Ostrinia nubilalis (Hbn.), in Massachusetts, 1971-73 (Lepidoptera, Pyralidae) 36

RAIMO, BERNARD. Infecting the Gypsy Moth, Porthetria dispar (L.) (Lepidoptera:
Lymantriidae) with Nuclear Polyhedrosis Virus Vectored by Apanteles melanoscelus
(Ratzeburg) (Hymenoptera: Braconidae) 246

RALPH, CAROL PEARSON. The Milkweed Pod as an Obstacle to the Large Milkweed
Bug, Oncopeltus fasciatus (Heteroptera: Lygaeidae) 260

RECHTORIS, CAROL and ARTHUR McINTOSH. A Toxic Factor from the Estab-
lished Cell Line, CP-169 (Hink) : Carpocapsa pomonella (Lepidoptera: Olethreutidae) 271

RICKLEFS, ROBERT E. Seasonal Occurrence of Night-Flying Insects on Barro Colo-
rado Island, Panama Canal Zone 19

ROB ACKER, DAVID, K. M. WEAVER and L. B. HENDRY. Visual Stimuli in the
Host Finding Mechanism of the Parasitic Wasp Itoplectis conquisitor (Say) (Hyme-
noptera: Ichneumonidae) 257

ROBERTS, JAMES E., SR. Control of External Parasites on Cattle by Means of
Dust Bags 253

ROLSTON, L. H. A New Species and Review of Sibaria (Hemiptera: Pentatomidae) 218

SCRIBER, J. MARK and PAUL P. FEENY. Growth Form of Host Plant as a De-
terminant of Feeding Efficiencies and Growth Rates in Papilionidae and Saturniidae

(Lepidoptera) 247

SIMMONS, G. A. and C. W. CHEN. Application of Harmonic Analysis and Polynomial
Regression to Study Flight Activity of Choristoneura fumijerana (Clem.) (Lepidoptera:

Tortricidae) in the Field 266

SMILOWITZ, ZANE, CAROL A. MARTINKA and EUGENE A. JOWYK. The In-
fluence of a Juvenile Hormone Mimic (JHM) on Trichoplusia ni (Hiibner) (Lepi-
doptera: Noctuidae) and Hyposoter exiguae (Viereck) (Hymenoptera: Ichneumoni-

dae) 262

SMITH, COREY W. and ELTON J. HANSENS. The Effect of Temperature and
Humidity on the Amount of Blood Ingested by the Stable Fly, Stomoxys calcitrans L.

(Diptera: Muscidae) 235

SURLES, W. W. and L. T. KOK. Sequential Releases of Rhinocyllus conicus Froelich

(Coleoptera: Curculionidae) for the Biocontrol of Carduus Thistles 250

TALLAMY, DOUGLASS W. and ELTON J. HANSENS. A Comparison of Malaise
Trapping and Aerial Netting for Research on Houseflies and Deerflies (Diptera:

Tabanidae) 245

TAUBER, CATHERINE A. and MAURICE J. TAUBER. Systematics and Ecology

of Chrysopidae (Neuroptera) : Theoretical and Applied Implications 277

THOMAS, J. H. and C. H. HILL. A Seasonal History of the Variegated Leafroller,
Platynota flavedana (Clemens) (Lepidoptera: Tortricidae), in Virginia Apple

Orchards 260

TINGEY, W. M., V. E. GRACEN, and J. M. SCRIBER. Leaf Feeding Resistance to
the European Corn Borer, Ostrinia nubilalis (Hiibner) (Lepidoptera: Pyralidae), in

Tropical Maize 256

TOPOFF, HOWARD. Behavioral Changes in the Army Ant Neivamyrmex nigrescens

during the Nomadic and Statary Phases 38

UEBEL, E. C., R. E. MENZER, P. E. SONNET, and R. W. MILLER. Identification
of the Copulatory Sex Pheromone of the Little House Fly, Fannia canicularis (L.)
(Diptera: Muscidae) 258

VANDENBERG, JOHN D. and RICHARD S. SOPER. Isolation and Identification of
Entomophthora spp. Fres. (Phy corny cetes: Entomophthorales) from the Spruce Bud-

worm Choristoneura fumiferana Clem. (Lepidoptera: Tortricidae) 254

VANDERLIN, ROBERT L. and FREDERICK A. STREAMS. Reproductive Diapause

in Notonecta undulata (Say) (Hemiptera: Notonectidae) 248

VOLNEY, J. The Role of Defoliators in the Arthropod Community of Red Maple

Crowns 283

WARD, R. H. and R. L. PIENKOWSKI. Cassida rubiginosa Muller (Coleoptera:

Chrysomelidae) , a Potential Biocontrol Agent of Thistles in Virginia 247

WEINER, THOMAS J. and ELTON J. HANSENS. Species and Numbers of Blood-
sucking Flies Feeding on Hogs and Other Animals in Southern New Jersey 198

WESELOH, R. M. Seasonal Variations in Activity of Apanteles melanoscelus Ratzeburg
(Hymenoptera: Braconidae) Adults as Related to Seasonal Variations in Age Structure

of its Host, Porthetria dispar (L.) (Lepidoptera: Lymantriidae) 242

WHALON, MARK E. and BRUCE L. PARKER. Oxygen Consumption of Coleomegilla
maculata lengi Timberlake (Coleoptera: Coccinellidae) Measured in a Differential

Respirometer 272

WOOLDRIDGE, DAVID P. A Phylogeny for Paracymus Thomson (Coleoptera: Hy-
drophilidae) Based on Adult Characters 273

' \ /

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The New York Entomological Society

Incorporating The Brooklyn Entomological Society

Incorporated May 21, 1968

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Organized June 29, 1892 — Incorporated February 25, 1893

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Founded in 1872 — Incorporated in 1885

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Fordham University, New York 10458

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American Museum of Natural History, New York 10024

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Secretary, Dr. Charles C. Porter

Fordham University, New York 10458
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Isaac Albert Research Institute, Brooklyn, N.Y. 11203
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Fairleigh Dickinson University, Madison, N.J. 07940
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Class of 1975

Trustees

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Dr. Howard Topoff

Class of 1976

Dr. David C. Miller

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Dr. Norman Platnick

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Mailed May 28, 1975

The Journal of the /New York Entomological Society is published quarterly for the Sjociety by Allen Press
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Journal of the

New York Entomological Society

Volume LXXXIII March 1975

No. 1

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Waksman Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lois J. Keller, RSM
Dr. Herbert T. Streu

Publication Committee

Dr. Kumar Krishna Dr. Ayodha P. Gupta

Dr. James Forbes, Chairman

CONTENTS

New or Little-Known Crane Flies from Iran II (Diptera: Tipulidae)

Charles P. Alexander 2

Notes on the Life Cycle and Natural History of Butterflies of El Salvador.

VI A. — Diaethria astala Guerin. (Nymphalidae-Callicorinae)

Alberto Muyshondt 10

Seasonal Occurrence of Night-Flying Insects on Barro Colorado Island, Panama
Canal Zone Robert E. Ricklefs 19

Differential Cold Survival of Two Sibling Species of Blow Flies, Phoenicia
sericata and Phoenicia pallescens Noreen Ash and Bernard Greenberg 33

Parasites Reared from Larvae of the European Corn Borer, Ostrinia nubilalis
(Hbn.), in Massachusetts, 1971—73 (Lepidoptera, Pyralidae)

F. B. Peairs and J. H. Lilly 36

Behavioral Changes in the Army Ant Neivamyrniex nigrescens during the
Nomadic and Statary Phases Howard Topoff 38

Mites (Aearina) Associated with Popilius disjunctus (Illiger) (Coleoptera:
Passalidae) in Eastern United States

Mercedes D. Delfinado and Edward W. Baker 49

The Relationship of Coleomegilla maculata (DeGeer) (Coleoptera :Coccinelli-
dae) to the Cocoon of its parasite Perilitus coccinellae (Schrank) (Hymenop-
tera:Braeonidae) Allen H. Benton and Andrew J. Crump 60

Proceedings of the New York Entomological Society: Abstracts 64

Announcement 67

Book Reviews 67

New York Entomological Society

New or Little-Known Crane Flies from Iran
II (Diptera: Tipulidae)1

Charles P. Alexander
Amherst, Massachusetts 01002

Received for Publication July 17, 1974

Abstract: The initial part of this short series of papers concerning the crane flies of Iran
was published in this Journal 82: 279, 1974 and concerned the tribe Pediciini. At this
time I am discussing the Eriopterine genus Gonomyia and provide descriptions of seven
undescribed species, Gonomyia ( Idiocera ) curticurva, G. (/.) laterospina, G. (/.) orthophallus,
G. (/.) spinistylata, G. ( Gonomyia ) basilobata, G. ( G .) elburzensis , and G. (G.) oxybeles,
from the Elburz Mountains in northern Iran.

As was discussed in the first part under this title the large series of Tipulidae
from northern Iran was collected by Dr. Fernand Schmid in 1955 and 1956 and
added greatly to the then poorly known Iranese crane fly fauna. I am very
indebted to Dr. Schmid for his efforts in making known the Tipulidae of virtually
all of southern Asia. The types of the new species are preserved in the Alexander
collection.

Gonomyia ( Idiocera ) curticurva, n. sp.

Mesonotal praescutum with three gray stripes, interspaces with two long pale brown
lines, pleura light brown above, below chiefly yellow; femora yellowed with a narrow
pale brown nearly terminal ring; wings faintly darkened, prearcular and costal regions pale
yellow, stigma pale brown; Sc short, Sci ending opposite origin of Rs, Sc2 far retracted;
m-cu more than its length before fork of M ; male hypopygium with three dististyles, all
terminating in blackened points; apex of aedeagus a very small curved hook.

Male. Length about 4 mm; wing 5 mm. Rostrum dark brown; palpi black. Antennae
with scape brown, pedicel yellow, flagellum brownish black. Head with anterior vertex
yellow, posterior vertex gray.

Pronotum brownish gray, scutellum and sides of scutum light yellow. Mesonotal prae-
scutum with three gray stripes, the interspaces appearing as two long pale brown lines,
pseudosutural foveae darkened; scutum gray, lobes slightly infuscated, posterior angles
yellowed; scutellum obscure yellow; postnotal mediotergite brownish gray, pleurotergite
light brown with a yellowed spot. Pleura light brown dorsally, sternopleurite and posterior
sclerites yellow, ventral sternopleurite pale brown. Halteres with stem obscure yellow,
knob pale brown. Legs with coxae yellow, bases of fore and middle pairs pale brown;
trochanters yellow; femora yellowed, with a narrow pale brown nearly terminal ring;
tibiae and basitarsi yellow, apices darkened, remainder of tarsi pale brown. Wings faintly
darkened, prearcular and costal regions pale yellow; stigma light brown. Longitudinal
veins beyond general level of origin of Rs with trichia, more sparse on vein Rs and tips of

1 Contribution from the Entomological Laboratory, University of Massachusetts.
New York Entomological Society, LXXXIII: 2-9. March, 1975.

Vol. LXXXIII, March, 1975

the Anals. Venation: Sc short, Sci ending opposite origin of Rs, Sc2 far retracted, Set and Rs
subequal in length ; distance on costa between Ri+2 and Rs about one-third the length of the
latter; m-cu more than its length before the fork of M.

Abdominal tergites brown, sternites paler; hypopygium brownish yellow. Male hypo-
pygium (Fig. 2) with outer lobe of basistyle, b, long, vestiture relatively short, inner lobe
lacking. Three dististyles, d, all terminating in blackened points; outer style curved,
narrowed gradually into a long slender spine ; intermediate style largest, its outer angle
a long gently curved spine, the inner angle short and stout; inner style straight, narrowed
gradually into a long straight blackened spine, several long pale setae at base. Aedeagus, a,
long, outer end slightly curved, apex a very small hook.

Holotype. $, Bar, Iran, June 30, 1956 (Schmid).

The species is generally similar to Gonomyia ( Idiocera ) orthophallus, n. sp., differing in
hypopygial structure, especially the intermediate dististyle and the aedeagus.

Gonomyia ( Idiocera ) laterospina, n. sp.

General coloration gray, patterned with brown ; femora yellow, tips narrowly brown ;
wings subhyaline, unpatterned except for the pale brown stigma, Sci very long; male hypo-
pygium with four dististyles, the outermost a narrow rod with a small blackened spine at
near midlength ; aedeagus with apex slightly curved, subtended by two low points.

Male. Length about 5 mm; wing about 5 mm. Rostrum gray; palpi black. Antennae
black. Head obscure gray.

Pronotal scutum dark gray, laterally light yellow, scutellum yellow. Mesonotal prae-
scutum with disk gray, with two intermediate more brownish gray longitudinal stripes,
humeral and lateral borders yellowed; scutum gray, centers of lobes vaguely more darkened;
scutellum grayish brown; postnotum gray. Pleura brownish gray above, lower half yellowed,
the ventral part slightly darker. Halteres with stem pale, knob dark brown. Legs with
coxae and trochanters yellow ; femora yellow, tips narrowly brown ; tibiae yellow, tips
very narrowly darkened; basitarsi yellowed, remainder of tarsi brown. Wings subhyaline,
stigma pale brown, inconspicuous; veins pale brown, Sc more yellowed. Macrotrichia on
most longitudinal veins beyond level of origin of Rs, lacking on R3, present on apices of
both Anal veins. Venation: Sci ending slightly beyond origin of Rs, Sc2 far retracted, Sci
nearly as long as Rs; distance on costa between veins Ri+2 and R.. about one-third to
one-half the latter vein.

Abdominal tergites dark brown, sternites and hypopygium slightly paler. Male hypo-
pygium (Fig. 3) with basistyle, b, produced into a longer outer and a small slender inner
lobe. Four dististyles, d, the outermost a long nearly straight rod, its basal half stouter,
at point of narrowing with a small blackened spine; second style largest, basal half slender,
apically dilated into a subcircular blade that terminates in an acute spine, at base of blade
with a smaller accessory projection that bears two unequal spines, the more basal one
smaller; third style a long very slender nearly straight blackened spine; inner style slightly
shorter, pale throughout, apex with a microscopic point. Aedeagus, a, with apex a slightly
curved spine that is subtended by two low points.

Holotype. $, Bar, Iran, June 30, 1956 (Schmid). Paratopotypes. 3 $$, with type.

From other regional species of Idiocera having the wings unpatterned except for the
stigmal darkening, the present fly is most readily told by the hypopygial structure,
including the four dististyles and the structure of the aedeagus. It is generally similar to
Gonomyia ( Idiocera ) alexanderiana (Lackschewitz) of Albania, differing in hypopygial
structure.

New York Entomological Society

Gonomyia ( Idiocera ) orthophallus, n. sp.

Size relatively large (wing about 6 mm) ; thorax dark grayish brown, patterned with
yellow, especially on pleura ; wings subhyaline, stigma scarcely darker ; Sc long, Sci
ending beyond one-third length of Rs ; male hypopygium with three dististyles, the inner
and outer similar to one another, their outer ends pointed; aedeagus long and straight,
apex not decurved.

Male. Length about 5 mm; wing 6 mm. Rostrum and palpi black. Antennae with scape
yellowed, flagellum black, the segments elongate. Front and orbits yellowed, vertex
brown.

Pronotal scutum dark grayish brown, margins broadly yellow; scutellum obscure
yellow. Mesonotal praescutum grayish brown with two intermediate brown stripes, lateral
borders yellow; scutal lobes brown, each with two vague darker brown spots, median
area anteriorly obscure yellow; scutellum obscure brownish yellow, base and a narrow
central area darkened; postnotum brownish gray, sides of anterior half of mediotergite
light yellow, pleurotergite light yellow above, lower third brownish gray. Pleura light
yellow, propleura, dorsal mesopleura and ventral sternopleurite dark gray, meron yellow,
narrowly darkened anteriorly. Halteres with stem yellow, knob brown. Legs with fore
coxae light yellow, darkened basally, mid-coxae similar, the darkened part restricted,
posterior coxae yellow; trochanters yellow; femora brownish yellow, tips darker; tibiae
and tarsi brown. Wings subhyaline, very faintly tinted, stigma scarcely darker than the
ground; veins light brown, Sc more yellowed. Longitudinal veins beyond general level
of origin of Rs with long trichia, including also outer ends of both anal veins. Venation:
Sc long, Sci ending shortly beyond one-third Rs, Sci long, about one-half Rs ; veins R i+2 and
Ra narrowly separated at costal border ; m-cu about its own length or slightly more
before the fork of M .

Abdomen brown. Male hypopygium (Fig. 4) with outer lobe of basistyle, b, large and
fleshy, with long setae, inner lobe small. Three dististyles, d, the outer and inner generally
similar in size and length, appearing as straight lobes, their pointed outer ends blackened, the
amount less on the inner style ; intermediate style much larger, with outer half more nar-
rowed, at its base with a blackened point. Phallosome with the aedeagus, a, long and
straight, apex not decurved; gonapophyses small, narrow, slightly curved outwardly,
separated by a low setiferous cushion.

Holotype. $, Durbadam, Iran, July 3, 1956 (Schmid). Paratypes. $, Bar, Iran,
June 30, 1956; S, Firouz Kuh, August 14, 1956 (Schmid).

Fig. 1. Gonomyia ( Gonomyia ) basilobata, n. sp.; venation.

Fig. 2. Gonomyia ( Idiocera ) curticurva, n. sp.; male hypopygium.

Fig. 3. Gonomyia ( Idiocera ) laterospina, n. sp. ; male hypopygium.

Fig. 4. Gonomyia ( Idiocera ) orthophallus, n. sp.; male hypopygium.

Fig. 5. Gonomyia ( Idiocera ) spinulistyla , n. sp.; male hypopygium.

Fig. 6. Gonomyia ( Gonomyia ) basilobata, n. sp.; male hypopygium.

Fig. 7. Gonomyia ( Gonomyia ) elburzensis, n. sp.; male hypopygium.

Fig. 8. Gonomyia ( Gonomyia ) oxybeles, n. sp.; male hypopygium.

(Symbols: Male hypopygium — a, aedeagus; b, basistyle; d, dististyles; id, inner dististyle;
md, middle dististyle; p, phallosome; t, 9th tergite).

Vol. LXXXIII, March, 1975

New York Entomological Society

The present fly is most readily distinguished from generally similar regional species by
hypopygial characters, especially the straight slender aedeagus and the conformation of
the three dististyles. Such similar regional species include Gonomyia ( ldiocera ) displosa
Alexander and G. (/.) phallostena Alexander, both from Kashmir, all differing among them-
selves in hypopygial structure.

Gonomyia ( ldiocera ) spinulistyla, n. sp.

General coloration of mesonotum brownish gray, praescutum with two longitudinal
brown stripes; pleura brownish black with a whitened longitudinal stripe; knob of halteres
black; femora yellow with a narrow pale brown nearly terminal darkening; wings whitened,
with a restricted dark brown pattern beyond the cord; veins Ri+2 and R3 contiguous at
margin closing the cell; male hypopygium with three dististyles, the inner one distinctive;
apex of aedeagus subtended by triangular points.

Male. Length about 6 mm; wing 5 mm. Rostrum and palpi black. Antennae broken.
Head above gray, extensively light yellow posteriorly.

Mesonotal praescutum light gray with two longitudinal brown stripes that are narrower
than the median interspaces; scutum brownish gray, each lobe with a large brown area;
scutellum brownish gray, vaguely paler medially; postnotal mediotergite dark brownish
gray, pleurotergite extensively light yellow, darker behind. Pleura brownish black,
ventrally sparsely pruinose with a conspicuous whitened longitudinal stripe extending
from fore coxae to the abdomen, widened behind, dorsopleural region whitish yellow.
Halteres long, stem yellow, knob black. Legs with fore coxae as described, middle coxae
blackened basally, remainder broadly white, posterior coxae obscure yellow, bases narrowly
brownish black ; trochanters yellow ; femora yellow with a narrow pale brown nearly terminal
darkening; tibiae and basitarsi yellow, extreme tips slightly darkened, remainder of tarsi
brown. Wings whitened, prearcular and costal fields light yellow; a restricted dark brown
pattern including h, Sc2 and base of Rs , tip of Sci, stigma, and a more extensive apical area
chiefly in cell Riy the center of the marking with a whitened spot ; other darkened seams
over cord, including m-cu , and at arculus; veins brownish yellow, clearer yellow in the
costal field, dark brown in the patterned areas. Venation: Sc long, ending nearly opposite
midlength of Rs, Sc2 retracted, placed shortly before Rs ; Ri+2 and R3 confluent or
contiguous at margin closing cell R^ ; vein Ri strongly upcurved on outer third ; m-cu
about one and one-half times its length before fork of M.

Abdominal tergites dark brown, incisures vaguely paler, sternites lighter brown. Male
hypopygium (Fig. 5) with three dististyles, d, outer style expanded outwardly, divided into
two major spines, the outer one long and slender with a smaller basal spinule, inner spine
shorter, strongly curved; intermediate style bifid at apex into a long slender spine and a
shorter spur; inner style distinctive, short and compact, terminating in a small spine, the
base of style with very numerous blackened spinules. Aedeagus, a, curved at apex, at
base subtended on either side by a small triangular point.

Holotype. $, Bar, Iran, June 30, 1956 (Schmid). Paratopotype, a fragmentary $, mounted
on slide.

Regional species that are generally similar to the present fly include Gonomyia ( ldiocera )
jucunda Loew, G. (/.) punctata (Lackschewitz) and G. (I.) schrenki Mik, all differing
among themselves in hypopygial structure, especially the dististyles.

Gonomyia ( Gonomyia ) basilobata, n. sp.

Rostrum light yellow; palpi and antennae black; mesonotal praescutum with disk
dark brown, lateral margins light yellow, pleura yellow, striped longitudinally with pale

Vol. LXXXIII, March, 1975

brown; wings with stigma slightly infuscated; male hypopygium with a small lobe at base
of inner dististyle; phallosome with a single blackened apophysis, the second one entirely
pale, apex of aedeagus obtusely rounded.

Male. Length about 5-5.5 mm; wing 4.5-5 mm. Rostrum light yellow; palpi black.
Antennae brownish black, pedicel more intensely darkened. Head gray.

Prothorax light yellow, darker on sides. Mesonotal praescutum with disk dark brown,
interspaces not or scarcely differentiated, humeral and lateral regions light yellow; scutum
with lobes blackened, pruinose, posterior angles slightly reddened, median area yellow;
scutellum brown, posterior border narrowly more darkened; postnotal mediotergite gray,
sides yellow; pleurotergite yellowed. Pleura yellow, striped with pale brown, the dorsal
area narrower and poorly delimited, sternal darkening more extensive. Halteres with
stem pale yellow, knob brown. Legs with coxae pale brown, middle pair more yellowed;
trochanters yellow; remainder of legs brown. Wings (Fig. 1) weakly darkened, stigma
slightly infuscated, large; veins medium brown. Venation: Sci ending shortly beyond origin
of Rs; i?2+3+4 long, gently arcuated; m-cu slightly before or beyond m-cu.

Abdomen brown, hypopygium slightly more yellowed. Male hypopygium (Fig. 6)
with outer dististyle, d, narrow, apical flange elongate; inner style with dorsal spine stout,
at base of style with a small lobe tipped with a strong seta. Phallosome, p , with two
gonapophyses, one blackened, the more slender lower spine entirely pale; apex of aedeagus
obtusely rounded.

Holotype. $, Mishgin, Iran, August 21, 1956 (Schmid). Paratypes. $, Bar, Iran,
June 30, 1956; $, Durbadam, Iran, July 3, 1956 (Schmid).

The present fly is most readily distinguished from other generally similar regional species by
the structure of the inner dististyle, with the reduced basal tubercle, and the single blackened
gonapophysis of the phallosome. The genotype of Gonomyia, tenella (Meigen), has the inner
dististyle generally similar but the phallosome has both gonapophyses blackened and the
apex of the aedeagus different.

Gonomyia ( Gonomyia ) elburzensis, n. sp.

General coloration of thorax yellow, praescutum with disk dark brown, scutal lobes and
mediotergite brown, pleura yellow, restrictedly patterned with brown; legs light brown;
male hypopygium with gonapophyses unequal, both heavily blackened, terminating in
slender spines, apex of aedeagus short and obtuse.

Male. Length about 4. 5-4. 6 mm; wing 5-5.3 mm. Rostrum, palpi and antennae brownish
black. Head gray.

Prothorax clear light yellow. Mesonotal praescutum with disk dark brown, the inter-
spaces concolorous with the stripes, lateral borders clear light yellow; scutal lobes dark
brown, median region light yellow, narrowly darkened behind; scutellum light yellow;
mediotergite brown, anterolateral portions and the pleurotergite light yellow. Pleura
chiefly light yellow, including the dorsopleural membrane; ventral sternopleurite and a
small area on lower anepisternum brown. Halteres with stem light yellow, apex of knob
brown. Legs with fore coxae weakly darkened, remaining coxae light yellow; trochanters
brownish yellow; remainder of legs light brown. Wings subhyaline, stigma not indicated;
veins pale brown. Venation: Sci ending about opposite one-fourth to one-fifth Rs ; m-cu at
fork of M.

Abdominal tergites brown, incisures pale; sternites light yellow medially and on extreme

New York Entomological Society

margins, sublateral areas narrowly brown. Male hypopygium (Fig. 7) with lobe of
basistyle, b, elongate, the apical glabrous flange elongate, outer setae long. Dististyle, d,
about as shown; outer basal lobe long and slender, tipped with a single long seta; rostral
prolongation short, the two modified setae elongate ; summit of style blackened. Phallosome,
p, with both gonapophyses blackened and extended into slender spines ; apex of aedeagus short
and obtuse, almost rounded.

Holotype. $, Lius, Iran, 2,200 meters, September 14, 1955 (Schmid). Paratopotype.
S, pinned with type. Paratypes, $, Waliabad, Iran, September 24, 1956; $, Nandeh, Iran,
June 10, 1956 (Schmid).

The most similar regional species include Gonomyia ( Gonomyia ) tenella (Meigen),
Europe, G. (G.) chalaza Alexander, Pakistan, and some others, all differing in hypopygial
details, particularly in the dististyles and phallosome.

Gonomyia ( Gonomyia ) oxybeles, n. sp.

Size large (wing over 6 mm); rostrum light yellow, palpi and antennae black; thoracic
dorsum grayish brown and yellow, pleura conspicuously patterned with brown ; wings
light brown, stigma very pale, Sex ending shortly beyond origin of Rs , cell Rz large;
male hypopygium with outer lobe of basistyle small; outer dististyle with lateral blade slightly
darkened, lateral spine of inner dististyle with a small point at base; phallosome distinctive,
especially the aedeagus beyond the gonapophyses.

Male. Length about 6.5 mm; wing 6.2 mm. Female. Length about 7.5 mm; wing 6.5 mm.
Rostrum light yellow; palpi black. Antennae black; flagellar segments long-oval. Head
light gray.

Pronotum light yellow, sides of scutum darkened. Mesonotal praescutum with disk
chiefly dark gray, stripes margined with brown, sides broadly light yellow; scutum yellow,
anterior and mesal parts of lobes dark brown, posterior ends yellowed; scutellum yellow;
postnotal mediotergite dark brown medially, sides yellow, pleurotergite yellow, above and
below vaguely margined with brown. Pleura chiefly light yellow, patterned with light
brown, including a narrow stripe on propleura and mesopleura, ventral sternopleurite more
extensively darkened. Halteres with stem light yellow, knob brown. Legs with coxae
and trochanters yellow; remainder of legs broken. Wings very light brown, prearcular
and costal fields light yellow, stigma very pale brown ; veins pale brown, Sc yellow.
Venation: Sci ending shortly beyond origin of Rs, in type about opposite one-fifth the length
of vein ; cell Rs large ; m-cu varying slightly in position, from before to beyond the fork of M.

Abdomen dark brown, lateral borders of tergites yellowed, the posterior margins more
narrowly so. Male hypopygium (Fig. 8) with outer lobe of basistyle, b, small. Outer
dististyle, d , a straight blade, outer lateral margin slightly darkened; inner style, id, with
outer spine large, slightly curved, with a small acute point at base; setae of rostral lobe
long. Phallosome, p, distinctive, with two unequal black apophyses; aedeagus, a, distinctive,
with a flattened rounded lobe near base and a larger outer lobe that bears a small darkened
point or short spine at outer end, distal end of aedeagus bent at a strong angle, as shown.

Holotype. $, Mughan, Iran, June 20, 1956 (Schmid). Allotopotype. ?, pinned with
type. Paratopotype. $ , pinned with types.

The present fly is similar to Gonomyia ( Gonomyia ) sibyna Alexander, of Sikkim and
Assam, differing most evidently in hypopygial structure, especially the inner dististyle and
phallosome.

Vol. LXXXIII, March, 1975

DISTRIBUTIONAL RECORDS

Gonomyia ( Idiocera ) similior Alexander

Gonomyia ( Idiocera ) similior Alexander; Ann. Mag. Nat. Hist. (12)9: 50-51; 1956.

Eastern Europe; southwestern Asia (Afghanistan, type).

Iran: Bagerabad, June 10, 1956; Kiakola, May 22, 1956; Persepolis, May 2, 1956;

Quattekas, 1800 meters, September 19, 1955; Tegan, July 5, 1956 (all Schmid).

Gonomyia ( Gonomyia ) abbreviata Loew
Gonomyia abbreviata Loew; Beschr. Europ. Diptera, 3: 58; 1873.

Gonomyia abbreviata de Meijere; Tijd. v. Ent., 63: 84, fig. 84 (venation, $ hyp); 1920.

Gonomyia ( Lipophleps ) abbreviata Edwards, Trans. Soc. Brit. Ent., 5: 112, pi. 5,

fig. 16 (wing) ; 1938.

Gonomyia abbreviata Lackschewitz ; Ann. naturhist. Mus. Wien; 50: 60, fig. 9, p. 57,

wing ; 1940.

Europe.

Iran: Ardehjan, September 9 and 11, 1956 (Schmid).

New York Entomological Society

Notes on the Life Cycle and Natural History of Butterflies
of El Salvador. VI A. — Diaethria astala Guerin.

( Nymplialidae-Callicorinae )

Alberto Muyshondt

101 Avenida Norte #322. San Salvador. El Salvador.

Received for Publication May 6, 1974.

Abstract: The results of observations carried on for a period of five years on one species of
the Callicorinae, Diaethria astala Guerin, are presented. An account is given of the external
morphological characteristics of the early stages, of the time elapsed in the metamorphosis,
of the progressive sizes of each stage, and of the foodplants in El Salvador. The evident
similarities between the early stages of this species and of Catagramma titania Salvin,
and C. pitheas Latreille, on one hand, and the early stages of species belonging to the
Catonephelinae are pointed out, suggesting a close phylogenetic relationship between
the three groups. The probability of the species having developed impalatability against
predators is deduced a priori from the noxious properties of the foodplants exploited
by the larvae, and a posteriori from the brilliant coloration exhibited by the adults.

INTRODUCTION

Through several series of articles my sons and I intend to divulge the results
of our observations on the early stages and adults of butterflies inhabiting the
neighborhood of San Salvador, capital city of El Salvador. The present one is
the sixth of the second series which was dedicated up to now to the Cato-
nephilinae. This one deals with a species of Callicorinae, in order to evidence
the close relation between the two groups which are widely accepted as Nym-
phalidae.

Even though two centuries ago Denis and Schiffermiiller (1775) were con-
scious of the importance of the characteristics of the larvae as well as these of the
butterflies when working up a system of the Lepidoptera, “Ein Auge auf den
Schmetterling, das andere auf die Raupe,” (one eye on the butterfly, the
other on the larva), and modern authors still accept the validity of that concept,
going even further: “any classification must take into account as many as

possible of the external and internal structures not only of the adults but of
the early stages” (Ford, 1945), it is evident that the early stages of many
Neotropical Rhopalocera are still little known. As a result, some groupings have
been arbitrarily made. We hope that our presentations will help, within their
limitations, to fill the existing gap of information.

Acknowledgments: Again we express our gratitude to Dr. Alexander B. Klots of
the American Museum of Natural History, New York, as without his valuable help and
advice this publication would not have been possible. We also thank Dr. Frederick D.
Rindge, of the same institution, who confirmed the tentative identification of the species.

New York Entomological Society, XXXIII: 10-18. March, 1975.

Vol. LXXXIII, March, 1975

We have reared Diaethria as tala Guerin a number of times since early 1968
from eggs collected immediately after oviposition. Photos have been taken of
them, of the subsequent larval instars and of the pupae. Record has been kept
of the time spent on each individual stage and their respective measurements.
Specimens of the early stages were perserved in alcohol and sent to the American
Museum of Natural History, New York, where they are available to students
of the group. In every instance we have reared this species, the eggs and larvae
were kept in transparent plastic bags which were cleaned daily and maintained
at all times under ambient light and temperature conditions. The identification
of the butterfly was tentatively made by Miguel Serrano, and confirmed
later by Dr. Frederick D. Rindge.

LIFE CYCLE STAGES

Egg. Truncated cone shaped. Green with 14 lighter green ribs running from base to
micropylar area. Ribs alternately reach the micropyle and vanish at the edge of the dome.
About .75 mm. long. Hatches in 4 days.

First instar larva. Head brown, roundish, naked. Body yellowish-green, cylindrical,
naked, with annulets between segments. 1.5 mm long when recently hatched, 3 mm
before moulting in 4 days.

Second instar larva. Head brown with short, stubby, knobbed horns on each epicranial
apex. Body yellowish-green profusely tuberculated by minute excrescences of lighter
color. A lateral spine, deflected caudad, at each side of the 9th abdominal segment.
5.5 mm long before moulting in 4 days.

Third instar larva. Head brown, cordiform, with two long (nearly % of body length)
slender, brown horns ornamented with three rosettes of accessory spines, bearing sparse
thin setae. Body light green, finely tuberculate, with minute subdorsal, black, tri-furcated
spines, from second abdominal segment to 8th abdominal segment. Lateral spines on
9th abdominal segment more developed and yellowish. About 10 mm long (not counting
the horns). Moults in 6 to 8 days.

Fourth instar larva. Head and body as in third stadium, but horns % of body length,
and subdorsal spines on yellow pinnacula. Grows to 15 mm in 5-7 days.

Fifth instar larva. Head reddish at base of horns and lateral margins, whitish in front.
Horn shafts alternately reddish brown and dirty white. Accessory spines on horns bearing
sparse dark setae at tips. Body light green with a scattering of tiny white tubercles and
three rows of yellow tubercules, one along meson from 1st to 7th abdominal segments,
and two subdorsally from 2nd thoracic segment to 8th abdominal segment. The subdorsal
tubercules bear each a small but prominent black spine and two smaller ones. The median
tubercules each bear one small black spine. The lateral furcated black spines on 9th ab-
dominal segment are very prominent now on yellow scoli. Grows to 25 or 27 mm in
6-7 days.

Prepupa. No changes in appearance, but shorter. One day.

Pupa. Abdomen thickening from brown flat cremaster to wing cases. Indentation separating
abdomen from humped and keeled thorax dorsally, terminating in bifid head. Color

New York Entomological Society

Figs. 1-8. Diaethria astala Guerin. 1. Egg. 2. First instar larva with frass pellets
stuck to its body. 3. Second instar larva on its perch, ready to moult. 4. Third instar
larva. Notice body parallel to leaf. S. Fifth instar larva with anterior part of the body
raised. 6. Pupa. Lateral view. 7. Pupa. Dorsal view. 8. Pupa. Ventral view.

green with brown lining laterally from cremaster, wing cases and head. Thin brown vein-
like markings ventrally on wingcases, and two dark spots about midway along the antennae.
Spiracula small and inconspicuous green. The whole dorsal surface covered by very short,
golden hair visible under a 10 X magnification only. Wingcases turning dark before adult
emergence. Duration 5-6 days.

Vol. LXXXIII, March, 1975

Adults. There is sexual dimorphism in this species, even if not so drastic as in some
Catonephelinae. The shape of the wings is the same in both sexes: front wing with

a slightly convex costal margin, rounded apex, almost straight outer margin, rounded
tornus and slightly concave inner margin. Hindwing almost round, with a humeral lobe
and a fold at inner margin. Dorsal ground color of both fore- and hindwings is in both
sexes velvety black, which in males gives a deep blue reflexion under direct sunlight.
In front wings of males there is an iridescent blue slanting bar arising from inner margin,
near tornus, towards mid-costal margin, disappearing around discal cell, and a white
spot subapically. On females the slanting bar is narrower, iridescent greenish-blue and
almost reaches the costal margin; there are two subapical white spots instead of one.
Ventrally both sexes have the same striking combination of colors: fore wing with a small
gray basal area, followed by a triangular red zone lined by dark gray band along inner
margin, then a thick, dull-black band from mid-costal margin to tornus, finally an apical
white triangle with two thin black lines parallel to outer margin. Hind wing mostly white
with the characteristic gray “89” surrounded by a thin black line ; another thin, red
line parallel to the black line midway between it and outer margin. Body black dorsally,
white ventrally. Dark brown eyes and black, white-ringed antennae. Wing span averaging
44 mm in males, 50 mm in females. Total developmental time varies from 35 to 41 days.

NATURAL HISTORY

Oviposition in this species occurs usually between 10 and 15 hours. The
females fly to the foodplant rather hesitantly. Once the foodplant has been
located, they fly around a few times until a suitable place is chosen and alight
on a mature leaf or a tender terminal. A single egg is deposited per location,
either on the edge of a mature leaf or on the tendrils or terminal bud of a young
shoot. Once the egg is deposited the females resume the circling flight and
the process is repeated several times before moving away. We have seen eggs
being laid from almost ground level on small rampant plants (which is the most
usual method), to about 16 m from the ground on the young terminals of
plants clinging to neighboring trees. This is done on vines belonging to the
Sapindaceae; the genera Serjania and Cardiospermum seem to be preferred,
even though we have collected eggs and larvae of D. as tala on Paullinia spp.
eventually. The eggs, due to their small size and green color, are rather hard to
find. The tiny hatching larvae eat an exit hole through a wall of the eggshell,
and at times eat afterwards a portion of the upper part of it, but always leaving
an identifiable remnant. The small larvae move later to the edge of the leaf
and feed on it, usually around a vein, which is prolonged with frass pellets
affixed with silk, and this is used by the larvae as a resting place while not
feeding. It is common to find small larvae with one or several pellets stuck to
their own bodies. This might function as camouflage or to have material at
hand to lengthen the perch as needed. The small larvae usually hold to the
perch with just the prolegs, raising the anterior part of the body, the head
pointing distally. Second instar larvae behave similarly. The larvae during these
stadia leave the perches only to feed, which is done early in the morning or

New York Entomological Society

Figs. 9-12. Diaethria astala Guerin. 9. Male dorsal side. Measures in cm. 10. Male
ventral side. 11. Female dorsal side. 12. Female ventral side.

late in the afternoon, and once this is done they crawl back to the resting places.
While walking, the larvae weave a foothold of silk, moving the head from side
to side. From third instar on, the larvae abandon the perch and wander about
the plant, usually on the upper surface of the leaves. Most of the time they
stay motionless adopting two peculiar attitudes, one with the whole body in
contact with the leaf surface, the head bent forward so that the horns are
parallel to the leaf surface; and a second with only the abdominal segments
parallel to the leaf surface, the thoracic segments raised, but as before the
head bent forward in a similar manner. When the observer blows on a larva
resting as described, it reacts by a continuous twitching motion of the thoracic
legs. If prodded with a sharp object, the larva strikes violently with its horns.
When by accident more than one larva move to the same leaf, a fight is certain
to occur as one larva touches the other. As a rule one or both contendants
will be punctured by the sharp spines of the horns, or their horns will lock in
such a way that both larvae will not be able to feed and therefore will starve.
One time we found a fifth instar larva moving about the plant with a dead

Vol. LXXXIII, March, 1975

third instar larva looped around its thoracic segments, the horns of both larvae
being firmly interlinked. The younger and weaker larva had succumbed to
starvation while still fighting to disentangle its horns. The bigger larva died
few days later as a result of an infection caused by the decaying body of the
smaller one, although due to its greater strength it could feed normally.

When ready to pupate, the larvae look for a convenient place on the same
vine or on a neighboring shrub or small tree and weave a silken pad usually
on the upper surface of a leaf, less commonly on the lower surface, clean the
digestive tract and hold to the silk with the anal prolegs. The larvae very
seldom hang to pupate. The pupae in consequence, may be on either surface
of a leaf, not hanging, but closely appressed to it. The pupae when disturbed
can produce a faint creaking sound by wiggling sidewise or moving accordion-
like. Shortly before the adult emergence the green pupae turn dark gray and
the dorsal colors of the wings are visible through the shell.

The emerging adult rapidly abandons the pupa shell and hangs from it until
the wings are rigid enough to fly, meanwhile expelling a rusty meconium. We
have never observed the adults while in copula, nor have we seen them feeding
on flowers nor on fermenting fruits, even though we suspect they do feed on the
latter; but very often we have collected adults on vertebrate excrements or
at mud puddles alongside creeks. When approached the butterflies fly swiftly
in circles, their bluish flash being very conspicuous.

The foodplants of Diaethria astala larvae we have found up to the present
all belong to the Sapindaceae, genera Paullinia ( P . pinnata), Serjania (several
species) and Cardiospermum (C. halicacabum) . Many plants belonging to the
genera Paullinia and Serjania are reported by various authors (Standley,
1924; Beille, 1909; Baillon, 1874) to contain poisonous or narcotic properties.
Cardiospermum halicacabum , according to Beille (1909), is rich in saponine.
All these plants are widely distributed in El Salvador. We have found them
mostly between 500 and 1500 m along ravines and creeks which harbor very
disturbed second growth plant communities in this densely populated country,
whose land is almost completely under intensive cultivation. It is within this
range (500-1200 m) that Diaethria astala is found. The adults favor the
neighborhood of coffee plantations, ravines and creeks with heavy vegetation.

When rearing this species we have lost many individuals due to parasitism,
usually by Tachinidae, but also by Hymenoptera. Others died when fed on
slightly decaying leaves of the foodplants, which seem to become more toxic
even for them.

DISCUSSION

Muller (1886) gives a description of the early stages of Callicore meridionalis
Bates, using Myscelia orsis Drury as comparison model; and of Catagramma

New York Entomological Society

pygas Godart comparing it with C. meridionalis which is cited by J. Rober
(1915). Muller in his work reports the foodplant for C. pygas to be Allophylus
petiolatus Radlkofer, (Sapindaceae), and amazingly Trema micrantha Dell,
(Ulmaceae he places under Urticaceae), for Callicore. This is repeated by
Rober (1915), by Bates (1923) for Diaethria clymena (Cramer) and quoted
by Kimball (1965). Trema micrantha , a small tree, is found in this country
in the same habitats in which we find the Sapindaceae vines used by Diaethria
as tala (as well as other species: D. Salvador ensis Franz, Catagramma titania
Salvin, C. pitheas Latreille) larvae as foodplants, yet not a single time have
we found, or have been able to make the larvae accept Trema micrantha as food.
Was a Sapindaceae tree misidentified? In any case, the species we have reared,
feed locally and exclusively on a variety of plants of the family Sapindaceae.
In our knowledge, this is the first time a complete description of the life cycle
of Diaethria astala , illustrated with photographs, is presented.

Ebert (1969) lists under Callicorinae: C. sorana Godart, Diaethria candrena
Godart, D. clymena , D. eluina Hewitson and Paulo gramma pyracmon Godart
as species existent in the Brazilian zone of Poqos de Caldas, Minas Gerais.
We do not find any Catagramma listed in that group, or in the closely related
Catonephelinae. After having reared Catagramma titania from the egg and

C. pitheas partially, we dare to suggest Callicorinae and Catagramminae are
at least as closely related as Catonephelinae and Callicorinae, (if they should
not be all aggregated into a single family, probably Catagrammidae, as Guenee
and Burmeister did, separating the groups into subfamilies or tribes), as there
is a great similarity between the eggs, larvae and pupae of Catagramma titania
(and what we have seen of C. pitheas ), and those of Diaethria astala and

D. Salvador ensis, (the latter using the same foodplants as D. astala but at
higher altitudes: 1200 m and up). One time we observed a C. titania ovipositing
on the young terminals of a Serjania vine high up in a supporting tree (16-20
m). The terminals were brought down and placed in a transparent plastic
bag. Some greenish eggs were found on the younger parts of the terminals,
along with some yellow ones, and under superficial examination were found
similar, the difference of coloration being attributed to different ages, and
all were reared to adult. To our surprise two kinds of larvae were noticed
when at third stadium: some typical Diaethria and others without the tiny
subdorsal spines, but with a thick scolus and with spines on meson of 8th
abdominal segment! The head and its horns, the body shape and color, and
the behavior of these larvae were almost the same as those of Diaethria. The
pupae formed later were all also very similar. Some of these produced adults
of C. titania others of D. astala. It is accepted that the egg, larval and pupal
characteristics are the ones which resist to a greater degree the changes induced
by divergent selection, and therefore are of extreme importance to determine

Vol. LXXXIII, March, 1975

phylogenetic relationships between species, genera and families. In this case
they seem to indicate the close relationship of Catagramma-Diaethria. As a
result of the comparison of the external characteristics of the eggs, larvae and
pupae of the two genera, reinforced by the similar behavior and the same
foodplant association, we conclude that the two groups also evidence a close
affinity with Catonephelinae. We refer to the descriptions of the early stages
and behavior of Catonephele numilia esite Felder (Muyshondt, 1973), Epiphile
adrasta adrasta Hewitson (Muyshondt, 1973a), Temenis laothoe liberia
Fabricius (Muyshondt, 1973b) Pseudonica jlavilla canthara Doubleday (Muy-
shondt, 1973c) and Pyrrhogyra hypsenor Godman & Salvin (Muyshondt,
1974) to support our contention, without having to be repetitious.

In our presentation of the Catonephelinae mentioned above we discussed
the probability that at least some of them (E. adrasta , T. laothoe , Pseudonica
jlavilla and Pyrrhogyra hypsenor ), which also feed on Sapindaceae, have
developed a more or less strong impalatability to predators, basing our asser-
tion not only on the poisonous properties of the foodplants, but on the gradually
showier colors and slower flights these species show, following the sequence as
above. Being that Diaethria astala larvae feed on the same plants the others
do, that they behave similarly during the early stages and most of all that the
adults have a brilliant coloration, we also suggest the probability of this species
being protected against predation for the same reasons. In addition to this
defense mechanism based on chemical properties, the adults of D. astala exploit
the contrasting dorsal and ventral bright colors to produce a bewildering effect
on attackers of “flash-and-substitute,” as the fast moving blue streak suddenly
disappears when the butterflies alight with their wings folded, and are replaced
by an altogether different and immobile object, however bright and gaudy their
coloration. In no case could these colors be considered cryptic or camouflaging,
but on the contrary they seem to advertise the noxious properties of the butter-
flies to their potential enemies.

Diaethria astala is another species which appears to support our hypothesis
that parasitizing Diptera and Hymenoptera will prefer hosts protected from
predation as a means to guarantee the survival of their progeny, (Muyshondt
1973b, c and 1974) because this species also is decimated mostly by tachinid
flies. We have found pupa shells in the fields also clearly showing exit holes
similar to the ones caused by Spilochalcis sp. on pupae of Pyrrhogyra hypsenor
in our insectarium.

Literature Cited

Baillon, H. 1874. Histoire des plantes. Hachette et Cie. Paris, 5: 389.

Bates, M. 1923. Notes on Florida Lepidoptera. Fla. Ent., 7: 42-43.

Beille, L. 1909. Precis de botanique pharmaceutique. A. Malone. Paris. 2: 645.
Denis and Schiffermuller. 1775. “Syst. Verz.” Schmett. Wien.

New York Entomological Society

Ebert, H. 1969. On the frequency of butterflies in Eastern Brazil with a list of the
butterfly fauna of Pogos de Caldas, Minas Gerais. Jour. Lep. Soc. 23, Sup. 3.
Ford, E. B. 1945. “Butterflies.” Collins. London.

Kimball, C. P. 1965. “Lepidoptera of Florida.” State of Florida Dept, of Agriculture.
Gainesville, Fla. p. 43.

Muller, W. 1886. Siidamerikanische Nymphalidenraupen. Zoologische Jahrbuch, 461-475.
Muyshondt, A. 1973. Notes on the life cycle and natural history of butterflies of El
Salvador. I A. — Catonephele numilia esite (Nymphalidae-Catonephelinae) Jour.
New York Entomol. Soc., 81: 164-174.

. 1973a. Notes on the life cycle and natural history of butterflies of El Salvador.

A. — Epiphile adrasta adrasta. (Nymphalidae-Catonephelinae) Jour. New York
Entomol. Soc., 81: 214-223.

. 1973b. Notes on the life cycle and natural history of butterflies of El Salvador.

III A. Temenis laothoe liberia (Nymphalidae-Catonephelinae) Jour. New York

Entomol. Soc., 81: 224-233.

. 1973c. Notes on the life cycle and natural history of butterflies of El Salvador.

IV A. — Pseudonica jlavilla canthar. (Nymphalidae-Catonephelinae) Jour. New

York Entomol. Soc., 81: 234-242.

. 1974. Notes on the life cycle and natural history of butterflies of El Salvador.

V A. — Pyrrhogyra hypsenor (Nymphalidae-Catonephelinae) . Jour. New York
Entomol. Soc. in press.

Rober, J. 1915. In Seitz’s Macrolepidoptera of the World. Vol. 5. Stuttgart.
Standley, P. C. 1924. Trees and shrubs of Mexico. Contrib. from the U.S. Nat. Herb.,
23, Part 3, 701-3.

Vol. LXXXIII, March, 1975

Seasonal Occurrence of Night-Flying Insects on
Barro Colorado Island, Panama Canal Zone

Robert E. Ricklefs

Department of Biology, University of Pennsylvania, Philadelphia, Pa. 19174
Received for Publication June 3, 1974

This report summarizes the seasonal occurrence of night-flying insects
attracted to ultraviolet-emitting fluorescent lamps (“black lights”) on Barro
Colorado Island. Observations were made between November 1967 and August
1968, during March 1970, and during June and July, 1971. Captures of insects
in a malaise trap, operated between November 1967 and June 1968 are also
tabulated for comparison.

Many ecologists who lack long-term experience in the tropics hold the common
misconception that the tropics are relatively aseasonal, but numerous reports
have demonstrated strong seasonal cycles in the occurrence of organisms or
aspects of their behavior. For examples, readers are referred to Skutch (1950),
Ricklefs (1966), Snow and Snow (1964), and Miller (1963) for reproductive
cycles in neotropical birds, to Janzen (1967) and Smythe (1970) for seasonal
patterns of flowering and fruiting in plants, to Wilson (1971) for the seasonal
occurrence of reproduction in bats, and to Fairchild (1942), Galindo et al.
(1956), Pipkin (1965), and Owen (1969), for seasonal cycles of abundance
in particular groups of insects. For the most part, seasonal cycles in the tropics
are closely tied to abrupt changes of rainfall associated with the onset of
marked wet and dry seasons, but even where rainfall is relatively abundant
throughout the year, biological seasonality is still a predominant feature.

Although evidence for seasonality in tropical faunas and floras is accumulating
rapidly, relatively little is known about year-to-year variation in population
sizes and reproductive activity. Collections of arboreal mosquitoes over a six
year period, reported on by Galindo et al. (1956), demonstrated considerable
year to year variation in individual species. Observations reported here further
substantiate this finding.

METHODS

Two fluorescent black lights were positioned over screened windows in the
laboratory clearing on Barro Colorado Island. The lights faced a ravine,
covered with tall second growth vegetation near its top and with relatively
mature forest farther down the slope. Thus the lights illuminated both canopy
and understory vegetation. The vegetation remained essentially unchanged
throughout the study.

New York Entomological Society, XXXIII: 19-32. March, 1975.

New York Entomological Society

Lights were turned on before dark and insects in several broad taxonomic
groups were counted on the four window screens directly under the lights
between 5 and 6 a.m. The numbers of individuals of several hundred species of
moths, identified with photographic keys made during November 1967, were also
recorded. Counts were made on 89 nights during the 10 months between
November 1967 and August 1968, an average of almost 9 nights per month.
The counts did not follow a regular schedule, and they varied between 4 and
12 per month in number.

A malaise trap with a cross-sectional area of 4 m2 was also employed for 39
night periods and 29 day periods between November and June. The trap was
located at ground level along a 5 m wide cleared path through second growth
vegetation attaining about 5-10 m in height. The collecting bottle on the trap
was usually emptied at dawn and dusk to separate day and night catches, but
it was occasionally emptied every two hours during the day to obtain diurnal
variation in flying insects. The wet weight of malaise trap collections was usually
the only measurement of abundance recorded, but individuals of several orders
were occasionally counted.

RESULTS

Major groups of insects. Monthly rainfall records for Barro Colorado Island,
averaged for both 44 years and for the years during which this study occurred,
are presented in Table 1. The climate is characterized by a rather severe dry
season that usually begins abruptly in late December and ends somewhat more
gradually in April. The timing and severity of the dry season vary considerably
from year to year. Between 1926 and 1967, the rainfall during the period
January through March varied by a factor of 27, between 0.6 and 16.3 in.
(1.5 and 41.4 cm).

The seasonal occurrence of several conspicuous groups of insects attracted to
the lights on Barro Colorado Island during the period November 1967 through
August 1968 are presented in Table 2. Moths are divided into two size groups
at a body length of 1 cm. Their seasonal occurrence will be discussed in detail
below, although it is clear from Table 2 that the abundance of large species
declined during the dry season months, and that the abundance of small species
was least during the early part of the rainy season (April-June) . Standard
errors of the mean for the moth samples vary between 10 and 20% of the mean.

Patterns of abundance for other groups appeared to vary greatly. Katydids
(Orthoptera: Tettigoniidae), beetles (Coleoptera), and both pentatomid and
reduviid bugs (Hemiptera) occurred in fairly regular numbers throughout the
year although katydids appeared to be more abundant during Feb- April,
beetles exhibited a peak of abundance in May 1968, and reduviid bugs were
relatively scarce in November 1967 and January 1968. Mantids (Orthoptera:
Mantidae) were also scarce during November and December. Few bees, wasps,

Vol. LXXXIII, March, 1975

New York Entomological Society

Table 2. Seasonal occurrence of certain groups of insects attracted to black lights on Barro
Colorado Island, expressed as number of individuals per 10 nights of observation.

MONTH

1967

1968

NOV

DEC

JAN

FEB

MAR

APR

MAY

JUN

JUL

AUG

Number of nights
Taxonomic group

Moths

Large

314

512

168

217

157

310

348

258

446

394

Small

404

655

413

912

616

243

219

175

321

310

Katydids

Mantids

Beetles

Bees, ants, and

wasps

251

Pentatomid bugs

Reduviid bugs

Cicadas

Owl- flies

and flying ants (Hymenoptera) were attracted to the lights during the dry
season months ( January-March) although tree flowering reaches a peak during
this period. A marked peak in the abundance of Hymenoptera at the lights
occurred during May 1968. Two smaller taxonomic groups, the cicadas
(Homoptera: Cicadidae) and the owl-flies (Neuroptera: Ascalaphidae) were
completely absent during large portions of the sample period and were most
abundant during the early part of the rainy season.

Moths (Lepidoptera) were attracted to the lights in far greater numbers
than any other group. Seasonal trends in their occurrence are shown in Figure 1.
Large moths were least numerous during the dry months, January through
March, and their numbers increased abruptly with the onset of the rainy period.
As a whole, small moths exhibited no decline in numbers during the dry season.
In fact, they appeared to attain peak abundance at that time. This peak con-
sisted mostly of individuals of one species that was present at no other time,
however; when this species was subtracted from the total, small moth abundance
can be seen to decline through the dry season, reaching low levels between
March and June (Figure 1).

The numbers of moths attracted to the lights varied greatly from night to
night. Coefficients of variation, calculated for each month’s counts and presented
in Figure 2, demonstrated that the magnitude of short-term variation in small
moths paralleled that in large moths and tended to decline slightly between
November and August.

Vol. LXXXIII, March, 1975

N D J F M A M J J A
MONTHS

Fig. 1. Monthly averages of the number of individuals of moths attracted to the black
lights on Barro Colorado Island between November 1967 and August 1968. Bars represent
standard deviations. Solid bars and lines represent large species (body length greater than
1 cm) ; open bars and dashed lines represent small species. Two sets of figures are presented
for small moths during February and March ; one set includes, and the other does not
include, a particularly abundant species present only during those months.

Daily records for the occurrence of moths at the lights indicate regular short-
term cycles, particularly for large moths during the dry season (Figure 3). It is
well known that the flight activity of moths varies more or less inversely with
the brightness of the moon (Willliams 1936, Brown and Taylor, 1971). The
periods of the abundance cycles do appear to be roughly four weeks, but peaks

New York Entomological Society

Fig. 2. Monthly coefficients of variation (standard deviation divided by the mean) of
counts of moths attracted to black lights on Barrro Colorado Island, November 1967 to
August 1968. Large and small species are distinguished. Data for 1970 are also indicated.

and troughs are not particularly well coordinated with new and full phases of the
moon. Peaks timed according to this same periodicity seem to occur during
November-December and April-June periods, but are out of phase and less
well marked during July and August.

Individual species of moths. Records were kept of the numbers of individuals of
several hundred species that were attracted to the lights each night. None of
the species were identified. Most of these species were too uncommon to discern
the presence or absence of marked seasonal trends, and many species were noted
only once. The monthly averages for several of the more common species, shown
in Figure 4, demonstrate a variety of seasonal patterns, ranging from relatively
uniform distribution throughout the study period to the occurrence of marked
peaks in abundance falling at different times of the year. All the species repre-
sented in Figure 4 appeared at least once during November, when a photographic
numbered key to the moth species was made. Other species clearly showed
narrow peaks of abundance during the dry season or early portion of the wet
season. For example, in Figure 5, the nightly abundance of one very abundant

Vol. LXXXIII, March, 1975

Fig. 3. Nightly occurrence of moths at the black lights on Barro Colorado Island, November
1967 through August 1968. Solid line in January through March suggests fluctuations in large
bodied species. Arrows represent extrapolation of peaks of abundance at 4 week intervals
throughout the sampling period.

New York Entomological Society

NDJ FMAMJJA
MONTHS

Fig. 4. Monthly average number of individuals of 13 selected species attracted to black
lights on Barro Colorado Island, November 1967 to August 1968. Numbers refer to the
photographic key used to distinguish the species.

small moth (unnumbered), present only during February and March, is com-
pared to the more uniform seasonal distributions of the species of small moths
numbered 17, 18, and 20. Fairchild (1942) also found great variety in the
seasonal distributions of species of tabanid flies in Panama. By contrast, all the

Vol. LXXXIII, March, 1975

Fig. 5. Nightly occurrence of four species of small moths at the black lights on Barro
Colorado Island, November 1967 to June 1968. Dots represent absence of a species on a
particular night. Species numbers 17, 18 and 20 correspond to those species in Figure 4.

species of arboreal mosquitoes studied by Galindo et al. (1956) in the same region
showed similar seasonal patterns of abundance, being almost completely absent
during the dry season (January- April) and most abundant during the early part
of the wet season ( May- August ) . The seasonal pattern of abundance in these
species is dictated by the fact that arboreal mosquitoes rely on the presence of
standing water in tree holes and bromeliads for reproduction.

Malaise trap samples. Wet weights of insects caught during the night period
were relatively high during November through January and about half as great
during February through June (Figure 6). Daytime catches did not exhibit any
marked seasonal pattern in total wet weight, however.

Most of the malaise trap sample collected during the night consisted of tiny
diptera, which were not represented at the lights. So we should not be concerned
over the lack of correspondence between the malaise trap samples and black
light counts. Most of the daytime samples consisted of relatively large species
of diptera and hymenoptera which reached peak abundance during midday
(Tables 3 and 4).

New York Entomological Society

Fig. 6. Monthly averages of the wet weight of insects caught in a malaise trap on
Barro Colorado Island. Daytime catches (0600-1800 hrs) and night-time catches (1800-
0600 hrs) are distinguished. Solid and open circles represent single day or night samples.
Bars represent standard deviations.

Y ear-to-year variation. Counts of insects attracted to the black lights during
March 1970 and June and July 1971 are compared to samples counted during
1968 in Table 5. Differences between years are conspicuous for several groups.
Comparing the month of March in 1968 and 1970, we note that large moths,
mantids, beetles, and reduviid bugs were more abundant in 1970; no group was
less abundant. The greatly increased number of beetles during 1970 was due to
one species that had not been abundant at any time during the 1967-1968
sampling period.

Although the number of small lepidoptera counted during March 1970 was

Vol. LXXXIII, March, 1975

New York Entomological Society

Table 4. Diurnal variation in the number of insects with body lengths exceeding 2 mm

caught in the malaise trap.

Hour of Day

Date

Group

6-8

8-10

10-12

12-14

14-16

16-18

Total

Feb. 23

LEP

DIP

HYM

COL

TOTAL

Feb. 29

LEP

DIP

HYM

COL

TOTAL

Apr. 30

LEP

DIP

HYM

COL

TOTAL

149

May 22

LEP

DIP

HYM

COL

TOTAL

May 24

LEP

DIP

HYM

COL

TOTAL

May 27

LEP

DIP

HYM

COL

TOTAL

Entire

LEP

Period

DIP

304

HYM

170

COL

TOTAL

130

141

545

similar to the number observed two years earlier, the particular species that
comprised more than two-thirds of the total sample in 1968 (see Figure 5),
accounted for less than one-third of the sample in 1970.

Differences in June and July samples between 1968 and 1971 were even more
striking. Considering only June, moths were 3 to 4 times as abundant in 1971 as
in 1968; numbers of katydids, hymenoptera were greater by factors of about 2,
and numbers of cicadas and beetles, by factors of 8 and 15, respectively. Only

Vol. LXXXIII, March, 1975

Table 5. Comparisons of insects attracted to black lights on Barro Colorado Island during

different years.

Month

and Year

March

June

July

1968

1970

1968

1971

1968

1971

Number of nights

Moths

large

157

421

258

822

446

761

small

616

654

175

730

321

654

Katydids

Mantids

Beetles

104

224

Bees, wasps and ants

Pentatomid bugs

Reduviid bugs

Cicadas

107

Owl-flies

Note: All figures are number of individuals per 10 nights.

reduviid bugs and mantids were less abundant. Differences between July 1968
and July 1971 were of a similar nature, but less marked in most groups.

It is tempting to relate the greater abundance of insects in the 1970 and 1971
samples, compared to 1968, to the unusually heavy rainfall during the months of
January 1970 (11.8 in. compared to 2.2 in. average) and May 1971 (22.6 in.
compared to 10.8). But since there are too few samples to treat the relationship
between abundance and rainfall statistically, and since so little is known about
the responses of populations to variation in rainfall in the tropics, it would be
unwise to pursue this apparent correlation here.

In summary, the numbers of insects attracted to black lights at the edge of a
lowland seasonally wet tropical forest exhibited marked fluctuation during the
course of one 10 month period. Different insect groups had different peak and
low periods of abundance, but the most conspicuous component of the samples,
the moths, were least abundant during the dry season months. In samples taken
at the same locality several years later, most groups exhibited strikingly greater
abundances although the character of the vegetation had not changed. It is
tempting to relate these increases to months of abnormally high rainfall just
preceding the samples, but regardless of their cause, year-to-year variations in
populations do occur in the tropics.

Literature Cited

Brown, E. S. and L. R. Taylor. 1971. Lunar cycles in the distribution and abundance
of airborne insects in the equatorial highlands of East Africa. J. Anim. Ecol. 40:
767-779.

New York Entomological Society

Fairchlld, G. B. 1942. The seasonal distribution of some Tabanidae (Dipt.) in Panama.
Ann. Entomol. Soc. Amer. 35: 85-91.

Galindo, P., H. Trapido, S. J. Carpenter, and F. S. Blanton. 1956. The abundance
cycles of arboreal mosquitoes during six years at a sylvan yellow fever locality in
Panama. Ann. Entomol. Soc. Amer. 49: 543-547.

Janzen, D. H. 1967. Synchronization of sexual reproduction of trees within the dry
season in Central America. Evol. 21: 620-637.

Miller, A. H. 1963. Seasonal activity and ecology of the avifauna of an American equa-
torial cloud forest. Univ. Calif. Publ. Zool. 66: 1-78.

Owen, D. F. 1969. Species diversity and seasonal abundance in tropical Sphingidae
(Lepidoptera) . Proc. R. Ent. Soc. London 44: 162-168.

Pipkin, S. B. 1965. The influence of adult and larval food habits on population-size of
neotropical ground-feeding Drosophila. Amer. Midi. Nat. 74: 1-2 7.

Ricklefs, R. E. 1966. The temporal component of diversity among species of birds.
Evolution 20: 235-242.

Skutch, A. F. 1950. The nesting seasons of Central American birds in relation to climate
and food supply. Ibis 92: 185-222.

Smythe, N. 1970. Relationships between fruiting seasons and seed dispersal methods
in a neotropical forest. Amer. Nat. 104: 25-35.

Snow, D. W. and B. K. Snow. 1964. Breeding seasons and annual cycles of Trinidad
landbirds. Zoologica 49: 1-39.

Williams, C. B. 1936. The influence of moonlight on the activity of certain nocturnal
insects, particularly of the family Noctuidae, as indicated by a light trap. Phil.
Trans. Roy. Soc. London (B) 226: 357-389.

Wilson, D. E. 1971. Ecology of Myotis nigricans (Mammalia: Chiroptera) on Barro

Colorado Island, Panama Canal Zone. J. Zool. Lond. 163: 1-13.

Vol. LXXXIII, March, 1975

Differential Cold Survival of Two Sibling Species of Blow Flies,
Phoenicia sericata and Phoenicia pallescens

Noreen Ash and Bernard Greenberg
Department of Biological Sciences, University of Illinois at Chicago Circle,

Chicago 60680

Received for Publication June 3, 1974

Abstract: The overwintering capabilities of sibling calliphorid species Phaenicia sericata

(Meigen) and Phaenicia pallescens (Shannon) are compared. P. pallescens is not capable
of overwintering in the Chicago region in an unheated shelter while P. sericata can overwinter
as larvae.

The synanthropic blow flies Phaenicia sericata (Meigen) and Phaenicia
pallescens (Shannon) are sibling species similar in appearance and general habits.
Within North America they differ in geographic range with P. pallescens a south-
ern species and P. sericata in nearly every part of the United States and southern
Canada. A comparative study was undertaken to determine if both species have
the same ability to overwinter in a northern temperate region.

Flies were trapped in early spring using a modified U.S.D.A. fly trap. P.
sericata was collected in Bensenville, Illinois, a western suburb of Chicago,
and P. pallescens in Bokeelia, Florida. Females of the two species were placed
in separate cages and allowed to lay eggs on raw hamburger. The colonies were
maintained in Chicago and were routinely kept at room temperature on sugar,
skim milk solution, and water. Maggots were raised on liver, hamburger, or dead
mice.

Two cages were set up outdoors in an open shelter in Bensenville, Illinois,
during the middle of August. Populations of P. sericata and P. pallescens were
derived from the laboratory populations. For two generations the colonies were
maintained in the usual manner and allowed to reproduce on hamburger in gallon
jars half-filled with sawdust. Maggots produced in early October were placed in
culture jars in an unheated closed shelter.

The maggots were checked on December 20. The P. sericata maggots were
constricted similar to the pupariation stage described by Fraenkel and Bhaskaran
(1973). Some of them moved slightly when the jar was disturbed. About 5%
of the maggots were dead and no pupae were seen. The P. pallescens were not
constricted and moved actively when disturbed; about 10% of these larvae were
dead.

January and February are typically the coldest months of the year in this
area of Illinois with night temperatures in the unheated shelter occasionally going
below 0°F. No pupae were observed during these months. On April 22

New York Entomological Society, XXXIII: 33-35. March, 1975.

New York Entomological Society

the jars were taken into the laboratory and examined. In the P. pallescens colony
there were 100% dead maggots (ca. 800) while in the P. sericata culture there
were 46 (5.5%) live maggots, 157 (18.7%) dead maggots, and 635 (75.8%)
pupae. After 6 days, eclosion began with most of the flies emerging. By June 1,
of the 46 live maggots, half of them had formed apparently normal pupae and
half had died; none of those that pupated from this latter group emerged.

Calliphoridae may overwinter in temperate regions as larvae, pupae, or
adults. The calliphorids most commonly found as overwintering adults include
Phormia regina (Dondero and Shaw, 1971), Protophormia terraenovae (Cousin,
19 32) , P ollenia rudis (Hall, 1948), and Calliphora species (Green, 1951 ; Sukhova,
1950). Most authors agree that P. sericata usually overwinters in the larval or
post-feeding larval stage as reported by Zumpt (1965) in South Africa, Green
(1951) in England, Norris (1959) in Australia, and James (1947) and Hall
(1948) in the United States. The overwintering stage or stages of P. pallescens
are less well documented.

Both P. pallescens and P. sericata have been reported to overwinter as larvae
at least as far north as Charleston, West Virginia (Mail and Schoof, 1954) and
diapausing P. sericata larvae have also been reported at New Brunswick, New
Jersey (Hagemann and Barber, 1948). The Florida strain of P. pallescens,
however, cannot overwinter in an unheated shelter in northern Illinois. Larvae
enter quiescence as described by Andrewartha (1971) rather than the cold-hardy
dormancy of true diapause. In this condition P. pallescens survives the milder
part of the winter but not the more severe cold of January and February. Hall
(1948) reports the fly to be numerous and active near Miami, Florida, in March
but it does not reach population peaks until July in Charleston, West Virginia
(Mail and Schoof, 1954), and the middle of August in Lawrence, Kansas (Schoof
and Savage, 1955). This could result from a high mortality among overwintering
larvae in areas with severe winters and annual re-colonization by incoming
migrants from the south. The primary screwworm, Cochliomyia hominivorax
(Coquerel), another sub-tropical calliphorid, is known to have this pattern.

By contrast, P. sericata becomes numerous in late spring and early summer
(Mail and Schoof, 1954; Schoof and Savage, 1955), has a facultative diapause
(Norris, 1965), and, as indicated in this study, is able to overwinter as larvae
in severe cold. Additional evidence that P. sericata overwinters in a pre-adult
stage is based on the earliest spring adults (about mid-April in the Chicago area)
which contain pupal fat balls in the hemolymph, unfrayed wings, and a complete
set of bulbous setae on the antennal pedicel (Greenberg, 1970). Observations of
numerous adults flying in the beginning of May in Lawrence, Kansas, and
Cohoes, New York (Schoof and Savage, 1955) suggest that P. sericata is capable
of overwintering in most of its range in the United States. Analysis of early
specimens should indicate the overwintering capability of this species in the
northernmost regions of its distribution.

Vol. LXXXIII, March, 1975

Literature Cited

Andrewartha, H. G. 1971. Introduction to the study of animal populations. The Univ.
of Chicago Press.

Cousin, G. 1932. Etude experimentale de la diapause des insectes. Bull. Biol. Fr. Belg.
Suppl. 15: 1-341.

Dondero, L., and F. R. Shaw. 1971. The overwintering of some muscoidean Diptera in
the Amherst area at Massachusetts. Proc. Entomol. Soc. Wash. 73: 52-53.

Fraenkel, G., and G. Bhaskaran. 1973. Pupariation and pupation in cyclorrhaphous
flies (Diptera): terminology and interpretation. Ann. Entomol. Soc. Am. 66:

418-422.

Green, A. A. 1951. The control of blowflies infesting slaughter-houses. I. Field obser-
vations of the habits of blowflies. Ann. Appl. Biol. 38: 475-494.

Greenberg, B. 1970. Species distribution of new structures on fly antennae. Nature 28:
1338-1339.

Hagemann, L. E., and G. W. Barber. 1948. Overwintering habits of Phaenicia sericata
(Mg.). J. Econ. Entomol. 41: 510.

Hall, D. B. 1948. The blowflies of North America. Thomas Say Foundation.

James, M. T. 1947. The flies that cause myiasis in man. U.S.D.A. Misc. Publ. No. 631.

Mail, G. A., and H. F. Schoof. 1954. Overwintering habits of domestic flies at Charleston,
West Virginia. Ann. Entomol. Soc. Am. 47: 668-676.

Norris, K. R. 1959. The ecology of sheep blowflies in Australia. In: Biogeography

and ecology in Australia. Monographiae Biologicae 8: 514-544.

Norris, K. R. 1965. The bionomics of blowflies. Ann. Rev. Entomol. 10: 47-68.

Schoof, H. F., and E. P. Savage. 1955. Comparative studies of urban fly populations
in Arizona, Kansas, Michigan, New York, and West Virginia. Ann. Entomol. Soc. Am.
48: 1-12.

Sukhova, M. N. 1950. Novye dannye po ekologii i epidemiologiches komu znacheniru
sinikh miasnykh mukh Calliphora uralensis Vill. and Calliphora erythrocephala
Meig. (Diptera, Calliphoridae) . Entomol. Obozrenie 31: 90-94.

Zumpt, F. 1965. Myiasis in man and animals in the Old World. Butterworths, London.

New York Entomological Society

Parasites Reared from Larvae of the European Corn Borer,
Ostrinia nubilalis (Hbn.), in Massachusetts, 1971—73
(Lepidoptera, Pyralidae) 1,2

F. B. Peairs1 2 3 and J. H. Lilly4

Department of Entomology, University of Massachusetts,

Amherst, Massachusetts 01002

Received for Publication June 5, 1974

Abstract: Three exotic Ostrinia nubilalis parasites; Eriborus terebrans , Macrocentrus grandii,
and Sympiesis viridula were detected in Massachusetts along with two native species ;
Aplomya caesar and Lixophaga sp. Of these, M. grandii was by far the most important,
accounting for over 92 per cent of the borers parasitized.

During a 1971-73 study of natural control of the European corn borer,
Ostrinia nubilalis (Hbn.), 1498 last instar borers were examined for parasites.
These borers were collected from the 10 Massachusetts localities listed in Table 1
and held individually in shell vials. The parasites and the percentages of borers
from which they emerged are summarized in Table 1.

Of the seven exotic O. nubilalis parasites listed as established in the United
States by Baker et al. (1949), only two, Eriborus terebrans (Grav.) (Ichneu-
monidae) and Macrocentrus grandii (Goid.) (Braconidae) were reared from
these borers. A third imported parasite, Sympiesis viridula (Thoms.) (deter-
mined by B. D. Burks) (Eulophidae), hitherto unreported from Massachusetts,
was found overwintering as pupae, three in Amherst and two in West Bridge-
water. Also E. terebrans was found only in two localities and only in limited
numbers (Table 1). Conversely, M. grandii was found in all 12 collections,
with percentages of parasitization ranging from 6.3 to 60. A number of colonies
of this polyembryonic wasp failed to produce adults. The successful ones
averaged 19.1 individuals for the 98 colonies containing only males, 18.0 for
the 111 containing only females, and 29.0 for the 21 colonies containing both
sexes.

Two native tachinid parasites were also reared. Aplomya caesar (Aid.) was
present in five collections but accounted for less than one per cent of the over-all
parasitization. A species of Lixophaga was found in one collection, killing at

1Part of a thesis submitted to the Graduate School of the University of Massachusetts,
in partial fulfillment of the requirements for the M. S. degree.

2 Published with the aid of a grant from the Guy Chester Crampton Research Fund of
the University of Massachusetts.

3 Formerly teaching assistant, Department of Entomology, University of Massachusetts.
Present address: Department of Entomology, Cornell University, Ithaca, New York 14850.

4 Professor

New York Entomological Society, XXXIII: 36-37. March, 1975.

Vol. LXXXIII, March, 1975

Table 1. Parasites reared from Ostrinia nubilalis in Massachusetts, 1971-73.

Location

Date

No.

borers

Per

cent

grandii

(a)

Per

cent

tere-

brans

(b)

Per

cent

caesar

(c)

Per

cent

Lixo-

phaga

sp.

(c)

Per

cent

uniden-

tified

(d)

Total

per

cent

parasit-

ization

Barnstable Co.

VII-23-73

30.0

(E. Sandwich)
Essex Co.

VII-24-73

9.1

27.3

(Danvers)

(Ipswich)

VII-12-73

25.0

(Waltham)

VII-30-73

11.7

8.3

20.0

Franklin Co.

X-l-71

300

25.7

1.0

26.7

(S. Deerfield)
Hampden Co.

VIII-4-73

8.3

3.3

8.3

30.0

(Holyoke)
Hampshire Co.

VII-28-72

15.0

(Easthampton)
Norfolk Co.

VII-16-73

28.3

1.7

30.0

(Attleboro)
Plymouth Co.

VII-30-73

6.3

(Bridgewater)

(W. Bridgewater)

VIII-3-72

10.0

X-15-72

675

21.0

0.7

21.7

VII-20-73

60.0

Over-all

1498

22.4

0.4

0.8

0.3

0.4

24.3

(a) Macrocentrus grandii (Goid) (= gijuensis) determined by P. M. Marsh.

(b) Eriborus terebrans (Grav.) (— Horogenes punctorius ) determined by R. W. Carlson.

(d) Six dipterous puparia, possibly additional Lixophaga.

least 8.3% (18.3% if additional similar pupae which failed to emerge were
Lixophaga) .

The 4 parasites listed in Table 1 killed 24.3% of the borers. However, M.
grandii accounted for over 92% of this mortality, with an over-all parasitization
of 22.4% of all borers examined. It is obvious from these data that M. grandii
is by far the most important parasite of the corn borer in Massachusetts.

Literature Cited

Baker, W. A., W. G. Bradley, and C. A. Clark. 1949. Biological control of the European
corn borer in the United States. USDA Tech. Bull. 983: 185 pp.

New York Entomological Society

Behavioral Changes in the Army Ant Neivamyrmex nigrescens
during the Nomarlic and Statary Phases

Howard Topoff

Department of Psychology, Hunter College of The City University of New York,
and Department of Animal Behavior, The American Museum of
Natural History, New York, N. Y. 10024

Received for Publication July 1, 1974

Abstract: The responses of workers of the army ant Neivamyrmex nigrescens to illumination
and to the presence of conspecifics were compared during the nomadic and statary phases.
During the statary phase the ants were more photonegative and exhibited a stronger tendency
to cluster together than during the nomadic phase. It is hypothesized that these differences
in the ants’ orientation are caused by corresponding changes in the level of the colony
excitation during the two phases of each behavioral cycle.

Introduction

Colonies of the army ant N eivamyrmex nigrescens Cresson exhibit cycles of
alternating nomadic and statary phases. The nomadic phase, which lasts for
17-20 days, is one of high colony activity, in which large nightly raids typically
end in emigrations to new nesting sites. During this phase raiding begins early in
the evening, and a considerable portion of the adult worker population partici-
pates (Schneirla, 1958, 1963, 1971). As the raid progresses, one or more
dendritic systems of interconnecting trails arise through the repeated division of
small terminal foraging groups of ants. The outward movement of the ants
from the nest may remain at a peak for up to several hours, because the ants’
high level of excitement persists both at the raiding fronts and on the basal
column extending to the nest (Schneirla, 1971).

The nomadic phase is followed by a statary interval of 17-20 days, charac-
terized primarily by the absence of emigrations. Raiding is also less vigorous,
with fewer individuals participating. Statary raids usually consist of a single,
long basal column which ends in a small and localized terminal branching system.
The outward surge of ants at the start of raiding usually peaks after only a few
minutes. As a result, the basal column remains relatively thin throughout the
night.

The cycles of activity in N . nigrescens are regulated by stimulative relation-
ships between the colony’s developing brood and the adult worker population.
During the nomadic phase, the adult workers are aroused to a high level of
excitement by stimuli originating from the newly eclosed callow workers and
from the maturing larval brood. When the larvae pupate the excitatory stimuli
decrease and the colony lapses into the “quieter” statary phase (Schneirla, 1957,
1971).

New York Entomological Society, XXXIII: 38-48. March, 1975.

Vol. LXXXIII, March, 1975

According to Schneirla’s theory, workers of N. nigrescens are aroused to very
different levels of excitement during the two phases of each behavioral cycle.
It is possible that many aspects of each ant’s physiological and behavioral con-
dition are affected by changes in the stimulative relationships among all indi-
viduals in the colony throughout each cycle. These may include changes in the
ants’ responsiveness to a variety of physical and biotic stimuli. Accordingly,
I conducted a series of tests designed to compare the responses of adult workers
of N. nigrescens to photic stimulation and to stimuli arising from other workers
during the nomadic and statary phases. The objective was to correlate phase-
specific differences in the behavior of the ants in the laboratory tests with our
observations of colony behavior in the field.

METHODS

Tests were conducted at the Southwestern Research Station of The American
Museum of Natural History, in Portal, Arizona. The apparatus used to measure
the ants’ responses to light consisted of a cylindrical arena (30.0 cm diam X
2.5 cm high) that was divided into five equal areas by a combination of opaque
rectangular partitions and a central cylindrical cartridge (Fig. 1). The arena
was illuminated from above by a 2 2 -watt fluorescent light ring, and the light
was diffused through a disc of neutral ground glass. Two neutral density filters
were placed on the lid of the arena to reduce the intensity of light in 2 opposite
chambers of the arena (Fig. 1). The intensity of illumination in the central
cartridge and in 2 opposite chambers of the arena was 16,000 lux; the intensity
of illumination in the remaining 2 chambers was reduced to 160 lux.

The central cartridge had 4 equidistant slit-like openings at the bottom,
which gave the ants simultaneous access to the brightly illuminated and dimly
illuminated arena chambers. The cartridge also functioned as an aspirator for
collecting the ants in the field (Fig. 2). As a result, the initial collection of the
test ants was the only manipulation they received. When used as an aspirator,
a tightly fitting plastic ring was slipped over the 4 exit slits. A piece of rubber
tubing was attached to the upper end of a central vent in the lid of the cartridge,
and an “L”-shaped tube was inserted into a hole near the edge of the lid.

For each test, 40-60 adult worker ants were collected from a raiding column
near the bivouac. The cartridge was transported to the laboratory in a dark
container. In the laboratory, the cartridge was lowered into the arena. Initially,
the cartridge was supported above the floor of the arena by 4 cubes of plastic
that projected centrally from the base of each vertical arena partition, far
enough to support the outer slip ring. To start the test the cartridge was pushed
down, which caused it to slide down through the slip ring, thus simultaneously
opening the 4 exit slits.

To test the ants’ responses to the presence of each other, another series of

New York Entomological Society

__ Fluorescent light

Filter

Ground glass

.—Arena cover

Fig. 1. Apparatus used to test responses of ants to illumination. The cartridge is shown
in place in the arena. To test the responses of ants to the presence of each other, the
fluorescent light was replaced with an infrared light source.

Vol. LXXXIII, March, 1975

Fig. 2. Central cartridge used for behavioral tests. When used in the field as an aspirator,
the plastic ring is slipped over the cartridge to seal the four exit slits. The rubber tubing
and “L”-shaped plastic tube in the lid of the cartridge are removable.

experiments was conducted in an identical arena, but no visible light was used.
Instead, illumination for photography was provided by 4 150-watt flood lamps
that were sealed behind gelatin filters that passed only wavelengths greater than
720 mfi.

New York Entomological Society

Table 1.

Percentage of ants in

central cartridge and arena

quadrants after

1 minute.

Colony #

Phase day1

% ants
bright

Bright-dim test

in % ants in

dim

% ants in
cartridge

Infrared test
% ants in
cartridge

66N-2

N-3

N-7

—

N-10

N-17

S-2

—

S-8

S-9

S-10

S-14

—

S-15

—

S-16

100

S-18

100

—

66N-7

N-4

N-ll

S-2

100

—

S-3

■ —

— ■

—

S-5

100

S-7

100

S-14

100

—

S-17

100

66N-13

N-10

N-16

S-l

100

S-7

S-10

100

S-13

100

72N-3

N-3

N-7

N-13

N-14

S-6

S-7

100

S-17

100

S-20

100

1N-nomadic; S-statary

Each series of tests lasted for 2 min. To record the position of the ants
throughout each test, a photograph was taken at 5 sec intervals from beneath
the apparatus. In the “bright-dim” tests, the fluorescent light remained on
throughout the test. In the “infrared” series, the infrared light source was
electrically programmed to be on for 1.5 sec during each 5 sec interval. Because
the infrared light was not visible to the experimenter, a buzzer that was syn-
chronized with the light provided the signal to take a photograph.

RESULTS

Results of tests conducted with 4 colonies of N. nigrescens are presented in
Table 1, which shows the percentage of ants in the central cartridge and arena
quadrants of the “bright-dim” and “infrared” tests after 1 min of each 2 min

Vol. LXXXIII, March, 1975

Fig. 3. Characteristic pattern of movement in “bright-dim” tests during nomadic phase.
The ants are in the central cartridge and all 4 arena quadrants. They are well spaced and
moving rapidly.

test. For reasons that will be discussed below, the best measure of the ants’
response to illumination is the percentage located in the 2 brightly illuminated
arena quadrants. The median percentage of ants taken from nomadic colonies is
31%, as compared to 0% for statary ants.

If we consider the percentage of ants in the dimly illuminated arena quadrants,
we find that the median for nomadic ants is 44%, whereas the value for statary
ants is again 0%. At first, this may seem to contradict the results obtained from
analyzing the percentage of ants in the brightly illuminated quadrants. The
discrepancy is resolved by considering the percentage of ants remaining in the
central cartridge. During the nomadic phase, the median percentage of ants
in the cartridge is 25%. During the statary phase, by contrast, the median is
100%. Thus, the data indicate that nomadic ants tend to leave the central
cartridge and enter into either the brightly or dimly illuminated quadrants of
the arena. Statary ants tend to remain in the central cartridge throughout the
test. If they leave the cartridge they invariably enter into the dimly illuminated
quadrants.

The large difference in the number of ants remaining in the central cartridge
during the nomadic and statary phases also existed when the tests were conducted
under conditions in which the central cartridge and all 4 arena quadrants were

New York Entomological Society

Fig. 4. Characteristic pattern of movement in “infrared” tests during the nomadic phase.
As in the “bright-dim” series, the ants occupy all areas of the apparatus.

uniformly illuminated with infrared light. In this series of tests, the median
percentage of ants remaining in the cartridge after 1 min was 28% for nomadic
ants, as compared to 77% for statary ants.

In addition to the quantitative data presented in Table 1, the photographs
used to record the location of the ants throughout each test also revealed striking
qualitative differences in the behavior of the ants during the 2 phases. Before
the start of a nomadic test, the ants were typically positioned uniformly around
the edge of the central cartridge. When the test began, the ants left the cartridge
and established columns in the arena quadrants. Regardless of which quadrants
they were in, the ants ran rapidly, were well spaced, and exhibited no tendency
to cluster. This pattern of behavior was exhibited by the ants during both the
“bright-dim” and “infrared” tests (Figs. 3, 4).

The behavior of ants taken from statary colonies was quite different (Figs. 5,
6). Before the start of a test, the ants were usually clustered together in one
small section of the cartridge. The clusters varied in degree, but the ants rarely
occupied the entire cartridge. In 12 out of 18 “bright-dim” tests conducted with
statary ants, the individuals formed into tight clusters that remained in the
cartridge throughout the entire test. In 8 out of 19 statary “infrared” tests,
similar clusters were formed. It is significant that no such clusters were ever
observed during tests conducted with ants taken from nomadic colonies.

Vol. LXXXIII, March, 1975

Fig. 5. Characteristic pattern of movement in “bright-dim” tests during statary phase.
The ants are clustered tightly at the edge of the cartridge.

DISCUSSION

The results of these behavioral tests indicate that workers of N. nigrescens
respond differently to illumination during the nomadic and statary phases.
In southeastern Arizona, N . nigrescens has adapted to conditions of high tempera-
tures and low humidity by conducting most of its raiding and emigration
activities at night. This correlates with my findings that the ants are always
photonegative, although the degree of their photonegativity shifts during the
2 phases of each behavioral cycle.

During the statary phase, the ants often exhibit a marked tendency to cluster
tightly together in the central cartridge of the experimental apparatus, regardless
of the intensity of illumination. Although the specific cause of the clustering
behavior is not known, a reasonable hypothesis is that it is due to the ants’
responsiveness to chemical and tactile stimuli arising from other workers. The
results of this experiment indicate that the response of the ants to the presence
of each other is so strong during the statary phase that it can often override
their negative reaction to illumination.

It is well known that the responses of many species of insects to stimuli of
constant physical intensity are influenced by environmental factors such as
temperature and humidity, and by internal factors, including age, sex, and
physiological condition. For example, studies of waterscorpions (Holmes, 1905),

New York Entomological Society

Fig. 6. Characteristic pattern of movement in “infrared” tests during the statary phase.
As in the “bright-dim” series, the ants are clustered in the cartridge and remain there
throughout the test.

mayflies (Allee and Stein, 1918), drone flies (Dolley and White, 1951), and
mosquitoes (Chiba, 1967) indicate that decreasing temperatures result in a
shift towards photonegativity. The general consensus of these investigators
is that any environmental factor that lowers the organism’s excitability tends
to increase its negative photoreactivity.

Changes in patterns of orientation with respect to light can also be caused
by corresponding changes in stimuli that originate within the organism. Newly
hatched larvae of the hawk moth are strongly photopositive, but just prior to
pupation the mature larvae become increasingly photonegative (Beetsma et al.,
1962). These investigators also showed that injection of the hormone ecdysone
could induce the photonegative response. Similar changes in response to illumina-
tion as a function of physiological condition have been found in tabanid flies
(Shamsuddin, 1966) and milkweed bugs (Barrett and Chiang, 1967).

Orientation towards chemical stimuli can also be influenced by external
environmental and internal physiological factors. An investigation that is
particularly relevant to the present study was conducted by Goldsmid (1967) on
the blue tick, Boophilus decolor atus. Newly hatched tick larvae exhibit a strong
negative reaction towards light. At this developmental stage, however, the
larvae also aggregate together in clusters by orienting towards chemicals secreted
by other larvae. This clustering tendency overrides the individuals’ negative

Vol. L XXXIII, March, 1975

response towards light. If the cluster is mechanically broken and the larvae
scattered within their container, they invariably reaggregate in approximately
the same location. After one week, changes in physiological conditions asso-
ciated with maturation eliminate the tendency to aggregate, and at this time
the larvae also become markedly photopositive.

The results of the present study show that workers of the army ant N. nigres-
cens respond differently to illumination and to the presence of conspecifics
during the nomadic and statary phases. Based on the studies cited above, it is
reasonable to hypothesize that the changes in the ants’ responsiveness may be
caused by corresponding changes in their degree of excitation during the 2 phases.
The nomadic phase is initiated by intense stimulation imparted to the adult
worker population by the eclosing callows, and is maintained by equivalent
stimulation derived from the developing larval brood (Schneirla, 1957). It is
possible that the resulting increase in adult worker excitation and activity
causes them to be less photonegative and less responsive to chemicals secreted
by other workers. When the larval brood completes its development and pupates,
there is a sharp decline in the intensity of social stimulation in the nest. The
overall level of excitation is lower, and this causes the workers to exhibit an
increase in their photonegativity and in their sensitivity to conspecifics. In the
case of the army ants, as in the blue tick, the ants’ increased sensitivity to other
ants seems to override their increased photonegativity.

Literature Cited

Allee, W. C. and Stein, E. R. 1918. Light reactions and metabolism in may-fly nymphs
Jour. Exp. Zool., 26: 423-458.

Barrett, R. W. and Chiang, H. C. 1967. Changes of behavior pattern within the fifth
nymphal instar of the Milkweed bug, Oncopeltus jasciatus (Dallas). Amer. Mid.
Nat., 78: 359-368.

Beetsma, J. L., deRuiter, L., and deWilde, J. 1962. Possible influence of neotenine
and ecdyson on the sign of phototaxis in the eyed Hawk caterpillar Smerinthus
ocellata L. Jour. Insect Physiol., 8: 251-257.

Chiba, Y. 1967. Activity of mosquitoes, Culex pipiens Pallens and Aedes japonicus
under a step-wise decrease of light intensity. Sci. Report, Tohoku Univ. ser. IV
(Biol.), 33: 7-13.

Dolley, W. L. and White, J. D. 1951. The effect of illumination on the reversal tempera-
ture in the drone fly Eristalis tenax. Biol. Bull., 100: 84-89.

Goldsmid, J. M. 1967. Observations on the behaviour of the blue tick, Boophilus
decoloratus (Koch) (Acarina: Ixodidae). Jour. Ent. Soc. S. Africa, 29: 74-89.

Holmes, S. H. 1905. The reactions of Ranatra to light. Jour. Comp. Neurol., 15:
305-349.

Schneirla, T. C. 1957. Theoretical consideration of cyclic processes in doryline ants.
Proc. Amer. Phil. Soc., 101: 106-133.

. 1958. The behavior and biology of certain nearctic army ants. Last part of

the functional season, southeastern Arizona. Insectes Sociaux, 5: 215-255.

. 1963. The behaviour and biology of certain nearctic army ants: springtime

resurgence of cyclic functions — southeastern Arizona. Anim. Behav., 11: 583-595.

New York Entomological Society

. 1971. “Army Ants: a Study in Social Organization.” W. H. Freeman & Co.,

San Francisco, Calif.

Shamsuddin, M. 1966. Behaviour of larval tabanids (Diptera: Tabanidae) in relation

to light, moisture, and temperature. Quaestiones Entomol., 2: 271-302.

Vol. LXXXIII, March, 1975

Mites (Acarina) associated with Popilius disjunctus (Illiger)
(Coleoptera: Passalidae) in Eastern United States1

Mercedes D. Delfinado

New York State Museum and Science Service, Albany, New York 12224

AND

Edward W. Baker

Systematic Entomology Laboratory, IIBIII U.S.D.A. Agricultural Research Service,

Beltsville, Maryland 20705

Received for Publication August 9, 1974

Abstract: Sixteen species of mites are reported associated with Popilius disjunctus

(Illiger) (Coleoptera: Passalidae) in eastern United States. Two new species are described:
Macrocheles disjunctus and M. whartoni. Changes in nomenclature are as follows:
Cosmolaelaps passali Hunter and Mollin = C. trifidus (Pearse and Wharton), new synonymy
and new combination; Dendrolaelaps passalorum (Pearse and Wharton), new combination.
Diagnostic features, as well as distributional and biological information are given for most
species.

A surprisingly large and somewhat heterogenous group of mites is found in
association with various species of passalid beetles (Coleoptera: Passalidae)
(Pearse et al., 1936; Tragardh, 1946, 1950; Womersley, 1957; Schuster and
Lavoipierre, 1970; Hunter and co-workers, 1964-1969). Either adults, immature
stages, or all developmental stages of certain species are found attached to
various parts of the beetle. The relationship between the mites and beetles is
undoubtedly one of phoresy, i.e., the mite utilizing the beetle as a means of
transport from one habitat to another. The attractiveness of the beetle Popilius
disjunctus (Illiger) to the mite has been observed by Mollin and Hunter (1964)
and Hunter and Davis (1965) working with Cosmolaelaps trifidus (Pearse and
Wharton) and Euzercon latus (Banks) respectively. They concluded that these
mites react to an attractant present on the external surface of their beetle host.
Cosmolaelaps trifidus reproduces only after a period of contact with the beetle.
The feeding habits of the majority of these mites are unknown. But certain
other species of Macrochelidae will feed on acarid mites ( Caloglyphus spp.),
fly eggs and larvae, and nematodes (Axtell, 1961, 1963, 1969). Probably many
other species will also feed on the same hosts.

This study is based on material taken by M. D. Delfinado from pinned
P. disjunctus beetles in the New York State Museum and Science Service
collection through the courtesy of John Wilcox, and from beetles sent by Marcel

1 Published by permission of the Director, New York State Museum and Science Service,
Journal Series No. 164.

New York Entomological Society, XXXIII: 49-59. March, 1975.

New York Entomological Society

Reeves, University of New Hampshire. Other mite specimens examined are from
the collection of Preston E. Hunter, University of Georgia at Athens.

This paper reports the following families of mites found on P. disjunctus in
eastern United States: Diarthrophallidae (1 sp.), Diplogyniidae (1 sp.), Euzer-
conidae (1 sp.), Megisthanidae (1 sp.), Digamasellidae (1 sp.), Laelapidae (2
spp.), Macrochelidae (3 spp.), and Heterocheylidae (1 sp.). Also listed here,
but not discussed, are immature specimens of 3 uropodine species described by
Pearse and Wharton (1936) and unnamed species of Anoetidae and Acaridae.
Cosmolaelaps passali (Hunter and Mollin, 1964), is synonymized with Cosmo-
laelaps trifidus (Pearse and Wharton, 1936), new combination. Two new species
of Macrocheles are described.

Family Diarthrophallidae
Diarthro phallus quercus (Pearse and Wharton)

Uroseius quercus Pearse and Wharton 1936: 478.

Diarthro phallus quercus , Tragardh, 1946: 371 (taxonomy) ; Hunter and Glover, 1968: 193
(re-description).

Passalobia duodecimpilosa Lombardini, 1938: 46. Synonymy by Hunter and Glover (1968).
Diarthro phallus similis Tragardh, 1946: 380. Synonymy by Womersley (1961).

Remarks : This unique species, upon which the genus and family was based (Tragardh,

1946), is distinguished in all stages by having very long, barbed adanal and body setae;
very short peritremes in the adult which are absent in the immature stages, by the sternal
shield lacking lateral endopodal projections, and the tarsus of leg I without caruncle or
claws and terminating in a series of short and long setae. All developmental stages are
found on the beetle. Pearse and Wharton (1936) observed that this “mite is usually found
on the outside of Passalus, where it lurks in the crevices between parts near the anterior
end, but sometimes it creeps under the elytra.” The present material was taken on the
coxal regions and under the elytra.

Distribution. This is one of the commonest species of mites found on the venter of head
and coxal regions and under the elytra of P. disjunctus. It is widely distributed in the
eastern United States and has also been recorded from Brazil on an unknown passalid
beetle and from Mexico on Proculus goryi Melly.

Family Digamasellidae

Dendrolaelaps passalorum (Pearse and Wharton). New combination.

(Figures 1-4)

Zercon passalorum Pearse and Wharton, 1936: 477.

Remarks. The type specimens of this species are presumably lost. Figures 28-30' (Pearse
and Wharton 1936: 477) on D. passalorum are undoubtedly based on nymphs. We have
adults of both sexes and nymphs taken from the beetle habitat and under the elytra of
P. disjunctus. A brief description of the adults is as follows:

Female. All dorsal and body setae simple; posterior end of dorsal plate sculptured,
punctate with scalloped margin; dorsal plate notched medially as shown on figure 1,
with 2 small platelets above the slit; infundibulum foraminis extending entire length of

Vol. LXXXIII, March, 1975

Figs. 1-4. Dendrolaelaps passalorum (Pearse and Wharton). 1, dorsum
2, venter of female; 3, venter of nymph with tritosternum ; 4, venter of male.

female ;

New York Entomological Society

trochanter and femur of Leg III; ventri-anal plate with 4 pairs of setae in addition to
anal setae. Male. Spermatodactyl straight, shorter than fixed chela; genital opening located
at anterior margin of sternal plate ; femur, genu and tibia of leg II with ventral protuberances ;
trochanter and femur of leg IV with small, lateral protuberances as in figure 4; posterior
end of dorsal plate as in female. Dorsal seta Si as long as seta Z5 but stronger; S5 longest and
most conspicuous of dorsal setae.

Distribution. Previously known only from North Carolina (type locality). We have
examined a series of immatures taken under the elytra of Popilius from New York (Lintner,
coll.), Ohio (P. Lowry, coll.), Iowa (L. C. Glover, coll.) and Virginia (E. W. Baker, coll.) ;
adult males and females were collected from Popilius habitat under a log pile in Virginia.

Family Diplogyniidae

Passalacarus sylvestris Pearse and Wharton

Passalacarus sylvestris Pearse and Wharton: 475; Tragardh, 1950: 369 (re-description,

taxonomy) .

Remarks. P. sylvestris was re-described and figured in detail by Tragardh (1950) who
placed it in the family Diplogyniidae. This species is distinguished in both sexes by
having the anal plate fused with the ventral plate; the female has a pair of sternal setae
placed close together at the middle near the posterior margin of the sternal plate, and a
pair of triangular plates hinged laterally to the ventral plate bearing 2 pairs of long setae
near the lateral margin. The male genital aperture is situated in front of the anterior
margin of the sternal plate. The biology is not known.

Distribution. P. sylvestris was previously known from North Carolina (type locality).
We have examined specimens taken in the anterior and hind coxal regions of P. disjunctus
from Iowa (L. C. Glover, coll.), New York (Moore, coll.) and from Virginia collected in
an alcohol jar with the beetles (E. W. Baker, coll.).

Family Euzerconidae
Euzercon latus (Banks)

Celaenopsis latus Banks, 1909: 135.

Euzercon latus , Hunter and Davis, 1965: 30 (biology).

Remarks. This euzerconid mite is characterized by having the lateral plates of the female
fused with the ventral plate, by having a T-shaped genital opening and by having the anal
plate separated from the ventral plate. No male or immature stage has been examined by us.
Biology and descriptions of both sexes, including the immature stages are given by Hunter
and Davis (1965).

Distribution. E. latus was originally found on a passalid beetle at Guelph, Ontario, Canada;
it has been recorded since from North Carolina and Georgia. We have examined females
taken on the anterior coxae of P. disjunctus from New York (Moore, coll.) and Iowa
(L. C. Glover, coll.).

Family Laelapidae

Cosmolaelaps trifidus (Pearse and Wharton). New combination.
Seiodes trifidus Pearse and Wharton, 1936: 474.

Cosmolaelaps passali Hunter and Mollin, 1964: 247. New synonymy.

Vol. LXXXIII, March, 1975

Remarks. We have remounted and examined the type specimens of Seiodes trifidus; it is
now obvious that this species is a typical laelapid mite. Both sexes can be distinguished
by the strong, lanceolate setae on the dorsal plate and by the short, strong ventral setae on
the posterior region of the body. Mollin and Hunter (1964) and Hunter and Mollin (1964)
gave detailed biology, descriptions and illustrations of this species as Cosmolaelaps passali,
a synonym of Seiodes trifidus. We cannot find characters to separate them.

Distribution. Hunter and Mollin (1964) reported that the adults were usually found
ventrally between the leg and. prothoracic regions, or attached to the setae in front of
legs I of Popilius from Georgia. One male was removed from the head region of P. disjunctus
from Louisiana (L. C. Glover, coll.). Previously known only from North Carolina
(type locality).

Hypoaspis ( Geolaelaps ) disjuncta Hunter and Yeh
Hypoaspis ( Geolaelaps ) disjuncta Hunter and Yeh, 1969: 97.

Remarks. H. disjuncta is a weakly sclerotized laelapid mite with simple body setae. It
may be distinguished readily by the shape of the sternal plate which has a rounded
posterior margin extending posteriorly to the region of coxae IV, and in that the dorsal
plate possesses 32 pairs of simple setae and completely covers the dorsum. The biology
of this mite was observed by Hunter and Yeh (1969).

Distribution. This species was found attached to the hairs on the venter
of Popilius from Georgia; it was also found in decayed frass mixed with
tunnels. We have females taken on the coxal region and mouthparts
from Ohio (P. Lowry, coll.) and Iowa (L. C. Glover, coll.).

Family Macrochelidae

Macrocheles tridentatus Pearse and Wharton
(Figures 5-11)

Macrocheles tridentatus Pearse and Wharton, 1936: 473.

Remarks. We have remounted and examined the type material (USNM Type No. 1172,
male and female) of M . tridentatus . The type specimen labelled male proved to be a female
as shown in figure 15 (Pearse and Wharton, 1936: 473). This mite represents a new

species of macrochelid. Figure 17 (Pearse and Wharton, loc. cit.) represents the type female
of M. tridentatus. Brief descriptions of the adults of M. tridentatus are as follows:

Female. Dorsal plate finely punctate and with reticulate pattern of punctate lines diminishing
at midregion. Vertical setae Di and all marginal (except M2) and lateral setae pectinate;
seta D8 and all other medial and dorsal setae simple, slender and lanceolate. Sternal
plate punctate with ridges of polygonal design. Genital and ventri-anal plates with reticulate
patterns of punctate lines. All setae on these plates simple. Fixed and movable chelae

of chelicerae with 4 and 2 teeth respectively; arthrodial brush reaching to % length of

movable chela. Leg I with tibia shorter than tarsus. Genu of leg IV with 6 setae pectinate
apically; all other leg setae simple.

Male. Dorsal plate ornamentations similar to those of female. Holoventral plate densely
punctate and with faint polygonal pattern, ventri-anal plate punctate, with reticulate
pattern of punctate lines. All setae on these plates simple. Fixed chelae with 4 teeth,

movable chelae with 1 tooth, spermatodactyl about the length of movable chela. Femur,
genu and tibia of leg II each with a spur, largest on femur. Femur of leg IV as figured.

of the prothorax
soil in the beetle
of P. disjunctus

New York Entomological Society

Figs. 5-12. Macrocheles tridentatus Pearse and Wharton. 5, venter of female; 6, dorsum of
female; 7, chelicera of female; 8, venter of male; 9, chelicera of male; 10, leg II of male;
11, leg IV of male; 12, ambulacra of leg III of male.

Distribution. M. tridentatus was previously known only from North Carolina (type
locality). We have examined females taken on the coxal region of P. disjunctus from
Ohio (P. Lowry, coll.) and Georgia (Y. T. Chiu, coll.) and males collected in beetle frass
from Georgia.

Vol. LXXXIII, March, 1975

Macrocheles disjunctus, n. sp.

(Figures 12-15)

Female. Length of body 735 microns. Dorsal plate heavily ornamented with circular and
polygonal pits of varying sizes; all dorsal setae clublike and strongly plumose, with most
marginal, lateral and anterior dorsal setae stoutest and longest; extra marginal (integumental)
setae also plumose; integument outside plate rugose, granular. Sternal plate pitted, all
setae pectinate. Genital plate small, rounded posteriorly, with pitted reticulate pattern;
setae pectinate. Ventri-anal plate small, longer than wide, pitted; all setae on this plate
pectinate; 3 anal setae simple; integument between these plates rugose; 2 pairs of sclerotized
platelets located between genital and ventri-anal plates. Metasternal plates very small,
each with a pectinate seta. Metapodals not seen. Chelicerae with movable chelae unidentate,
fixed chelae tridentate ; arthrodial brush reaching to *4 length of movable chela. Legs
rugose; most setae strong, either plumose or pectinate. Tarsus of leg I longer than tibia.

Male. Not known.

Holotype. Female, Duncan Falls, Ohio, June 18, 1916 (P. Lowry, coll.), taken on venter
of P. disjunctus , deposited in the New York State Museum and Science Service at Albany.

Paratypes. 1 female with same data as holotype; 1 female, Ft. McPherson, Georgia,
July 22, 1946 (no coll.); 1 female, McRae, Georgia, November 7, 1959 (H. O. Lund, coll.),
both taken on passalid beetle, deposited in the U.S. National Museum and New York
State Museum and Science Service collections.

Remarks. The pitted ornamentation of the dorsal plate and the plumose setae, the rugose
legs and the small genital and ventri-anal plates are distinctive for this species.

Macrocheles whartoni, n. sp.

(Figures 16-18)

Female. Length of body 370 microns. Dorsal plate finely punctate and weakly ornamented
with a network of fine punctate lines. Vertical setae Di short, spinelike, simple and close
to each other; setae D8 finely pectinate; remainder of setae on dorsal plate simple, long,
thin, and lanceolate; extra marginal setae on integument simple. Sternal plate with
characteristic granular and knobby ornamentation; all sternal setae simple. Genital
plate truncate posteriorly, and nearly touching anterior margin of ventri-anal plate, with
concentric pattern of small granular lines. Ventri-anal plate truncate anteriorly, with
punctate polygonal and concentric ornamentation, all setae on this plate simple. Meta-
sternal setae simple, located on small elongate plates. Metapodal plates very small, elongate
and weakly sclerotized. Chelicerae with bidentate fixed chelae and unidentate movable
chelae ; arthrodial brush reaching to % length of movable chela. Leg I with tibia shorter than
tarsus. All leg setae simple.

Male. Not known.

Holotype. Female, Athens, Georgia, August 10, 1965 (Y. T. Chiu, coll.), on passalid
beetle, deposited in the New York State Museum and Science Service at Albany. Paratypes.
6 females with same data as holotype ; 1 female, Clarke Co., Georgia, October 4, 1960,
inside rotten log, and 1 female, Athens, Georgia, University Farm, January 12, 1961, in tunnel
of passalid beetle, both collected by P. E. Hunter; 1 female, Duke Forest, Durham, North
Carolina, June 12, 1933, “on Passalus cornutus Fabr.” (A. S. Pearse, coll., labelled USNM
type no. 1172); 3 females, Stafford, Virginia, August 23, 1973, in Popilius habitat under
oak log, and 2 females on Popilius same locality, all collected by E. W. Baker, deposited
in the U.S. National Museum and New York State Museum and Science Service collections.

New York Entomological Society

Figs. 13-16. Macrocheles disjunctus, n. sp. 13, dorsum of female; 14, venter of female;
15, leg III of female; 16, chelicera of female.

Figs. 17-19. Macrocheles whartoni, n. sp. 17, dorsum of female; 18, venter of female;
19, chelicera of female.

Vol. LXXXIII, March, 1975

Remarks. M. whartoni is distinguished by having only the D8 setae pectinate, the
remainder of body setae being simple, and by the characteristic knobby and granular
ornamentation of the sternal plate. It is the common macrochelid species found on P.
disjunctus.

This mite is named for G. W. Wharton of the Ohio State University.

Family Megisthanidae
Megisthanus floridanus Banks

Megisthanus floridanus Banks, 1904: 145; Baker and Wharton, 1952: 45; Krantz, 1971: 130.

Remarks. Pearse and Wharton (1936) reported that this mite has never been taken in
abundance and none has actually been taken on the beetles which were examined each month.
Our present collection contains 2 females and 1 male collected in P. disjunctus habitat
under a log pile in Virginia. None was found on the beetle. Baker and Wharton (1952), and
recently Krantz (1971), figured M. floridanus. It is distinguished by its unique genital
opening: crescent shaped in the female and placed just below the sternal plate, whereas
the male genital opening is located in the sternal plate aperture.

Distribution. Florida, Georgia and North Carolina. It was collected in Virginia in the
beetle habitat by E. W. Baker.

Family Uropodidae

In the collection are immatures of 3 species of uropodine mites which were named
by Pearse and Wharton (1936). Adults are not known.

Uroobovella spinosa Pearse and Wharton, 1936: 480. Genus uncertain.

Distribution. North Carolina, Ohio, New York, Iowa and Georgia. These mites were
found on the front and hind coxal region of the beetles and under the elytra.

Uroobovella setosa Pearse and Wharton, 1936: 479. Genus uncertain.

Distribution. North Carolina, Louisiana, Ohio, New York and Georgia. The specimens
were taken on the front coxal region of Popilius.

Uroobovella levis Pearse and Wharton, 1936: 481. Genus uncertain.

Distribution. This species is the most common of the uropodine mites found on the beetle
in North Carolina, New York, Ohio, Iowa, Virginia, Georgia and Connecticut. It is found
attached to the hollow areas beneath the head of Popilius and on the front coxal region.

Family Heterocheylidae

Heterocheylus proximus Schuster and Lavoipierre
Heterocheylus proximus Schuster and Lavoipierre, 1970: 26.

Heterocheylus fusiformis Lombardini, of Pearse and Wharton, 1936: 747. Misidentification.

Remarks. H. proximus is obviously the heterocheylid mite reported by Pearse and Wharton
(1936) from North Carolina. It is distinguished from other North and South American
species in that tarsus IV has 4 setae ; the dorsal seta of tarsus IV is located on the distal
portion of the segment which lacks the small basal posterior seta. Seta Im is anterior to
seta IM (after Schuster and Lavoipierre, 1970: 22).

The biology is not known.

New York Entomological Society

Distribution. H. proximus is found under the elytra of P. disjunctus, and is common and
widely distributed in the eastern United States. It has also been recorded on a number of
passalid species from Mexico, Central and South America.

Family Anoetidae

Histiostoma sp.

Remarks. The figure of the hypopus by Pearse and Wharton (1936) readily places this mite
in the above family. They gave it no generic name and placed it in the family ‘Tyroglyphidae.’
The mites collected on the beetles are Histiostoma sp., and probably represent an undescribed
species.

Distribution. North Carolina, Connecticut and Iowa. The mites were found on the coxal
region and under the elytra of the P. disjunctus.

Family Acaridae

Genus nr. Forcellinia.

Remarks. Only the hypopial forms were found, and they are probably an undescribed
genus. Pearse and Wharton (1936) gave a rough figure of this form.

Distribution. North Carolina, New York, Iowa, Connecticut, Virginia and Louisiana.
They were found on the front and hind legs, coxal region and under the elytra of Popilius.

Literature Cited

Axtell, R. C. 1961. New records of North American Macrochelidae (Acarina: Meso-
stigmata) and their predation rates on the house fly. Ann. Entomol. Soc. Amer.
54: 748.

. 1963. Manure inhabiting Macrochelidae (Acarina: Mesostigmata) predaceous on

the house fly. Adv. Acarology 1: 55-59.

. 1969. Macrochelidae (Acarina: Mesostigmata) as biological control agents for

synanthropic flies. Proc. 2nd Int. Cong. Acarology (1967): 401-416.

Baker, E. W., and G. W. Wharton. 1952. Introduction to Acarology. The Macmillan
Co. New York. 465 pp.

Banks, N. 1904. The Arachnida of Florida. Proc. Acad. Nat. Sci. Phil.: 120-147.

. 1909. New Canadian Mites [Arachnoidea, Acarina]. Proc. Entomol. Soc. Wash.

9: 133-143.

Evans, G. O., and E. Browning. 1956. British Mites of the subfamily Macrochelinae
Tragardh (Gamasina-Macrochelidae) . Bull. Br. Mus. (Nat. Hist.) Zool. 4(1):
3-55.

Hunter, P. E., and K. Mollin. 1964. Mites associated with Passalus beetle I. Life
stages and seasonal abundance of Cosmolaelaps passali n. sp. (Acarina: Laelaptidae).
Acarologia 6: 247-256.

, and R. Davis. 1965. Mites associated with Passalus beetle III. Life stages and

observations on the biology of Euzercon latus (Banks) (Acarina: Euzerconidae) .

Acarologia 7: 30<-42.

, and L. Butler. 1966. New Klinckowstroemia mites from Costa Rican passalid

beetles (Acarina: Klinckowstroemiidae) . J. Georgia Entomol. Soc. 1: 24-30.

, and S. Glover. 1968. The genus Passalobia Lombardini 1926, with descriptions of

a new species (Acarina: Diarthrophallidae) . Proc. Entomol. Soc. Wash. 70: 193—
197.

Vol. LXXXIII, March, 1975

, and W. M. Yeh. 1969. Hypoaspis ( Geolaelaps ) disjuncta n. sp. (Acarina: Lae-

lapidae) associated with horned Passalus beetles. J. Georgia Entomol. Soc. 4:
97-102.

Krantz, G. E. 1962. A review of the genera of the family Macrochelidae Vitzthum
1930 (Acarina: Mesostigmata) . Acarologia 4: 143-173.

Lombardini, G. 1938. Acari nuovi. Mem. Soc. Entomol. Ital. 17: 44-46.

Mollin, K. and P. E. Hunter. 1964. Mites associated with Passalus beetle II. Biological
studies of Cosmolaelaps passali Hunter and Mollin (Acarina: Laelapidae). Acarologia
6: 421-431.

Pearse, A. S. and G. W. Wharton in Pearse, A. S., M. T. Patterson, J. S. Rankin, and
G. W. Wharton. 1936. The ecology of Passalus cornutus Fabricius, a beetle which
lives in rotting logs. Ecolog. Monogr. 6: 455-490.

Schuster, R. O., and M. M. J. Lavoipierre. 1970. The mite family Heterocheylidae
Tragardh. Occ. Pap. Calif. Acad. Sci. 85: 1-42.

Tragardh, I. 1946. Diarthrophallina, a new group of Mesostigmata found on passalid
beetles. Entomol. Medd. 24: 369-394.

. 1950. Studies on the Celaenopsidae, Diplogyniidae and Schizogyniidae. Arkiv

Zool. 1(25): 361-451.

Womersley, H. 1957. On some acarina from Australia and New Guinea paraphagic upon
millipeds and cockroaches, and on beetles of the family Passalidae. Trans. R. Soc.
N. S. W. 81: 13-29.

. 1961. On the Family Diarthrophallidae (Acarina-Mesostigmata-Monogynaspida)

with particular reference to the genus Passalobia Lombardini 1926. Trans. Roy. Soc.
S. Austral. 84: 27-46.

New York Entomological Society

The Relationship of Coleomegilla maculata (DeGeer)
(Coleoptera:Coccinellidae) to the Cocoon of Its parasite
Perilitus coccinellae (Schrank) (Hymenoptera:Braconidae)

Allen H. Benton and Andrew J. Crump
Department of Biology, State University College, Fredonia, New York, 14063

Received for Publication July 5, 1974

Abstract: Evidence is presented to indicate that clasping of the occupied cocoon of the
parasitic wasp, Perilitus coccinellae (Schrank) by adult ladybird beetles, Coleomegilla
maculata (DeGeer) is voluntary. There appears to be an attraction of the occupied cocoon
for the adult beetle.

The braconid wasp, Perilitus coccinellae (Schrank) (Fig. 1) is a common
parasite of many beetles, including a variety of Coccinellidae. The distribution,
host records and ecology of this wasp have been studied in some detail by Balduf
(1926), Smith (1953), Sluss (1968) and others. The adult wasp parasitizes
adult or larval beetles (David and Wilde, 1973) and the larva feeds upon the
fat bodies and gonads of its host. It emerges through the suture between pos-
terior abdominal tergites, and upon emergence immediately spins a cocoon.

Many workers have noted that an adult beetle is often found clasping the
cocoon of this parasite (Fig. 2). Several of them (e.g. Balduf, 1926; Smith,
1960) have noted that the larval wasp, as it spins its cocoon, often entangles the
legs of the beetle, either by intent or by accident. Recent observations in our
laboratory and in the field suggest that this interpretation is in error. The
cocoon is often, if not always, attached to the substrate (usually a leaf) and
the beetle clasps it voluntarily.

Our belief that the association of the adult beetle with the cocoon is voluntary
is based upon studies of specimens observed or collected in a corn field
near Fredonia, Chautauqua County, New York, from July to September, 1973.
In the field, and later in the laboratory, we found adult beetles clasping a small
cocoon, which, upon emergence of its occupant, proved to be that of P. coc-
cinellae. Adults found clasping a cocoon usually died soon after emergence of
the wasp, or even before emergence in a few cases. Several workers have reported
that parasitized beetles survived, but we suspect that these reports are based on

Acknowledgments: This research has been supported by National Science Foundation

Grant B036435. We are grateful to Dr. W. R. M. Mason and Dr. C. C. Loan, Biosystematics
Research Institute, Ottawa, Ontario, Canada, for identification of the parasite. Dr. Bernard
C. Smith, Research Station, Harrow, Ontario, Canada, gave us helpful comments on the
first draft of this paper.

New York Entomological Society, XXXIII: 60-63. March, 1975.

Vol. LXXXIII, March, 1975

Fig. 1. Perilitus coccinellae (Schrank) Newly emerged adult.

cases in which the cocoon was clasped by a beetle other than the parasitized
individual. Thus, after emergence of the wasp, the beetle was sufficiently
healthy to leave the cocoon and go its way. Sluss (1968), tracing the life cycle
of the parasite in Hippodamia convergens Guerin, reported that parasitized
individuals died within 3 to 4 days of emergence of the wasp.

Three sorts of observations suggest that the association of the beetle with the
cocoon is voluntary. First, we have observed adult beetles abandoning a cocoon,
and we have found abandoned cocoons in the field. On two occasions, abandon-
ment occurred while the pupa was still in the cocoon, but this occurred only while
we were collecting and transporting beetles from field to laboratory, indicating
that it resulted from disturbance. On a few other occasions, the beetle abandoned
the cocoon after the emergence of the wasp. This would be impossible if the
cocoon were attached to the beetle.

Second, we observed, on one occasion, one beetle clasping a cocoon and
another beetle trying to grasp it from the other side. For some time, both
beetles held the cocoon at opposite ends, but eventually the “intruder”, which
may have been unparasitized and hence stronger, took over the cocoon. This

New York Entomological Society

Fig. 2a. Lateral view of adult Coleomegilla maculata clasping a cocoon of P. coccinellae.
Note both here and in Fig. 2b that the beetle’s legs appear to be actively clasping the cocoon
and that there is no noticeable entanglement of the legs in threads of the cocoon.

may explain previous observations that “parasitized” beetles lived after the
parasite emerged.

Third, if the cocoon is indeed attached to the beetle by the larval wasp, it
would not be attached to the substrate. We found, in the field, two cocoons*
from which the wasp had emerged. Both were attached to corn leaves. In one
case the attaching threads were primarily at one end, while the other cocoon
was attached by threads which extended over the leaf in all directions. There
was no doubt that the threads of the cocoon were firmly attached to the leaf.
We were never able to find any evidence that a cocoon was actually attached to
a beetle. We regularly observed beetles shifting their legs about on the cocoon,
but their legs were never entangled to any significant degree.

It appears, therefore, that the clasping of the cocoon of P. coccinellae is a
voluntary act on the part of adult C. maculata. The occupied cocoon seems to

* While this paper was in press, Mr. Jules Silverman conducted further field studies. He
found numerous cocoons attached to corn leaves, several of them with dead beetles still
clasping the cocoon.

Vol. LXXXIII, March, 1975

Fig. 2b. Ventral view of adult Coleomegilla maculata clasping a cocoon of P. coccinellae.

have a positive attraction for the adult beetle, but this attraction is apparently
lost when the wasp emerges. It would seem most likely that this attraction is
chemical in nature, although we have thus far no direct evidence for this.
Further olfactory experiments are planned.

Literature Cited

Balduf, W. V. 1926. The bionomics of Dinocampus coccinellae Schrank. Ann. Entomol.
Soc. Amer., 19: 465-489.

David, Menter H., and Wilde, Gerald. 1973. Susceptibility of the convergent lady beetle
to parasitism by Perilitus coccinellae (Schrank) (Hymenoptera:Braconidae) . J. Kan-
sas Entomol. Soc., 46: 359-362.

Sluss, R. 1968. Behavioral and anatomical responses of the convergent lady beetle to
parasitism by Perilitus coccinellae (Schrank) (Hymenoptera:Braconidae) . J. Invertebr.
Pathol., 10: 9-27.

Smith, B. C. 1960. Note on parasitism of two coccinellids, Coccinella trifasciata perplexa
Muls. and Coleomegilla maculata lengi Timb. (Coleoptera:Coccinellidae) in Ontario.
Canad. Entomol., 92: 652.

Smith, Owen J. 1953. Species distribution and host records of the Braconid genera
Microctonus and Perilitus (Hymenoptera:Braconidae) . Ohio J. Sci., 53: 173-177.

New York Entomological Society

Proceedings of the New York Entomological Society
Abstracts of Talks Presented at Meetings

FEEDING IN COCKROACHES

The environment of the cockroach was structured so that they could alter their eating
when the environment changed. Three parameters of the environment were altered: tempera-
ture, water supply, and light. Two of these, temperature (35°, 30°, and 25°), and water
supply, are conditions which directly effect energy usage of the animal. The third environ-
mental parameter, light, has no direct energy usage relationship. Within the light parameter,
the light-dark cycle was changed from 16:8 to constant light, and a change from a transparent
to an opaque retreat greatly affected cockroach behavior.

Higher temperatures decreased meal length, but decreased meal number. Females demon-
strated this increase more than did males; however, males decreased the amount they ate
at low temperatures much more than did females. Patterns of feeding varied at different
temperatures. Males particularly reacted more to the light-dark cycle at 35° than at 30°
or at 25°. The animals showed some evidence of acclimation to higher and lower tempera-
tures. The relationship between temperature and meal parameters (meal length, meal
number, intermeal interval) were real, but less pronounced in animals maintained three
weeks at the new temperature.

Light-dark cycles and nature of the provided retreat greatly effected feeding patterns.
When provided an opaque retreat, cockroaches eat most frequently during scotophase.
When given a transparent retreat, the animals eat a number of meals during photophase.
In constant light, with an opaque retreat, both sexes ate irregularly. Meal number is
greatly reduced in constant light.

To determine the effects of a limited water supply on cockroaches, they were given a
specified quantity of water at various intervals from one to four days. Water was given only
during photophase. It was found that female cockroaches will eat in the light, often after
drinking. A few males will eat like females, but most will feed only during the dark.

Betty Faber

New York Entomological Society, XXXIII: 64-68. March, 1975.

Vol. LXXXIII, March, 1975

“Invertebrate Tissue Culture: Applications in Medicine,
Biology and Agriculture”

is the theme of the IV International Conference on Invertebrate Tissue Culture
to be held at Mont Gabriel, Quebec, Canada, June 5-8, 1975. For details,
write to the Chairmen of the Conference: Prof. E. Kurstak (Department of
Microbiology, Faculty of Medicine, University of Montreal, P.O. Box 6128,
Montreal 101, Canada) or Prof. K. Maramorosch (Waksman Institute of Micro-
biology, Rutgers University, New Brunswick, N.J. 08903).

Travel support is available to U.S. scientists through a grant from NIH.
Submit applications to Professor Maramorosch, specifying: (1) cost of economy
round-trip plane fare from airport nearest your home to Dorval Airport, Mon-
treal; (2) mode of participation (main speaker, discussion leader, session chair-
man; whether presenting a paper — indicate title; or other); (3) date of birth;
(4) citizenship; (5) area of special interest (endocrinology, parasitology,
genetics, virology, embryology, neurophysiology, plant pathology, etc.). Applica-
tions will be evaluated by an outside committee and the awards made before
the conference. Those interested in invertebrate tissue culture, and particularly
young scientists and graduate students are urged to apply. Federal employees
are not eligible.

Abstracts of papers will be required by May 1, 1975. The Proceedings will
be published.

BOOK REVIEW

Family Sphingidae of the Palaearctic and Chinese-Himalayan Faunas. N. Ya.

Kuznetsova. 1972. Amerind Publishing Co., Ltd., New Delhi. 43 pp., 1 table.

This is a translation from the Russian of an article originally published in Horae
Societatis Entomologicae, Vol. 37, pp. 293-346, in 1916. The translation, by Dr. M. M.
Haque, was published for the Smithsonian Institution in agreement with the National Science
Foundation. In 1916 the classification of the Sphingidae, largely by Rothschild and Jordan,
and Tutt, was not generally known, so that one purpose of the author was to make this
readily available to Russians. The author consequently prepared an outline of the world
classification. However considerable additional material was added, based in part, on
specimens (especially southern Asiatic) not previously studied. Nomenclature and classifica-
tion changes were suggested. Keys for all taxa from species-level up are given, as well as
much discussion of Palaearctic subspecies. Generic and specific synonymies are given and
discussed. This little known work was considered important enough to warrant its translation
and modern publication. Although the classification of the Sphingidae has come a long way
since 1916, this work has enough value in itself, as well as historical interest, to be a
necessity even for modern and future workers in the group, especially in the higher taxa.

Alexander B. Klots

American Museum of Natural History

New York Entomological Society

BOOK REVIEW

The Physiology of Insecta. 2nd ed. Vol. I. Morris Rockstein, ed. Academic Press, Inc.
1973. $38.

Eight years after the publication of the impressive 3 volumes of the ‘Physiology of Insecta’
the first of a 6-volume new and vastly expanded, second edition appeared. In recent years
there has occurred a very rapid increase in the knowledge of insect biology and physiology,
fully justifying this ambitious venture. The first new volume, a superb book, is completely
revised and up to date. In its preface, Prof. Rockstein reveals the secret of the success of
this impressive treatise: it brings together not only the known facts about insects, but also
discusses the controversial subjects, and many still unsolved and unsettled problems of insect
physiology.

In the introductory chapter on the biology of insects, Rockstein calls attention to the nearly
one million species of insects and their successful evolution, as well as their adaptation to
diverse habitats. The second chapter, by de Wilde and de Loof, deals with the male and
female reproductive systems. The same authors discuss endocrine control of reproduction in
the third chapter. Physiological and biochemical changes during development are dealt with
by Agrell and Lundquist, and the endocrine aspects of growth and development by Lawrence
Gilbert and David Shaw King. The last chapter, by Rockstein and Jaime Miguel, includes
physiological, histological, and microanatomical interpretations of the aging process in insects.
Those interested in gerontology of vertebrates will also be interested in this chapter and the
fascinating basic problems presented.

All authors display impressive expertise in describing diverse and intriguing phenomena,
and they do a very good job in presenting complex problems so that not only insect
physiologists, but also general biologists can profit from their discussions. The book is well
illustrated and produced, and it can be highly recommended. It is a very useful addition to
the entomological literature. Understandably this new, revised edition will become a major
reference work, indispensable to teachers and students alike, and a source of information and
inspiration to all engaged in insect physiology research.

Karl Maramorosch

BOOK REVIEW

A Systematic Monograph of New World Ethmiid Moths (Lepidoptera, Gelechioidea).

Jerry A. Powell. 1973. Smithsonian Contrib. Zool. No. 120. iv + 302 pp., 294 figs., 22 pis.
Superintendent of Documents, U.S. Govt. Printing Office, Washington, D.C. $3.85.

The Ethmiidae are a small, but distinctive, worldwide family of small moths, consisting of
about 250 described species. They show their greatest diversity in the Neotropical region. A
majority of the species occupy relatively small niches, chiefly in xeric areas, and feed mostly
on plants of the large family Boraginaceae and, to a lesser extent, the North American
Hydrophyllaceae. Only 48 species are Nearctic, of which one is a recent introduction from
the Palaearctic and one in California is a probable introduction. The life histories of many
of the Nearctic species are at least partially known, those of the Neotropical ones far less so.
They have essentially no economic importance, although one Hawaiian species is at times a
pest on ornamentals.

Vol. LXXXIII, March, 1975

In all 133 species are covered, of which 49 are described as new. These are placed in only 3
genera (none new) since the author very wisely decided that the present knowledge of the
Neotropical fauna is too scanty to justify a generic classification. The great majority of the
species are placed in the genus Ethmia, as they have always been. All available material was
studied (including the majority of the types) and all discoverable taxonomic characters were
used and are fully illustrated. Numerical taxonomic analyses were extensively made.

The extensive studies of geographical distribution are especially interesting, although
handicapped by a paucity of Neotropical material. The eastern Nearctic is depauperate,
the majority of the species occurring in California and the Southwest. The largest number
of Neotropical species occur in the northern mainland region, with a goodly number in the
Greater Antilles. These West Indian species suggest a strong faunal connection with the
Yucatan Peninsula. Some especially interesting features appear in the life histories of South-
western and Californian species. Such features as diurnal adults flying during only very short
periods around midday (correlated with small eyes) apparently enable some species to
occupy niches in regions probably unsuitable for them at other times of year because of xeric
conditions or high altitudes.

Professor Powell has produced an excellent monograph on a group heretofore little (and
chaotically) known. It will serve as a firm basis for the great amount of work needed in
the Neotropics.

Alexander B. Klots

American Museum of Natural History

New York Entomological Society

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American Museum of Natural History, New York 10024

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Fordham University, New York 10458
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Isaac Albert Research Institute, Brooklyn, N.Y. 11203
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Fairleigh Dickinson University, Madison, N.J. 07940
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Journal of the

New York Entomological Society

Volume LXXXIII June 1975 No. 2

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Waksman Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lois J. Keller, RSM
Dr. Herbert T. Street

Publication Committee

Dr. Kumar Krishna Dr. Ayodha P. Gupta

Dr. James Forbes, Chairman

CONTENTS

Revision of the Genus Endeodes LeConte with a Tabular Key to the Species
(Coleoptera: Melyridae) .... Ian Moore and E. F. Legner 70

Comparative Behavior of Wasps in the Genus Lindenius (Hymenoptera :
Sphecidae, Crabroninae) Richard C. Miller and Frank E. Kurczewski 82

New or Little-Known Crane Flies from Iran. Ill (Diptera: Tipulidae)

- Charles P. Alexander 121

New or Little-Known Crane Flies from Iran. IV (Diptera: Tipulidae)

Charles P. Alexander 129

Book Reviews

81, 128, 139

New York Entomological Society

Revision of the Genus Endeodes LeConte with a Tabular Key to the
Species (Coleoptera: Melyridae)

Ian Moore and E. F. Legner

Division of Biological Control, University of California, Riverside 92502
Received for Publication June 17, 1974

Abstract: Species of the genus Endeodes are known only from the seashore of Pacific

North America. E. fasciatus, E. rothi and E. intermedius, n. spp., are from the upper
Gulf of California. A tabular key to the nine known species is given. The pronotum and
elytron of each species is illustrated.

LeConte (1859) included three species from California when he described
the genus Endeodes. Blackwelder (1932) reviewed the genus and added two
more California species, Moore (1954) reviewed the genus, added a new species
from the Pacific Coast of Baja California Norte, Mexico and reduced one of
LeConte’s species to synonymy. Marshall (1957) described a new species from
the south end of the Gulf of California, and Moore (1964) reviewed the genus
again adding another species from Sonora, Mexico.

The species of this genus are rather unusual in their intertidal habitat. Some
are found on the sandy Pacific beaches of southern California and Baja Cali-
fornia where they are usually encountered under debris, often patches of dried
seaweed. Other species occur from California northward on reefs exposed at
low tide. The Gulf of California supports another group of species which is
also found on rocky shores at low tide.

The genus Endeodes may be distinguished from other members of the family
Melyridae by the combination of the following characters (Arnett 1962): first
sternite not keeled between the coxae, eye simple, head neither rostrate nor exca-
vated, protrusible vesicles present on prothorax and between metathorax and
abdomen, abdomen without bristles, elytra strongly abbreviated, protarsus 5-
segmented, antenna 1 1 -segmented.

Males of Endeodes may be distinguished from females by the presence on
the protarsus of an elongated swollen second segment which terminates in a
comb of thick black setae.

Acknowledgments: We are indebted to Paul Arnaud and David Kavanaugh of the
California Academy of Sciences, San Francisco for loan of a holotype and several paratypes,
to Milton Campbell of the Entomology Research Institute, Ottawa, Canada, for loan and
gift of material and to R. E. Orth, of the University of California, Riverside, for criticism and
technical help. We particularly thank Vincent D. Roth of the Southwestern Research Station
of the American Museum of Natural History, Portal, Arizona, for allowing us to study
intertidal beetles collected by him in the Gulf of California.

New York Entomological Society, LXXXIII: 70-81. June, 1975.

Vol. LXXXIII, June, 1975

The larva of insularis was described by Moore (1956) and a key to the larvae
of three species was given by Moore (1964). A pupa doubtfully identified as
insularis was figured by Moore (1954).

In this paper we describe three new species from the upper Gulf of California,
reduce one species to synonymy and present a tabular key to the species.
Drawings are given of the pronotum and an elytron of each species. The pronota
and elytra offer the best characters for specific separation.

The construction and use of tabular keys, developed by I. M. Newell, were
discussed in two recent papers (Newell 1970, 1972).

STATEMENT OF CHARACTERS

1. Ratio of length to width of elytron = RAT. LEN. WID. ELY.

(5.4 to 12.4)

2. Shape of apex of elytra = SHAPE APEX ELY.

ARCU = arcuate

TRUN = subtruncate, apex straight centrally with the angles broadly rounded

3. Color of elytra = COL. ELY.

PICE = piceus

FERR = entirely ferruginous

BASE = piceus with the base ferruginous

APEX = piceus with the apex ferruginous

MIX rz general mixture of piceus and ferruginous

4. Sculpture of elytra = SCUL. ELY.

ROUG = surface rough and microreticulate

SMOO = surface not rough except for microreticulation

5. Ratio of width to length of pronotum = RAT. WID. LEN. PRON.

(6.4 to 4.4)

6. Shape of pronotum = SHAPE PRON.

TRAN = transverse, not constricted at base

CORD = cordate, not or hardly transverse, constricted at base

7. Color of head = COL. HEAD

PICE r= entirely piceus
FERR = yellow to ferruginous

VARI = variable from ferruginous to ferruginous with dark areas
Distribution = DISTR.

NoPac = California and Pacific Northwest
Cal =z California

C & B = California and Pacific Baja California
Baja = Pacific Baja California

Son = Sonora, Mexico

Gul = Baja California gulf coast

Source

SPM = specimen
Par = paratype
Hoi = holotype

New York Entomological Society

Tabular key to the species of Endeodes

12 3 4 5 6 7

RAT. SHAPE COL. SCUL. RAT. SHAPE COL.

LENG. APEX ELY. ELY. WID. PRON. HEAD

WID. ELY. LEN.

ELY.

PRON.

Distr.

Source

Name

12.4

Trun

Mix

Smoo

5.4

Tran

Ferr

Son

Hoi

faseiatus

12.4

Arcu

Apex

Smoo

5.4

Tran

Ferr

Gulf

Spm

terminalis

9.4

Arcu

Mix

Roug

4.4

Cord

Ferr

Son

Hoi

rot hi

8.4

Arcu

Ferr

Smoo

41/2.4

Tran

Ferr

Son

Hoi

sonorensis

8.4

Arcu

Base

Roug

4.4

Cord

Ferr

C&B

Spm

basalis

8.4

Trun

Ferr

Roug

4.4

Cord

Ferr

Baja

Par

blaisdelli

7.4

Arcu

Mix

Smoo

41/2.4

Tran

Ferr

Son

Hoi

intermedius

5.4

Arcu

Pice

Roug

6.4

Tran

Pice

NoPac

Spm

collaris

5.4

Arcu

Ferr

Roug

6.4

Tran

Ferr

Cal

Par

insularis

Endeodes faseiatus n. sp.

Description of holotype, male.

Color. Head, pronotum and appendages ferruginous; clypeus testaceus; eyes black; elytra
ferruginous with a common piceus spot at inner apical angles which also embraces apical
two-thirds of scutellum, and a piceus band across just below the middle leaving the apices
bright ferruginous; beneath largely dark except head which is ferruginous.

Head. Oval, about as wide as long, tempora about as long as eye; surface rather strongly
microreticulate, very finely punctured and pubescent, the punctures generally separated by
more than their diameters; antennae semi-monilliform, second segment about as long as
third, tenth segment very little longer than wide.

Pronotum. About one-fourth wider than long, widest centrally; apex and base each evenly
arcuate into sides so that the angles are not prominent; surface rather strongly microreticu-
late, very finely and sparsely punctured and pubescent, the punctures separated by more
than twice their diameters.

Elytra. Each elytron a little more than twice as long as wide ; humerus rather narrowly
rounded, sides straight to the just perceptibly inflated apex; outer apical angles broadly
rounded into the briefly truncate apex; inner apical angles more narrowly rounded. Surface
smooth except for a dense but fine microreticulation. Pubescence very fine, short and sparse.

Abdomen. Upper surface concealed by the elytra either due to deformity or damage, the
abdomen being displaced forward so that the basal segments override the metathorax.

Length. About 2.5 mm. This specimen would probably be about 3 mm long except for the
abnormal abdomen.

Specimen described. Holotype, male, Mexico, Sonora, Punta Cirio (29.53°-112.50°) 20
March 1974, from seaweed lying on a 2" to 6" boulder strewn beach, V. Roth and W. Brown
collectors. Deposited in American Museum of Natural History, New York City.

Notes. This species is distinct in its small size, relatively long semitruncate elytra and the
color pattern of the elytra.

Endeodes terminalis Marshall

Endeodes terminalis Marshall 57-13; Moore 64-58; Moore 71-278.

Color. Head ferruginous with the disc near base infumate; pronotum yellow; elytra piceus

Vol. LXXXIII, June, 1975

in basal three-fourths with apex abruptly yellow, dividing line between the two colors
oblique; abdomen and scutellum piceus; legs and antennae ferruginous and infumate.

Head. Oval, about one-fourth wider than long; tempora about as long as eye; surface
densely microreticulate, pubescence fine and moderately dense, punctures imperceptible ;
antenna with second segment not quite as long as third, tenth segment short, as wide as long.

Pronotum. About one-fifth wider than long; apex arcuate, evenly rounded into the broadly
rounded apical angles, sides briefly straight and convergent, basal angles broadly rounded
into the arcuate base, base narrower than apex. Surface sculpture and pubescence very
similar to that of head.

Elytra. Each elytron about two and two-thirds times as long as wide; humerus broadly
rounded, sides straight and somewhat divergent, outer apical angles broadly rounded into
the arcuate apex, inner apical angles broadly rounded. Surface sculpture and pubescence very
similar to that of foreparts.

Abdomen. As finely but not as densely sculptured and pubescent as elytra.

Length. About 3.5 mm.

Specimen examined. One female, Mexico, Baja California Norte, Puertocito, 31 May 1963,
T. Palmer collector. This specimen is accompanied by a larva and a pupa.

Notes. The elytra are the longest in the genus leaving little more than two abdominal seg-
ments exposed. The type locality was given as “Baja California, Mexico, S.E. and Isla
Caballo, III-30-53.” This locality proved to be an error for Isla Ceralbo at the very
southern part of the Gulf of California (Moore 1971). The new locality for the specimen
described above is five hundred and fifty miles north in the upper Gulf of California.

Endeodes rothi n. sp.

Description of holotype, female.

Color. Head, pronotum and appendages bright ferruginous; clypeus testaceus; eyes black;
elytra with base, apex and marginal beeding ferruginous, disc piceus; abdomen largely
piceus with the basal, lateral and apical margins of anterior segments ferruginous; beneath
ferruginous except for patches of piceus on terminal abdominal segments.

Head. Oval, a little longer than wide ; tempora about one and one-half times as long as
eye ; surface finely microreticulate, very finely punctured and pubescent, the punctures gen-
erally separated by more than their diameters; antennae with all the segments longer than
wide, second segment almost as long as third, tenth segment half again as long as wide.

Pronotum. About as wide as long, widest at apical third; apex broadly rounded, evenly
rounded into apical angles, thence sharply constricted in basal third to the narrowly rounded
basal angles ; base gently arcuate ; surface somewhat impressed in center of base ; base four-
fifths as wide as pronotum. Surface very finely microreticulate and shining. Punctures very
fine, generally separated by about twice their diameters.

Elytra. Each elytron a little more than twice as long as wide; humerus broadly rounded,
sides thence nearly straight for a short distance, thence widened and broadly arcuate to the
widest point at about four-fifths of the length, thence broadly rounded into the evenly
arcuate apex. Elytra conjointly appear sharply constricted at basal third and widely inflated
in basal two-thirds, the surface flattened in basal third. Sculpture rough and rather strongly

New York Entomological Society

1 mm

Figs. 1-9, elytra of Endeodes :

1. terminalis
3. jasciatus
5. basalis
7. intermedins
9. collaris

2. rothi
4. blaisdelli
6. sonorensis
8. insularis

Vol. LXXXIII, June, 1975

microreticulate with fine punctures separated generally by at least twice their diameters.
Pubescence of fine decumbent pale hairs and sparse coarse long dark setae.

Abdomen. About as wide as elytra. Microreticulation a little finer than on elytra. Punctures
very sparse. Pubescence very fine.

Length. About 4.8 mm.

Specimen examined. Holotype, female, Mexico, Sonora, Punta Cirio (29.53°-112.50°) , March
20, 1974, from seaweed on 2"-6" boulder strewn beach, V. Roth and W. Brown collectors,
in collection of The American Museum of Natural History, New York City.

Notes. This is the most distinctive species in the genus. It differs from the other species
particularly in its longer tempora, elongate antennomeres and the fact that the elytra are
conjointly constricted at the base and inflated in the apical two-thirds. The shape of the
pronotum is similar to that of the Pacific Coast species basalis and blaisdelli. This species
is named in honor of one of its collectors, Vincent D. Roth.

Endeodes sonorensis Moore
Endeodes sonorensis Moore 64-57, 58.

Color. Head and pronotum ferruginous; legs ferruginous with the femora infumate, antennae
ferruginous becoming darker apically; elytra piceus with humerus very narrowly ferruginous;
abdomen largely ferruginous with a piceus central cloud on each segment, last segment mostly
piceus; beneath ferruginous except for piceus metasternum and last abdominal segment.

Head. Oval, a little wider than long; tempora about as long as eyes; surface without
ground sculpture, finely punctured, the punctures generally separated by about their diam-
eters; pubescence very fine. Antennae semi-monilliform, second segment slightly shorter than
third, tenth a little longer than wide.

Pronotum. A little wider than long, widest at apical fourth ; apex arcuate, anterior angles
broadly rounded, sides thence straight and convergent to broadly rounded basal angles, base
straight, narrower than apex; without ground sculpture; punctures and pubescence very
similar to that of head.

Elytra. Each elytron about twice as long as wide; humeral angle narrowly rounded, sides
gently arcuate into broadly arcuate apex. Surface sculpture a dense but fine microreticulation,
otherwise smooth except for a small central slightly rough patch.

Abdomen. A little wider than conjoint elytra; punctures and pubescence similar to that of
pronotum.

Length. About 4 mm.

Specimen examined. Holotype, female, Mexico, Sonora, Punta De Los Cuervos, San Carlos
Bay, near Guaymas, 18 November 1962, intertidal reef, Ian Moore collector, in collection
of California Academy of Sciences, San Francisco.

Notes. This species resembles intermedins but differs in its longer elytra and other details.
We have also seen one male and one female from Mexico, Sonora, Kino Bay, 21-22 Sep-
tember 1973, V. Roth and W. Brown collectors. These specimens are similar to the holotype
except that the abdomen and metasternum of one are entirely ferruginous and in the other
slightly infumate.

New York Entomological Society

Endeodes basalis (LeConte)

Atelestus basalis LeConte 52-168.

Atelestus abdominalis LeConte 52-168, New Synonym. (This synonymy was suggested by
Moore, 1954, but not clearly indicated.)

Endeodes basalis LeConte 59-122; Horn 72-112; Blackwelder 32-134; Moore 54-196, 197;
Moore 57-140; Moore 64-158.

Endeodes abdominalis LeConte 52-122; Horn 72-112; Blackwelder 32-134.

Color. Head, pronotum and antennae ferruginous, legs ferruginous with the femora infumate,
elytra largely piceus with the base ferruginous, abdomen piceus.

Head. Oval, about as wide as long; tempora about one and one-half times as long as eye;
surface very finely, densely microreticulate ; pubescence fine, short and moderately dense,
punctures imperceptible ; antenna with second segment slightly more than half as long as
third, tenth segment slightly longer than wide.

Pronotum. About as long as wide ; widest at about apical third ; apex arcuate into the broadly
rounded apical angles; sides briefly constricted just before the rounded basal angles; base
slightly emarginate. Sculpture and pubescence very similar to that of head.

Elytra. Each elytron about twice as long as wide, humeral angle narrowly rounded, sides
straight and diverging to the broadly rounded outer apical angles, apex arcuate, inner apical
angles narrowly rounded. Surface rough and with dense microreticulation. Punctures and
pubescence much as on the elytra with added long scattered erect setae.

Abdomen. Sculpture, punctures and pubescence very similar to that of elytra.

Length. About 3.5 mm.

Specimen described. Female, California, San Luis Obispo County, Cambria, 21 August 1972,
under dry seaweed on berm of beach, Ian Moore collector.

Notes. This species is distinct in the combination of its long piceus elytra with the base pale
and its cordate pronotum. The color of the abdomen is variable, ranging from entirely
ferruginous to entirely piceus with many intergrades (Moore 1954) which led LeConte to
describe one color form under the name abdominalis. It is reported from Ensenada, Baja
California, Mexico to Monterey County. It is most commonly found under dried seaweed
and other debris on the berm of the sandy beaches.

Endeodes blaisdelli Moore
Endeodes blaisdelli Moore 54-196; Moore 64-58.

Color. Entirely dull ferruginous except eyes, tips of mandibles black and abdomen above
and beneath piceus.

Head. Oval, about as wide as long; tempora slightly longer than eye; finely densely micro-
reticulate throughout ; finely pubescent but not perceptibly punctured. Antenna with the
second segment about half as long as third, tenth segment slightly longer than wide.

Pronotum. About as wide as long, widest at about apical third; apex arcuate into the broadly
rounded apical angles, sides briefly constricted just before the rounded basal angles base
very slightly emarginate. Sculpture and pubescence similar to that of head.

Elytra. Each elytron about twice as long as wide ; humeral angles narrowly rounded, sides
straight and slightly diverging to the broadly rounded outer apical angles; apex straight,

Vol. LXXXIII, June, 1975

1mm

10. terminalis
12. jasciatus
14. b as alls
16. intermedins
18. collaris

11. rothi
13. blaisdelli
15. sonorensis
17. insularis

New York Entomological Society

inner apical angles broadly rounded. Surface rough and with dense microreticulation. Pu-
bescence fine and sparse with a few long, erect, pale setae.

Abdomen. Microreticulation more sparse than that of foreparts, punctures and pubescence
very fine and sparse.

Length. About 3 mm.

Specimens examined. Seven paratypes, Mexico, Baja California, Colonia Guerrera, 19 August
1950, Ian Moore collector.

Notes. This species is easily known from all the others except fasciatus by its long, truncate
elytra; it differs from fasciatus in its cordate pronotum and concolorous elytra. It is known
only from the type locality.

Endeodes intermedius n. sp.

Description of holotype, female.

Color. Head, pronotum, legs and under surface of abdomen ferruginous; eyes and tips of
mandibles black ; antenna largely ferruginous becoming darker apically ; elytra piceus on
disc with a narrow ferruginous rim, a little widest as base ; abdomen largely piceus with
edges of basal segments testaceous.

Head. Oval, very slightly longer than wide ; tempora about one-half longer than eyes ; surface
just perceptibly microreticulate, shining; moderately, coarsely punctured, the punctures gener-
ally separated by about their diameters ; antennae semi-monilliform, second segment two-thirds
as long as third, tenth segment hardly longer than wide.

Pronotum. Slightly wider than long, widest at apical third, apex gently arcuate into the
broadly arcuate anterior angles, sides thence convergent and nearly straight to the more
narrowly rounded basal angles, base nearly straight but slight emarginate centrally; base
about four-fifths as wide as apex. Surface without microsculpture; punctures dense, sepa-
rated by less than their diameters; pubescence short and dark.

Elytra. Each elytron a little less than twice as long as wide: humerus narrowly rounded,
sides briefly straight, thence arcuate into the very broadly rounded outer apical angles;
surface vaguely impressed near scutellum ; surface finely microreticulate ; punctures and
pubescence much like those of pronotum.

Abdomen. Reticulation a little finer than on elytra; punctures and pubescence a little more
dense and fine than on elytra.

Length. About 4 mm.

Specimen described. Holotype, female, Mexico, Sonora, Punta Cuevas (29.42 °-l 12. 35°) , 24-5
September, 1973, V. Roth and W. Brown collectors, on algae covered pitted ryolite, at night
during low tide, in the collection of The American Museum of Natural History, New York
City.

Allotype, male, Mexico, Sonora, Puerto de Lobos (30.16°-112.50°), March 18-19, 1974, V.
Roth and W. Brown collectors.

Paratype, one female, same data as holotype.

Notes. The name intermedius was chosen for this species because the elytra are intermediate
in length between those of collaris and insularis and those of the other species. The shape of
the pronotum along with that of sonorensis is intermediate between that of basalis and

Vol. LXXXIII, June, 1975

blaisdelli and that of the other species. The lengths of the antennal segments along with those
of sonorensis are intermediate between those of rothi and the other species.

Endeodes collar is (LeConte)

Atelestus collaris LeConte 52-168.

Endeodes collaris LeConte 59-122; Horn 72-112, Blackwelder 32-134; Moore 54-196; Moore
56-220 (Larva) ; Moore 64-58.

Color. Piceus with the bases of the antennae, bases of the tibiae, the tarsi and trophi paler and
the pronotum testaceus.

Head. Oval, one-third wider than long; tempora almost twice as long as eyes; surface densely,
finely microreticulate ; punctures moderately large, separated by less than their diameters;
impressed in center of apical half ; second antennal segment almost as long as third, tenth
segment almost as wide as long.

Pronotum. One-third wider than long; widest in the middle; apex arcuate, apical angles
broadly rounded, sides arcuate into the broadly rounded basal angles, base nearly straight but
perceptibly emarginate in center. Surface without microreticulation, punctures feeble, generally
separated by about their diameters, with a few dark scattered setae.

Elytra. One-fifth wider than long; humeral angles obsolete, sides arcuate into the broadly
rounded outer apical angles and arcuate apex, inner apical angles broadly rounded. Surface
rough and densely microreticulate, with fine pubescence and scattered erect setae. Punctures
not apparent.

Abdomen. Feebly microreticulate with punctures generally separated by a little more than
their diameters, with sparse pale decumbent pubescence.

Length. About 4.75 mm.

Specimen described. Female, Nanaimo, British Columbia, Canada, 16 June 1927, L. G.
Saunders collector.

Notes. This species is distinct in the combination of the very small elytra and the piceus head.
The elytra are very much smaller in relation to the pronotum than those of the other species
except insularis. It is known from Vancover Island, British Columbia to San Mates County,
California. It is usually taken under drift on the beach below high tide mark.

Endeodes insularis Blackwelder

Endeodes insularis Blackwelder 32-134; Moore 54-196, 198; Moore 56-229 (Pupa?); Moore
64-58.

Endeodes rugiceps Blackwelder 32-135; Moore 54-196; Moore 56-220 (Larva) ; Moore 64-58

New Synonym.

Color. Entirely pale ferruginous except tip of mandibles black, eyes grey and abdomen
entirely piceus.

Head. Oval, two-fifths wider then long; tempora slightly shorter than eye; surface very
densely finely microreticulate; pubescence short, dense and fine with scattered short dark
setae in basal half ; flattened in center of apical half ; antenna with second segment a little
more than half as long as third, tenth about as wide as long.

New York Entomological Society

Pronotum. One-half wider than long, widest at apical fourth, apex arcuate, apical angles
broadly rounded, sides rounded into broadly rounded basal angles, base arcuate ; surface
feebly microreticulate, punctures dense, separated by less than their diameters, pubescence
as on head with scattered short dark setae throughout.

Elytra. Each elytron very small, about one-fourth longer than wide, conjointly much narrower
than pronotum; humeral angle hardly apparent, sides very weakly arcuate, apex broadly
arcuate; surface rough, sculpture and pubescence similar to those of head; numerous short
dark erect setae throughout.

Abdomen. Sculpture a fine dense microreticulation; pubescence fine, pale, decumbent,
moderately dense.

Length. About 4.5 mm.

Specimen described. Paratype female, San Miguel Island, Santa Barbara County, California,
20 June 1910, V. W. Owen collector.

Notes. This species is distinct in its very small elytra combined with its red head. The elytra
are very much smaller in relation to the head than any other species in the genus except collaris.

We have also examined one male and one female paratype with the same data as above
and one male paratype from Prince Island, Santa Barbara County, California, 19 May 1919,
E. P. Van Duzzee collector.

We have seen four paratypes of E. rugiceps Blackwelder from Carmel, Monterey County,
California taken in March, May and June from 1912 to 1923. Two of these are males and
two females. We can find no difference between these and the paratypes of insularis except
that the femora and antennae of two of them are somewhat darker than those parts in
insularis, a character that is variable in some species of the genus. Blackwelder (1932, p. 135)
said of rugiceps “Male genitalia as in collaris.” However, we have dissected a male paratype
(Carmel, III-25-23, Blaisdell collection) and find the aedeagus to be the same as that of
insularis as figured by Blackwelder (1932, p. 133, figs. 3F). Therefore, we conclude that
rugiceps is a synonym of insularis.

Literature Cited

Arnett, Ross H. Jr. 1962. The Beetles of the United States (A Manual for Identification).
Pt. V, pp. 648-850. The Catholic University of America Press. Washington.

Blackwelder, Richard E. 1932. The genus Endeodes LeConte. Pan-Pac. Ent., 8: 128—
136, 3 Figs.

Horn, George H. 1872. Synopsis of the Malachiidae of the United States. Trans. Ameri-
can Ent. Soc., 4: 109-127.

LeConte, John L. 1852. Catalogue of the Melyrids of the United States, with descrip-
tion of new species. Proc. Acad. Nat. Sci. Philadelphia, 6: 163-171.

. 1859. Descriptions of some species of Coleoptera from the vicinity of the southern

boundary of the United States of America. Arcana Naturae, 1: 121-128.

Marshall, M. Y. 1957. Studies in Malachiidae VII. Coleopt. Bull., 11: 13-16.

Moore, Ian. 1954. Notes on Endeodes with a description of a new species from Baja
California (Coleoptera: Malachiidae). Pan-Pac. Ent., 30: 195-198.

— . 1956. Notes of some intertidal Coleoptera with descriptions of the early stages

(Carabidae, Staphylinidae, Malachiidae). Trans. San Diego Soc. Nat. Hist., 12: 207-
230, pis. 14-17.

. 1957. A northern extension of range for Endeodes basalis LeConte. Pan-Pac.

Ent., 33: 140.

Vol. LXXXIII, June, 1975

. 1964. A new species of Endeodes from Sonora, Mexico. (Coleoptera: Melyridae).

Pan-Pac. Ent., 40: 57-58.

. 1971. The type locality of Endeodes terminalis Marshall (Coleoptera: Malachi-

idae). Pan-Pac. Ent., 47: 278.

Newell, Irwin M. 1970. Construction and use of tabular keys. Pac. Insects 12: 25-37,
3 Figs.

. 1972. Tabular keys, further notes on their construction and use. Trans. Conn.

Acad. Arts Sci., 44: 259-267. 4 Figs.

BOOK REVIEW

PEST CONTROL: A Survey. Arthur Woods, Halsted Press, John Wiley & Sons, New
York, 407 p. $29.50. 1974.

This book can be recommended highly to all interested in the principles and methods of
pest control. It is written as an introductory text, with carefully chosen examples of insect
pests, diseases of plants, technological advances, and biological means of control. The author
first defines pests and their control, as well as the economics of pest control and of methods
used. Factors such as population density, death rate, and community stability are analyzed
in the second chapter. The third chapter deals with the uses of chemical pesticides, as well
as the economics of their production. Drawbacks of chemical control receive due attention
in the fourth chapter. Biological control, including the use of insects, bacteria, viruses, fungi,
higher plants, to mention the main ones, are outlined in the following 3 chapters, followed
by newer approaches, such as sterilization, genetic control, pheromones, attractants, repel-
lents, and the use of miscellaneous other control methods. Finally, integrated control is
presented in proper perspective. The book is so written that it can be used not only by
the professional scientist, interested in biology, agriculture, entomology, or ecology, but
also by the general reader. To achieve this and not to oversimplify has been a difficult
task solved by the author admirably. The book can be used as a text in university and
college courses on pest control, conservation, and courses dealing with the impact of man
on his environment. References are well chosen and the index is divided into 3 parts,
listing separately diverse names and subjects, scientific names of species, including viruses,
and Latin names.

I admired the readability of this excellent volume, a feature seldom found in technical
books. Perhaps the fact that the author has written numerous articles for technical journals
and that he has produced 6 courses on biological topics for the Australian television network
have been responsible for this feature. The book would make a valuable addition to school
libraries and public libraries everywhere.

Although the author modestly states that a book like his rapidly becomes out of date,
this volume contains so much valuable information and presents it so well that it will be
used as a reference in the years to come. Efficient pest control is urgently needed and if
adequate awareness of population control becomes a reality, mankind might find a way of
survival. Otherwise, even the best control methods will merely postpone the doom. Pro-
ducing more food is a necessity but producing adequate amounts of food for an uncon-
trolled population of the world is an impossibility.

Karl Maramorosch

Waksman Institute of Microbiology

Rutgers University

New Brunswick, New Jersey

New York Entomological Society

Comparative behavior of wasps in the genus
Lindenius (Hymenoptera: Sphecidae, Crabroninae)

Richard C. Miller1 and Frank E. Kurczewski

Department of Entomology, State University of New York, College of
Environmental Science and Forestry, Syracuse, New York 13210

Received for Publication June 17, 1974

Abstract: Observations on the nesting behaviors of the Nearctic Lindenius armaticeps, L.

buccadentis, and L. columbianus errans are presented and the world literature on the
ethology of the Palaearctic L. albilabris , L. panzeri, and L. pygmaeus is reviewed. Be-
havioral features shared by all species include construction of nests 3-12 cm deep in firmly-
packed sand or fine gravel, presence of a vertical main burrow leading frequently to a
short horizontal passage, absence of a temporary nest closure during provisioning, attach-
ment of the egg to the neck of a prey along the ventral midline, and distribution of the
prey remains evenly over the surface of the cocoon. Valuable characteristics for separating
species or species groups include the kinds, proportions, and stages of the prey, chronological
placement of cells in the nest, method of prey storage, absence or presence of impalement
of the prey during transport, exiting behavior, and number of prey per cell.

INTRODUCTION

The genus Lindenius currently includes 48 species of small ground-nesting
wasps, 37 in the Palaearctic and 11 in the Nearctic and Neotropical regions
(DeBeaumont, 1956; Court and Bohart, 1958; Leclercq, 1954, 1959, 1960).
The majority of the Palaearctic species are found along the Mediterranean,
especially at the western end (DeBeaumont, 1956). The southwestern U.S. is
a secondary stronghold for the genus because all the New World species except
L. montezuma (Cameron), known only from Mexico, occur there (Muesebeck

Acknowledgments: We are grateful to D. L. Dindal and E. O. Price for critically reading
parts of the manuscript, and to S. E. Ginsburg and C. J. Lane for their assistance in the
field. We are indebted to R. A. Norton and D. J. Peckham for many of the photographs,
and to H. E. Evans for prey records from Blackjack Creek, Kansas, and nest provisions
from Lexington, Massachusetts. We are especially grateful to the following persons for
their help in identifying the unusually large number of prey insects: E. F. Cook, Univer-
sity of Minnesota; K. V. Krombein, Smithsonian Institution; B. D. Burks, R. W. Carlson,
R. J. Gagne, J. L. Herring, L. V. Knutson, P. M. Marsh, A. S. Menke, L. M. Russell,
C. W. Sabrosky, R. I. Sailer, D. R. Smith, G. C. Steyskal, and W. W. Wirth, all of the
Systematic Entomology Laboratory, USDA, ARS. This work was supported by a Grant-
in-Aid from the Research Foundation of the State University of New York (No. 10-7116-A)
and an NSF Undergraduate Research Participation Program Grant, summer 1970 (No.
GY-7286).

Present address: Department of Entomology, Cornell University, Ithaca, New York

14850.

New York Entomological Society, LXXXIII: 82-120. June, 1975.

Vol. LXXXIII, June, 1975

et al., 1951; Leclercq, 1954; Krombein, 1958; Krombein and Burks, 1967).
The only species with ranges extending into the eastern U.S. are the 3 treated
in this paper: L. armaticeps (Fox), L. buccadentis Mickel, and L. columbianus
errans (Fox).

The nesting behaviors of the Nearctic species are essentially unknown. In
contrast, the Palaearctic L. albilabris (Fabricius), L. panzeri (Van der Linden),
and L. pygmaeus (Rossi) have been studied over a period of 90 years. The
purpose of this paper is to review the world literature on the nesting behavior
of Lindenius , to present original results on L. armaticeps, L. buccadentis, and
L. c. errans, and to determine the ethological characters most useful in distin-
guishing species and those of potential value in separating Lindenius from
other sphecid genera. A detailed account of male behavior and intraspecific
interactions in aggregations of Lindenius has been presented elsewhere (Miller
and Kurczewski, 1973).

LITERATURE REVIEW

Lindenius ( Lindenius ) albilabris (Fabricius)

Kohl (1915) reported this species as the most common and widespread
member of the genus in the Palaearctic region. Nests have been found from
June to October in level, often compact, sand, sandy-gravel, loess, or chalk
of paths and sandpits (Adlerz, 1903, 1910; Gronblom, 1925; Minkiewicz,
1931, 1933; Chambers, 1949; Bliithgen, 1955; Bonelli, 1967). The wasps
often aggregated with as many as 16 nests per sq yd (Nielsen, 1900).

The circular nest entrance, 2-3.5 mm in diameter, was surrounded by a
transitory tumulus, 30 mm wide X 10 mm high, that was periodically renewed
during burrow construction (Minkiewicz, 1931, 1933; Bonelli, 1967). The
main burrow, straight or somewhat curved depending on subterranean ob-
stacles, descended vertically 3-10 cm and usually turned horizontally into a
passage, 1-10 cm long (Nielsen, 1900; Adlerz, 1910; Minkiewicz, 1931, 1933;
Bristowe, 1948). From 1 to 7 cells, 5-6 X 6-13 mm, were positioned around
the horizontal burrow or the lower half of the vertical shaft at depths of 4-
10 cm (Adlerz, 1910; Minkiewicz, 1931, 1933). Bonelli (1967) described an
unusual nest in which the vertical burrow forked into 2 branches, each ter-
minating in 3 cells. Straight or winding side burrows, 1-6 cm long, were
plugged with sand and led from the main burrow to completed cells (Minkie-
wicz, 1933). The cells were either strongly inclined (Bonelli, 1967) or nearly
horizontal (Minkiewicz, 1933; Bristowe, 1948), and were occasionally sepa-
rated by only 1-2 mm (Minkiewicz, 1931).

In the first nest figured by Minkiewicz (1931, Tab. XI, Fig. 6) the incom-
pletely provisioned cell was the shallowest, whereas in his second example
(1933, Tab. XII, Fig. 7) it was the deepest one in the nest. Bonelli (1967)

New York Entomological Society

excavated a nest having the older cells nearer the entrance. Newly-captured
prey were stored in open, incompletely provisioned cells rather than in the
burrow (Adlerz, 1910; Minkiewicz, 1931, 1933).

One female observed in Italy by Bonelli (1967) opened her nest entrance
at 0830 hrs but did not exit until 0900 hrs, whereupon she made an orientation
flight facing the entrance. Such orientation was not repeated subsequently
unless the female experienced difficulty in entering the nest or was about to
begin provisioning a newly-excavated cell. After the full complement of prey
had been gathered, the female closed the entrance with sand, sealed off the
completed cell, and constructed a deeper new cell. The sand was pushed out-
side, increasing the size of the tumulus. Within 30-45 min after plugging the
entrance, the female emerged and began provisioning the new cell. She made
15 trips for prey before plugging the entrance again and spending the night
in the burrow. Two cells were completed per day, the excavation and pro-
visioning of each taking about 3 hrs. Adlerz (1910) noted that the female
rested in the burrow not only at night but also at mid-day and during bad
weather. He observed that the nest entrance was plugged only when the
female was excavating a new cell. Minkiewicz (1931) believed that each
female constructed only a single nest.

Provisioning wasps returned to their nests in flight, diving in without hesi-
tation (Adlerz, 1910; Bristowe, 1948; Bluthgen, 1955; Bonelli, 1967). Adlerz
(1910) reported that the prey was held with the middle legs when the wasp
was on the ground, whereas Bristowe (1948), Hertzog (1954), and Bluthgen
(1955) observed impaling of the prey on the sting. Bonelli (1967) believed
that the female held the prey with the posterior claws. Bristowe (1948) and
Bluthgen (1955) observed the impaling only after capturing the wasp and its
prey in a container. The former noted that the [ tibial ] spurs of the middle
legs were used to hold the impaled prey in position under the abdomen.

The average time required to capture a prey and return to the nest, calcu-
lated from Bonelli’s (1967) data, was 4.3 min (1-12, N = 19). The average
time needed to store a prey and return to the surface was only slightly less
(x = 4 min, 2-6, N = 18). Hamm and Richards (1926) and Bristowe (1948)
found prey discarded near the nest entrances but disagreed on the underlying
causes.

Although L. albilabris stores mainly adult and nymphal Miridae, it occa-
sionally captures adult flies of the families Empididae, Dolichopodidae, and
Chloropidae (Table 1). Twenty-two genera of mirids made up 95% of all
prey records. Possibly the numerous undetermined Diptera found by Adlerz
(1903, 1910) included representatives of families other than those mentioned.
In 13 of 20 areas where L. albilabris was studied, only mirids were found as
prey. Unfortunately, detailed records of nest contents were not made in the

Vol. LXXXIII, June, 1975

Table 1. Prey of Lindenius albilabris.

Family

Species

Source

Miridae

Hemiptera

Adelphocoris sp.

Hamm and Richards, 1926

Amblytylus nasutus (Kirschbaum)

Hamm and Richards, 1926;

[af finis Fieber]1

Bristowe, 1948

Calocoris norvegicus (Gmelin)

Hamm and Richards, 1926;

Calocoris roseomaculatus (DeGeer)

Bonelli, 1967

Hamm and Richards, 1926

Conostethus roseus (Fallen)

Hamm and Richards, 1926

Globiceps flavomaculatus (Fabricius)

Bouwman, 1911

Halticus apterus (Linnaeus)

Gronblom, 1925

Heterotoma merioptera (Scopoli)

Bristowe, 1948

Hoplomachus thunbergi (Fallen)

Nielsen, 1900

Leptopterna ferrugata (Fallen)

Gronblom, 1925

Lygocoris pabulinus (Linnaeus)

Bristowe, 1948

Lygus pratensis (Linnaeus)

Minkiewicz, 1931, 1933

Lygus sp.

Gronblom, 1925

Megaloceroea recticornis (Geoffroy)

Bristowe, 1948

[ linearis Fuessly]

Megalocoleus molliculus (Fallen)

Hamm and Richards, 1926

Miris sp.

Adlerz, 1910; Hamm and

Notostira erratica (Linnaeus)

Richards, 1926
Hamm and Richards, 1926

Orthotylus ericetorum (Fallen)

Hamm and Richards, 1926

Orthotylus sp.

Bristowe, 1948

Pithanus maerkeli (Herrich-Schaeffer)

Bristowe, 1948

Plagiognathus sp. nr. albipennis

Bluthgen, 1955

(Fallen)

Plagiognathus arbustorum (Fabricius)

Bristowe, 1948

Plagiognathus chrysanthemi (Wolff)

Gronblom, 1925; Hamm and

Polymer us unifasciatus (Fabricius)

Richards, 1926; Bristowe,
1948

Hamm and Richards, 1926

Stenodema calcaratum (Fallen)

Bouwman, 1911; Gronblom,

Stenotus binotatus (Fabricius)

1925

Hamm and Richards, 1926;

Trigonotylus ruficornis (Geoffroy)

Bristowe, 1948

Gronblom, 1925; Hamm and

Empididae

Diptera

Empis aestiva Loew

Richards, 1926; Bristowe,
1948

Bristowe, 1948

Empis albinervis Meigen

Bristowe, 1948

Dolichopodidae

Diaphorus latifrons Loew

Sickmann, 1893

Chloropidae

Meromyza pratorum Meigen

Hamm and Richards, 1926

Meromyza saltatrix (Linnaeus)

Hamm and Richards, 1926;

[laeta Meigen]

Bristowe, 1948

1 [ ] indicates synonyms given by original authors.

3 areas where only flies were reported so that one may doubt whether L.
albilabris ever stores flies exclusively. In 1 area of Poland L. albilabris ap-
peared to be prey-specific, taking only Lygus pratensis (Linnaeus) (Minkie-
wicz, 1933), whereas in England it captured 10 species of mirids as well as

New York Entomological Society

3 species of chloropid and empidid flies (Bristowe, 1948). Bonelli (1967)
observed prey specificity in Italy on a different mirid, Calocoris sp., probably
norvegicus (Gmelin). The claims of certain compilers (Iwata, 1942; Dupuis,
1947) that L. albilabris preys on Hymenoptera are in error.

The most extensive observations on the number of prey per complete cell
were made in Sweden by Adlerz (1910) who found 6-23 (x = 16.1) prey in 23
cells. Minkiewicz (1931) found as few as 4 per complete cell, whereas Bonelli
(1967) found as many as 25. Data on the number of genera and species of
prey per nest and cell were lacking except in the 2 instances of prey specificity
cited above. Hamm and Richards (1926) and Bristowe (1948) found more
female than male mirids. The prey were stacked one atop another in the cell,
all heads facing in the same direction (Adlerz, 1910). They were paralyzed
(Nielsen, 1900; Adlerz, 1910; Bouwman, 1911; Minkiewicz, 1933; Bristowe,
1948; Bonelli, 1967), and the egg-bearer was not mutilated (Minkiewicz,
1931).

The egg was attached to the neck of a mirid along the ventral midline and
extended obliquely backward at an angle of 45° to the body axis of the prey
(Minkiewicz, 1931, Tab. XIV, Fig. 7; Bonelli, 1967, Tab. II), or was nearly
transverse (Adlerz, 1910). According to Minkiewicz (1931), the egg was
white, 2-3 X 0.6 mm, and followed the curvature of the bug’s prosternum.
It was laid after the full complement of prey had been gathered (Minkiewicz,
1933) and was borne by 1 of the first prey placed in the cell (Bonelli, 1967).

The development of the larva and the construction of the cocoon were
discussed by Bonelli (1967). The egg hatched in about 40 hrs in an artificial
cell in the laboratory and the larva reached maturity in 4 days, consuming
even the more sclerotized parts of the nymphal mirids. The larva enveloped
itself in a silken cocoon in 12 hrs and, during the next 24 hrs, spread a secre-
tion over the inside that cemented the threads and sand grains together. The
yellow-brown cocoons, covered only with sand, were 4 X 7-8 mm (Fig. II).
Gronblom (1925) found smaller, darker cocoons in Finland and noted that
they were covered with prey remains as well as with larval excrement and
gravel particles. According to Bonelli (1967), L. albilabris is monovoltine in
at least part of Italy and overwinters in the larval stage.

Lindenius ( Trachelosimus ) panzeri (Van der Linden)

Kohl (1915) recorded this species from central and southern Europe,
England, northern Africa and western Asia. Nests were found from July to
September in flat, hard-packed sand and loess of garden walks and woods’
paths or, less commonly, in sloping banks (Marchal, 1893; Bouwman, 1911;
Kohl, 1915; Hamm and Richards, 1926; Minkiewicz, 1932, 1933; Guichard
and Yarrow, 1947; Abrahamsen, 1950).

Vol. LXXXIII, June, 1975

Table 2. Prey of Lindenius panzeri.

Family

Species

Source

Simuliidae

Simulium ornatum Meigen

Sickmann, 1893

Tephritidae

Dithryca guttularis (Meigen)

Orellia jaceae (Robineau-Desvoidy)
Tephritis sp.

Trupanea stellata (Fuessly)

Sickmann, 1893; Kohl, 1915
Minkiewicz, 1932
Bouwman, 1911
Minkiewicz, 1932

Milichiidae

Madiza glabra Fallen

Minkiewicz, 1932

Chloropidae

Chlorops hypostigma Meigen
Chlorops pumilionis (Bjerkander)

[ lineata Fabricius, taeniopus Meigen]
Chlorops rufina (Zetterstedt)

Chlorops troglodytes (Zetterstedt)
Chlorops sp.

Meromyza saltatrix (Linnaeus)

[ laeta Meigen]

Thaumatomyia glabra (Meigen)
undetermined sp.

Bouwman, 1911
Marchal, 1893 ; Hamm and
Richards, 1926
Hamm and Richards, 1926
Hamm and Richards, 1926
Abrahamsen, 1950
Hamm and Richards, 1926

Minkiewicz, 1932
Guichard and Yarrow, 1947

The circular entrance, 3-3.5 mm in diameter, was surrounded by a small
tumulus of soil that soon eroded (Minkiewicz, 1932, 1933). The main bur-
rows were vertical, slightly inclined, or tortuous as a result of stones in the
soil, and descended to depths of 4-12 cm (Bouwman, 1911; Minkiewicz, 1932;
Abrahamsen, 1950; Olberg, 1959). Abrahamsen (1950) noted that nests were
shallower in a heavily used, somewhat clayey footpath than in the sand along-
side a forest road. From 2 to 9 cells per nest were found at depths of 3-11 cm
and were joined to the main burrow by short, nearly horizontal burrows
(Bouwman, 1911; Abrahamsen, 1950, Figs. 10-11). Bouwman (1911) sche-
matically illustrated a nest having 9 cells in 2 horizontal planes cutting through
a vertical burrow, the lower level of cells being constructed before the upper
one. He believed that the sand from the excavation of a new cell was used
to plug the side burrow leading to the previously provisioned cell.

Prey-laden females returned to their open nest entrances in flight, carrying
the prey head-forward beneath them. The prey was held with the middle
and hind legs of the wasp before landing, but only the middle legs were used
on the ground (Olberg, 1959, p. 372). Tn contrast, Hertzog (1954) observed
L. panzeri carrying its prey impaled on the sting.

With one exception, the prey of L. panzeri were flies of the acalyptrate
muscoid Cyclorrhapha (Table 2). Chloropidae was the family most commonly
stored, but it was not represented by as many genera as the Tephritidae.
Hamm and Richards (1926) found only Chloropidae as prey in 3 areas in
England, whereas Minkiewicz (1932) obtained Chloropidae, Tephritidae, and
Milichiidae from nests in Poland. Bouwman (1911) and Minkiewicz (1932)
indicated that non-chloropid flies were seldom captured. The fact that only
2 species of prey have been reported from more than 1 locality seems to

New York Entomological Society

indicate that additional prey records are necessary for a more realistic appraisal
of the prey limits of L. panzeri.

The number of prey per complete cell ranged from 11 (Abrahamsen, 1950)
to 22 (Bouwman, 1911). Hamm and Richards (1926) found that 85 (98%)
of 87 chloropids were males. Although Marchal (1893) believed the prey
were killed outright, Abrahamsen (1950) asserted that they were paralyzed
and moved their legs for several days after capture. Minkiewicz (1933) noted
that the egg was attached ventrally to the neck of 1 of the prey, but only
after the full complement had been gathered. In the Netherlands, the larva
consumed the provisions in 1-5 days (Bouwman, 1911).

Lindenius ( Trachelosimus ) pygmaeus (Rossi)

Leclercq (1954) recorded this species from central and southern Europe,
England, northern Africa, and southern Russia. Nests were found from June
to September in level or gently sloping sand, compact sandy-clay, and loess
(Ferton, 1901; Grandi, 1928, 1961; Minkiewicz, 1932, 1933; Maneval, 1937).
The circular entrance, 2 mm in diameter, was surrounded by a small tumulus
of soil that was often obliterated by wind and rain (Minkiewicz, 1932, 1933).
The main burrows, vertical or slightly inclined, descended 8-10 cm and were
seldom winding (Ferton, 1901; Grandi, 1928, 1961; Minkiewicz, 1932, 1933;
Maneval, 1937). The burrow figured by Minkiewicz (1932, Tab. VI, Fig. 1)
ended blindly, whereas that described by Grandi (1928, 1961) bent to the
side after descending. Two cells excavated by Grandi (1928, 1961) were
4X7 mm and only 1 cm apart at a depth of 10 cm. Minkiewicz (1932)
stated that the cells were all at about the same [unspecified] depth.

Ferton (1901) observed a female of L. pygmaeus capture a braconid on a
flower, sting it for a long time, and carry it away. According to Minkiewicz
(1932, 1933), females were slow in provisioning their nests and hovered cau-
tiously above the open entrances before diving in. Olberg (1959, p. 375)
photographed females holding the prey ventral-side-up and head-forward, using
the middle pair of legs on the ground and both middle and hind legs in flight.

At least 95% of the prey of L. pygmaeus were Chalcidoidea and Ichneu-
monoidea (Hymenoptera) although nematocerous and acalyptrate flies were
occasionally captured (Table 3). Chalcidoids were the most common provi-
sions, constituting 88% of all prey records. The most common prey family,
Pteromalidae, was represented by 15 genera, whereas the second in importance,
Eulophidae, was represented by only 3 genera. A winged ant (Maneval, 1937)
was an exceptional prey from the order Hymenoptera. The same author ob-
tained 2 nematocerous flies, a sciarid and a ceratopogonid from the cell con-
taining the ant, thereby confirming Grandi’s (1928) observation that L. pyg-
maeus hunts Diptera as well as Hymenoptera. Olberg (1959) also found flies

Vol. LXXXIII, June, 1975

Table 3. Prey of Lindenius pygmaeus.

Family

Species

Source

Braconidae

Hymenoptera

Apanteles sp.

Ferton, 1901; Minkiewicz, 1932;

Ichneumonidae

Ophioninae sp.

Maneval, 1937
Ferton, 1901

Chalcidoidea

undetermined sp.

Ferton, 1901; Maneval, 1937

Eulophidae

Euplectrus bicolor (Swederus)

Minkiewicz, 1932

Necremnus hippia (Walker)

Minkiewicz, 1932

Tetrastichus sp.

Grandi, 1928, 1961 ;

Torymidae

Torymus verbasci Ruschka

Minkiewicz, 1932
Minkiewicz, 1932

Pteromalidae

Cecidostiba collaris Thomson

Minkiewicz, 1932

Coelopisthia cephalotes (Walker)

Grandi, 1928, 1961;

Coelopisthia spp.

Minkiewicz, 1932
Grandi, 1928, 1961

Conomorium eremita Forster

Grandi, 1928, 1961;

Cyclogastrella deplanata (Nees)

Minkiewicz, 1932
Minkiewicz, 1932

[ domesticus Walker]
Dibrachys cavus (Walker)

Minkiewicz, 1932

[ boucheanus Ratzeburg]
Diglochis silvicola (Walker)

Minkiewicz, 1932

[ complanatus Thomson]
Habrocytus artemisiae Forster

Minkiewicz, 1932

Habrocytus psittacinus Forster

Minkiewicz, 1932

Habrocytus spp.

Grandi, 1928, 1961;

Homoporus sp.

Minkiewicz, 1932
Minkiewicz, 1932

Mesopolobus modestus (Walker)

Minkiewicz, 1932

Mesopolobus sp. [ Eutelus Walker]

Grandi, 1928, 1961

Pachyneuron formosum Walker

Minkiewicz, 1932

Pteromalus sp.

Marchal, 1893

Sphegigaster sp.

Grandi, 1928, 1961

Stenomalina subfumatus Thomson

Minkiewicz, 1932

Sy stasis encyrtoides Walker

Minkiewicz, 1932

Systasis longicornis Thomson

Grandi, 1928, 1961

Trichomalus punctinucha Thomson

Minkiewicz, 1932

Trichomalus sp.

Grandi, 1928, 1961;

Formicidae

Leptothorax nylanderi Forster

Minkiewicz, 1932
Maneval, 1937

Ceratopogonidae

Diptera

Forcipomyia bipunctata Linnaeus

Maneval, 1937

Sciaridae

Sciara sp.

Maneval, 1937

acalyptrate muscoid Cyclorrhapha

Olberg, 1959 (in photograph,

undetermined specimen

not stated)
Grandi, 1928, 1961

as prey but neglected to point out that the ones being carried by the females
he photographed (p. 375) included acalyptrates, a group not previously re-
ported as prey of L. pygmaeus.

The number of prey per complete cell ranged from 17 (Minkiewicz, 1932)
to 42 (Grandi, 1928, 1961), whereas the number of prey families per cell
ranged from 3 (Grandi, 1928, 1961) to at least 5 (Maneval, 1937). The only

New York Entomological Society

cell whose contents were completely identified (Minkiewicz, 1932) held 12
species belonging to 10 genera. Of the 45 prey Chalcidoidea listed by Minkie-
wicz (1932), 78% were females. Grandi (1928) also found that a majority
of the chalcidoids were females. Marchal (1893), Ferton (1901), and Grandi
(1928, 1961) found the prey to be paralyzed to varying degrees.

Minkiewicz (1933) noted that the egg was placed on the prey only after
the provisions had been gathered. The elongate egg was attached ventrally
to the neck of a chalcid along the midline, and curved obliquely backwards
at an angle of 30-45° to the longitudinal axis of the prey’s body (Minkiewicz,
1932, Tab. VI, Fig. 2). Grandi (1928, 1961) found the egg-bearer lying
supine near the surface of the mass of provisions. The egg, 1.6 X 0.4 mm,
was curved and off-white in color. Marchal (1893) and Grandi (1928) ob-
served that the cocoon was covered with chalcid remains and had a metallic
sheen. It was elongate, 4X8 mm, and had the aboral end more acute
(Grandi, 1928).

RESULTS

Lindenius ( Trachelosimus ) armaticeps (Fox)

This species occurs in southern Canada and the U.S. east of the Cascade
and Sierra Nevada Mountains (Muesebeck, et al., 1951). Twenty-five nests
were excavated as follows: 17, Selkirk Shores St. Pk., Oswego Co., N.Y., 15
June-4 August 1971-72; 5, Penny Settlement Rd., Lewis Co., N.Y., 18 August-
19 September 1971; 2, Medford Lakes, Burlington Co., N.J., 22-23 July 1972;
and 1, Great River, Suffolk Co., N.Y., 7 August 1972. Nests were usually
found in level, firm and heavily-vegetated or extremely hard-packed and bare
sandy roadbeds. The roads were bordered by wild grasses and annual herbs.
At Selkirk, nests were located in a 6-ft-high sand cliff (Fig. 1).

The entrances of most active nests in level sand were surrounded by roughly
circular tumuli, averaging 21 mm in diameter and 3 mm high (15-25 X 2-5,
N = 13). Tumuli were often lacking in older nests. The burrows (Figs. 2A-H),
2 mm wide, entered the roadbed perpendicularly and descended more or less
vertically to depths of 4.0-9. 0 cm (x = 6.1, N = 25). Pebbles and rocks were
responsible for deviations of burrows from the vertical. Terminal horizontal
passages, 1. 0-3.0 cm long (x = 1.6), were present in 18 of 23 nests in flat sand.
The cliff nests at Selkirk entered the bank at nearly right angles and des-
cended obliquely for 6.5-10.0 cm (Fig. 2H). One of 2 nests possessed a
1-cm-long horizontal passage extending parallel to the cliff face from the apex
of the main burrow. Tumuli were not present around such entrances because
the sand fell down the slope.

All nests had the cells arranged radially around the lower half or two-thirds
of the main shaft, the lowest cell never being deeper than the open burrow.
With one exception, the oldest cell in a nest was the shallowest and the

Vol. LXXXIII, June, 1975

Fig. 1. Six ft high sand cliff at Selkirk Shores St. Pk., Oswego Co., N.Y. Females of
Lindenius armaticeps nested in the lower 4 ft.

New York Entomological Society

newest, the deepest (Figs. 2A-H). In level sand the distance of the cells
from a vertical line through the entrance ranged from 0.3-4. 2 cm (x = 1.8,
N = 48) ; pairs of cells separated by 1 cm or less were common. Cells in flat
sand were unearthed at an average depth of 5.2 cm (2. 7-8.0, N = 57), whereas
in cliffs they were 2-9 cm (x = 4.9, N = 7) below the level of the entrance and
3. 5-7. 5 cm (x = 4.8) into the bank from a vertical line through the entrance.
An average cell measured 3.2 X 7.4 mm (2. 5-4.0 X 6. 0-9.0, N = 47).

Temporary storage of prey in an un widened section of the burrow was
common in nests being actively provisioned. However, in two nests, prey
were stored in a slightly or conspicuously widened area at the end of the
burrow. Storage of prey at the bottom of the vertical shaft was most common
(Figs. 2A-D, F), but sometimes the flies were stored beyond the bend (Fig.
2G) .

The nest resident plugs the upper burrow with sand when she has obtained
enough prey to complete a cell. In such nests, a terminal horizontal passage
has usually already been constructed and the female forms a cell at its apex.
By opening nests at the appropriate time, it was ascertained that the female
takes the prey from burrow storage and places them head-inward in the cell
before ovipositing. Whether or not she removes them again before affixing
an egg to the innermost fly is unknown. The vertical burrow is extended
deeper and a new side passage is usually constructed prior to the female’s
reappearance at the surface. The entire process from entry to reappearance,
including prey positioning, oviposition, and construction of a new side burrow
took 88 min for one female and 108 for another.

Completed nests contained from 3 to 11 cells. Many nests were abandoned
after periods of rain, and the entrances were subsequently obliterated. Females
took 60 to 100 min to finally close their nest. The wasp removed sand from
the upper walls of the vertical burrow with the mandibles and used its forelegs
to push this sand down the burrow. One female exited every 5-15 sec during
the final stage of nest closure and pushed sand from around the entrance
backward into the burrow. When nest closure was complete, all that remained
was a small conical depression, 5-7 mm wide and 5-15 mm deep. Each female
then made an orientation flight above her closed nest before flying off and
beginning a new nest elsewhere.

The prey were placed head-inward in the cells, but at least 1 fly was usually
oblique or head-outward. The majority of prey on the bottoms of the cells
were ventral-side-up while those on top were commonly dorsum-up. An egg-
bearer, either male or female, was found at the inner end of the cell and was
positioned dorsum-up, on its side, or venter-up. The white, slightly curved egg,
1.5 X 0.4 mm, was attached to the neck of a fly along the ventral midline
and was directed obliquely backward at an angle of 30-50° to its longitudinal

Vol. LXXXIII, June, 1975

Fig. 2. Top (t) and side (s) views of nests of Lindenius armaticeps. Stippling indicates
tumulus and sand fill; •, burrow storage. Cells are numbered in apparent chronological
order, according to contents.

New York Entomological Society

Table 4. Prey of Lindenius armaticeps and their relative importance as provisions.

Species N % total $ $ $ $

Selkirk Shores St. Pk., Oswego Co., N. Y.

Diplotoxa versicolor (Loew)
Meromyza sp. nr. pratorum Meigen
Parectecephala eucera (Loew)
Parectecephala sanguinolenta (Loew)
Thaumatomyia glabra (Meigen)

Penny Settlement Rd., Lewis Co., N. Y.
Parectecephala eucera (Loew)

Great River, Suffolk Co., N. Y.
Parectecephala eucera (Loew)

Medford Lakes, Burlington Co., N. J.
Chlorops sp.

Diplotoxa versicolor (Loew)

0.7

3.1

341

76.6

228

113

18.0

1.6

163

100.0

107

100.0

27.3

72.7

axis. It extended across either side of the fly with equal frequency, and the
wing of the prey nearest its caudal end was spread as often as not.

The prey comprised flies of the family Chloropidae (Table 4). Parecte-
cephala eucera (Loew) was especially prominent, making up 77% of the pro-
visions at Selkirk and 100% of the provisions at Penny Sett. Rd. and Great
River. At Selkirk, 228 (67%) of 341 prey of this species were males compared
to 107 (66%) of 163 flies at Penny Sett. Rd. On the other hand, only 26
(33%) of 80 Parectecephala sanguinolenta (Loew) from nests at Selkirk were
males. P. sanguinolenta was abundant in the fresh, low grass in the road
between the nests, whereas P. eucera was common in the older, taller grass
to the sides. The former species was captured as prey only in June and early
July, while the latter was obtained from nests throughout the summer. Prey
capture was observed several times within 15 ft of the midline of the road.
During hunting, the wasps either walked on the grass blades or flew in circles
around the stems and darted at dark objects on the green background including
imperfections of the plants.

The average weight of a single prey was 1.1 mg (0.4-5.1, N = 202), the
lightest being a male of P. eucera and the heaviest a female of P. sanguinolenta.
The prey were rarely as heavy as their female captors which weighed 2. 8-5. 2 mg
(x = 4.4, N = 9). The average total weight of the flies in a fully-provisioned
cell was 12.1 mg (7.4-18.8, N = 19). The number of prey per complete cell
ranged from 3 to 15 (x = 9.9, N = 39). The average number of genera per
nest and cell were 1.5 (1-3, N = 20) and 1.2 (1-2, N = 46) at Selkirk and
Penny Sett. Rd., while the average number of species per nest and cell were
1.8 (1-4) and 1.5 (1-3), respectively. Except for the nests in New Jersey,
each cell contained some Parectecephala.

Vol. LXXXIII, June, 1975

Two females were observed digging nests from the sand surface in mid-
June. Each burrowed inward in a slow spiral, pushing up sand for slightly
more than an hour before their tumuli remained stationary. These wasps
stayed inside for the rest of the day and started provisioning the next morn-
ing. One female completed 2 cells and began provisioning a third during
that day. She provisioned cells much slower a week later, and during her
third and final week she brought no prey to the nest. She opened the nest
each morning and closed it every evening but was rarely seen in the nest
vicinity. A similar senescence was observed in another female who opened
and closed her nest which contained only cocoons! Two other females showed
a different kind of aging: newly-captured flies walked or flew out of the

entrance shortly after the female dove down the burrow with them. After the
non-paralyzed flies escaped, the wasp began backing up and down the burrow
or repeatedly exiting and reentering in flight.

Females made orientation flights above their nests before the first provi-
sioning trip of the day, after completion of a cell, or after disturbances such
as a passing insect kicking sand into the entrance or an ant entering the
burrow. In flat sand, an orientation lasted 3-15 sec and consisted of 1 to
many obverse and reverse half-circles, 15 cm in diameter, above the nest.
During lengthy orientations, the semi-circular flights changed to linear, spring-
like movements which gradually increased in length from the entrance. Fe-
males inhabiting cliffs made transverse flights above and in front of their
entrances, the flights gradually increasing in length up to 20 cm.

Provisioning females returned to their nests in flight, holding the prey
ventral-side-up and head-forward with the middle legs (Fig. 3). A female
approached her nest with little or no hovering and dove into the open entrance
from a distance of 3-4 cm. Occasionally a female with prey landed in front
of her entrance and simply walked into the burrow. Abandoned prey were
not found near L. armaticeps nests.

Many females displayed an unusual method of prey transport — impalement
of the fly on the sting — if they returned to their nests and found the entrances
obstructed. Obstructed entrances often occurred naturally; however, compara-
ble observations were made by artificially blocking openings with sand or bits
of leaves. In either case, the female approached the nest holding the prey
with her middle legs but then landed and impaled the fly in the region of the
mesosternum. One wasp walked around looking for her naturally obstructed
entrance for several minutes with the fly trailing from the end of her abdomen.
She even made a brief, wavering flight with the prey still impaled. After
locating the entrance, she walked into her nest with the prey extending head-
forward from the under-curved tip of the abdomen. Significantly, wasps began
carrying their prey into the nests in the usual manner on later trips when
the entrances were unobstructed. A few females never impaled flies at arti-

New York Entomological Society

ficially-blocked entrances but continued to hold their prey with the middle
legs as they dug in with the forelegs.

Females provisioned nests from 0900 to 1950 hrs, but the majority of wasps
were active between 1030 and 1600 hrs. At night, they plugged the upper
burrows with sand and remained inside. The plug was removed in the morn-
ing, often as much as 1 hr before the first provisioning trip, and fresh sand
was pushed out of the entrance if the nest was relatively new.

The average time required by 8 females at Selkirk to capture a fly and
return to the nest was 11.3 min (0.8-35.0, N = 50). One female averaged only
9.0 min (1.7-35.0, N = 10), whereas a second took 12.5 min (2.1-21.5, N =
13). The average time needed to store a prey in the burrow and return to the
surface at Selkirk was 2.2 min (0. 8-8.0, N = 52), 1 female averaging only 1.8
min (1.0-5.1,N = 17) and another 2.9 min (1.5-6. 6, N =11). Females usually
spent several seconds looking out of the entrance upon returning to the surface
(Fig. 4). The average time spent in the entrance before flying off to obtain
prey was 10 sec (0-30, N = 46) at Selkirk, the extremes being 0-15 sec (x = 6,
N = 10) for one individual and 4-30 sec (x = 17, N = 19) for another. With
rare exception, L. armaticeps walked out of the nest, partly or entirely, before
taking flight (Fig. 5).

One individual required 7 days to develop from an egg to a mature larva
ready to spin a cocoon. The same rate of development was observed for
specimens reared in sand-filled plastic containers in the laboratory. Mature
larvae were positioned head-inward in the cells, facing away from the side
passages leading to other cells.

The brown, ellipsoidal cocoons, averaging 2.5 X 6.0 mm (2-3 X 5-7, N = 12),
were completely and evenly covered with the wings, legs, and thoracic sclerites
of the prey. These remains were firmly attached to the exterior in a matrix
of silk and sand. Cocoons removed from cells provisioned in late June and
early July contained pupae, not resting larvae. These individuals would have
comprised the August-September generation of L. armaticeps.

The sarcophagid Phrosinella fulvicornis (Coquillett) was observed larviposit-
ing in closed nest entrances and was reared from a puparium found in a cell
at Selkirk.

Lindenius ( Trachelosimus ) buccadentis Mickel

This species has been recorded from New York, Pennsylvania, Virginia, Iowa,
Nebraska, Kansas, and Arizona (Muesebeck, et al., 1951; Krombein and Burks,

Fig. 3-5. Female of Lindenius armaticeps carrying a chloropid fly with the middle legs;
4. pausing in entranceway before exiting; and, 5. walking out of nest.

Vol. LXXXIII, June, 1975

New York Entomological Society

1967). The Cornell University Museum contains specimens from Vermont,
New Hampshire, New Jersey, and Texas. Two nests were excavated at Med-
ford Lakes, Burlington Co., N.J. on 8 August 1971, and 6 were dug at Bo-
hemia, Suffolk Co., N.Y. from 7 to 9 August 1972. In both areas, nests were

Vol. LXXXIII, June, 1975

found in the pine barrens in level hard-packed roadbeds of coarse sand mar-
gined by patches of wild grasses.

Nest entrances were surrounded by roughly circular tumuli, averaging 22
mm in diameter and 3 mm high (18-25 X 2-5, N = 8). The main burrows
descended vertically or in weak spirals to depths of 4. 8-9.0 cm, and often
curved 5-6 cm below the surface (Figs. 6A-G). Five nests had short hori-
zontal passages, 1.0-2. 5 cm long, at the apices of the main burrows. Nest
entrances and burrows were 1.75-2.00 mm in diameter.

The oval cells were arranged radially around the lower third of the main
burrow and were 0.2-3 .8 cm from a vertical line through the entrance (x = 1.9,
N = 21). The average cell depth was 6.0 cm (3.5-10.5) and the average cell
size, 2.9 X 6.8 mm (2. 5-3. 5 X 5. 5-8.0). Side burrows could not be traced.
Active nests usually contained recently captured prey in temporary storage
at the bottom of the main burrow (Figs. 6A, C, E-G), and these were occa-
sionally sealed off by a loose plug of sand (Figs. 6C, E, G). Sections con-
taining prey were no wider than other parts of the burrow. However, 2 nests
at Bohemia (Figs. 6E, F) had prey temporarily stored in an open cell at the
end of a short side passage. Neither nest contained enough prey for the female
to complete the cell. The maximum number of cells per nest was 4, the cells
decreasing in age with increasing depth.

Provisioning females returned to their nests in flight, holding the prey tightly
against the sternum with the middle legs. They dove into their entrances from
distances of 3-5 cm. The average time required to capture a prey and return
to the nest, based on the activities of 2 females at Bohemia, was 4.4 min
(0.7-13.5, N = 34). One female took an average of only 3.7 min (0.9-10.6,
N = 23), whereas the other required an average of 5.7 min (0.7-13.5, N = 11).
The average time needed to store a prey was 41 sec (12-360, N = 38), one
female taking 36 sec (12-80, N = 13) and the other 44 sec (16-360, N = 25).
Storage times did not exceed 80 sec with the exception of one 360 sec. This
unusually long storage time occurred when a worker ant, Monomorium mini-
mum (Buckley), fell into a nest seconds before the wasp returned with prey.
The ant failed to reappear and, when the nest was dug open, 2 such worker
ants lay paralyzed with other prey at the bottom of the main burrow. Exiting
behavior of females varied greatly with the same individual either pausing or
failing to pause in the entrance and either walking out of the nest or not
before taking flight. The average time spent looking out of the entrance before
flying off was 6 sec (0-55, N = 34). Females made brief orientation flights
over their nests when disturbed by male or female searchers (see Miller and
Kurczewski, 1973), and they sometimes made 2-3 sec orientations without
apparent cause.

After gathering sufficient prey to complete a cell, the female plugged the

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Table 5. Prey of Lindenius buccadentis and their relative importance as provisions.

Family

Species

% total

$ $

Medford Lakes, N. J.

Diptera

Empididae

Drapetis sp. nr. divergens Loew

1.7

Hymenoptera

Braconidae

A gat his sp.

1.7

Apanteles paralechiae Muesebeck

61.7

Orgilus sp.

10.0

Eulophidae

Achrysocharella silvia Girault

8.3

Euplectrus sp.

1.7

Tetrastichus sp.

3.3

Pteromalidae

Pteromalinae sp.

3.3

Eurytomidae

Bruchophagus sp.

1.7

Ormyridae

Ormyrus brunneipes Provancher

1.7

Cynipidae

Charips sp.

1.7

Bethylidae

Apenesia parapolita Evans

1.7

Formicidae

Monomorium minimum (Buckley)

1.7

Bohemia, N. Y.

Hymenoptera

Braconidae

A gat his sp.

0.7

Apanteles sp.

0.7

Bracon sp.

1.1

Diaeretus sp.

0.7

Pauesia sp.

0.4

Phanerotoma sp.

0.4

Rhaconotus cressoni Muesebeck & Walkley

0.4

Ichneumonidae

Acrolytina, n. gen.

0.4

Mesochorus sp.

0.4

Toxophoroides scitulus (Cresson)

0.4

Eulophidae

Achrysocharella silvia Girault

0.4

Chrysocharis sp.

0.7

Closterocerus tricinctus (Ashmead)

1.1

Euderus sp.

0.7

Eulophus anomocerus (Crawford)

0.4

Hyssopus sp.

0.7

Tetrastichus whitmani (Girault)

0.4

Tetrastichus sp.

1.4

Perilampidae

Perilampus fulvicornis Ashmead

1.1

Perilampus robertsoni Crawford

0.4

Ormyridae

Ormyrus brunneipes Provancher

2.9

Pteromalidae

Capellia sp.

0.4

Erythromalus sp.

1.8

Gastrancistrus aphidis (Girault)

216

77.7

175

Pachyneuron siphonophorae (Ashmead)

0.4

Eurytomidae

Eudecatoma sp.

0.4

Chalcididae

Spilochalcis sp.

0.4

Cynipidae

Charips sp.

0.4

Cynipinae sp.

1.4

Bethylidae

Goniozus sp.

0.7

Formicidae

Monomorium minimum (Buckley)

0.7

Tapinoma sessile (Say)

0.4

Worker.

Vol. LXXXIII, June, 1975

101

upper burrow with sand. She did not push quantities of sand out of the
entrance during extension of the burrow. One female required 75 min to
complete a cell and reopen the burrow. The number of cells constructed and
provisioned each day varied between individuals. One female completed a cell
containing 29 prey between 0930 and 1300 hrs whereas another female re-
quired most of 2 days for completion of 2 cells containing 77 and 74 prey.

Over 92% of the prey, comprising 1 family of Diptera and 11 families of
Hymenoptera (Table 5), were from the superfamilies Ichneumonoidea and
Chalcidoidea. Braconidae was the main prey family at Medford Lakes, whereas
Pteromalidae was the main source of prey at Bohemia. Selection of exclusively
one sex of prey was not apparent, but high percentages of male pteromalids
and female eulophids were noted. The average number of prey per complete
cell was 42 (22-77, N = 7) and the average weight of the cell contents, at
Medford Lakes, was 7.25 mg (6.45-8.05, N = 2). A minute eulophid, Achryso-
charella silvia Girault, was the lightest prey while a braconid, Agathis sp., was
the heaviest. The average weight of an individual prey was 0.30 mg (0.10-
1.15, N = 49), whereas the average weight of 5 female wasps was 3.1 mg (2.6-
3.4). The average number of prey families per nest and cell were 7.0 (4-10,
N = 4) and 5.4 (3-9, N = 7), while the average number of genera and species
per cell was 8.4 (4-14, N = 7).

Most of the larger prey were positioned head-inward in the cell, away from
the main burrow, whereas smaller prey, especially chalcids, were found in
various positions. The egg-bearer was at the innermost end of the cell and
was ventral-side-up, if large, but randomly-positioned, if small. The off-white,
slightly curved egg, 1.5 X 0.3 mm, was attached to the neck of a braconid
or chalcid along the ventral midline and extended longitudinally backward at
an angle of 10° to the body axis of the prey. The brown, ellipsoidal cocoons,
averaging 2.5 X 6.0 mm (2. 0-3.0 X 5. 5- 7.0, N = 5), were completely covered
with the heads, thoraces, legs, and wings of the prey. These remains were
firmly embedded in the cocoon.

Lindenius ( Trachelosimus ) columbianus errans (Fox)

L. c. errans is found in southern Canada and the U.S. east of the Rockies,
while the nominate subspecies occurs in B.C., Washington, Idaho, Utah, and
Wyoming (Muesebeck, et al., 1951; Evans, 1970). In Massachusetts L. c.
errans provisioned nests with minute Diptera, Hymenoptera, and Hemiptera
(Evans, 1963), whereas in Wyoming the nominate subspecies stored a chalcid
wasp (Evans, 1970).

Our behavioral studies were made in sandpits in 4 areas of central New
York: Auburn and Sennett, Cayuga Co.; Chittenango, Madison Co.; and

Selkirk Shores St. Pk., Oswego Co. The major part of the study was per-

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Fig. 7. Level area of compact, sparsely-vegetated sand at Chittenango, Madison Co.,
N.Y. in which Lindenius columbianus errans nested.

formed during the summers of 1969-71 at a large commercial sand and gravel
pit near Auburn where a dense aggregation dotted a 3 X 14 ft ridge of firm
clayey-sand (see Miller and Kurczewski, 1973, Fig. 1). A shallow pond 50 ft
southwest of the ridge, annual herbs, grasses, and eastern cottonwood seedlings
bordered the nesting site. The nests at Chittenango were located in a flat
4 X 20 ft area of sparsely vegetated sand compacted by a payloader and
surrounded by a dense growth of annual herbs and grasses (Fig. 7). For
other characteristics of this site see Kurczewski, et al. (1969). At Selkirk
Shores, nests were found not only in the steep bank (Fig. 1) but also in a
level area of sand packed down intentionally by the senior author. The nests
at Sennett were scattered in tire ruts among the annual herbs and grasses
that had overgrown a recently bulldozed area of level sand.

Entrances to newly-constructed nests were surrounded by roughly circular
tumuli averaging 19 X 2.5 mm (12-26 X 1-4, N = 17). Rain and wind rap-
idly weathered these tumuli so that only vestiges remained around the 1.75-
2.00-mm-wide openings. In nests constructed in flat sand, the main burrow
descended vertically for 3.5-11 cm (x = 6.8, N = 40) and, in 31 (78%) of 40
nests, turned into a horizontal passage, 1-5 cm long (x = 2.2, N = 29) (see Figs.
8A-H). Bank nests, such as that depicted in Fig. 8H, were uncommon. The
ellipsoidal cells were arranged radially around the lower half of the vertical

Vol. LXXXIII, June, 1975

103

shaft (Fig. 8F) or, less commonly, around the apex of the horizontal burrow
(Fig. 8E). They averaged 2.8 X 6.8 mm (2. 5-3. 5 X 3. 5-8.0, N = 60) and
were separated from the main burrow by straight or winding side burrows,
1-5 cm long (Fig. 8B). Cell depths ranged from 3.5 to 11.5 cm (x = 6.7,
N = 131), the extreme means being 5.8 cm (4. 5-8.0, N = 11) at Sennett and

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6.9 cm (3.5-11.5, N = 101) at Auburn. This difference reflected the greater
compaction of the sand at Sennett.

The first cell provisioned in a nest was usually the shallowest (Figs. 8B,
C, E-H). Cell 3 was generally deeper than cell 2 but there was no discernible
pattern in the depths of cells beyond the third. Although as many as 10 cells
were found in a single nest, it was not possible to determine whether this
was the work of one female. For example, the nest in Fig. 8G was taken over
and provisioned for a day by a second female after the original resident was
removed. Cell 4, constructed by the new female, was at almost the same
depth as cell 1 made by the original female but was on the opposite side of
the main burrow.

Burrow storage of prey was observed at all 4 areas in 36 of 39 nests which
were being provisioned. Recently-captured prey were found half-way down
against the walls of the vertical shaft (Figs. 8A-C), near the juncture of the
vertical and horizontal burrows (Figs. 8B, C, E, F, H), or in the apical half
of the horizontal gallery (Figs. 8B, D, F). As indicated, several nests con-
tained prey in 2 or more storage locations in the burrow. Occasionally, a
loose plug of sand was found in front of prey stored in the horizontal passage
(Fig. 8F) .

The resident plugs the entrance and upper burrow with sand when she has
gathered enough prey for a cell. Before ovipositing, she positions the burrow
storage prey in a cell at the end of the horizontal burrow and then excavates
a new side passage. She may or may not excavate a cell at the end of this
passage before ovipositing. The passage is usually unmodified during pro-
visioning (Figs. 8A, B, D-H), but occasionally a completely-formed empty
cell, temporarily sealed off by a loose plug of sand, is found at the end of
the new burrow (Fig. 8C). The sand blocking the vertical burrow during cell
completion is apparently used to fill the passage leading to the completed cell
because, when the entrance is reopened, the female can be seen working her
mandibles against the walls of the burrow and pushing sand downward. The
entire process, from the time of entry with prey to reappearance at the surface,
required 45-90 min (N = 3).

Upon removing the sand plug from the entrance in the morning or after
completing a cell, the female exited and, while facing the entrance, made a
5-10 sec orientation consisting of increasingly longer, transverse flights. On
subsequent trips, except as indicated below, the wasp did not reorient but flew
directly away from the nest. The entrance remained open during her absence
except when a searching female entered and plugged the nest from inside.
After being disturbed by searchers or passing insects, the wasp usually made
a short orientation flight prior to hunting. Final nest closure was not observed
but it was noted that inactive nests always had the horizontal passage filled in.
The vertical burrows remained open until the first hard rain.

Vol. LXXXIII, June, 1975

105

Although some females dug their entire nests beginning from the sand sur-
face, it was not clear whether all females were able to do so. One female,
working in short spurts interspersed with 2-3 min rests, dug an L-shaped
burrow in 25 min. During interruptions, the wasp flew around the entrance
or landed on the sand nearby. She appeared to dig with the front legs in
unison, her body moving into the sand in a slow spiral. Upon completing
the nest, she plugged the entrance by pushing up columns of sand with her
abdomen. The entrance remained closed throughout the afternoon.

Some females began hunting as early as 0900 hrs on sunny days at Auburn,
but the majority were not active until 1030 hrs. Provisioning reached a peak
between 1100 and 1400 hrs, decreasing noticeably after 1430. A few nests
remained active all afternoon and, on occasion, provisioning continued until
1830 hrs. Females spent the night inside their closed nests while males rested
in vacant nests or abandoned burrows of other insects (see Miller and Kurc-
zewski, 1973). Nesting females were seldom active on damp or cloudy days.
Although females were collected from June to October, the most intensive
provisioning took place during July and August.

Females were observed hunting at Auburn in a patch of white sweet clover,
Melilotus alba Desvaux, growing at the edge of the nesting ridge. They hov-
ered slowly around the stems, maintaining their body in a horizontal position.
At Sennett, many females similarly circled umbels of Queen Anne’s lace,
Daucus carota (Linnaeus), occasionally darting at small objects. The flowers
were only a few feet from the nest entrances. One wasp caught a small fly
as it landed on an umbel, taking only 3 sec to sting it, position it, and fly
away. The prey were paralyzed and moved their legs or antennae when they
were removed from the cell.

Provisioning females returned to their nests in flight and dove rapidly into
their entrances (Fig. 9). On windy days wasps carrying heavy prey often
landed near their entrances until the wind subsided. In flight, large prey such
as Chironomus midges were held ventral-side-up and head-forward with the
wasp’s middle and hind legs. Small prey, especially Chalcidoidea, were held
obliquely with only the middle legs. On the ground, all prey were held with
the middle legs (Fig. 10). Some Chironomus were so large that they became
lodged in the entrance when the female dove in. From 2-5 sec were required
for the wasps to turn around and pull such flies inside. Abandoned prey were
common around nest entrances (see Miller and Kurczewski, 1973).

Before departing to hunt, the female usually spent several seconds in the
entrance looking out with the head or head and upper thorax exposed (Fig. 11).
The average time so spent was 8 sec (0-60, N = 77), the extremes in means
for 2 Auburn females being <1 sec (0-1, N = 24) and 22 sec (10-60, N = 7).
If a male or searching female interfered with the nesting female at this time,
she would back down into the nest or even plug the entrance with sand. The

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New York Entomological Society

female having the longest mean provisioning and prey storing times also spent
the longest mean time in the entrance. The resident with the shortest mean
provisioning time spent the shortest mean time in the entrance but did not
have the shortest mean storing time. After looking around the entrance, nest-
ing females of L. c. errans did not walk out on the sand but flew directly out
of the burrow.

The average time taken to capture a prey and return to the nest was 3.0
min (0.4-12.6, N = 93). The extremes in mean provisioning times for 2 Auburn
females were 7.2 (5.0-12.6, N = 9) and 1.5 min (0.4-3.4, N = 24). Provision-
ing times tended to vary more between different females than between successive
hunting trips of a single female. On several occasions, provisioning females
continually returned to hunt in the same vegetation. The average time spent
inside the nest between trips was 42 sec (5-250, N = 95). One Auburn female
took an average of only 22 sec (5-50, N = 13) to store a prey while another
from the same area required an average 71 sec (30-250, N = 10). The dif-
ferences may be attributed to the varying distances of burrow storage areas
from the entrance.

L. c. errans provisioned nests with 29 families of Diptera, Hymenoptera,
Hemiptera, and Homoptera. Table 6 lists the prey families and indicates their
relative importance as provisions in terms of numbers of individuals captured
and percent total catch. The species of prey and the areas from which they
were obtained are given in Table 7. The data from Lexington, Mass, and
Blackjack Creek, Kans. were provided by H. E. Evans. Overall, Diptera was
the order most commonly preyed upon, making up 29-90% of the total catch
depending on the area. Hymenoptera, comprising 9-59% of the prey, was more
important than Diptera as a source of provisions only at Chittenango, N.Y.
and Lexington, Mass. Nevertheless, more families and genera of Hymenoptera
were captured than Diptera. Hemiptera was regularly preyed upon in small
numbers, never exceeding 17% of the total catch, whereas Homoptera (Aphidi-
dae) was represented from only 2 areas.

Chironomidae was the most common prey family at Auburn, Selkirk Shores,
and Blackjack Creek. Chalcids of the family Pteromalidae were the main
source of provisions at Chittenango while Scatopsidae and Chloropidae (Dip-
tera) were the most important prey at Sennett and Lexington, respectively.
Other families making up 10% or more of the prey in an area during a given
season were, in decreasing order of their significance, Eulophidae, Ceratopo-

Fig. 9-11. Provisioning female of Lindenius columbianus errans diving into nest; 10.
holding prey with middle legs as she enters partly closed nest; and, 11. pausing in entrance
before leaving.

Vol. LXXXIII, June, 1975

107

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New York Entomological Society

Table 6. Families of Prey of Lindenius columbianus errans and
their relative importance as provisions.

Family

1969

Auburn

1970

Auburn

1971

Auburn

DIPTERA

448

79.29

850

76.85

196

76.87

Ceratopogonidae

1.24

104

9.40

9.02

Chironomidae

421

74.51

667

60.31

155

60.78

Sciaridae

0.18

Scatopsidae

0.09

0.39

Cecidomyiidae

0.18

1.81

1.96

Empididae

0.71

0.27

0.39

Chamaemyiidae

0.18

Milichiidae

Ephydridae

0.09

0.78

Chloropidae

1.24

2.35

0.78

Agromyzidae

1.41

2.17

2.75

HEMIPTERA

0.18

3.71

0.78

Anthocoridae

0.18

3.53

0.39

Miridae

0.09

0.39

Lygaeidae

0.09

HOMOPTERA

0.18

Aphididae

0.18

1970

1962

1971

Chittenango

Lexington

Sennett

Family

DIPTERA

105

29.17

40.74

170

84.58

Ceratopogonidae

1.39

21.39

Chironomidae

1.39

0.50

Sciaridae

1.00

Scatopsidae

4.72

103

51.24

Cecidomyiidae

3.89

7.41

1.49

Empididae

Chamaemyiidae

Milichiidae

0.28

1.49

Ephydridae

Chloropidae

16.67

33.33

7.46

Agromyzidae

0.83

HEMIPTERA

11.67

16.67

1.99

Anthocoridae

10.00

7.41

1.99

Miridae

1.67

9.26

Lygaeidae

HOMOPTERA

Aphididae

gonidae, Milichiidae, Cecidomyiidae, Eucharitidae, and Anthocoridae. Only 4
families, Cecidomyiidae, Braconidae, Eulophidae, and Anthocoridae were re-
corded as prey at all 6 areas.

It is significant that the prey families from Massachusetts and Kansas were
represented among the New York prey and that the relative importance of

Vol. LXXXIII, June, 1975

109

Table 6 (cont.). Families of Prey of Lindenius columbianus errans and
their relative importance as provisions.

Family

1971

Selkirk Shores

1952

Blackjack Cr.

Grand Totals

DIPTERA

219

89.75

84.62

2043

72.34

Ceratopogonidae

5.33

195

6.90

Chironomidae

40.16

58.97

1370

48.51

Sciaridae

0.41

0.18

Scatopsidae

0.41

123

4.36

Cecidomyiidae

18.03

15.38

3.43

Empididae

2.46

2.56

0.53

Chamaemyiidae

0.07

Milichiidae

18.85

7.69

1.88

Ephydridae

0.11

Chloropidae

4.10

138

4.89

Agromyzidae

1.49

HEMIPTERA

1.23

5.13

104

3.68

Anthocoridae

1.23

5.13

3.19

Miridae

0.46

Lygaeidae

0.04

HOMOPTERA

0.41

0.14

Aphididae

0.41

0.14

1969

1970

1971

Auburn

Family

HYMENOPTERA

115

20.35

213

19.26

22.34

Braconidae

0.71

6.06

9.02

Ichneumonidae

Mymaridae

0.18

0.09

Eulophidae

1.59

6.24

4.31

Encyrtidae

0.27

Eupelmidae

0.18

Eucharitidae

12.57

1.45

2.35

Torymidae

0.53

0.54

0.39

Pteromalidae

4.25

3.98

6.2 7

Eurytomidae

Chalcididae

Ceraphronidae

0.18

0.36

Formicidae

Sphecidae

0.18

0.09

Total prey

565

1106

255

the prey orders was not very different from that of certain New York areas.
Lexington, Mass, and Chittenango, N.Y., the 2 areas where Hymenoptera ex-
ceeded Diptera as a source of provisions, were remarkably alike in terms of
prey families captured and percent total catches, suggesting that these habitats
are rather similar. Likewise, the prey tallies from Blackjack Creek, Kans. and
Selkirk Shores, N.Y. were very similar. The 3 areas where Chironomidae

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New York Entomological Society

Table 6 (cont.). Families of Prey of Lindenius columbianus errans and
their relative importance as provisions.

Family

1970

Chittenango

1962

Lexington

1971

Sennett

HYMENOPTERA

213

59.17

42.59

13.43

Braconidae

6.67

1.85

1.49

Ichneumonidae

Mymaridae

Eulophidae

11.94

27.78

5.47

Encyrtidae

0.56

Eupelmidae

2.50

Eucharitidae

0.28

Torymidae

1.94

Pteromalidae

120

33.33

12.96

4.98

Eurytomidae

0.56

1.49

Chalcididae

Ceraphronidae

0.28

Formicidae

1.11

Sphecidae

Total prey

360

201

1971

1952

Selkirk Shores

Blackjack Cr.

Grand Totals

Family

HYMENOPTERA

8.61

10.25

673

23.83

Braconidae

1.64

2.56

127

4.50

Ichneumonidae

0.04

Mymaridae

0.04

Eulophidae

3.69

7.69

170

6.02

Encyrtidae

0.18

Eupelmidae

0.42

Eucharitidae

3.33

Torymidae

0.60

Pteromalidae

2.05

226

8.00

Eurytomidae

0.82

0.42

Chalcididae

0.41

0.04

Ceraphronidae

0.04

Formicidae

0.18

Sphecidae

0.04

Total prey

244

2824

formed the main

prey were near sizable bodies

of water.

Despite the great

differences in prey sample sizes during the 3 years of investigations at Auburn,
the relative importance of each of the prey orders remained very stable.
Non-specificity in prey selection was quantified by determining the numbers

of orders and families per nest and per cell. The average number of prey
orders per nest and per cell were, respectively, 2.4 (1-3, N = 39) and 2.1
(1-3, N = 64). Cells containing a single order of prey were found only at
Auburn and were rare. The average number of prey families per nest and

Vol. LXXXIII, June, 1975

111

Table 7. Prey of Lindenius columbianus errans.

Family

Species

Area

Ceratopogonidae

DIPTERA

Dasyhelea grisea (Coquillett)

A,C,D

Dasyhelea spp.

A,SX

Forcipomyia brevipennis (Macquart)

A,C,D,SX

Jenkinshelea magnipennis (Johannsen)

Chironomidae

Chironomus spp.

Cricotopus sp.

C,S

Orthocladius spp.

A,C,D2

Paratendipes subaequalis (Malloch)

Pentaneura sp.

Procladius spp.

A,D

Psectrocladius sp.

T any tarsus sp.

Sciaridae

Bradysia sp.

A,D,S

Scatopsidae

Swammerdamella obtusa Cook

A,C,D,SX

Swammerdamella sagittata Cook

Scatopse fuscipes Meigen

Cecidomyiidae

Anarete johnsoni (Felt)

A,C

Anarete pritchardi Kim

A,C

Anarete spp.

A,B,L,S

Aster omyia carbonifera (Osten Sacken)

Clinodiplosis sp.

A,C

Dasineura sp.

A,C

Mayetiola sp.

A,C

Neolasioptera spp.

A,S

Ozirhincus millefolii (Wachtl)

Porricondyla sp.

Procystiphora n. sp.

Empididae

Drapetis septentrionalis Melander

Drapetis sp.

Platypalpus trivialis Loew

Platypalpus sp.

Rhamphomyia sp.

Chamaemyiidae

Chamaemyia juncorum (Fallen)

Leucopis sp.

Milichiidae

Madiza parva (Adams)

Leptometopa halteralis (Coquillett)

D,SX

Leptometopa latipes (Meigen)

C,S

Paramyia nitens (Loew)

Ephydridae

Philygria debilis Loew

Hydrellia sp.

Chloropidae

C onioscinella melancholica (Becker)

A,D

C onioscinella minor (Adams)

L,S

C onioscinella triorbiculata (Sabrosky)

Diplotoxa versicolor (Loew)

Hippelates bishop pi Sabrosky

Hippelates n. sp. nr. bishoppi

Meromyza sp.

Area Code:

A Auburn, N. Y.

B = Blackjack Creek, Kans.

C = Chittenango, N. Y.

D = Selkirk Shores St. Pk., N. Y.

L = Lexington, Mass.

S = Sennett, N. Y.

1 Indicates species or genus made up 1-4% of total number of prey (2824) from all areas.

2 Indicates 4% or more.

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Table 7. (cont.)

F amily

Species

Area

Olcella cinerea (Loew)

Olcella parva (Adams)

Olcella quadrivittata Sabrosky

Olcella trigramma (Loew)

D,S

Oscinella carbonaria (Loew)

Oscinella frit (Linnaeus)

A,D,L

Oscinella luteiceps Sabrosky

A,L

Oscinella soror (Macquart)

A,S

Oscinella umbrosa (Loew)

Oscinella sp.

Siphonella nigripalpis (Malloch)

A,C,S

Agromyzidae

Agromyza sp.

Cerodontha ( Cerodontha ) dorsalis (Loew)

Cerodontha ( Cerodontha ) sp.

Liriomyza sp.

Ophiomyia sp.

Phytoliriomyza arctica (Lundbeck)

Pseudonap omyza lacteipennis (Malloch)

HEMIPTERA

Anthocoridae

Orius insidiosus (Say)

AjBjCjDjS1

Orius tristicolor (White)

A,D,L,S

Miridae

Chlamydatus associatus (Uhler)

C,L

undetermined nymphs

Lygaeidae

undetermined nymph

HOMOPTERA

Aphididae

Aphis sp.

Capitophorus elaeagni (Del Guer.)

Rhopalosiphum maidis (Fitch)

Schizaphis sp.

HYMENOPTERA

Braconidae

A gat his spp.

Apanteles limentidis Riley

Apanteles xylinus (Say)

Apanteles spp.

A^CAS1

Aphidius obscuripes Ashmead

Aphidius spp.

A,C,V

Bracon sp.

Chelonus {Micro chelonus) sp.

Dacnusa sp.

Diaeretiella spp.

Elasmosoma sp.

Euphoriana uniformis Gahan

Lysaphidus sp.

Lysiphlebus spp.

A,C

Microplitis sp.

Orgilus gelechiae (Ashmead)

Orgilus sp.

Praon spp.

Trioxys spp.

AjC1

Ichneumonidae

Adelognathus flavopictus Davis

Mymaridae

Polynema sp.

Eulophidae

Aprostocetus sp.

Chrysocharis sp.

Vol. L XXXIII, June, 1975

113

Table 7. (cont.)

Family

Species

Area

Diaulinopsis callichroma Crawford

Euderus subopacus (Gahan)

Euderus sp.

A,C

Hemiptarsenus americanus (Girault)

Hyssopus novus Girault

Necremnus sp.

N otanisomorpha ainsliei Crawford

Pnigalio sp.

A,S

Sympiesis bimaculatipennis (Girault)

Tetrastichus bruchophagi Gahan

A,D,S

Tetrastichus chlamytis Ashmead

A,C

Tetrastichus fumipennis (Girault)

Tetrastichus incertus (Ratzeburg)

Tetrastichus semilongifasciatus (Girault)

Tetrastichus tesserus Burks

Tetrastichus spp.

A,C,D,L,S2

Entedontini

Encyrtidae

Copidosoma sp.

A,C

Anagyrini

Bothriothoracini

Eupelmidae

Eupelmella vesicularis (Retzius)

Eupelmus allynii (French)

Eupelmus sp.

A,C

Eucharitidae

Pseudometagea schwarzii (Ashmead)

A,CX

Torymidae

Eridontomerus isosomatis (Riley)

Pseudotorymus lazulellus (Ashmead)

A,C

Pteromalidae

Asaphes lucens (Provancher)

A,C

Ecrizotes sp.

Erixestus winnemanna Crawford

Habrocytus sp.

C,D,L

Halticoptera patellanna (Dalman)

Halticoptera sp.

A,C

Heteroschema sp.

Homoporus chalcidiphagus (Walsh & Riley)

AjCjLjS1

Homoporus febriculosus (Girault)

A,C

Mesopolobus sp.

C,L,S

Pachyneuron allograptae Ashmead

Pachyneuron siphonophorae (Ashmead)

A,C

Pachyneuron sp.

Parecrizotes marylandenis Girault

Pteromalus puparum (Linnaeus)

Pteromalus vanessae Harris

Sy stasis sp.

Tridymus sp.

A,D

Pirenini

A,S

Pteromalini

Tridymini

Eurytomidae

Bruchophagus sp.

Eudecatoma sp.

D,S

Eurytoma sp.

A,C

Harmolita sp.

A,D

Systole sp.

Chalcididae

Spilochalcis albifrons (Walsh)

Ceraphronidae

Lygocerus sp.

Formicidae

Lasius sp. ( $ $ )

A,C

Sphecidae

Spilomena pusilla (Say)

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per cell were 6.9 (1-13, N = 39) and 4.6 (1-13, N = 64), while the average
number of prey genera per cell was 6.0 (2-15, N = 60). One cell from Selkirk
containing 15 genera, 13 families and 3 orders of prey represented the extreme
in non-specificity. The other extreme was exemplified by a 2-celled nest at
Auburn containing only 2 genera of chironomids, mostly of a single species.

Selection of a particular sex of prey was marked in only a few instances.
At Auburn, 98% of the Tany tarsus midges and 88% of the ceratopogonid
Jenkinshelea magnipennis Johannsen were males. At Selkirk Shores, 100% of
the cecidomyiid Procystiphora n. sp. and 95% of an undetermined chironomid
were females. All prey were invariably smaller and lighter than their captors,
weighing an average of 0.24 mg (.05-1.05, N = 1100) compared to 2.90 mg
(1.25-4.15, N = 65) for the wasps. Minute ceratopogonids of the genus Dasy-
helea were the smallest prey, whereas the largest were females of the genera
Chironomus (Chironomidae) and Forcipomyia (Ceratopogonidae) . The weight
of the provisions in a fully-provisioned cell ranged from 2.0 to 13.3 mg (x = 7.2,
N = 33). From 8 to 76 prey (x — 25.5, N = 77) were stored per cell.

Large prey were stored head-inward in the cell, facing away from the
burrow, whereas tiny prey were stacked more randomly, their bodies often
turned obliquely or backwards. The egg-bearer was 1 of the innermost prey
and was usually ventral-side-up. The off-white, slightly curved egg was 1.4-1. 6
mm long and 0.3-0. 4 mm wide in the middle. Its larger cephalic end was
attached to the neck of the prey along the ventral midline. The direction in
which the egg extended varied with the prey. Eggs on chironomids and
ceratopogonids extended transversely across the prosternum at nearly right
angles to the body of the prey (Fig. 12). Eggs were more obliquely -placed on
chloropids and agromyzids and were distinctly longitudinally-placed on small
pteromalids and eulophids. The caudal end was normally free but occasionally
appeared tightly fastened to the spread wing of the prey. The egg-bearer was
usually a common prey in the nest, regardless of size. For example, egg-
bearing Hemiptera and Homoptera were not found.

The egg hatched in 1-2 days and the larva grew to a length of 3 mm during
the first 3-4 days. At this time the larva was relatively slender and had
consumed only a single prey. During the next 2 days it increased rapidly in
girth and devoured all of the provisions, facing away from the main burrow
and pushing the discarded sclerites into a compact mass at the other end.
Within 7-9 days after the egg was laid the larva had spun a cocoon of silk
and sand, distributing the excrement and sclerotized prey remains evenly over
the surface. The resultant cocoons were brown, ellipsoidal, and averaged 2.3
X 5.8 mm (2-3 X 4-7, N = 21). The species overwinters in the cocoon as a
diapausing larva, pupates in late spring, and the adults begin to emerge in
early summer. Although pupae and teneral adults were dug up in early Au-

Vol. LXXXIII, June, 1975

115

Fig. 12. Egg of Lindenius columbianus errans on male F orcipomyia brevipennis (Mac-
quart) (Ceratopogonidae).

gust, it is uncertain whether they represent a second generation or late emerg-
ing individuals from cells provisioned the year before.

DISCUSSION

Viewed collectively the behavioral features possessed by species of Lindenius
distinguish this genus from other sphecid genera, although these features may
be modified extensively when more of the 48 described species are studied.
The 6 species examined in this paper construct nests 3-12 cm deep in sand,
fine gravel, loess, or chalk. They prefer hard-packed ground such as garden
paths, roads, and compacted areas. The nests have a vertical or slightly in-

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dined burrow which often leads to a short horizontal passage whose distal end
marks the location of the next cell.

The sequence of nest orientation behaviors is essentially identical for L.
albilabris, L. armaticeps , and L. c. errans. The females orient in flight, facing
the entrance, before the first provisioning trip but not prior to subsequent
trips unless the entrance has been obstructed on the return flight. Normally,
the nests of all 6 species remain open during provisioning trips. In aggrega-
tions of L. c. errans , however, searching conspecific females may enter a nest
and plug the entrance with sand, creating difficulties for the provisioning
female when she returns. Also, the resident may plug the entrance from
within if searchers interfere with her when she is preparing to exit. Otherwise,
the entrances are closed only at night and when the females are excavating
new passages and cells, ovipositing, or filling burrows.

The 6 species of Lindenius store adult insects, but L. albilabris and L. c.
errans sometimes capture nymphal Hemiptera. Diptera is the only order
preyed upon by all. In each species, the egg is not laid until the full com-
plement of prey for a cell has been gathered. The heads of most of the prey
face in the same direction and the egg is attached to 1 of the first prey placed
in the cell. Egg placement is similar in all species, the egg being fastened to
the neck of a prey along the ventral midline.

L. albilabris , L. armaticeps , L. c. errans , and L. panzeri develop from egg
to mature larva in 5-8 days. All 6 species spin cocoons of silk and sand and
distribute the sclerotized remains of the prey evenly over the surface. Cocoons
of the mirid-hunter L. albilabris occasionally lack prey remains as a result of
utilizing nymphal prey (Bonelli, 1967).

Other behavioral features have potential value in separating species or
species groups. In nests of L. armaticeps and L. buccadentis, the oldest cells
were the shallowest and the newest cells, the deepest (Figs. 2, 6). The nests
of L. c. errans were more irregular, with the first cell usually being the shal-
lowest but the third and fourth being located either above or below the second
(Figs. 8B-H). A similar lack of uniformity in cell placement was reported
for L. albilabris by Minkiewicz (1931, 1933), who found an incompletely pro-
visioned cell to be the shallowest in 1 nest but the deepest in another. Bouw-
man (1911) schematically illustrated a 9-celled nest of L. panzeri in which a
lower level of cells was constructed before an upper one. If the latter arrange-
ment is characteristic of this species, it is one of the few clearcut differences
between L. panzeri and its Nearctic relative L. armaticeps.

Transport of prey with the middle legs is probably the common method in
Lindenius. Many of the conflicting reports on prey transport in L. albilabris
and L. panzeri may be attributed to capturing prey-laden wasps in boxes or
vials and expecting them to show normal prey transport behavior. For ex-

Vol. LXXXIII, June, 1975

117

ample, Crossocerus maculiclypeus (Fox) always carries its prey with the mid-
dle legs but, when enclosed in a vial, sometimes impales its prey on the sting
and walks around on all 6 legs. Other conflicting reports may result from the
investigator tampering with the nest entrance in an attempt to slow down
the wasp for closer observation. L. armaticeps normally carries its prey with
the middle legs but will often impale the prey on the sting at an obstructed
nest entrance. In contrast, L. c. errans was never observed impaling its prey
at an obstructed entrance during 3 summers of extensive observation.

A comparison of mean provisioning times and number of prey per cell may
be of value in separating species. On the average, L. armaticeps stores fewer
prey per cell than the other species and has the longest mean provisioning time.
The mean numbers of prey per cell for both L. buccadentis and L. c. errans
are well over twice that for L. armaticeps , and their mean provisioning times
are well under half that of L. armaticeps. However, this inverse relationship
between number of prey per cell and provisioning time is not perfect. L.
buccadentis stores, on the average, more prey per cell than L. c. errans, yet
the former species has a longer mean provisioning time.

A possible group difference in prey storage behavior is strengthened by
a comparison of prey-storing times for L. albilabris, L. armaticeps, L. bucca-
dentis, and L. c. errans. The last 3 species usually store newly-captured prey
head-inward in an unmodified part of the burrow whereas L. albilabris is re-
ported to place its prey directly in a cell at the end of a side passage. Per-
haps, as a result, the average storing time is longer in L. albilabris than in

L. armaticeps, L. buccadentis, or L. c. errans. The last species often releases

its prey only part way down the vertical burrow so that a “line” of prey is
gradually formed. This type of storage has not been reported for the other
species.

After storing the prey and returning to the surface, L. armaticeps, L. bucca-
dentis, and L. c. errans often spend several seconds in the entrance looking

around. L. c. errans typically flies out of the burrow directly from the head-

in-entrance position, whereas L. armaticeps, in both sloping and horizontal
sand, walks out and then flies away. L. buccadentis is intermediate in this
respect because the same individual may walk out of the entrance on one trip
and fly out on the next.

Weights of cell contents and of individual prey are available only for the 3
Nearctic species. The average weight of the contents of 19 cells of L. armati-
ceps was 12.1 mg, whereas that of 33 cells of L. c. errans was only 7.2 mg.
Two cells of L. buccadentis were similar in weight to an average cell of L. c.
errans. The average weight of a single prey of L. armaticeps was about 4 times
that of either L. c. errans or L. buccadentis . Only the first species was observed
carrying prey heavier than itself. L. albilabris is the largest species yet studied

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and undoubtedly has greater mean cell and individual prey weights than any
of the others.

One may readily distinguish the 6 species by the kinds and proportions
of prey stored. The Palaearctic L . albilabris is a mirid-hunter which at times
stores empidid, dolichopodid, and chloropid flies. Both the Nearctic L. ar-
maticeps and the Palaearctic L. panzeri are chloropid-hunters, the latter occa-
sionally capturing simuliid, tephritid, and milichiid flies. The Nearctic L.
buccadentis and the Palaearctic L. pygmaeus store mainly chalcidoids and
braconids but occasionally hunt flies. The Nearctic L. c. errans is the most
polyphagous species yet studied, preying on 11 families of Diptera, 14 of
Hymenoptera, 3 of Hemiptera, and 1 of Homoptera. Overall, Diptera was
the most important food source of this species. Although the specific dif-
ferences in degree of polyphagy are quite useful, individual females of a given
species may be very specific in their choice of prey. Instances approaching
host-specificity can be cited for all but L. pygmaeus whose nesting behavior
has not been investigated thoroughly.

The use of Hymenoptera as provisions by 3 species of Lindenius deserves
special attention because the behavior is not common in the Crabroninae.
Encopognathus and Tracheliodes are soil-nesting ant hunters, the latter prey-
ing specifically on workers of the genera Liometopum and Tapinoma (Muese-
beck, et al., 1951). Krombein (1958) noted an undescribed species of Crosso-
cerus from Utah nesting in twigs and storing chalcidids, while Hamm and
Richards (1926) found a tenthredinid as exceptional prey of the wood-nester
Crossocerus leucostomoides (Richards). Janvier (1928) reported winged ants
among the dipterous and lepidopterous prey of the sand-nester Euplilis rujo-
taeniatum (Kohl). Thus, L. buccadentis , L. c. errans , and L. pygmaeus are
the only crabronines known to prey on diverse groups of Hymenoptera.

In summary, the 6 species of Lindenius exhibit similar behavior in con-
structing nests with vertical burrows in compact sand or sandy-gravel, leaving
the entrance open during provisioning trips, diving into the entrance with
prey in flight, including Diptera among the provisions (but not necessarily in
each nest or study area), ovipositing only after the full complement of prey
has been gathered, affixing the egg to the neck of a prey along the ventral
midline, and distributing the prey remains evenly over the surface of the
cocoon in a matrix of silk and sand. Collectively, these features distinguish
Lindenius from all other sphecid genera.

The 6 species differ most clearly in the kinds and proportions of prey.
Other facets of behavior useful in separating the species but generally more
difficult to measure include sequence/ depth of cell placement, method of prey
transport at obstructed entrances, provisioning and storing times, number of
prey per cell, type of prey storage, exiting behavior, stages of prey, weights

Vol. LXXXIII, June, 1975

119

of cell contents and individual prey, frequency of discarded prey, male be-
havior, and kinds of intraspecific interactions. As the remaining 42 species
are investigated, these other features must be relied upon increasingly. The
inevitable overlap in prey families, already so conspicuous between L. armati-
ceps and L. panzeri, indicates the importance of having the prey thoroughly
identified as well as the precariousness of depending solely on prey differences
to distinguish species.

Literature Cited

Abrahamsen, S. E. 1950. Gravehvepsen Crabro Lindenius panzeri v.d. Lind. Redekolonier
fundet i Danmark. Flora og Fauna 56: 125-130.

Adlerz, G. 1903. Lefnadsforhallanden och Instinkter inom Familjerna Pompilidae och
Sphegidae. I. K. Svenska Vet.-Ak. Handl. 37(5): 1-181.

. 1910. Same title. III. K. Svenska Vet.-Ak. Handl. 45(12): 1-75.

Bluthgen, P. 1955. Zur Biologie von Lindenius albilabris (F.) (Hym., Sphecidae).
Deutsche Ent. Zeitschr. (N.F.) 2: 158.

Bonelli, B. 1967. Osservazioni biologiche sugli Imenotteri melliferi e predatori della Val
di Fiemme. XXV. Boll. 1st. Ent. Bologna 28: 291-303.

Bouwman, B. E. 1911. Crabro’s. Levende Natuur 16: 121-126, 173-177, 199-204.
Bristowe, W. S. 1948. Notes on the habits and prey of twenty species of British hunting
wasps. Proc. Linn. Soc. London 160: 12-37.

Chambers, V. H. 1949. The Hymenoptera Aculeata of Bedfordshire. Trans. Soc. Brit.
Ent. 9(4) : 197-252.

Court, H. K. and R. M. Bohart. 1958. New species of Lindenius from western North
America (Hymenoptera: Sphecidae). Pan-Pacif. Ent. 34(3): 161-167.

De Beaumont, J. 1956. Notes sur les Lindenius palearctiques (Hym. Sphecid.) Mitt.
Schweiz. Ent. Ges. 29(2) : 145-185.

Dupuis, C. 1947. Les Proies des Sphegides chasseurs d’Heteropteres. Feuille Nat. N.S.
2(1947): 111-113.

Evans, H. E. 1963. Predatory wasps. Sci. Am. 208 (4) : 145-154.

. 1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming.

Bull. Mus. Comp. Zool. 140(7): 451-511.

Ferton, C. 1901. Notes detachees sur l’instinct des Hymenopteres melliferes et ravisseurs
avec la description de quelques especes. (1° serie). Ann. Soc. Ent. Fr. 70: 83-148.
Grandi, G. 1928. Contributi alia conoscenza biologica e morfologica degli Imenotteri
melliferi e predatori. VII. Bull. Lab. Ent. Bologna 1: 259-326.

. 1961. Studi di un entomologo sugli imenotteri superiori. Boll. 1st. Ent. Univ.

Bologna 25: 1-659.

Gronblom, T. 1925. Bidrag till kannedom om Levnadssattet hos vara rovsteklar (Hy-
menopt., Sphegidae). I. Notulae Ent. 5: 1-9.

Guichard, K. M. and I. H. H. Yarrow. 1947. The Hymenoptera aculeata of Hampstead
Heath and the surrounding district, 1832-1947. London Nat. (1947): 81-111.
Hamm, A. H. and O. W. Richards. 1926. The biology of the British Crabronidae. Trans.
Ent. Soc. London 74: 297-331.

Hertzog, L. 1954. Hymenopteres predateurs et melliferes de Camargue. Terre et Vie
101(1): 95-110.

Iwata, K. 1942. Comparative studies on the habits of solitary wasps. Tenthredo 4:
1-146.

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Janvier, H. 1928. Recherches biologiques sur les predateurs du Chili. Ann. Sci. Nat.
Zool. 11(10): 67-207.

Kohl, F. F. 1915. Die Crabronen (Hymenopt.) der palaarktischen Region. Ann. K. K.
Nat. Hofmus. 29: 1-453.

Krombein, K. V. 1958. Hymenoptera of America north of Mexico. Synoptic catalog.

First supplement. U.S.D.A., Agri. Mono. 2: 1-305.

Krombein, K. V. and B. D. Burks. 1967. Hymenoptera of America north of Mexico.

Synoptic catalog. Second supplement. U.S.D.A., Agric. Mono. 2: 1-584.
Kurczewski, F. E., N. A. Burdick and G. C. Gaumer. 1969. Observations on the nest-
ing behavior of Crossocerus (C.) maculiclypeus (Fox) (Hymenoptera: Sphecidae).
J.N.Y. Ent. Soc. 77(2) : 92-104.

Leclercq, J. 1954. Monographie systematique, phylogenetique et zoogeographique des
Hymenopteres Crabroniens. These, Fac. Sci. Univ. Liege. 371 p.

. 1959. Lindenius ceballosi sp. nov., Crabronien nouveau d’Espagne. Eos. 35:

267-268.

. 1960. Crabroniens d’Espagne appartenant aux genres Crabro , Lindenius et

Entomognathus (Hym. Crabronidae) . Eos. 36: 417-426.

Maneval, H. 1937. Notes sur les Hymenopteres (5° serie). Rev. Fr. Ent. 4(3): 162-181.
Marchal, P. 1893. Observations biologiques sur les Crabronides. Ann. Soc. Ent. Fr. 62:
331-338.

Miller, R. C. and F. E. Kurczewski. 1973. Intraspecific interactions in aggregations of
Lindenius (Hymenoptera: Sphecidae, Crabroninae) . Insectes Sociaux 20(4): 365-
378.

Minkiewicz, R. 1931. Nids et proies des Sphegiens de Pologne. Fragments ethologiques
(1° serie). Polskie Pismo Ent. 10: 196-218.

. 1932. Same title. (2° serie). Polskie Pismo Ent. 11: 98-112.

. 1933. Same title. (3° serie). Polskie Pismo Ent. 12: 181-261.

Muesebeck, C. F. W., K. V. Krombein and H. K. Townes. 1951. Hymenoptera of
America north of Mexico. Synoptic catalog. U.S.D.A., Agri. Mono. 2 : 1-1420.
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Olberg, G. 1959. Das Verhalten der Solitaren Wespen Mitteleuropas. Berlin, VEB Deutsch.
Verb Wiss. 402 p.

Sickmann, F. 1893. Die Hymenopteren-fauna von Iburg und seiner nachsten Umgebung,
mit biologischen und kritischen Bemerkungen. I. Abteilung die Grab wespen. Jahresber.
Nat. Ver. Osnabriick 9: 39-112.

Vol. LXXXIII, June, 1975

121

New or Little-Known Crane Flies from Iran. Ill
(Diptera: Tipuliclae)1

Charles P. Alexander
Amherst, Massachusetts 01002

Received for Publication July 23 , 1974

Abstract: Part II of this series of papers concerning the crane flies of Iran was published
in this Journal (82: 279-284, 1974). In that report various species of the Eriopterine genus
Gonomyia were considered and in the present paper further new species and records in the
Eriopterini are provided. The species here described are Lipsothrix iranica, Cheilotrichia
( Empeda ) gnoma, Erioptera ( Pseuderioptera ) schmidi, E. ( Psiloconopa ) cancriformis,
and Molophilus ( Molophilus ) pallidipes, all from the Elburz Mountains in northern Iran.
Additional to the above novelties, 13 further previously described European species are
added to the list of species of Tipulidae from Iran.

In the preceding two reports on the crane flies of Iran that were collected by
Dr. Fernand Schmid in 1955 and 1956 a portion of the species belonging to the
tribes Pediciini and Eriopterini were treated. At this time I am discussing the
remaining members of the Eriopterini contained in the collection and supplying
several records of previously described species hitherto known from Europe.
I again extend my deepest thanks to Dr. Schmid for his work in collecting this
valuable series of crane flies from a scarcely known area of southern Asia.

One of Schmid’s important papers on the Trichoptera of Iran provides full
information concerning the various stations in the Elburz Mountains where the
present series of flies was taken and should be consulted (Trichopteres d’lran.
Beitrage zur Entomologie, 9: 200-219, 376-389; 1959). This paper includes a
map showing itinerary and collecting stations, and complete geographical data
for this expedition, September 1955 and April to October 1956, with eight
photographs showing especially important collecting localities.

Lipsothrix iranica, n. sp.

Mesothorax orange, pronotum yellow, narrowly brownish black medially; legs yellow,
femoral tips narrowly black, tibiae yellow, extreme bases and tips darkened; wings pale
yellow, stigma dark brown, conspicuous, vein R2 + 3 + 4 short and straight, longer than the
strongly arcuated basal section of R:, ; abdomen yellowed, patterned with black, outer two
segments yellow.

Female. Length about 11 mm.; wing 9.5 mm.; antenna about 1.8 mm.

Rostrum orange ; palpi brownish black, unusually long, nearly one-half the antennae ;
terminal segment about one-third longer than the more slender third segment. Antennae

1 Contribution from the Entomological Laboratory, University of Massachusetts.
New York Entomological Society, LXXXIII: 121-128. June, 1975.

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with scape and pedicel brownish yellow, flagellum light yellow, outer segments pale brown;
segments long-oval with a circlet of about six black setae that are shorter than the segment.
Head light brown ; posterior vertex with long black setae.

Pronotum yellow, anteriorly narrowly brownish black. Mesothorax almost uniformly
orange, without dark pattern. Halteres with stem whitened, knob slightly more yellowed.
Legs with coxae orange; trochanters yellow; femora yellow, tips abruptly brownish black,
including about the outer twelfth of segment; tibiae yellow, extreme bases and tips dark
brown; tarsi yellow, outer two segments light brown. Wings (Fig. 1) pale yellow, prearcular
and costal fields clearer yellow; stigma dark brown, conspicuous; veins of base and costal
region yellowed, remaining veins brown. Longitudinal veins beyond general level of origin
of Rs and cord with strong trichia, lacking on both Anals except for a very few at tips.
Venation: Rs straight, R2+3+i short and straight, slightly longer than the strongly arcuated
basal section of R-,; veins beyond cord straight, generally parallel.

Basal abdominal segment yellowed, tergites two to six obscure yellow medially, lateral and
posterior borders more blackened, seventh segment black; sternites yellowed medially,
blackened on sides, especially posteriorly, seventh sternite black, remainder, including
ovipositor, yellow.

Holotype. $, Ardehjan, Iran, September 11, 1956 (Schmid).

The most similar European species that have the apices of the femora blackened and the
stigma of the wing dark are Lipsothrix nobilis Loew, L. nervosa Edwards and L. nigristigma
Edwards, all with the thoracic dorsum conspicuously patterned with black. Of the above,
nervosa has the darkened wing pattern somewhat less conspicuous, including the stigma,
differing from the present fly in other characters, including the venation, the longitudinal
veins beyond the cord being much shorter, with vein i?2+3 + 4 long, about two-thirds Rs.

Cheilotrichia ( Empeda ) gnoma, n. sp.

Size very small (wing about 3-3.5 mm.); head and thorax dark gray; halteres yellow;
legs brown; wings faintly tinted, stigma scarcely indicated; cell Rs small, triangular in
outline, cell 1st M2 closed; male hypopygium with both dististyles uniformly pale, outer style
bifid, both arms expanded outwardly, inner style a long slender pale rod.

Male. Length about 3-3.3 mm.; wing 2. 8-3.4 mm.

Female. Length about 3.4-3. 7 mm.; wing 3-3.2 mm.

Rostrum and palpi black. Antennae black; pedicel much enlarged, verticils of basal
flagellar segments very long. Head dark gray.

Thorax almost uniformly dark gray, praescutal stripes slightly darker. Halteres yellow.
Legs with coxae and trochanters light brown; remainder of legs brown. Wings (Fig. 2)
faintly tinted, stigmal darkening scarcely indicated; veins pale brown. Longitudinal veins
beyond general level of origin of Rs with small trichia, including also about the outer half
of 2nd A. Venation: Sci ending about opposite one-third to one-half Rs, Sci about four to
five times Sc2 ; cell R?. triangular in outline, vein Rs oblique, straight or nearly so; cell 1st M2
closed; m-cu shortly beyond fork of M.

Abdomen brown, pleural region slightly darker. Male hypopygium (Fig. 5) with both
dististyles, including the vestiture, pale; outer style large, bifid, the arms longer than the
base, inner blade more cleaver-shaped, as shown, outer arm more oval; inner style about
four-fifths as long, appearing as a long, very slender pale rod. Phallosome, p, about as
figured, the aedeagus with an erect lateral darkened lobe near apex.

Holotype. $, Ardehjan, Iran, September 11, 1956 (Schmid).

Vol. LXXXIII, June, 1975

123

Allotopotype. $ . Paratopotypes. 14 $ $ , on five pins.

The present fly appears certainly to belong to Empeda despite the venation which is very
similar to that of Gonempeda flava (Schummel) and certain species in the typical subgenus
Cheilotrichia. However the structure of the male hypopygium, including the dististyles, are
much as in Empeda and I consider the reference to this subgenus to be correct. The fly is
readily told by the very small size, venation, and in hypopygial details.

Erioptera {Pseuderioptera) , n. subgen.

Wing (Fig. 3) with vein R2 before the outer radial fork, leaving an element R2 + i; cell
1st Mo closed; vein 2nd A with a low terminal bend. Trichia of wing veins very short and
sparse, including the costal fringe; marginal setae of proximal two-thirds of posterior wing
margin long and delicate; legs with elongate pale scales additional to the normal setae. Male
hypopygium (Fig. 6) with both dististyles simple, subterminal; gonapophyses appearing as
flattened paddlelike blades.

Type of subgenus. Erioptera ( Pseuderioptera ) schmidi, n. sp.

Other subgenera of Erioptera having interpolated scales on the legs include Meterioptera
Alexander, Tasiocerodes Alexander, and Teleneura Alexander, all having the venational de-
tails and wing trichiation distinct.

Erioptera ( Pseuderioptera ) schmidi, n. sp.

Mesonotal praescutum yellow with a cinnamon brown median stripe, posterior sclerites of
mesonotum and the pleura chiefly light yellow; femora yellow, tips brownish black; wings
pale yellow with a very restricted pale brown pattern that includes the cord and apices of
outer radial veins; vestiture of veins unusually short, including the costal fringe, lacking on
nearly the basal third of wing; R2 before fork of cell i?3, cell 1st M2 present; abdominal
tergites light brown basally, yellowed posteriorly ; male hypopygium with two simple
dististyles; gonapophyses appearing as flattened blades, apices with microscopic spines.

Male. Length about 4 mm.; wing 4 mm.; antenna about 0.75 mm.

Rostrum and palpi light yellow. Antennae with scape and pedicel brownish black to
black, flagellum light brown ; flagellar segments oval, progressively smaller outwardly,
verticils subequal in length to the segments. Front and anterior vertex silvery white, posterior
vertex abruptly light brown ; anterior vertex broad.

Pronotum light yellow, scutellum narrowly more darkened medially. Mesonotal praescutum
with a cinnamon brown central stripe that ends some distance before suture, lateral stripes
short and narrow, sides broadly light yellow; scutum light yellow, lobes chiefly cinnamon
brown; scutellum and anterior mediotergite light yellow, posterior parts light brown. Pleura
light yellow. Halteres yellow. Legs with coxae and trochanters light yellow; femora yellow,
the relatively broad tips brownish black; remainder of legs yellow, outer tarsal segments
darkened; legs with long narrow interpolated pale scales among the normal setae. Wings
(Fig. 3) pale yellow, with very small and inconspicuous pale brown spots at Sc2, Ri, R2+3,
Rs and cord; veins yellow, darker in the patterned areas. Venation: Sc2 far retracted, Sci
about opposite two-thirds Rs; R2 before the radial fork, R?J + 4 variable in length, in the
holotype longer than R2, shorter in the paratype ; m-cu before fork of M ; vein 2nd A con-
spicuously sinuous on outer fifth. Vestiture of veins unusually short, including the costal
fringe; trichia of veins short and inconspicuous as compared with the normal condition in
Erioptera , lacking on veins of about the basal third of wing.

Abdominal tergites bicolored, basally light brown, lateral and posterior borders light yellow,

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Fig. 1. Lipsothrix iranica, n. sp. ; venation.

Fig. 2. Cheilotrichia ( Empeda ) gnoma, n. sp.; venation.

Fig. 3. Erioptera ( Pseuderioptera ) schmidi, n. sp.; venation.

Fig. 4. Erioptera ( Psiloconopa ) cancrif ormis , n. sp.; venation.

Fig. 5. Cheilotrichia ( Empeda ) gnoma , n. sp.; male hypopygium.

Fig. 6. Erioptera ( Pseuderioptera ) schmidi , n. sp.; male hypopygium.

Fig. 7. Erioptera ( Psiloconopa ) iranica Alexander; male hypopygium.

Fig. 8. Erioptera ( Psiloconopa ) cancrif ormis, n. sp.; male hypopygium.

Fig. 9. Molophilus ( Molophilus ) pallidipes , n. sp.; male hypopygium.

(Symbols: Male hypopygium — a, aedeagus; b, basistyle; d, dististyles; g, gonapophysis ;
p , phallosome; t, 9th tergite.)

Vol. LXXXIII, June, 1975

125

sternites more uniformly yellow. Male hypopygium (Fig. 6) with the simple dististyles
subterminal, apex of basistyle, b, short, narrowly obtuse, with long yellow setae, outer face
of style subglabrous, mesal face with abundant shorter pale setae. Outer dististyle, d, a
nearly straight slender rod, apex blackened; inner style subequal in length, appearing as a
slightly curved flattened yellow blade, the apex a short acute point. Gonapophyses, g, ap-
pearing as a pair of flattened blades, apices with a row of microscopic spines; aedeagus, a,
divided into paired rods, tips recurved into points.

Holotype. $, on slide, Dashte Maghan, Iran, September 29, 1956 (Schmid).

Paratopotype. Broken $ , with the type.

This distinct fly is named for the collector of this fine series of Iranian Tipulidae, Dr.
Fernand Schmid. It is readily separated from other generally similar members of the genus
by the subgeneric characters as listed above, especially the retracted vein R2 and the
hypopygial structure.

Erioptera ( Psiloconopa ) cancriformis, n. sp.

General coloration of thorax light yellow, patterned with brown, pleura with a very
narrow brown central stripe; knobs of halteres brown; wings whitened, without a stigmal
darkening, veins light brown, Sc white ; R2 about one-half its length beyond the basal fork
of Rs, cell 1st M2 closed; male hypopygium with outer dististyle bilobed, the lobes blackened
and pointed, together suggesting a crabs claw; gonapophyses appearing as slender blackened
rods.

Male. Length about 5 mm.; wing 4.2 mm.

Female. Length about 5.5-6 mm.; wing 5-5.2 mm.

Rostrum light brown ; palpi black. Antennae light brown ; flagellar segments oval, verticils
short. Head buffy yellow, vertex more darkened medially, more intensely on anterior vertex.

Prothorax light yellow. Mesonotal praescutum very light brown, darker medially, with
a still darker central vitta, humeral region light yellow; scutum brown, narrowly more
darkened medially, the outer parts of lobes more diffusely darkened; scutellum light yellow,
in male with a narrow darker central line. Pleura light yellow, ventral sternopleurite light
brownish gray, central area of pleura with a very narrow brown line extending from base
of fore coxa to beneath the root of haltere. Halteres with stem yellow, knob brown. Legs
with coxae and trochanters light yellow ; femora and tibiae obscure yellow, apices pale brown ;
tarsi light brown. Wings (Fig. 4) whitened, without a stigmal darkening; veins light brown,
Sc whitened. Venation: Sci ending about opposite or slightly before R2 , Sci subequal to
Rs; R2 about one-half its length beyond the radial fork; cell 1st M2 closed; m-cu at or
shortly before fork of M ; vein 2nd A virtually straight to slightly extended on distal fifth.

Abdomen yellow, tergites striped longitudinally with dark brown, posterior borders of
segments narrowly yellow, sides more broadly so. Male hypopygium (Fig. 8) with the
tergite, t, having the posterior border produced into two small triangular lobes, subequal in
size to the median emargination. Outer dististyle, d, conspicuously bilobed into blackened
points, the two lobes taken together suggesting a crabs claw, the outer part more obtuse
with a smaller lobule on inner margin; inner style pale and fleshy, with abundant setae.
Phallosome, p , including slender blackened rodlike apophyses; aedeagus slender, straight.

Holotype. $ , Tegan, Iran, July 5, 1956 (Schmid).

Allotype. $ , Durbadam, Iran, July 3, 1956.

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Paratype. $ , with the allotype.

Erioptera ( Psiloconopa ) idiophallus (Savtchenko) , described as an Ilisia (1973) from the
district Irshava, Transcarpathia, U.S.S.R., is generally similar but differs evidently in
hypopygial structure.

Molophilus (M olophilus) pallidipes, n. sp.

General coloration of head light gray ; thorax brownish gray, pleura brown ; antennae
short, brown; halteres light yellow; legs with femora and tibiae yellow, tips narrowly light
brown; wings brownish yellow; male hypopygium with outer lobe of basistyle extended
into a narrow pale plate ; a single long slender dististyle.

Male. Length about 4.5-5 mm.; wing 4.5-5 mm.; antenna about 1.2-1.3 mm.

Rostrum and palpi dark brown. Antennae brown, scape and pedicel more yellowed;
flagellar segments oval. Head light gray.

Pronotal scutum brown, scutellum light yellow. Mesonotal praescutum light to darker
brownish gray, with darker brown stripes, lateral pair short and ill-defined; scutum brownish
gray; scutellum yellowed, postnotum brownish gray. Pleura brown, dorsopleural membrane
yellowed. Halteres light yellow. Legs with coxae and trochanters yellow; femora and tibiae
yellow, tips narrowly light brown; tarsi brown. Wings brownish yellow, veins slightly
darker, the vestiture darker brown.

Abdomen medium brown. Male hypopygium (Fig. 9) with outer lobe of basistyle, h,
extended into a narrow pale plate, the apical margin farther produced into a point; inner
lobe of style small and narrow, apically with long pale setae. A single dististyle, d, appear-
ing as a long rod, gradually narrowed and curved to the acute twisted tip, apex acute.
Aedeagus, a , subequal in length and diameter to the dististyle, outer fourth more narrowed.

Holotype. $ , Pul-i-Zoghal, Iran, May 18, 1956 (Schmid).

Paratopotype. $ , pinned with type.

Paratypes. 2 $ $, Barajan, Iran, 2000 meters, September 15, 1955; $, Mughan, June 20,
1956; $ , Luis, September 14, 1955 (all Schmid).

The most similar regional species is Molophilus ( Molophilus ) stroblianus Nielsen (Zeitschr.
Wien. Ent. Gesell., 38: 36, figs.; 1953), known from Austria and Czechslovakia, a dark
colored fly with uniformly black legs, differing further in details of the male hypopygium.

DISTRIBUTIONAL RECORDS

Cheilotrichia ( Empeda ) cinerascens (Meigen)

Erioptera cinerascens Meigen; Klass., 1: 114; 1804.

Cheilotrichia ( Cheilotrichia ) cinerascens Edwards; Trans. Soc. Brit. Ent., 5: 119, pi. 5,
fig. 12 (wing) ; text fig. 23b (hypopygium) ; 1938.

Europe. Iran : Kamalabad, October 1955 (Schmid).

Ormosia bivittata (Loew)

Rhypholophus bivittatus Loew; Beschr. Eur. Dipt., 3: 41; 1873.

Rhypholophus bivittatus de Meijere; Tijd. v. Ent., 63: 50, fig. 40 (hypopygium); 1920.
Rhypholophus ( Rhypholophus ) bivittatus Lackschewitz ; Ann. naturhist. Mus. Wien, 50:
28; 1940.

Central Europe. Iran : Pul-i-Zoghal, October 12, 1956 (Schmid).

Erioptera ( Erioptera ) fuscipennis Meigen

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127

Erioptera fuscipennis Meigen; Syst. Beschr. 1: 111; 1818.

Erioptera fuscipennis de Meijere; Tijd. v. Ent., 63: 75, fig. 70 (hypopygium) ; 1920.
Erioptera ( Erioptera ) fuscipennis Edwards; Trans. Soc. Soc. Brit. Ent., 5: 124, text fig.
24 g (hypopygium) ; 1938.

Europe. Iran : Babal, May 21, 1956; Emaret, May 21, 1956; Lius, 2200 meters, September
14, 1955; Quattekas, 1800 meters, September 19, 1955; Zanus, 2000 meters, September 21,
1955 (Schmid).

Erioptera ( Erioptera ) trivialis Meigen

Erioptera trivialis Meigen; Syst. Beschr. 1: 112; 1818.

Erioptera trivialis de Meijere; Tijd. v. Ent., 63: 75, 76, fig. 71 (hypopygium); 1920.
Erioptera ( Erioptera ) trivialis Edwards; Trans. Soc. Brit. Ent., 5: 125, text fig. 24 n
(hypopygium) ; 1938.

Europe. Iran : Baranjan, 2000 meters, September 15, 1955; Lius, 2200 meters, September
14, 1955 (Schmid).

Erioptera ( Symplecta ) hybrida (Meigen)

Limnobia hybrida Meigen; Klass., 1: 57; 1804.

Symplecta punctipennis de Meijere; Tijd. v. Ent., 63: 77, 78, fig. 75 (hypopygium); 1920.
Erioptera ( Symplecta ) hybrida Edwards; Trans. Soc. Brit. Ent., 5: 126, pi. 5, fig. 5 (wing);
text fig. 24 A, g (hypopygium) ; 1938.

Europe ; Asia; Northwestern North America. Iran : Bar, June 30, 1956; Barajan, 2000

meters, September 15, 1955; Gurgan, April 1, 1956 (Schmid).

Erioptera ( Symplecta ) stictica (Meigen)

Limnobia stictica Meigen; Syst. Beschr. 1: 158; 1818.

Symplectomorpha stictica de Meijere; Tijd. v. Ent., 63: 78, fig. 76 (hypopygium); 1920.
Erioptera ( Symplecta ) stictica Edwards; Trans. Soc. Brit. Ent., 5: 128, pi. 5, fig. 4 (wing);
1938.

Eurasia; western North America. Iran: Cheshme, Ali, April 23, 1956; Ghulaman, July 8,
1956; Marus, June 28, 1956; Sefid Khok, June 1, 1956 (Schmid).

Erioptera ( Ilisia ) maculata Meigen

Erioptera maculata Meigen; Klass., 1: 61; 1804.

Acyphona maculata de Meijere; Tijd. v. Ent., 63: 67, 68, fig. 62 (hypopygium); 1920.
Erioptera ( Ilisia ) maculata Edwards; Trans. Soc. Brit. Ent., 5: 130-131, pi. 5, fig. 21 (wing) ;
text fig. 25 a (hypopygium) ; 1938.

Europe, widespread. Iran: Ardehjan, September 9, 1956; Bar, June 30, 1956; Mughan,
June 20, 1956; Pul-i-Zoghal, May 18, 1956 (Schmid).

Erioptera ( Psiloconopa ) czizeki (Bangerter)

Ilisia czizeki Bangerter; Mitteil. Schweiz. Ent. Gesell., 20: 353-354; 1947.

Erioptera ( Ilisia ) czizeki Stary; Casopis Moravskeho Musee, 55: 165, 166, fig. 19 (hypo-
pygium) ; 1971 (not 1970, as printed).

Central and Eastern Europe. Iran: Nandeh, May 29, 1956 (Schmid).

Erioptera ( Psiloconopa ) iranica Alexander

Erioptera ( Psiloconopa ) iranica Alexander; Jour. New York Ent. Soc., 81: 83-85; 1973.
Iran: Zanus, Mazanderan, September 21, 1955 (Schmid). Male hypopygium (Fig. 7).

M olophilus ( Molophilus ) bifidus Goetghebuer

Molophilus bifidus Goetghebuer; Bull. Soc. Ent. Belgique, 2: 135-136, fig. 9 (hypopygium);
1920.

Europe. Iran: Ochrid, 800 meters, August 9, 1955 (Schmid).

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New York Entomological Society

Molophilus ( M olophilus ) pleuralis de Meijere

Molophilus pleuralis de Meijere; Tijd. v. Ent., 63: 60-61, fig. S3 (hypopygium) ; 1920.
Molophilus pleuralis Edwards; Trans. Soc. Brit. Ent., 5: 144, text fig. 29 g (hypopygium);
1938.

Europe. Iran : Bar, June 30, 1956; Emaret, May 21, 1956; Lius, 2200 meters, September
14, 1955; Quattekas, September 19, 1955; Zanus, 2000 meters, September 21, 1955 (Schmid).

BOOK REVIEW

INSECT PHYSIOLOGY. Vincent B. Wigglesworth, 7th ed., 166 p. 1974. John Wiley &
Sons. $4.95 paperbound ($8.95 cloth).

Sir Vincent’s paperbound 7th edition is a real bargain at current book prices. Nearly 40
years after publishing the first edition, this new, revised little book is as readable an intro-
ductory account of insect physiology as only this masterful author can present. The brief,
but complete survey of the subject can be recommended as a stimulating introduction to
insect physiology for naturalists, biology students in high schools and colleges, and to
scientists in other disciplines who would like to become acquainted with an authoritative
and clear treatment of insect physiology. Each chapter is followed by a list of references,
from Dietrich Bodenstein to J. de Wilde, with a fair sprinkling of Wigglesworth’s own
contributions to almost all subjects. There are adequate drawings, illustrating anatomical
details. A subject index completes the book.

Karl Maramorosch

Waksman Institute of Microbiology

Rutgers University

New Brunswick, New Jersey

Vol. LXXXIII, June, 1975

129

New or Little-Known Crane Flies from Iran. IV
(Diptera: Tipulitlae)1

Charles P. Alexander
Amherst, Massachusetts 01002

Received eor Publication July 23, 1974

Abstract: Part III of this series of papers that concern the crane flies of Iran was published
in this Journal. In this paper species belonging to the tribe Eriopterini were treated. In the
final report here provided I am treating species in the major tribes Limoniini and Hexatomini.
The new species here described are Limonia ( Dicranomyia ) nigritorus, L. ( D .) schmidiana,
L. ( D .) subdidyma, Dicranota ( Dicranota ) ophidia, and Limnophila ( Elaeophila )
albofascia. In addition to the above novelties about a score of other species in these two
tribes are added to the previous limited list of crane flies presently known from Iran.

The three preceding papers on the crane flies of Iran that were collected by
Dr. Fernand Schmid in 1955 and 1956 discussed species in the tribes Pediciini
and Eriopterini. In this final report the tribes Limoniini and Hexatomini are
stressed and rather numerous species are added to the poorly known fauna of
Iran. The detailed report by Schmid on the Trichoptera of Iran was cited in
the previous paper in this series. It includes an excellent account of the various
collecting stations where he studied the chiefly aquatic groups of insects in 1955
and 1956 and the paper should be consulted by all students working on this
particularly interesting part of southwestern Asia. As had been stressed before,
the Schmid collections of crane flies made in southern Asia have provided the
great majority of the species presently known. All types of Tipulidae from
these collections are preserved in the Alexander Collection.

LIMONIINI

Limonia ( Dicranomyia ) decemmaculata (Loew)

Limnobia decemmaculata Loew; Berlin. Entomol. Zeitschr., 17: 35; 1873.

Dicranomyia decemmaculata Lackschewitz ; Ann. naturhist. Mus. Wien; 42: 205-206, pi.. 5, fig.
6 (hypopygium) ; 1928.

Limonia ( Dicranomyia ) decem-maculata Edwards; Trans. Soc. Brit. Ent., 5: 30-31, pi. 2,
fig. 12 (wing) ; 1938.

Wide-spread in Europe. Iran: Lius, 2200 meters, September 14, 1955; Ramsar, October 2,
1956 (Schmid). Wing (Fig. 1) ; hypopygium (Fig. 3).

It should be noted that Lackschewitz in the above reference shows the male hypopygium
with two rostral spines on the ventral dististyle, presumably in error. All other specimens
known to me have the single spine as shown in the figure. The Oriental Limonia
( Dicranomyia ) flavocincta (Brunetti), L. (D.) vibishana Alexander and L. ( D .) whitei

1 Contribution from the Entomological Laboratory, University of Massachusetts.

New York Entomological Society, LXXXIII: 129-138. June, 1975.

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New York Entomological Society

Alexander, of South India and Ceylon evidently are allied despite the virtually unpatterned
wings.

Limonia ( Dicranomyia ) nigritorus, n. sp.

Size medium (wing of male 8.5 mm); mesonotum gray, patterned with darker; knobs of
halteres dark brown; legs yellow, tips of femora narrowly dark brown; wings whitish, con-
spicuously patterned with brown, Sc long, Sci ending opposite one-third Rs ; abdomen dark
brown, posterior borders of segments broadly light yellow; male hypopygium with dorsal
dististyle long and slender, sinuous; ventral dististyle with two separated rostral spines, face
of style near base of prolongation with a subconical black lobe that is provided with numerous
black spines, mesal face of style at apex with dense microscopic setae.

Male. Length about 8.5 mm; wing 8.5 mm; antenna about 1.6 mm.

Rostrum and palpi dark brown. Antennae with scape and pedicel dark brown, flagellum
brown, the bases of proximal two segments narrowly yellowed; proximal flagellar segments
oval with truncated ends, outer ones elongate; verticils shorter than the segments. Head
dark brown.

Pronotal scutum dark brown, scutellum paler. Mesonotal praescutum gray with darker
stripes; scutum blackened, gray pruinose, lobes darker; scutellum brownish gray, narrowly
more blackened medially, parascutella light yellow; postnotum brown, light gray pruinose.
Pleura light gray, dorsopleural membrane more brownish yellow. Halteres with stem light
yellow, knob large, dark brown. Legs with coxae and trochanters light yellow; femora
yellow, tips narrowly dark brown; tibiae yellow, extreme tips darkened; tarsi yellow, outer
segments darkened. Wings whitened, conspicuously patterned with brown; cells C and Sc
with three darker areas, placed at base, near midlength and at outer end of vein Sc; stigma
brown, confluent with a large concolorous spot; other major paler brown clouds in cell Mi
and at ends of both anal veins, with a further broken series in cell M, chiefly along vein
Cu; smaller darkened seams over cord, outer end of cell 1st M 2, and at tips of veins Rz and
Mz; veins chiefly brown, darker in the more heavily patterned areas, yellowed in the costal
interspaces. Venation: Sc long, Sci ending opposite one-third Rs; cell 1st M2 subequal to
vein Mi + 2; m-cu shortly before fork of M.

Abdomen dark brown, posterior borders of segments broadly light yellow. Male
hypopygium (Fig. 4) with tergite, t, transverse, posterior border shallowly emarginate, the
low lobes more thickened, with long setae. Basistyle, b, and ventral dististyle nearly sub-
equal in area or the latter slightly larger. Dorsal dististyle, d, very long and slender,
sinuous; ventral style with rostral prolongation slender, with two separated subequal spinoid
setae, the outer one about one-half its length from apex of rostrum; apex of mesal face of
body of style with a concentration of very short blackened erect setulae, much smaller than
the normal setae ; face of style near base of the prolongation with a subconical black lobe,
its apex provided with several strong spines, the lobe placed in the curvature of the outer
style. Gonapophyses, g, with mesal-apical lobe slender. Aedeagus, a, narrow, especially
the lateral flanges, apex simple.

Holotype. $, Barajan, Iran, 2000 meters, September 15, 1955 (Schmid).

The present species is readily told from other regional members of the subgenus that have
patterned wings and long Sc by the body coloration and especially the hypopygial structure.
The distinctive blackened lobe on the ventral dististyle is particularly noteworthy and has
suggested the specific name. Limonia ( Dicranomyia ) modesta (Meigen), widely distributed
throughout the Holarctic region, has the hypopygium with somewhat similar but longer

Vol. LXXXIII, June, 1975

131

modified setulae on the ventral dististyle but differs in many other regards, including the
unpatterned wings, short Sc, and details of the hypopygium.

Limonia ( Dicranomyia ) schmidiana, n. sp.

Allied to mitis; general coloration of thoracic dorsum brownish gray, pleura yellow;
rostrum light yellow, antennal scape brownish yellow, remainder brown; legs light brown;
wings subhyaline, virtually unpatterned, stigmal region scarcely darker; Sc2 retracted, at
near two-thirds Sc; male hypopygium with ninth tergite pale, posterior border with two
broadly rounded lobes, the setae short; ventral dististyle small and rounded, only slightly
larger than the basistyle; rostral spines long, about twice the prolongation; mesal-apical
lobe of gonapophyses small, slender.

Male. Length about 7.5-8 mm; wing 7-8 mm; antenna about 1.2 mm.

Rostrum clear light yellow, palpi and mouthparts dark brown. Antennae with scape
brownish yellow, remainder of antenna brown, the extreme bases of proximal flagellar
segments more yellowed; segments short-oval, the outer ones longer, terminal segment
strongly narrowed on outer third. Anterior vertex yellowed, remainder of head chiefly
light gray, paler behind ; anterior vertex relatively broad.

Pronotum obscure yellow. Mesonotal praescutum with three brownish gray stripes that
virtually cover the dorsum ; scutal lobes chiefly brownish gray, central area pale ;
scutellum pale yellow; postnotal mediotergite yellowed, pleurotergite more whitened or
light gray. Pleura yellow, including the dorsopleural membrane. Halteres with stem yellow,
the large knob brown. Legs with coxae and trochanters yellow; femora light brown, tips
not darker; tibiae and tarsi light brown. Wings subhyaline, virtually unpatterned, the
stigmal region scarcely darker than the remainder; veins pale, Sc2 and base of Rs slightly
darker. Vein Sc without trichia; sparse trichia at end of 2nd A. Venation: Sci ending
about opposite origin of Rs, Sc2 retracted, at near three-fifths to two-thirds Sc; m-cu at or
shortly before fork of M.

Abdominal tergites yellowish brown, sternites and hypopygium yellow. Male hypopygium
(Fig. 5) combining the characters of mitis (long rostral spines) and chorea (small ventral
dististyle), differing in details. Ninth tergite, t, pale, posterior border with two broadly
rounded lobes, the median emargination acute; setae short and pale. Basistyle, b , in area
slightly less than the ventral dististyle; ventromesal lobe with moderately long setae.
Dorsal dististyle, d, long and slender, sickle-shaped, curved and narrowed to the acute
more or less recurved apex: ventral style short-oval to rounded; rostral prolongation small,
the two spines approximated, long and straight, about twice the length of the prolongation.
Gonapophysis, g, with mesal-apical lobe small, relatively slender. Aedeagus, a, with apertures
subterminal, median lobe conspicuous.

Holotype. S, Darband, Iran, April 22, 1956 (Schmid). Paratopotypes, 2 $ $, pinned with

type.

The species is dedicated to the collector, Dr. Fernand Schmid. It is most nearly related to
Limonia ( Dicranomyia ) mitis (Meigen) and allied species, as shown by the hypopygial
structure, especially the relative lengths of the rostral spines of the hypopygium. In the
small ventral dististyle it agrees more nearly with L. ( D .) chorea (Meigen) but is quite
distinct from this and other members of this group as delimited by Lackschewitz (Ann.
naturhist. Mus. Wien, 42: 209-217; 1928). Other particularly important papers on this
group of flies include Edwards (Trans. Soc. Brit. Ent., 5: 28-44; 1938) and de Meijere
(Tijd. voor Ent., 62: 65-90; 1919). Attention may be called to the species L. ( D .) lutea
Meigen (see Edwards, above, p. 37; de Meijere, p. 78) that was placed as a race or variety

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of mitis , differing in coloration of the body and wings and in hypopygial structure, including
the ventral dististyle, as discussed. Lackschewitz (1928, above) considered this as being a
yellowish autumnal form of chorea.

Limonia ( Dicranomyia ) subdidyma, n. sp.

General coloration of mesonotal praescutum and scutal lobes dull light brown, posterior
sclerites of notum and the pleura more yellowed; knobs of halteres blackened; legs with
femora yellow, tips very narrowly dark brown; wings pale yellow with a pale brown pattern
including four costal areas that are narrower than the interspaces, other darkenings on wing
paler; Sc2 far retracted, at near midlength of Sc; male hypopygium with rostral prolongation
of dististyle triangular in outline, with two short spines near base on face of style.

Male. Length about 8 mm; wing 9 mm.

Rostrum and palpi brown. Antennae brown; flagellar segments oval with truncated ends;
terminal and penultimate segments subequal in length. Head brown.

Pronotal scutum pale brown, scutellum more yellowed. Mesonotum dull light brown with
vague indications of light gray stripes; scutal lobes light brown, median area and the
scutellum pale yellow; postnotum brownish yellow. Pleura dull brownish yellow, dorsopleural
region clearer yellow. Halteres with stem yellow, knob almost black. Legs with coxae and
trochanters yellow; femora obscure yellow, tips very narrowly dark brown; tibiae brownish
yellow, extreme tips faintly darkened; tarsi brownish yellow, outer segments darker. Wings
pale yellow with a relatively inconspicuous pale brown pattern that includes four costal areas,
much narrower than the interspaces, the first area above the arculus, second at Sc2, the third
slightly larger, involving the tip of Sci and origin of Rs ; fourth darkening comprises the pale
brown stigma, nearly confluent with a smaller area over the fork of Rs ; further more re-
stricted markings over remainder of cord, outer end of cell 1st M2 and tip of Rs; still smaller
marginal clouds at ends of veins Ms to 2nd A, the last more extensive and slightly paler;
veins yellow, light brown in the clouded areas. Venation: Sci ending opposite origin of Rs,
Sc2 far retracted, close to midlength of Sc; free tip of Sc2 and R2 in transverse alignment;
m-cu shortly before the fork of M .

Abdomen pale brown, the extreme posterior borders of segments more yellowed;
hypopygium with basistyle darker brown. Male hypopygium (Fig. 6) with tergal lobes, t ,
rounded, vestiture pale and inconspicuous. Basistyle, b, with ventromesal lobe rounded,
vestiture long. Dorsal dististyle, d, bent at midlength, the narrowed outer end slender, tip
decurved; ventral style about twice the basistyle, rostral prolongation triangular in outline;
spines short, placed on face of prolongation near base. Gonapophyses, g, with mesal-apical
lobe erect.

Holotype. $, Darband, Iran, April 22, 1956 (Schmid).

The most similar Palaearctic species include Limonia ( Dicranomyia ) chorea (Meigen),
L. ( D .) didyma (Meigen) and L. ( D .) mitis (Meigen), all with the wings patterned and with
Sci very long. The single most similar species is didyma which differs evidently in wing

Fig. 1. Limonia ( Dicranomyia ) decemmaculata (Loew) ; venation.

Fig. 2. Dicranota ( Dicranota ) ophidia , n. sp.; venation.

Fig. 3. Limonia ( Dicranomyia ) decemmaculata (Loew) ; male hypopygium.
Fig. 4. Limonia ( Dicranomyia ) nigritorus , n. sp.; male hypopygium.

Fig. 5. Limonia ( Dicranomyia ) schmidiana, n. sp.; male hypopygium.

Vol. LXXXIII, June, 1975

133

Fig. 6. Limonia ( Dicranomyia ) subdidyma , n. sp.; male hypopygium. Subfigures: A,

didyma (Meigen) ; B, chorea (Meigen) ; C, mitis (Meigen).

Fig. 7. Dicranota ( Dicranota ) ophidia, n. sp.; male hypopygium.

Fig. 8. Limnophila ( Elaeophila ) albofascia, n. sp. ; male hypopygium.

(Symbols: Male hypopygium — a, aedeagus; b , basistyle; d, dististyles; g, gonapophysis;
z, interbase; p, phallosome; t, 9th tergite.)

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New York Entomological Society

pattern, as the undarkened arcular region, and especially in hypopygial details. I have
provided illustrations of the rostral prolongations of the above three species in subfigures, -
didyma, 6 A; chorea , 6B; mitis, 6C.

PEDICnNI

Dicranota ( Dicranota ) ophidia, n. sp.

Size relatively large (wing 7-8 mm); antennae short; general coloration of thorax light
brownish gray, praescutum with a darker central stripe, pleura brownish yellow, sparsely
pruinose ; halteres and legs yellow ; wings faintly infuscated, stigma only slightly darker,
R 2 + 3+4 subequal to or shorter than basal section of male hypopygium with lateral tergal
blades short, posterior border truncate, setae short; dorsal lobe of basistyle with short
blackened subspinoid setae; interbase a sinuous snakelike rod, slightly constricted before the
pointed head.

Male. Length about 7-8 mm; wing 7-8.5 mm; antenna about 1-1.2 mm.

Female. Length about 8.5-9 mm; wing 7-8 mm.

Rostrum brownish gray, palpi black. Antennae short, brownish black, scape pruinose.
Head brownish gray.

Pronotum light brown, pretergites and posterior borders of scutum and scutellum light
yellow. Mesonotal praescutum light brownish gray with a conspicuous darker brown central
stripe, lateral stripes much narrower; scutum light gray, centers of lobes extensively light
brown; scutellum brown, light gray pruinose; postnotum brownish yellow, slightly pruinose.
Pleura brownish yellow, slightly pruinose. Halteres light yellow. Legs with coxae yellow,
fore and middle pairs slightly pruinose ; trochanters yellow ; remainder of legs brownish
yellow, outer tarsal segments slightly darker; claws long, gently curved. Wings (Fig. 2)
faintly infuscated, stigma only slightly darker; veins light brown. Macrotrichia of veins
beyond cord long and delicate ; basal veins, including M and both Anals with much shorter
trichia on outer two-thirds or more, less extensive on Cu. Venation: Ri + 3 + 4 variable in
length, in cases subequal to or shorter than basal section of Ro, in cases twice as long; cell
Mi present.

Abdominal tergites light to darker brown, sternites slightly paler, hypopygium darker
brown. Male hypopygium (Fig. 7) with tergite, t, truncate, with abundant short setae,
lateral tergal ends produced into short slender blades. Basistyle, b, with dorsal lobe stouter,
vestiture short, blackened, subspinoid; ventral lobe glabrous above, lower margin with
numerous pale setae, the more basal ones shorter. Interbase, i, distinctive, appearing as a
long sinuous snakelike rod, slightly constricted before the long pointed head. Phallosome,

with details generally as figured, aedeagus short and slender.

Holotype. $ , Zanus, Iran, 2000 meters, September 21, 1955 (Schmid).

Allotopotype, $, pinned with type. Paratypes, $ 2, Lius, 2200 meters, September 14, 1955;

Rayne, 1800 meters, September 2-5, 1955; Waliabad, September 16-24, 1956 (Schmid).

The more similar species include Dicranota ( Dicranota ) capillata Lackschewitz and the
larger D. (D.) fuscipennis Lackschewitz, of central Europe, especially the former. I possess
a paratype of capillata received in an exchange with Lackschewitz (Salzburg, Gastein, 1879,
collected by Joseph Mik). This differs from the present fly in the very distinct venation of
the radial field but this feature may well represent an individual variation only. In the
specimen r-m is before the fork of Rs that forks into a trident and cell Mi is very reduced.
The details of the male hypopygium are quite distinct, especially the produced tergal border

Vol. LXXXIII, June, 1975

135

and the shape and vesiture of the dististyle. The interbase is generally as in the present fly,
differing in the outer conformation.

HEXATOMINI

Limnophila ( Elaeophila ) albofascia, n. sp.

General coloration of head and thorax brownish gray, praescutum conspicuously patterned
with darker brown ; knobs of halteres dark brown ; wings with distinctive pattern, including
darker costal areas, with a complete unbroken pale band at midlength between the third and
fourth darkened areas; no darkened spots or dots on veins; male hypopygium with outer
dististyle narrowed at apex into a short curved point ; dorsal crest long and low ; gonapophyses
large ; aedeagus long and slender.

Male. Length about 7.5 mm; wing 6.5 mm.

Female. Length about 8 mm; wing 7 mm.

Rostrum light brown; palpi black. Antennae with scape and pedicel light brown;
flagellum broken. Head brownish gray.

Pronotum brownish gray. Mesonotal praescutum brownish gray, patterned with brown,
including six longitudinal lines, intermediate pair broader on anterior half, directed laterad
anteriorly to form lateral marginal stripes, posterior halves of intermediate stripes much
narrower and paler, sublateral darkenings broader; pseudosutural foveae brownish black;
posterior sclerites of notum pale yellowish gray with very inconspicuous darker markings on
scutal lobes and at median line. Pleura pale brown, vaguely patterned with darker. Halteres
with stem yellow, outer end of the large knob dark brown. Legs with coxae and
trochanters yellow; remainder of legs broken. Wings with ground pale yellowish white, the
darkened areas subequal in size ; a series of about six darker brown marks in costal field,
with a very broad continuous ground crossband at midlength of wing that completely
divides the darkened areas; costal darkenings darker, with three marks before the dividing
ground band, the basal one at and near the arcular area; second darkening small, in cell R
narrowed; third band complete, in the type narrowed at the supernumerary crossvein in cell
M , behind expanded and more or less divided at end of vein 2nd A ; beyond the intermediate
ground band with the fourth darkened area in costal field expanded to include the forks of
veins Sc and Ri, almost confluent, narrowed posteriorly over r-m and posterior cord, more
or less confluent with darkenings at end of cell 1st M 2; two outer darkened areas at ends of
veins Rs and Ri, more extensive in holotype, in outer radial cells separated by a circular
ground mark in outer end of cell R3 ; behind the dark pattern more extensive in the holotype,
involving much of outer ends of cell Ri and R5, with only the tip pale ; in the female the
dark pattern at wing apex more restricted, appearing as seams over fork of Mi + 2 and the
apices of all longitudinal veins excepting R5 ; no supplementary spots or dots on longitudinal
veins as in several species in this subgenus; veins yellow in the ground areas, darkest in the
costal darkenings, paler behind. Venation: Supernumerary crossvein in the third darkened
area; R2+ 3+4 about one-half longer than basal section of R3.

Abdomen yellow, patterned with brown, most evident as darkenings at posterior ends of
segments, broader outwardly. Male hypopygium (Fig. 8) with outer dististyle, d, as shown,
the apex narrowed into a short curved point; outer margin with a row of small appressed
spines, dorsal crest long and low. Gonapophyses, g, appearing as long paddles that are
more than one-half the length of the long slender aedeagus.

Holotype. $ , Durbadam, Iran, July 3, 1956.

Allotype. $ , Chenes, Iran, May 19, 1956 (Schmid).

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New York Entomological Society

From other regional members of the subgenus Elaeophila with somewhat comparable
wing patterns, especially with no darkened spots or dots along the veins, the present fly is
most readily told by the broad continuous pale ground fascia at near midlength of the wing
and in details of hypopygial structure, especially the outer dististyle and phallosome. The
most similar such species is Limnophila ( Elaeophila ) submarmorata (Verrall) which differs
in the above respects. The great variation found in the wing pattern in submarmorata has
been discussed by Edwards and various forms have been named by him (Trans. Soc. Brit.
Ent., 5 : 81-84, pi. 4, figs. 1-7, wings ; 1938) .

DISTRIBUTIONAL RECORDS
LIMONDNI

Helius ( Helius ) hispanicus Lackschewitz

Helius hispanicus Lackschewitz; Ann. naturhist. Mus. Wien, 42: 242-243; 1928.

Described from Algeciras, Andalusia, Spain (Hanns Zerny). Iran : Mughan, June 20,

1956 (Schmid) .

Helius ( Helius ) pallirostris Edwards; Trans. Ent. Soc. London 1921: 206; 1921.

Europe : Britain; Sweden; Denmark, and others. Iran: Kia Kola, May 22, 1956 (Schmid).

Antocha ( Antocha ) libanotica Lackschewitz

Antocha ( Antocha ) libanotica Lackschewitz; Ann. naturhist. Mus. Wien, 50: 8, pi. 1, figs.
4; 1939 (1940).

Type from Libanon ; paratype in Alexander Collection.

Iran: Ardehjan, September 11, 1956; Dazdban, May 18, 1956; Lius, 2200 meters, September
14, 1955; Meyur, August 23, 1956; Mishgin, August 21, 1956; Rayne, 1800 meters, September
2, 1955; Shirgah, May 23, 1956; Zanus, 2000 meters, September 21, 1955; Zirab, May 23,
1956 (Schmid) .

Limonia ( Limonia ) hercegovinae (Strobl)

Limnobia Hercegovinae Strobl; Glasnik Zem. Mus. Bosni i Hercegov., 10: 610; 1898.

Central and Eastern Europe. Iran: Barajan, 2000 meters, September 15, 1955; Hassankif,
September 28, 1956; Khazlak, June 6, 1956; Lius, 2200 meters, September 14, 1955; Zanus,
2000 meters, September 21, 1955 (Schmid).

Limonia ( Limonia ) neonebulosa Alexander

Dicranomyia nebulosa Alexander; Canad. Ent., 45: 203; 1913 (preoccupied by Zetterstedt,
1838).

Limonia ( Dicranomyia ) neonebulosa Alexander; Philippine Jour. Sci., 24: 555; 1924.

Eastern Asia; Eastern North America. Iran: Baharistan, August 20, 1956 (Schmid).

Limonia ( Melanolimonia ) morio (Fabricius)

Tipula morio Fabricius; Mantissa Ins., 2: 324; 1787.

Europe (widespread). Iran: Daiband, April 22, 1956; Khozlok, June 6, 1956 (Schmid).

Limonia ( Dicranomyia ) chorea (Meigen)

Limnobia chorea Meigen; Syst. Beschr. 1: 134; 1818.

Dicranomyia chorea de Meijere; Tijd. v. Ent., 62: 74, fig. 7 (hypopygium) ; 1919.
Dicranomyia chorea Lackschewitz; Ann. naturhist. Mus. Wien, 42: 211, fig. 8 (hypopygium) ;
1928.

Limonia ( Dicranomyia ) chorea Edwards; Trans. Soc. Brit. Ent., 5: 35, pi. 2, fig. 20 (wing) ;
text fig. 5 a (hypopygium) ; 1938.

Europe (widespread). Iran: Lius, 2200 meters, September 14, 1955 (Schmid).

Vol. LXXXIII, June, 1975

137

Limonia ( Dicranomyia ) didyma Meigen
Limonia didyma Meigen; Klass., 1: 55; 1804.

Europe (widespread). Iran : Aliabad, 1800 meters, September 7, 1955; Ardehjan, Sep-
tember 11, 1956; Barajan, 2000 meters, September 15, 1955; Lius, 2200 meters, September
14, 1955; Waliabad, September 16 and 24, 1956 (Schmid).

Limonia ( Dicranomyia ) fusca (Meigen)

Limnobia fusca Meigen; Syst. Beschr. 6: 274; 1830.

Widespread in Europe and North America. Iran: Baharistan, 2000-3000 feet, September
10, 1956; Barajan, 2000 meters, September 15, 1955; Harandan, 100 feet, September 11,
1956 (Schmid).

Limonia ( Dicranomyia ) longipennis (Schummel)

Limnobia longipennis Schummel; Beitr. zur Ent., 1: 104; 1829.

Holarctic; widespread. Iran : Barajan, 2000 meters, September 15, 1955; Gulugah, Sep-
tember 8, 1956; Mishgin, August 21, 1956; Javardi, 4000 feet, October 7, 1956; Quattekas,
1800 meters, September 19, 1955 (Schmid).

Limonia ( Dicranomyia ) modesta (Meigen)

Limnobia modesta Meigen; Syst. Beschr., 1: 134; 1818.

Europe. Iran: Gach-i-Lai (name faulty), May 17, 1956; Lius, 2200 meters, September
14, 1955; Mishgin, 4500 feet, August 21, 1956; Ziarat, 2000 feet, July 13, 1956 (Schmid).

Limonia ( Dicranomyia ) ventralis (Schummel)

Limnobia ventralis Schummel; Beitr. zur Entomol., 1: 136; 1829.

Limonia ( Dicranomyia ) pristomera Alexander; Oriental Insects, 1: 204-205, fig. 8 (hypo-
pygium) ; 1967 (synonym).

Europe; Afghanistan; South India (Kerala; Madras; Mysore). Iran: Fumen, 50 feet,

August 18, 1956; Pul-i-Zoghal, 1760 feet, October 12, 1956 (Schmid).

HEXATOMINI

Paradelphomyia ( Oxyrhiza ) czizekiana Stary

Paradelphomyia ( Oxyrhiza ) czizekiana Stary; Casopis Moravskeho Musee, 55: 135-137,
figs.; 1971.

Types from Moravia, Czechoslovakia. Iran: What appears to be this species from

Ardehjan, September 11, 1956; Baharistan, circa 2000-3000 feet, September 10, 1956; Lius,
circa 7000 feet, September 14, 1955 (Schmid). The most important difference from Stary’s
description and figures is in the hypopygium, especially the ventral fork where the two
spines are widely separated basally, being placed on a horizontal connecting rod. Despite
this difference I believe the identification is correct.

Paradelphomyia ( Oxyrhiza ) fuscula (Loew)

Cladura fuscula Loew; Berlin. Ent. Zeitschr., 17: 35; 1873.

Europe. Iran: Quattekas, circa 4500 feet, September 19, 1955 (Schmid).

Austrolimnophila ochracea (Meigen)

Limonia ochracea Meigen; Klass., 1: 55; 1804.

Europe. Iran: Ardehjan, September 11, 1956; Bozak, 1800 meters, September 11, 1955
(Schmid). The present fly and the Nearctic Austrolimnophila toxoneura (Osten Sacken)
are very similar and perhaps will be found to be identical.

Pseudolimnophila lucorum (Meigen)

Limnobia lucorum Meigen; Syst. Beschr. 1: 125; 1818.

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New York Entomological Society

Europe. Iran : Ardehjan, September 9, 1956; Baharistan, September 10, 1956; Barajan,
2000 meters, September 15, 1955; Chalus, May 19, 1956; Zanus, 2000 meters, September 21,
1955 (Schmid).

Limnophila {Elaeophila) submarmorata (Verrall)

Ephelia submarmorata Verrall; Ent. Mo. Mag., 23: 264; 1887.

Limnophila ( Elaeophila ) submarmorata Edwards; Trans. Soc. Brit. Ent., 5: 81-84, pi. 4,
figs. 1, 2; text fig. 15 b; 1938.

Europe. Iran : Gurgan, April 1, 1956; Zanus, 2000 meters, September 21, 1955 (Schmid).
Pilaria discicollis (Meigen)

Limnobia discicollis Meigen; Syst. Beschr., 1: 125; 1818.

Europe. Iran: Quattekas, 1800 meters, September 19, 1955 (Schmid).

Pilaria scutellata (Staeger)

Limnophila scutellata Staeger; in Krojer, Naturhist. Tidsskr. 3: 34; 1840.

Europe. Iran: Chalus, May 19, 1956 (Schmid).

TIPULINAE

Tipula ( Acutipula ) maxima transcaucasica Savtshenko

Tipula ( Acutipula ) maxima transcaucasica Savtshenko; Fauna U. S. S. R., Diptera II, No.
3: 413; 1961.

U.S.S.R.: Transcaucasia. Iran: Lius, 2200 meters, September 14, 1955 (Schmid).

Tipula ( Tipula ) orientalis Lackschewitz

Tipula ( Tipula ) orientalis Lackschewitz; Konowia, 9: 272-273, fig. 7; 1930.

South Europe; Egypt; Kurdistan; Caucasus. Iran: Rayne, 1800 meters, September 5,

1955 (Schmid).

Vol. LXXXIII, June, 1975

139

BOOK REVIEW

The Pest War. W. W. Fletcher, Halsted Press, John Wiley & Sons, N. Y. 218p., 1974.
$11.95.

The problems associated with control of insects and other pests are complex. The author
discusses man’s war against the major pests that threaten human health and the supply of
food. The book is primarily dealing with insects, weeds, fungi and certain vertebrates.
Methods of pest control, from mechanical ones, crop rotation, quarantine and eradication,
to biological and chemical methods are outlined. The development of insecticides from its
early days, through the dramatic period following the application of DDT, cyclodienes,
carbamates, organophosphorus and systemic insecticides, the resistance to these compounds,
the synergistic effects, as well as diverse uses of herbicides are discussed in brief. I was
intrigued by the description of the discovery of 2,4 D in this book, since it differed
strikingly from the story I knew. According to Fletcher, 3 scientists at Rothamsted Ex-
perimental Station, Nutman, Thorton and Quastel hit upon 2,4 D, and the result of their
preliminary work was communicated in 1942 to the Agricultural Research Council, who
asked Prof. G. E. Blackman of Oxford University to initiate a program of field trials.
These results appeared as late as 1945 in NATURE, having been held up until then for
security reasons. The author then mentions that in 1942 two Americans, Zimmerman and
Hitchcock, described the use of 2,4 D as a plant growth regulator, but not as a herbicide.
Also in the United States, Marth and Mitchell, as well as Hamner and Tukey described the
herbicidal uses in 1944. It might be difficult to establish precedence for the precise dis-
covery of the herbicidal activity from these quotations, but I recall that the patent was
applied for, and given, to Zimmerman and Hitchcock at Boyce Thompson Institute. It
was not contested by the workers at Rothamsted, and the American scientists deserve full
credit for this discovery. The various fungicides, including thiram, captan, quinones, as
well as antibiotics such as streptomycin and griseofulvin are briefly mentioned. A whole
chapter is devoted to methods of application of insecticides and herbicides.

Among the vertebrate pests, the rabbit eradication attempts in Australia and Europe by
the myxomatosis virus are described. The resistance to the virus forced the reintroduction
of effective chemical control methods. While rabbits seem to be regarded with some af-
fection, rats are generally despised and the most drastic eradication methods are sometimes
proposed, and used. Among them is the application of anticoagulant agents, to which,
unfortunately, rats can develop resistance. Several species of birds, such as pigeons, gulls,
and others also are listed as pests, and their control discussed. The impressive success of
biological control methods, as well as integrated biological and chemical control, and novel
methods of control are presented in a very stimulating manner. Sterilization by chemicals
and radiation, pioneered by Knipling, use of sex attractants, repellents, electromagnetic
energy, ionizing radiation, as well as the use of insect hormones as insecticides are all
briefly presented. A whole chapter is devoted to the effects of pesticides on the environ-
ment. The book ends with an appendix, listing additional sources of information for in-
terested readers, as well as a list of common and scientific names of pests, and of pesticides.
A good index, on 18 pages, is provided.

The author should be congratulated for his comprehensive and well balanced presentation
of this complex subject, in a manner understandable by laymen as well as by experts.

Karl Maramorosch

Waksman Institute of Microbiology

Rutgers University

New Brunswick, New Jersey

140

New York Entomological Society

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Vol. LXXXIII

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Devoted to

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The New York Entomological Society

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Incorporating The Brooklyn Entomological Society

Incorporated May 21, 1968

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The New York Entomological Society

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Organized June 29, 1892 — Incorporated February 25, 1893

Reincorporated February 17, 1943

The Brooklyn Entomological Society

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Officers for the Year 1975

President, Dr. Daniel J. Sullivan, S.J.

' r II

Fordham University, New York 10458

Vice-President, Dr. Peter Moller

American Museum of Natural History, New York 10024
Secretary, Dr. Charles C. Porter

Fordham University, New York 10458

Assistant Secretary, Dr. Louis Trombetta

Isaac Albert Research Institute, Brooklyn, N.Y. 11203

Treasurer , Dr. Ivan Huber

Fairleigh Dickinson University, Madison, N.J. 07940

Assistant Treasurer , Ms. Joan DeWind

American Museum of Natural History, New York 10024

Trustees

Class of 1975

Dr. Howard Topoff
Class of 1976

Dr. David C. Miller

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Dr. Pedro Wygodzinsky

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Dr. Norman Platnick

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Mailed September 26, 1975

Known office of publication: Waksman Institute of Microbiology, New Brunswick, New Jersey 08903.

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Journal of the

New York Entomological Society

Volume LXXXIII September 1975 No. 3

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Waksman Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lois J. Keller, RSM
Dr. Herbert T. Streu

Publication Committee

Dr. Kumar Krishna Dr. Ayodha P. Gupta

Dr. James Forbes, Chairman

CONTENTS

An Annotated List of New York Siphonaptera

Allen H. Benton and Danny L. Kelly 142

Notes on the Life Cycle and Natural History of Butterflies of El Salvador.

I B. — Hamadryas februa (Nymphalidae-Hamadryadinae)

Alberto Muyshondt and Alberto Muyshondt, Jr. 157

Notes on the Life Cycle and Natural History of Butterflies of El Salvador.

II B. — Hamadryas guatemalena Bates (Nymphalidae-Hamadryadinae)

Alberto Muyshondt and Alberto Muyshondt, Jr. 170

Notes on the Life Cycle and Natural History of Butterflies of El Salvador.

III B. — Hamadryas amphinome L. (Nymphalidae-Hamadryadinae)

Alberto Muyshondt and Alberto Muyshondt, Jr. 181

Notes on the Male Reproductive System in Ants (Hymenoptera : Formicidae)

A. C. F. Hung and S. B. Vinson 192

Species and Numbers of Bloodsucking Flies Feeding on Hogs and Other Ani-
mals in Southern New Jersey Thomas J. Weiner and Elton J. Hansens 198

Speleognathinae Collected From Birds In North America (Acarina: Erey-
netidae) A. Fain and K. E. Hyland 203

Elliptochthoniidae, a New Mite Family (Acarina: Orihatei) From Mineral Soil
in California Roy A. Norton 209

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New York Entomological Society

An Annotated List of New York Siphonaptera

Allen H. Benton and Danny L. Kelly
Department of Biology, State University College, Fredonia, New York. 14063

Received for Publication July 29, 1974

Abstract: Geary (1959) listed 42 species of Siphonaptera from New York. The present
list includes numerous additional distributional records, and adds three species to Geary’s
list: Peromyscopsylla h. hamifer (Rothschild), Ceratophyllus dif finis Jordan (previously

reported but missed by Geary), and Epitedia ( cavernicola Traub?). Geary also removed
from the state list Echidnophaga gallinacea (Westwood), but we consider the published
record to be a valid one.

New York is one of the most thoroughly studied of states with respect to
its flea fauna. Many of the specimens studied by Carl F. Baker near the
turn of the century were from New York, largely through the collecting of
G. S. Miller, Jr., whose home was in Madison county. The type localities of
four fleas described by Baker are in that vicinity, and seven other forms have
been described from type localities in New York.

The first state list of fleas from any eastern state was that of Stewart (1928),
listing 26 species from New York. Jordan (1929) made some corrections which
reduced Stewart’s list to 22, and added nine more species, bringing the list
to 31 forms. He predicted that about 50 species would eventually be discovered
in the state, and presented a hypothetical list comprising 11 species. Of these,
seven have since been recorded in the state, two are not to be expected in
light of current knowledge, and two remain on the hypothetical list.

Stewart (1933) revised the list once again, recording 36 forms and including
one which Jordan had discredited in 1929. Fox (1940) based his New York
list primarily on the 1933 list of Stewart, adding three species and eliminating
one species and one subspecies. His list thus totalled 37 forms.

The most recent summary of New York collections was that of Geary (1959),
which included 42 forms with definite records and four species which had been
previously recorded but which Geary considered to be of doubtful validity.

Since 1960, we have had access to more than 4,000 flea specimens from
New York, covering many areas which had been poorly represented in col-
lections up to that time. We are grateful to the New York State Museum
and Science Service and Dr. Paul Connor for the use of collections from Lewis,
Otsego, Schoharie, St. Lawrence and Suffolk counties. In addition, more than
a thousand specimens from the Catskill Mountain area have been loaned by
Daniel Smiley, John New and Robert Fisher. Numerous students and friends
have supplied additional collections and have assisted in the preparation of
specimens. Part of the work has been supported by grants from the Research

New York Entomological Society, LXXXIII: 142-156. September, 1975.

Vol. LXXXIII, September, 1975

143

Foundation of State University of New York, the Atmospheric Sciences Re-
search Center of State University of New York, and Health Research Incor-
porated, Albany, N. Y.

The present list is undoubtedly incomplete, and it is likely that Jordan’s
estimate of about 50 species is extremely accurate. Although much remains
to be learned about local and ecological distribution, it is unlikely that more
than a half-dozen species remain to be discovered within the state.

Nomenclature of the Siphonaptera follows the classification of Hopkins and
Rothschild (1953 et seq.) so far as available, except for a few taxonomic
changes which have occurred since the pertinent volume was published. Mam-
mal names follow Hall and Kelson (1959). Host relationship data follow the
plan of Sakaguti and Jameson (1962): true hosts permit the flea to carry out
its life cycle indefinitely; secondary hosts are commonly parasitized, but are
not considered biologically adequate as permanent hosts; accidental hosts are
those which result from accidental coming together of host and parasite, and
such relationships are not likely to occur frequently. Since our knowledge of
host relationships is far from complete, these designations should be taken as
considered opinions, which may prove to be wrong in the light of further data.

Host records listed are those which are known from New York. In most
cases, the true host is the same throughout the range of the species, but this
is not always true. In New York, for example, Monopsyllus vison is rarely
found on any host except the red squirrel. In Minnesota, however, large
numbers occur on the eastern chipmunk, and the species is known from
southern Illinois, beyond the range of the red squirrel. It is evident that
host relationships as they occur in our area are not necessarily the same
throughout the range of the flea in question.

The most serious gap in our knowledge of flea distribution is the almost
complete lack of information about the influence of factors other than the
presence of the host. The puzzling distribution patterns of such species as
Stenoponia americana, Peromyscopsylla scotti, Peromyscopsylla hamifer and
others cannot be explained on the basis of host distribution. Detailed study
of the life history and ecology of almost any flea species would be extremely
rewarding, but such studies have been undertaken for only a few species which
have great medical or veterinary importance.

FAMILY PULICIDAE

Echidnophaga gallinacea (Westwood)

This species is included on the basis of a record reported by Fox and
Sullivan (1925). While there is probably no resident population within the
state, we see no reason to doubt the validity of this record, or to doubt that
the species may occasionally be brought into the state on rats or domestic
animals.

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New York Entomological Society

Fig. 1. Map of New York State showing counties. Numbers are the code used for
distributional data under species accounts.

Host: Rattus sp.

County: 561.

Range within the state: Probably restricted to occasional accidental intro-
ductions

Pulex irritans Linnaeus

Although human fleas are undoubtedly brought into the state often, records
are few. We have not seen specimens from the state, and thus cannot evaluate
the possibility that some or all of the records may refer to P. simulans Baker.

True host: Homo sapiens
Counties: 22, 24, 26

Range within the state: Unknown; possibly there is no permanent pop-

ulation

1 The map of New York, Figure 1, has the counties coded by number for economy of
space. Please refer to this map for identification of the counties indicated in the species
accounts.

Vol. LXXXIII, September, 1975

145

Ctenocephalides canis (Curtis)

True hosts: Canis familiaris, Vulpes fulva, Urocyon ciner e oar gent eus

Accidental hosts: Mephitis mephitis , Rattus norvegicus

Counties: 8, 20, 24, 25, 50, 51, 56

Range within the state: Probably all of the state, with the possible ex-

ception of the highest elevations

Ctenocephalides felis felis (Bouche)

True hosts: Felis domestica , Canis familiaris, Vulpes fulva, Urocyon cin-
ereoargenteus

Secondary host: Homo sapiens

Accidental hosts: Blarina brevicauda, Didelphis marsupialis, Procyon lotor ,
Rattus norvegicus , Sylvilagus floridanus, Tamiasciurus hudsonicus

Counties: 2, 7, 11, 13, 17, 19, 20, 21, 24, 25, 26, 28, 35, 36, 38, 42, 47,
50, 53, 56

Range within the state: Throughout the state.

Cediopsylla simplex (Baker)

This species occurs in great numbers on all species of Leporidae occurring
within the state. In the higher mountains of the Adirondacks, however, where
Sylvilagus floridanus does not occur, we have been unable to find this flea.
Whether its absence is due to the absence of the cottontail, or whether it is
due to some other ecological factor, is not yet clear.

True hosts: Sylvilagus floridanus, S. transitionalis , Lepus americanus, L.
europaeus

Secondary and accidental hosts: Canis familiaris, Didelphis marsupialis,

Felis domestica, Mustela frenata, Rattus norvegicus, Tamiasciurus hud-
sonicus

Counties: 2, 9, 10, 19, 20, 21, 22, 23, 25, 26, 27, 30, 31, 35, 36, 42, 44,
45, 46, 47, 48, 49, 50, 52, 56, 61

Range within the state: Throughout the state, except for the higher moun-
tain areas. Additional collecting at high elevations is needed to determine
whether it is indeed absent from those areas.

Xenopsylla cheopis (Rothschild)

The Oriental rat flea is the major carrier of bubonic plague, and is there-
fore of great medical importance. Undoubtedly, the species is brought into
the state occasionally on rats, but the records are so few as to suggest that
the species is only a sporadic entrant, but permanent populations may persist
in the extreme southeastern counties.

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New York Entomological Society

True host: Rattus norvegicus
Counties: 20, 56

Range within the state: Uncertain; possibly there is no permanent pop-
ulation

FAMILY VERMIPSYLLIDAE

Chaetopsylla lot oris (Stewart)

Although named for its type host, the raccoon, this species is also very
frequently taken from foxes (Zeh, 1973).

True hosts: Procyon lotor, Vulpes julva, Urocyon cinereoargenteus
Secondary hosts: Martes pennanti, Didelphis marsupialis
Counties: 7, 15, 17, 18, 19, 25, 26, 29, 30, 42, 45, 46, 49, 50, 51, 52
Range within the state: Throughout the state

FAMILY HYSTRICHOPSYLLIDAE

Hystrichopsylla tahavuana Jordan

True hosts: Parascalops breweri, Condylura cristata
Secondary host: Blarina brevicauda

Accidental hosts: Microtus pennsylvanicus, M. pinetorum , Peromyscus leu-
copus

Counties: 1, 5, 8, 17, 23, 25, 26, 42, 48

Range within the state: Probably wherever its true hosts occur; because

moles are not usually taken in large numbers by collectors, records are
relatively few

Atyphloceras bishopi Jordan

This is a winter flea, most commonly taken from nests, and hence rather
rare in collections. Most New York records are from the meadow vole, but it
is taken on the red-backed vole in more northern areas.

True hosts: Microtus pennsylvanicus , Clethrionomys gapperi
Secondary hosts: Microtus pinetorum, M. chrotorrhinus
Accidental hosts: Peromyscus leucopus, Blarina brevicauda
Counties: 17, 22, 28, 34, 42, 48

Range within the state: Probably throughout the state; it may have eco-
logical limits as yet unknown

Stenoponia americana (Baker)

This species shows little host specificity, occurring on a variety of small
mammals. Since it occurs up the Atlantic coast as far as New Brunswick,

Vol. LXXXIII, September, 1975

147

there seems no reason why it should not occur in parts of New York where
it has not yet been recorded.

Hosts: Peromyscus leucopus , Blarina brevicauda, Microtus pennsylvanicus ,
M. pinetorum, Clethrionomys gapperi , Sorex cinereus , Scalopus aquaticus
Counties: 25, 48, 61

Range within the state: So far as known, confined to Long Island and

the Hudson valley.

Tamiophila grandis (Rothschild)

True host: Tamias striatus

Accidental hosts: Tamiasciurus hudsonicus, Vulpes julva
Counties: 1, 5, 8, 12, 19, 22, 26, 33, 35, 36, 42, 45, 48
Range within the state: Throughout the state

Catallagia borealis Ewing

True host: Clethrionomys gapperi

Secondary and accidental hosts: Microtus chrotorrhinus , Napeozapus in-

signis, Peromyscus maniculatus , Blarina brevicauda
Counties: 8, 22, 42, 46, 48, 50

Range within the state: Throughout the state where its host occurs
Epitedia ( cavernicola Traub?)

From Pennsylvania to Alabama, Epitedia cavernicola occurs as a nest para-
site of the eastern woodrat, Neotoma jlondana. We have examined one female
Epitedia from a woodrat, collected by Daniel Smiley in Ulster county. While
it does not agree perfectly with E. cavernicola, it is obviously not E. wenmanni,
so we tentatively assign it to this species until further specimens can be
secured. It is possible that the population of woodrats in eastern New York
is sufficiently isolated to have permitted the development of a distinct species
or subspecies of flea.

Host : Neotoma jloridana
County: 48

Range within the state: Unknown. Woodrats occur only in southeastern

counties, so far as known.

Epitedia jaceta (Rothschild)

True hosts: Glaucomys volans , G. sabrinus
Secondary host: Tamiasciurus hudsonicus
Accidental host: Mustela sp.

Counties: 23, 26, 33, 42, 48

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New York Entomological Society

Range within the state: Probably throughout the state; it is rarely col-
lected, being primarily a nest flea

Epitedia wenmanni wenmanni (Rothschild)

Some authorities have expressed doubt of the validity of the division of this
species into two subspecies, whose distribution is unlike that of any other
American form. However, the differences in the male genitalia are quite dis-
tinct, and both forms, along with intergrades, occur along a line at least to
the Rocky Mountains. In New York, the nominate subspecies occupies most
of the state, with E. w. testor occurring in Long Island, the Hudson valley
as far north as Albany county and Rensselaer county. The type locality of
testor is at Lansingburg, Rensselaer county, an unfortunate occurrence, since
intergrades are found only a few kilometers away.

True hosts: Peromyscus leucopus, P. maniculatus

Secondary hosts: Microtus pennsylvanicus , M. chrotorrhinus, Clethrionomys
gapperi , Napeozapus insignis, Parascalops breweri

Accidental hosts: Didelphis marsupialis, Mustela erminea, Sylvilagus sp.,

Urocyon ciner e oar gent eus

Counties: 1, 4, 5, 7, 8, 19, 20, 22, 23, 28, 34, 35, 44, 46, 48, 51

Range within the state: All of the state except the Hudson valley and its
tributaries and Long Island

Epitedia wenmanni testor (Rothschild)

True host: Peromyscus leucopus

Secondary and accidental hosts: Blarina brevicauda, Sorex jumeus, Clethri-
onomys gapperi , Glaucomys volans, Mustela sp.

Counties: 23, 25, 26, 48, 61

Range within the state: Hudson valley and valleys of its tributaries, and
Long Island

Corrodopsylla hamiltoni (Traub)

True host: Cryptotis parva

Accidental host: Microtus pennsylvanicus

Counties: 35, 42

Range within the state: The true host is known from the lake plain of
Lake Erie and Lake Ontario, as far north as Oswego county, from parts
of the Finger Lakes region, and from Long Island. The flea should be
expected throughout these areas.

Corrodopsylla curvata curvata (Rothschild)

True hosts: Shrews of the genus Sorex ; possibly also Blarina brevicauda

Accidental host: Zapus hudsonius

Vol. LXXXIII, September, 1975

149

Counties: 1, 5, 12, 23, 26, 48, 50

Range within the state: Probably throughout the state, though possibly

confined to higher elevations or colder sections

Ctenophthalmus pseudagyrtes pseudagyrtes Baker

This species appears to be completely non-specific in its choice of hosts,
occurring on virtually every mammalian species in the area. It shows some
preference for rodents and insectivores, carnivores being, perhaps, accidental
hosts.

Hosts: Sorex fumeus, S. cinereus, Blarina brevicauda, Condylura cristata,
Scalopus aquaticus, Parascalops breweri, Peromyscus leucopus, P. manicu-
latus, Clethrionomys gapperi, Microtus pennsylvanicus, M. chrotorrhinus,
M. pinetorum, Synaptomys cooperi, Ondatra zibethica, Erethizon dorsatum,
Tamias striatus, Tamiasciurus hudsonicus, Glaucomys volans, Rattus nor-
vegicus, N apeozapus insignis, Sylvilagus jloridanus, Mustela erminea, M.
frenata , Mephitis mephitis, Vulpes fulva

Counties: 1, 2, 4, 5, 7, 8, 9, 12, 17, 19, 21, 22, 23, 25, 26, 27, 28, 35,
36, 40, 42, 46, 48, 50, 53, 61

Range within the state: Throughout the state, though scarce or absent at
the highest elevations

Doratopsylla blarinae C. Fox

True host: Blarina brevicauda

Secondary and accidental hosts: Condylura cristata, Parascalops breweri, Sorex
fumeus, S. dispar, Microtus pinetorum, Peromyscus leucopus, Clethriono-
mys gapperi, N apeozapus insignis, Tamiasciurus hudsonicus

Counties: 1, 4, 5, 7, 8, 17, 19, 20, 22, 23, 25, 26, 27, 35, 36, 42, 46, 48,
50, 51, 61

Range within the state: Throughout the state
Nearctopsylla genalis genalis (Baker)

The taxonomy of this genus has been a point of disagreement for many
years, and many earlier records were referred to N. g. laurentina.

True hosts: Scalopus aquaticus, Parascalops breweri, Condylura cristata,

Blarina brevicauda

Secondary and accidental hosts: Sorex fumeus, Clethrionomys gapperi,

Synaptomys cooperi

Counties: 5, 8, 23, 25, 42, 48, 50, 61

Range within the state: Probably throughout the state. No specimens have

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New York Entomological Society

been taken in the western counties, but there are records from adjacent
counties in western Pennsylvania (Plolland and Benton, 1968).

Conor hino psylla stanfordi Stewart

Although the type specimen was found on the red squirrel, this species is
typically a parasite of the flying squirrels, Glaucomys volans and G. sabrinus.
Its rarity in collections is probably due to the fact that it is primarily a nest
flea, seldom staying on the host when it leaves the nest.

True hosts: Glaucomys volans , G. sabrinus

Secondary hosts: Sciurus carolinensis, Tamiasciurus hudsonicus

Accidental host: Vulpes julva

Counties: 19, 23, 42, 48

Range within the state: Unknown at present: its rarity in collections makes
it impossible to determine its range accurately

FAMILY CERATOPHYLLIDAE

Ceratophyllus diffinis Jordan

Jordan (1937) reported this species from “Long Island,” and Parkes (1954)
collected one from a robin in Hamilton county, but Geary (1959) was unaware
of these earlier records. In addition, we have seen two females from Essex
county, in the collection of the New York State College of Environmental
Science and Forestry, and one in our own collection taken from a deer mouse
in Franklin county.

True hosts: Hylocichla ustulata, Turdus migratorius
Accidental host: Peromyscus maniculatus
Counties: 2, 7, 8, 61 ( ? )

Range within the state: Unknown at present

Ceratophyllus gallinae (Schrank)

True hosts: G alius gallus, Passer domesticus, Troglodytes aedon
Accidental hosts: Tamias striatus, Canis familiaris, Homo sapiens , Rattus

norvegicus, Peromyscus maniculatus
Counties: 8, 12, 15, 17, 20, 29, 35, 42, 47, 48, 50, 53, 61
Range within the state: Throughout the state

Ceratophyllus celsus celsus Jordan

True hosts: Petrochelidon pyrrhonota , Riparia riparia
Counties: 21, 25, 26

Range within the state: Probably wherever cliff swallows nest, although

none were present in numerous nests from Essex county

Vol. LXXXIII, September, 1975

151

Ceratophyllus idius Jordan and Rothschild

True hosts: Progne subis, Iridoprocne bicolor
Accidental host: Troglodytes aedon

Counties: 2, 4, 25, 21, 35

Range within the state: Probably throughout the state

Ceratophyllus styx riparius (Jordan and Rothschild)

True hosts: Riparia riparia, Stelgidopteryx rufipennis

Secondary and accidental hosts: Hirundo rustica, Megaceryle alcyon, Sturnus
vulgaris

Counties: 2, 8, 18, 19, 21, 27, 35, 36, 42
Range within the state: Throughout the state

Megabothris acerbus (Jordan)

True host: Tamias striatus

Secondary host: Tamiasciurus hudsonicus

Accidental hosts: Napeozapus insignis, Microtus pennsylvanicus, Sciurus

carolinensis, Marmota monax, Sylvilagus floridanus
Counties: 1, 5, 7, 8, 12, 19, 20, 23, 26, 27, 36, 42, 48
Range within the state: The true host occurs throughout the state, but we
have made or examined extensive collections in several counties which
failed to produce this species. Apparently ecological factors limit its
distribution

Megabothris asio asio (Baker)

True host: Microtus pennsylvanicus
Secondary host: Microtus chrotorrhinus

Accidental hosts: Mustela erminea, Blarina brevicauda, Zapus hudsonius ,
Synaptomys cooperi, Clethrionomys gapperi, Sylvilagus floridanus, Vulpes
fulva

Counties: 1, 4, 8, 17, 19, 20, 23, 25, 26, 27, 35, 42, 50, 53, 61
Range within the state: Throughout the state

Megabothris quirini (Rothschild)

Although its host is common in suitable habitats throughout the state, this
species is unaccountably rare in western counties. The species was also absent
from extensive collections in western Pennsylvania (Holland and Benton,
1968). A single collection from a gray fox in Livingston county is the only
record from the western half of the state.

True host: Clethrionomys gapperi

Secondary and accidental hosts: Microtus chrotorrhinus, M. pennsylvanicus,

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New York Entomological Society

Peromyscus leucopus, P. maniculatus, N apeozapus insignis, Urocyon cin-
ereoargenteus

Counties: 5, 7, 8, 26, 30, 46, 48, 50

Range within the state: Would be expected throughout the state, but see
above

Monopsyllus vison (Baker)

True hosts: Tamiasciurus hudsonicus, Tamias striatus
Accidental hosts: Mustela vison, M. frenata, Procyon lotor, Peromyscus

maniculatus, Vulpes fulva

Counties: 1, 2, 5, 7, 8, 9, 19, 20, 21, 22, 23, 26, 33, 36, 42
Range within the state: Throughout the state except for Long Island and
the lower Hudson valley

Nosopsyllus jasciatus (Bose)

True host: Rattus norvegicus

Accidental hosts: Microtus pennsylvanicus, Mustela frenata, Vulpes fulva,
Tamiasciurus hudsonicus, Didelphis marsupialis
Counties: 19, 21, 25, 26, 35, 42, 56, 61
Range within the state: Throughout the state

Orchopeas caedens durus (Jordan)

This transcontinental species is a northern form, and has thus far been
taken only in the Adirondack counties in this state. It might reasonably be
expected in the high Catskills and in the Alleghenies, but no records are cur-
rently available from these areas.

True host: Tamiasciurus hudsonicus
Secondary host: Tamias striatus
Accidental host : Peromyscus maniculatus
Counties: 1, 2, 3, 4, 5, 6, 7, 8

Range within the state: Adirondack and Tug Hill counties at elevations
above 1000 feet

Orchopeas howardii howardii (Baker)

True hosts: Sciurus carolinensis, S. niger

Secondary hosts: Tamias striatus, Tamiasciurus hudsonicus, Glaucomys

volans, G. sabrinus

Accidental hosts: Blarina brevicauda, Procyon lotor, Urocyon cinereoar-

genteus, Mustela erminea, M. vison, M. frenata, Didelphis marsupialis,
Myocastor coypu, Marmota monax, Peromyscus maniculatus, P. leucopus,

Vol. LXXXIII, September, 1975

153

Synaptomys cooperi, Clethrionomys gapperi, Sylvilagus floridanus, Vulpes
julva , Urocyon ciner e oar gent eus

Counties: 1, 2, 4, 6, 7, 8, 9, 13, 19, 20, 22, 23, 25, 26, 33, 35, 36, 42,
45, 47, 48, 49, 50, 53, 56, 61

Range within the state: Throughout the state except at high elevations
where its principal hosts do not occur

Orchopeas leucopus (Baker)

True hosts: Peromyscus leucopus, P. maniculatus

Secondary and accidental hosts: Microtus pennsylvanicus, M. pinetorum,
Clethrionomys gapperi , Neotoma floridana, Zapus hudsonius, N apeozapus
insignis, Tamias striatus, Tamiasciurus hudsonicus, Marmota monax,
Sciurus carolinensis, Glaucomys volans, Didelphis marsupialis, Blarina
brevicauda, Mustela erminea, Urocyon ciner e oar gent eus, Sylvilagus flori-
danus

Counties: 1, 2, 3, 5, 7, 8, 12, 17, 18, 19, 20, 21, 22, 23, 25, 26, 27, 33,
34, 35, 36, 42, 46, 47, 48, 49, 50, 51, 53, 61
Range within the state: Throughout the state

Orchopeas sexdentatus pennsylvanicus (Jordan)

True host: Neotoma floridana
Counties: 48, 51

Range within the state: Those areas of southeastern New York where the
wood rat occurs

Opisodasys pseudarctomys (Baker)

True hosts: Glaucomys volans, G. sabrinus

Secondary and accidental hosts: Tamiasciurus hudsonicus, Marmota monax
Counties: 1, 6, 8, 21, 22, 23, 26, 42, 48

Range within the state: Throughout the state

Oropsylla arctomys (Baker)

True host: Marmota monax

Secondary and accidental hosts: Didelphis marsupialis, Vulpes fulva, Uroc-
yon ciner e oar gent eus , Mephitis mephitis, Canis latrans, Sylvilagus flori-
danus, Erethizon dorsatum, Tamiasciurus hudsonicus, Dama virginiana,
Homo sapiens

Counties: 1, 2, 3, 4, 8, 10, 17, 18, 19, 20, 21, 23, 24, 25, 26, 27, 30, 31,
32, 33, 35, 36, 39, 42, 45, 47, 48, 49, 50, 51, 52, 53, 61
Range within the state: Throughout the state

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FAMILY LEPTOPSYLLIDAE

Leptopsylla segnis (Schonherr)

The only records of this species are very old, and it seems unlikely that
there is any permanent population in the state, though it must often be brought
in by its hosts.

True host: Mus musculus
Secondary host: Rattus norvegicus
County: 56

Range within the state: Thus far collected only in New York City

Peromyscopsylla catatina (Jordan)

True host: Clethrionomys gap peri

Secondary hosts: Microtus chrotorrhinus , M. pennsylvanicus
Accidental hosts: Blarina brevicauda , Parascalops breweri
Counties: 1, 5, 7, 8, 21, 23, 26, 42, 45, 46, 48, 50

Range within the state: Probably throughout the state, although not yet

taken in western counties, where its host occurs

Peromyscopsylla hamijer hamifer (Rothschild)

True host: Syria ptomys coo peri
County: 1

Range within the state: Should occur on Microtinae throughout the state,
but evidently has precise ecological requirements (Miller and Benton,
1970)

Peromyscopsylla hesperomys hesperomys (Baker)

True hosts: Peromyscus leucopus, P. maniculatus

Secondary hosts: Microtus pennsylvanicus, M. chrotorrhinus, Clethrionomys
gap peri, Blarina brevicauda

Accidental hosts: Neotoma jloridana, Tamiasciurus hudsonicus
Counties: 4, 8, 12, 17, 19, 20, 22, 23, 26, 27, 35, 36, 40, 42, 46, 48, 50
Range within the state: Throughout the state except on Long Island, where
extensive trapping has not yet revealed it

Peromyscopsylla scotti (I. Fox)

True host: Peromyscus leucopus

Secondary and accidental hosts: Blarina brevicauda, Microtus pennsylvanicus
Counties: 25, 42, 48, 61

Range within the state: Long Island and the Hudson valley, with an

isolated record from Tompkins county; apparently has precise ecological
requirements, replacing the previous species in appropriate areas.

Vol. LXXXIII, September, 1975

155

Odontopsyllus multispinosus (Baker)

True host: Sylvilagus floridanus
County: Suffolk

Range within the state: Long Island, probably lower Hudson valley

FAMILY ISCHNOPSYLLIDAE

Nycteridopsylla chapini (Jordan)

True host: Eptesicus fuse us
County: 62

Range within the state: Unknown; its host occurs throughout the state,
but this flea occurs only in certain types of caves

Myodopsylla insignis (Rothschild)

True host: Myotis lueijugus

Secondary hosts: Myotis subulatus, Eptesicus juscus

Counties: 7, 11, 20, 23, 33, 36, 42

Range within the state: Throughout the state

Hypothetical List

The following species have been taken from states or provinces adjacent to
New York, on hosts which occur in New York. Thus it is reasonable to
expect that they may eventually be found within the state.

PULICIDAE

Hoplopsyllus glacialis lynx (Baker). Taken in Vermont from the snowshoe
hare, Lepus americanus

HYSTRICHOPSYLLIDAE

Rhadinopsylla orama Smit. Taken in Pennsylvania and Connecticut from the
pine vole, Microtus pinetorum

CERATOPHYLLIDAE

Ceratophyllus rossitensis swansoni (Liu). Taken in Ontario, Canada from
nests of the crow, Corvus brachyrhynchos

Ceratophyllus garei Rothschild. Taken in Quebec, Canada, from “eider down,”
which probably simply means a duck nest. It occurs in dry or bulky nests,
most often on the ground.

Literature Cited

Fox, Carroll, and E. C. Sullivan. 1925. A comparative study of rat-flea data for
several seaports of the United States. Publ. Health Repts., 40: 1909-1934.

156

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Fox, I. 1940. The fleas of eastern United States. Iowa State College Press, Ames, Iowa.
191 pp.

Geary, John M. 1959. The fleas of New York. Cornell Univ. Agric. Exp. Sta. Memoir
3 55: 104 pp.

Hall, E. Raymond, and Keith Kelson. 1959. The mammals of North America. The
Ronald Press Co., N. Y. Vol. 1 : 1-546; Vol. 2: 547-1083.

Holland, G. P., and A. H. Benton. 1968. Siphonaptera from Pennsylvania mammals.
Amer. Midland Naturalist, 80: 252-261.

Hopkins, G. H. E., and Miriam Rothschild. 1953-1971. An illustrated catalogue of the
Rothschild collection of fleas (Siphonaptera) in the British Museum (Natural His-
tory). Vol. 1: Tungidae and Pulicidae, 1953, xv -f- 361 pp.; Vol. 2: Vermipsyllidae
to Xiphiopsyllidae, 1956, xi -f- 560 pp.; Vol. 3: Hystrichopsyllidae (in part), 1961,
ix + 560 pp.; Vol. 4: Hystrichopsyllidae (in part), 1966, viii + 549 pp.; Vol. 5:
Leptopsyllidae and Ancistropsyllidae, 1971, viii + 530 pp. British Museum Nat. Hist.,
London.

Jordan, Karl. 1929. On a small collection of Siphonaptera from the Adirondacks with
a list of the species known from the state of New York. Novit. Zool., 35: 168-177.

. 1937. On some North American Siphonaptera. Novit. Zool., 40: 262-271.

Miller, Donald H., and Allen H. Benton. 1970. Ecological factors in the distribution
of Peromyscopsylla h. hamifer (Rothschild). Amer. Midland Naturalist, 83: 301-303.

Parkes, Kenneth C. 1954. Notes on some birds of the Adirondack and Catskill Moun-
tains, New York. Annals Carnegie Mus., Art. 7: 178 pp.

Sakaguti, K., and E. W. Jameson, Jr. 1962. The Siphonaptera of Japan. Bernice P.
Bishop Mus., Pac. Insects Monograph 3 : 169 pp.

Stewart, M. A. 1928. Siphonaptera. In “A list of the insects of New York,” by M. D.
Leonard. Cornell Univ. Agric. Exp. Sta. Memoir 101: 1868-1869.

. 1933. Revision of the list of Siphonaptera from New York State. J. N. Y.

Entomol. Soc., 41: 253-262.

Zeh, John B. 1973. A survey of the ectoparasitic fauna of the red fox, Vulpes fulva
(Desmarest) and the gray fox, JJrocyon cinereoargenteus (Schreber). Unpublished
thesis, State Univ. College, Geneseo, New York: 71 pp.

Vol. LXXXIII, September, 1975

157

Notes on the Life Cyele and Natural History of Butterflies of
El Salvador. I B. — Hamadryas februa (Nymphalidae-Hamadryadinae)

Alberto Muyshondt and Alberto Muyshondt, Jr.

101 Ave. N. 322, San Salvador, El Salvador

Received for Publication August 23, 1974

Abstract: Observations carried on in the neighborhood of San Salvador since 1970 on

eggs, larvae, pupae and adults of Hamadryas februa Hiibner are presented, giving an ac-
count of the early stage characteristics and developmental times, with photographic illus-
trations. The foodplants of this and related species are recorded for El Salvador. The
behavior of the species is compared with the behavior of other local and South American
Hamadryas spp. emphasizing the progressive change from solitary to gregarious behavior
which the whole group exhibits, with the corresponding adaptations that such a change
requires. Impalatability of the species to predators is suggested by the larval foodplant
characteristics and the typical non-palatable way the larvae behave.

This is the first article of a third series in which we present our observations
on the early stages and adult behavior of butterflies of El Salvador, Central
America. Elsewhere a first series has been presented dealing with the local
Charaxinae, and in this same journal a second on the Catonephelinae-Calli-
corinae, all of them subfamilies of the Nymphalidae. Even though there have
been earlier descriptions and some illustrations of the early stages of species
belonging to this group, e. g. Muller’s (1886) and Friihstorfer’s (1916), we
expect our contribution will be of interest, since new elements are presented.

Our studies on this species started on August 1st, 1970, when one of us
(A. M., Jr.) while walking down a road bordering pasturelands, observed a
female Hamadryas februa Hiibner ovipositing on a vine, near the village of
Zaragoza (some 15 km SSW of San Salvador). Some eggs were collected and
eventually two adults were obtained the 1st and 2nd of September. Since then
the species has been reared from the egg a number of times. The eggs and
larvae have been put in transparent plastic bags, fresh leaves of the foodplant
have been supplied every three days, and the bags cleaned of old leaves and
frass every day until pupation. The pupae were transferred to a mosquito-net
covered cage until adults emerged. Measurements of each stage and the times
elapsed in each phase were recorded, and photos taken of the whole process.

Acknowledgments: We are deeply grateful to Dr. Alexander B. Klots for dedicating his

time to read and criticize this paper and for giving his valuable advice to improve it. We
are thankful also to Dr. A. H. B. Rydon for the wealth of information obtained from
his correspondence and from the reference material kindly supplied by him. Our gratitude
also to Dr. F. D. Rindge, of the American Museum of Natural History, New York, for de-
termining many of the species mentioned and to Dr. C. W. Sabrosky, of the USDA, for
determining the tachinid parasite.

New York Entomological Society, LXXXIII: 157-169. September, 1975.

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During the development of the larvae, and during pupation, the bags and cage
were kept at all times under ambient light and temperature conditions. Sam-
ples of eggs, larvae in the different instars and pupae have been preserved in
alcohol to be sent to the American Museum of Natural History, New York,
where the adults were determined.

LIFE CYCLE

Egg. Roughly spherical with flattened base and irregular sculpturings ; white when recently
deposited, darkening before hatching in 5 days. About 1 mm.

First instar larva. Head faintly cordiform, naked, shiny black. Body cylindrical, brown with
scattered white tubercles, legs and prolegs dark brown. About 2.5 mm when recently
hatched, 4 mm in 3 days when ready to moult.

Second instar larva. Head shiny black with tiny white spines on anterior and lateral areas
of epicrania, sparse short setae on frons and thick, short horns on epicranial apices. The
horns are armed basally by 4 tiny spines, and thicken distally. Body brown with transverse
rows of furcated spines, very short and alternately dark and light colored. About 7 mm
before moulting in 3-5 days.

Third instar larva. Head black with two long black spines laterally and several light and
short spines anterad and between the base of the long and slender horns (about three times
as long as head), which show basally two short accessory spines directed forward, two
longer spines a little higher on the horn shaft and directed outwards, and about the middle
of the horn shaft still two other longer spines directed inwards. The horns are each armed
distally with a spiny sphere with short setae. Body black with a longitudinal, broken,
orange stripe supraventrally, and a profusion of black, forked spines arranged in the
following order: on first thoracic segment (T-l) a black cervical shield with two short

white spines, one long black forked spine supraspiracularly and a shorter black simple spine
subspiracularly ; on T-2, a 5-forked black spine subdorsally, a 4-forked supraspiracular spine
and a simple spine supraspiracularly; on T-3 a prominent 5-forked subdorsal spine, a 4-
forked supraspiracular spine and a simple spine subspiracularly. On first abdominal segment
(A-l) a dorsal 3-forked spine, a subdorsal 4-forked spine, a supraspiracular simple spine,
a subspiracular 3 -forked spine, a supraventral simple spine and finally a simple spine in
line with legs and prolegs; A-2 presents one 5-forked dorsal spine, a 5-forked subdorsal
spine, a 3-forked supraspiracular spine, a 3-forked subspiracular spine sided by a simple
spine, supraventrally one simple spine and two simple spines in line with prolegs. From
A-3 to A-6, a 3-forked spine dorsally, a 4-forked subdorsal spine, a 3-forked supraspiracular
spine, a 3 -forked subspiracular spine sided by a simple spine, and a row of 3 simple spines
over the proleg. A-7 as A-6 but with two dorsal forked spines (one behind the other),
the first one similar to the preceding ones, the second twice as big and 4-forked. A-8, as
A-6 but with only one dorsal, prominent, 5-forked spine deflected posterad. A-9 has only
two lateral 5-forked spines directed posterad. Grows to 12 mm in 3-4 days.

Fourth instar larva. Head as in third instar with longer horns. Body as in third instar also,
but with several yellow, longitudinal, broken lines dorsally and subdorsally, and orange
spots between dorsal and subdorsal spines. Subspiracular and supraventral spines light
colored, the rest black. Grows to 18-20 mm in 4-5 days.

Fifth instar larva. Head, when recently moulted, red, turning usually to black after a time.
Body mostly black with light green spines, and six yellow, longitudinal, thin stripes from

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159

Figs. 1-6. Hamadryas februa Hiibner. 1. Egg, about 1 mm. 2. First instar larva, 3.5 mm.
3. Second instar larva, 6 mm. 4. Third instar larva, 12 mm. 5. Fourth instar larva, 18 mm.
6. Fifth instar larva, 30 mm.

thorax to abdominal tip, located one at either side of each row of spines; two orange spots
at either side of abdominal median spine. At times during this instar, some individuals
keep the head red permanently. When this occurs, body shows a reduction of the basic
black color, which is substituted by orange. Subdorsal spines prominent, especially T-2, T-3
and A-2, where they have developed a number of small accessory spines on the shaft of
the scolus. The same happens on the dorsal spines A-7 and A-8. Grows to 30-32 mm in
4-6 days.

Pre-pupa. Considerably shorter than 5th instar and showing a notable discoloration of the
darker shades. In the case of the “orange” morph, becomes almost solid orange. Hangs
incurved during 1 day.

Pupa. Variable in color, depending on individuals: some are light brown, some, greenish-
brown, some, reddish-brown, with darker touches dorsally and fine, vein-like lines of darker
shade ventrally on wingcases. The body thickens gradually from the flat cremaster to the
posterior end of the wingcases, which point is the thickest part of the body, then has a
dorsal and lateral concavity, then thickens again laterally and forms a slightly keeled hump
dorsally. From there the body narrows down to the head, which terminates in two flat,
partly fused, then diverging, prolongations, which follow the longitudinal axis of the body,

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Figs. 7-8. Hamadryas februa Hiibner. 7. Pupa, lateral view, 30 mm. long. 8. Pupa,
ventral view.

being about % of the total length of the body. Ventrally three small warts are noticeable
along either side of the antenna-cases. Measures about 30 mm long by 8.5 mm laterally
and 7.5 mm dorso-ventrally at widest points. Durations 6-9 days.

Adults. No sexual dimorphism has been noticed in this species. Forewing shape: costal

margin slightly convex, rounded apex, almost straight but sinuose outer margin directed a
little inwards, rounded tornus and straight inner margin. Hindwing: almost straight costal
margin, rounded outer angle continuing in a convex and faintly sinuose outer margin,
rounded anal angle and straight, folded inner margin.

Color dorsally, dominantly light grayish-brown, lighter towards forewing apical zone,
with darker brown, sinuose, broken lines and a few circles, forming all a very complicated,
practically undescribable pattern. Ventrally mostly whitish-gray with fewer dark brown
markings and circles in both wings distally, more so in front wings. Faint orange and
yellow bordering basally the crescent shaped spots in the two circles closer to anal angle.
Wingspan averages 65 mm in males, 72 mm in females. Complete metamorphosis took
from 29 to 38 days.

NATURAL HISTORY

The females of Hamadryas februa search for the foodplant along low brushed,
open land, flying close to the ground in the neighborhood of wooded areas,
until one vine is located. A mature leaf is chosen where the female alights,
usually on its underside, and one egg is deposited around the middle of the

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Figs. 9-12. Hamadryas februa Hiibner. 9 and 10. Male, dorsal and ventral sides.
11 and 12. Female, dorsal and ventral sides. Black bar 10 mm.

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Figs. 13-17. Hamadryas jebrua Hiibner. Sequence showing phases of emergence of adult.

leaf. Several eggs are thus deposited on the same plant. It is not unusual
that two eggs are laid one on top of the other by the same female (and
eventually three), but never have we seen the same female deposit two eggs
side by side on the same leaf. In some instances the female alights on top of

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Figs. 18-20. Hamadryas februa Hiibner. Sequence showing gradual expansion of wings
following emergence of adult.

a leaf to oviposit. When this occurs, the female moves near the edge of the
leaf, and incurving the abdomen, deposits one egg on the underside of the
leaf, close to the border.

The tiny larvae, upon hatching, eat from the eggshell just an exit hole,
located at the side of it, and leave the rest untouched. They then move to the
edge of the leaf and start feeding around a vein (usually at the apex), baring
it. The larvae prolong the vein affixing to it their own small frass pellets using
their silk as agglutinant. Quite often first, and eventually second instar larvae,
are found with several excrement pellets adhering to their own bodies. This
behavior might serve a double purpose: camouflage and reserve of materials
to lengthen the resting vein when needed. First and second instar larvae are
seen through the daylight period resting on their prolonged veins, motionless,
head pointing outwards. They only move back to the leaf, late in the afternoon
to feed on it. From third instar on, the larvae abandon their perch and move
slowly about the plant, on the upper surface of the leaves, weaving a foothold
of silk as they crawl, but staying motionless most of the time, with the third
thoracic segment humped and the head bent so as to place the horns parallel
to the leaf surface. It is not unusual to find more than one larva on the same
leaf, but no interaction has been noticed even when they accidentally come in
contact. If the larvae are prodded with a blunt object, or teased with a thin
flexible one, they end by striking with their horns. When ready to enter
pupation, which is announced by a shortening and decoloration of the body,

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the larva weaves a mat of silk usually around a thin twig of the foodplant,
but at times under a leaf, cleans the digestive tract and hangs from the silken
mat by its anal prolegs with head and thorax incurved. After a day the larval
skin splits dorsally behind the head and after many body contractions the
larval skin is shed and the cremaster anchored firmly by actively wiggling. At
first the capital prolongations are limply apposed to the thorax, but after a
few minutes they expand forward and position themselves along the longi-
tudinal axis of the body, fusing together their midpart, separated at the base
and diverging from each other at the tips. The pupae, usually hanging straight
down, wiggle laterally and violently when molested. Their wiggling lasts a few
seconds and they then stay bent to one side or the other for a long period of
time afterwards. The cremaster of this species (and other related species of
the genus), has a flattened base, somewhat as in Catonephelinae, so that it is
rigidly applied to the silk button. When the twig is turned upside down, the
pupa turns with it staying standing upwards, straight or bent to one side.
The day the adult is ready to emerge, the pupa becomes dark gray and the
pattern of the wing colors is visible through the pupashell.

The adults rapidly emerge from the pupashell and hang from it ejecting a
reddish meconium and unfolding their wings, which are held close together
until rigid. It is after their first flight that they learn to keep them spread
open most of the time. The adults of Hamadryas jebrua fly actively in, or in
the neighborhood of, wooded areas, from about 500 to 1500 m altitude mostly,
alighting on tree trunks with their wings spread open and tightly hugging the
tree surface, where their gray and brown complicated pattern merges perfectly
with the lichen growths which spatter the bark. Males can produce a peculiar
loud clicking sound as they fly when meeting other males in flight near their
chosen tree, and after an interchange of excited and repeated click-clicks plus
many swift circumvolutions, without actually coming in contact, the intruder
is chased and the defendant goes back to rest on his lookout, most commonly
with the head down. This clicking sound, combined with the aggressive flight,
is used also to pursue other approaching butterflies of diverse species, and
even dogs, as per repeated observations of one of us (A. M., Jr.). The adults
in our cages keep their wings folded at night. In the fields we have seen indi-
viduals resting among shrub leaves with their wings folded also, but when
alerted by the sound or the shadow of the observer they immediately spread
them open. When the adults come to the ground to feed on fermenting fruits,
(mangoes, guayavas, rubber-tree fruit, etc.) they sometimes keep their wings
open, but at other times the wings are folded. When they feed at wounds on
tree-trunks, the wings are always kept spread. We have not noticed the males’
clicking sound when they court females, although there is a lot of chasing
around.

The preferred local larval foodplant, Dalechampia scandens L. and the very

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165

seldom used alternate foodplant, Tragia volubilis L. are scandent vines belong-
ing to the Euphorbiaceae, and both have urticant properties when touched
with the back of the hand. Either plant is used by other local species of but-
terflies as larval food. Dalechampia besides at least two other species of
Hamadryas, (probably all of them), is used by Dynamine spp., Catonephele
nyctimus Westwood and Mestra amymone Menetries. Tragia is used by Biblis
hyperia Cramer, Mestra amymone and Dynamine spp.

The Dalechampia scandens vine has coarse, slightly pubescent, alternate,
petiolate, tri-lobate and cordate at the base of the leaves; the flower is monoe-
cious, apetalous, with 3 long stemmed pistils surrounded by a host of short
stemmed stamens, inside two tri-lobed bracts. The schizocarpous fruits split
into three carpels, each bearing one black and brown seed. The fruits are
surrounded by long and thin segments covered by a profusion of sharp, fulvous
spinulets, which penetrates the skin at the faintest touch.

The Tragia volubilis vine shows alternate, petiolate, conspicuously dentate,
densely pubescent, shallowly cordate, long leaves. The flowers grow in racemes,
and are small, apetalous and produce 3-lobate capsules.

Both plants grow on sunny ground, in low brushed habitats near ravines,
road fences and abandoned pieces of land, mostly near wooded areas or coffee
plantations. We have found them from about 500 m to about 1500 m altitude.
People use them for folklore medicine.

DISCUSSION

The butterflies pertaining to this group have been called at different times
by different authors Peridromia (Boisduval, 1836), Ageronia (Hiibner, 1810)
and Hamadryas (Hiibner, 1806), as per information supplied by Dr. A. H. B.
Rydon (personal communication). We use the last, which is the older of the
three, even if according to Hemming (1967) all three are available generic
names.

We are aware of at least two authors describing the early stages of several
species belonging to this group (Muller, 1886 and Friihstorfer, 1916), under
the name of Ageronia. They mention as foodplants for the species they de-
scribe, other species of Dalechampia. Still, to our knowledge, this is the first
description with photographic illustrations of Hamadryas februa.

Modern authors usually lump the genus Hamadryas within the Nymphalidae,
whether in the subfamily Ergolinae (Klots, 1960); in the tribe Ergolini of the
subfamily Nymphalinae (Ehrlich & Ehrlich, 1961); or under the subfamily
Hamadryadinae (Ebert, 1969). If we were to follow the rule of thumb, which
uses egg characteristics to separate the families, we would advocate a family
status for this group, as was done, using various nominations, by several
authors: Seitz (1915), group K, Ageronidae, which was first used by Double-
day in 1847, and Peridromiidae used by Boisduval in 1836, the latter thus

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having priority. The egg shape of the butterflies of the group is quite different
from any other egg of the local butterflies. The larvae also are easily recog-
nized, from the third instar on, just by examining the horns of their head.
The pupae, even if there are differences in the shape of the head prolongations
and the colors between the several species we have seen, are easily identified as
Hamadryas upon seeing them on account of their peculiar shape.

The females of H. jebrua and its close relative H. guatemalena have the
same ovipositing behavior: they usually lay one egg at a time, but quite often
two, and seldom even three are deposited one on top of the other. According
to Muller (1886) (observations on one undetermined species of Ageronia and
on A. arete Doubleday) some species deposit one egg exclusively at a time.
His own observations on A. fornax were that this species forms a single string
of several eggs, one under the other, so that the whole string hangs perpen-
dicularly from the underside of the leaf. Our own observations (and Muller’s
as well) on H. amphinome show that the eggs of this species are also deposited
on long strings, forming groups of 3 to 6 strings. It is our feeling that this
group of species ( Hamadryas spp.) is a living example of how natural selection
operates: some of the species still have solitary behavior {arete) ; some are
in the very process of changing from solitary to gregarious behavior {jebrua
and guatemalena ); one has partially adopted it, forming small groups of 5 to
10 individuals {fornax) and another {amphinome) has fully adopted gregari-
ousness, with groups up to 45 individuals. Ford (1945) states: “the gregarious
habit, determined by the method of egg-laying, is exceptional and carries with
it certain noteworthy adaptations.” We feel that before the egg-laying method
can lead the species to gregarious behavior, there must be a prerequisite sine
qua non in the larval behavior: the larvae must first abandon the habit of
devouring the eggshell upon hatching. If that condition did not pre-exist to
the oviposition in clusters, the first individual hatched could destroy at least
some, if not many, of the adjacent eggs of the same group, as is the case in
solitary species of Lepidoptera, such as Heliothis zea , when several females
deposit one egg each on the silks of a single corn-ear and the first larva hatched
dispose of many eggs close by and even later larvae, (personal observations).
We have noticed that this necessary trait of eating just an exit hole from the
eggshell is existent in all the local species with gregarious behavior that we
have reared: Battus polydamas L., B. laodamas Felder, Papilio anchisiades
idaeus Fabr. (Papilionidae) ; Chlosyne spp., Phyciodes spp., Microtia elva
Bates, Thesalia theona Menetries (Melitaeinae) ; Actinote spp. (Acraeinae) ;
Mechanitis isthmia isthmia Bates (Ithomiidae) ; Caligo memnon Felder
(Brassolidae) ; Manataria maculata Hopff. (Satyridae ?); Dione juno hua-
scama Reakirt (Muyshondt, Young & Muyshondt, 1973) (Heliconiidae) ;
Gynaecia dirce L. (Coloburinae) ; Theritas lisus (Stoll) (Lycaenidae) and
Hamadryas amphinome L. In all of these species the hatching larvae eat

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167

just an exit from the micropylar zone of the egg. The exception is H . am-
phinome, which cuts it from the side of the egg, which is of great importance
as the eggs are deposited one on top of the other, not side by side as the
others do. The same characteristic of not eating the eggshell has also been
adopted by several species, which, if not strictly gregarious, have learned to
live in a loose community without bellicose interaction, as a result of the need
of ovipositing on such restricted areas of the plant as the tender shoots. Even
if the females lay only a single egg per location, several females often visit
the same plant and thence a number of eggs from several females are accu-
mulated within a very small area. In this group we mention: Dircenna klugii
Geyer, Hyposcada virginiana nigricosta Forbes & Fox, Tithorea harmonia
salvadores Staudinger, Hymenitis oto oto Hewitson (Ithomiidae) ; Narope
cyllastros testaceae Godman & Salvin (Brassolidae), all the local species of
Heliconiidae (except Dione juno , which is gregarious), all the local species of
Danaidae (except Anetia tkirsa , which we have been unable to rear) ; Precis
genoveva Stoll, Anartia jatima Fab., A. jatrophae luteipicta Friih., Hypanartia
lethe Fab. (which at times deposits one egg on top of another also, as per
own observations) (Nymphalidae) ; Biblis hyperia Cramer (Biblinae); Mar -
pesia spp. (Marpesiinae) ; Pyrrhogyra hypsenor Godman & Salvin (Muy-
shondt, 1974 a) (Catonephelinae) ; Dynamine spp. and several species of
Adelpha (some Adelpha very occasionally deposit one egg on top of another
also) (Limenitinae), and many species of Pieridae and Lycaenidae. All of
these eat the exit hole from the eggshell, and at the most consume a small
portion of its wall. Still there are other species which do not devour the egg-
shell, but have not acquired the tolerance towards other individuals and main-
tain strict individualistic behavior as larvae, fighting when coming in contact
with a neighboring one, with often fatal results to one or the other.

The adaptation of not feeding on the eggshell mentioned, is more remarkable
in Hamadryas februa, H. guatemalena, Hipanartia lethe and some Adelpha sp.
where two or more eggs are deposited one on top of the other, as they make
the exit hole through the side of the eggshell, which is very important to pre-
vent damage for the egg on top. When two or three eggs are thus laid, the
ensuing larvae might at times be found living in the same leaf, even if each
one has made its own perch, until the last instar. However they pupate
separately.

The habit of baring a vein and prolonging it with excrement pellets (a
characteristic shared with several other Nymphalidae) is shown only by the
species of Hamadryas which deposit the eggs singly (occasionally in twos and
threes). This habit has been lost by the gregarious Hamadryas , which have
developed in place of the cryptical defense against predation that characterizes
individualistic species, a rather unpleasant odor, enhanced by the multitude of
larvae living in community, plus an irritable temperament which causes the

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whole congregation to wiggle convulsively at the faintest provocation. These
peculiarities are evidently some of the “noteworthy adaptations” Ford (1945)
refers to, that the gregarious species have gone through.

The need to conceal themselves shown by first and second instar larvae of
H . februa as a defense against predation seems to cease being indispensable
to their survival when they reach the third stadium, as evidenced by the
desertion of their resting place and crawling openly on the leaves. This un-
concerned exposure of themselves seems to indicate that a new means of
protection against predation has been acquired in the meantime. Since their
foodplants are Dalechampia scandens (and Tragia volubilis ) Euphorbiaceae,
which are renown as containing considerable amounts of fluids with poisonous
or very caustic properties in many cases, it would not be too daring to deduce
that H. februa larvae derive from their foodplant predator-deterrent qualities,
which take some time to become effective (the time to reach third stadium).
Another factor sustaining this notion is the fact that the species is subject to
massive parasitism, mostly by Tachinidae. Muyshondt (1973 a, b, 1974, 1975)
has pointed out the apparent relationship which exists between parasites and
chemically protected larvae, and the behavior of the early stages and adults.
Specimens of one tachinid sent to the USDA was determined by Dr. C. W.
Sabrosky as “Tachinidae — Gen. sp. — intermediate Eryciinae and Sturniini” with
the comment: “Odd species!!”. We have obtained up to 4 parasites out of a
single larva or pupa.

Even if these factors suggest impalatability, the adult color pattern can be
considered a very effective camouflage when they are perched on the sides of
tree-trunks with the wings spread open. They seem to have thus a dual de-
fense mechanism: chemical and cryptic.

One of Muller’s observations (mentioned also by Seitz) has failed to prove
true in our experience. Muller states that the pupae of several Ageronia-
Hamadryas in his experience are light-sensitive, and would bend their bodies
to one side or the other depending on the direction of the sunlight. We have
tried several times, at different hours of the day, to induce a reaction from
many pupae of various species of Hamadryas, including februa, exposing them
to direct sunlight, then masking the light, for various periods of time, with
negative results. What we have noticed is that upon the slightest disturbance
the pupae violently wiggle laterally for a short time, and then stop moving
with the body bent to one side or the other, which position is kept for long
periods of time afterwards.

Literature Cited

Ebert, H. 1969. On the frequency of butterflies in Eastern Brasil with a list of the
butterfly fauna of Poqos de Caldas, Minas Gerais. Jour. Lep. Soc., 23, Suppl. 3.
Ehrlich, P. R. and A. H. Ehrlich. 1961. How to know the butterflies. Wm. C. Brown
Co. Publishers. Dubuque, Iowa.

Vol. LXXXIII, September, 1975

169

Ford, E. B. 1945. Butterflies. Collins, London.

Fruhstorfer. 1916. In Seitz’s Macrolepidoptera of the World. Vol. 5. Stuttgart.
Hemming, F. 1967. Generic names of the butterflies and their type-species. Bull. Br.
Mus. Nat. Hist. (Ent.), Suppl. 9.

Klots, A. B. 1960. A field guide to the butterflies. Riverside Press. Cambridge, Mass.
Muller, W. 1886. Siidamerikanische Nymphalidenraupen. Versuch eines naturlichen
Systems der Nymphaliden. Zoologische Jahrbuch. 453-461.

Muyshondt, A. 1973a. Notes on the life cycle and natural history of butterflies of El
Salvador. Ill A. — Temenis laothde liberia Fabricius (Nymphalidae-Catonephelinae) .
Jour. N. Y. Ent. Soc. 81: 224-233.

. 19736. Notes on the life cycle and natural history of butterflies of El Salvador.

IV A. Pseudonica flavilla canthara (Nymphalidae-Catonephelinae). Jour. N. Y. Ent.
Soc. 81: 234-242.

. 1974. Notes on the life cycle and natural history of butterflies of El Salvador.

V A. — Pyrrhogyra hypsenor (Nymphalidae-Catonephelinae). Jour. N. Y. Ent. Soc.
82: 163-172.

. 1975. Notes on the life cycle and natural history of butterflies of El Salvador.

VI A. — Diaethria astala Guerin (Nymphalidae-Callicorinae) . Jour. N. Y. Ent. Soc. 83:
10-18.

, A. M. Young and A. Muyshondt, Jr. 1973. The biology of the butterfly Dione

juno huascama (Nymphalidae-Heliconiinae) in El Salvador. Jour. N. Y. Ent. Soc.
81: 137-151.

Seitz, A. 1915. Macrolepidoptera of the World. Vol. 5. Stuttgart.

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Notes on the Life Cycle and Natural History of Butterflies of
El Salvador. II B. — -Hamadryas guatemalena Bates
( Nymplialidae-Hamadryadinae )

Alberto Muyshondt and Alberto Muyshondt, Jr.

101 Ave. N. #322, San Salvador, El Salvador

Received for Publication October 1, 1974

Abstract: Observations on the adults and early stages of Hamadryas guatemalena Bates
have been carried out in the vicinity of San Salvador, El Salvador, for a period of 4 years.
In this article the results are presented for the first time, with a detailed account of the
life cycle, illustrated with photographs, of the larval behavior and the plant used as food.

The characteristics of the species are compared with the characteristics of other closely
related species. The contention that there should be several genera within the group is
discussed.

As in other Nymphalidae, in this species the gaudy coloration and daring behavior of the
larvae, and the use of a foodplant belonging to the Euphorbiacae ( Dalechampia scandens L.)
reputedly poisonous, suggest impalatability of the adults.

This is the second article of a series dealing with butterflies belonging to
the genus Hamadryas, found in El Salvador. In this article we give an account
of our observations on the early stages and adults of Hamadryas guatemalena
Bates carried on since August 1970 in various zones of the country, mostly
within 15 km from the capital city San Salvador. The first time we found
larvae of this species was shortly after we started studies on a close relative,
H. jebrua Hiibner, during August 1970. As both species feed on the same
plant, we ended up studying the two species simultaneously, which caused at
first some confusion, as eggs collected produced at times two different kinds
of larvae. The problem was solved when a female of Hamadryas guatemalena
was observed ovipositing also on the same plant as H. jebrua . It is practically
impossible to tell apart the eggs of one species from the other. As usual,
eggs were collected just after oviposition and put in transparent plastic bags
fastened with rubber bands. Emerged larvae were fed on fresh leaves of the
foodplant replaced every three days until pupation. The bags were cleaned
every day of excrement and excess humidity. The pupae were transferred to
a wooden cage with mosquito-net covering, where the adults emerged. Bags
and cage were kept indoors at all times under ambient light and temperature

Acknowledgments: We express our gratitude to Dr. Alexander B. Klots, for the great
help he has given us to make this article presentable, criticizing our manuscript and suggest-
ing valuable improvements to it. We are also thankful to Dr. A. H. B. Rydon and Col.
C. F. Cowan for their information on this group of butterflies and for supplying reference
literature; to Dr. F. D. Rindge of the American Museum of Natural History for kindly
determining the adults of H. guatemalena.

New York Entomological Society, LXXXIII: 170-180. September, 1975.

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conditions. Notes were kept of the measurements and the duration of each
phase of the metamorphosis. Specimens in alcohol were sent to the American
Museum of Natural History, New York, where the adults were determined.

LIFE CYCLE STAGES

Egg. Pure white, almost round with small flat base and sculpturings starting basally with
thick ribs which disappear about a third from the base and are substituted by irregular,
rounded or sharp protuberances covering part of the wall and the micropylar area. About
1 mm diameter. Hatches in 3-5 days.

First instar larva. Head shiny black, slightly cordiform, naked. Body cylindrical, greenish-
brown with lighter tubercules and sparse short setae. Legs and prolegs dark brown. About
3 mm when recently hatched, about 4 mm before moulting in 3 days.

Second instar larva. Head black with small white spines on lateral margins and frontal
area. Short thick horns on apices of epicrania. Body dark brown with longitudinal rows
of furcate short spines and four rows of white dots, two subdorsally and two supraspiracu-
larly. About 7 mm long before moulting in 4 days.

Third instar larva. Head black with long and slender horns on epicrania, two spines be-
tween their bases, three long spines on lateral margin of head and several short spines
frontally; ocelli black, surrounded by sparse, short golden setae. The horns have basally
on the shaft two accessory spines directed forward, a little higher two spines directed later-
ally and about the middle of the shaft two more spines directed inwards; the horns are
tipped by a sphere armed with tiny spines. The body’s ground color is black with longi-
tudinal rows of yellow spots subdorsally and supraspiracularly. The spine arrangement is
as follows: on first thoracic segment (T-l): one bifurcate subdorsal spine, one bifurcate
supraspiracular spine and one simple subspiracular spine ; on T-2 : one prominent subdorsal
6-furcate spine, and 6-furcate supraspiracular spine, one small spiracular simple spine and
one longer spine subspiracularly ; on T-3 : one most prominent 6-furcate subdorsal spine,

one 5-furcate supraspiracular spine, one small, spiracular simple spine and two simple spines
subspiracularly. On first abdominal segment (A-l): one 4-furcate subdorsal spine, one sim-
ple supraspiracular spine, one 4-furcate subspiracular spine and two supraventral simple
spines. On A-2: one prominent 5-furcate subdorsal spine, one 3-furcate supraspiracular spine,
one 4-furcate subspiracular spine sided by a simple spine, 3 supraventral simple spines. From
A-3 to A-6: one 5-furcate subdorsal spine, one 3-furcate supraspiracular spine, one 4-furcate
subspiracular spine sided by a simple spine and 2 simple spines supraventrally. On A- 7 one
very prominent dorsal 3-furcate spine, one 6-furcate subdorsal spine, one 3-furcate supra-
spiracular spine, one 5-furcate subspiracular spine sided by a simple spine and two simple
supraventral spines. On A-8: one prominent 6-furcate dorsal spine, one prominent 6-

furcate subdorsal spine, one 3-furcate supraspiracular spine, one 5-furcate subspiracular
spine sided by a simple spine and two simple supraventral spines. On A-9: one 8-furcate
subdorsal spine deflected caudad. On A- 10: two simple spines, side by side, on anal plate.
Grows to 1.3 cm in 4 days.

Fourth instar larva. Head as in third instar, with longer horns. Body ground color black
with light yellow, very conspicuous dorsal oval patches forming an irregular and broken
stripe from T-l to A-9, and two supraspiracular light yellow dots on each abdominal seg-
ment. The shafts of the prominent subdorsal spines on T-2, T-3, A-2 and A-8, and of the
median spines on A- 7 and A-8 are armed by a host of small brown spinulets directed dis-
tally. Grows to 2.5 cm in 3-4 days.

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Figs. 1-8. Hamadryas guatemalena Bates. 1. Two eggs side by side. Notice the left is
grayish, the other pure white. One hatched one day before the other. 2. Two eggs one on
top of the other. Both hatched the same day. 3. Eggshell showing the exit hole on the
side. 4. First instar larva. Notice frass pellets stuck on the body. 5. Second instar larva.
6. Third instar larva. 7. Fourth instar larva. Notice spinulets on some spines. 8. Fifth
instar larva.

Fifth instar larva. The only change is that the body markings become bright deep yellow,
and the horns on the head and spines on the body are dull yellow. Prominent subdorsal
spines on T-2, T-3, A-2 and A-8 and dorsal spines on A-7 and A--8 look “hairy” due to
the profusion of dark accessory spines on the shaft of the scoli. Grows to 4.2 cm in 4-5
days.

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Figs. 9-11. Hamadryas guatemalena Bates. 9. Pupa, dorsal view. 10. Pupa, side view.
11. Pupa, ventral view.

Prepupa. Does not change in aspect, but for slight shortening of the body. Hangs from
anal prolegs, with thorax incurved ventrally, for one day.

Pupa. Hangs rigidly anchored from flat cremaster. Abdomen thickens abruptly from cre-
master and then gradually to base of wings, then narrows laterally and dorsally, forming a
slight indentation, thickening again on thoracic segments, then narrows abruptly to head,
which terminates in two flat prolongations diverging laterally from each other and incurved
dorsally. The edges of the wingcases get very close to each other dorsally around the union
of the thoracic with the abdominal segments, which is the narrowest point. Color light
green ventrally with fine criss-crossing, vein-like pattern, darker lines on wing cases. Along
each antenna there are two lighter warts. Dorsally light green also with a subdorsal dark
green longitudinal stripe on either side from cremaster to distal end of wingcases, giving the
impression of a partly rolled leaf. Measurer 3.8 cm long, 1 cm laterally at widest point
and .8 cm dorso-ventrally at widest point. Lasts 11 days.

Adults. No noticeable sexual dimorphism in this species. Shape of forewing: slightly convex
costal margin, rounded apex, almost straight but faintly sinuose outer margin, rounded
tornus and straight inner margin. Hindwing with almost straight costal margin, rounded
outer angle, continuing in the rounded and faintly sinuose outer margin, rounded anal angle
and almost straight and folded inner margin.

Colors dorsally mostly dark gray with bluish tinge on forewing apex and along hindwing
outer margin, with whitish markings, mostly on forewing, forming a complicated pattern
of bars, lines and circles. There is a conspicuous S-shaped reddish marking at the mid-

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costal margin directed towards the center of the forewing. Ventrally the dominant color
is beige, which covers the basal third of forewing and two-thirds of the hind wing. There
is a replica of the dorsal pattern of dark brown and yellowish white on the apical zone of
both wings. The reddish S marking is also present, even more conspicuous, due to the sharp
contrast of the reddish color with the ventral lighter coloration. The body is dark gray
dorsally, cream colored ventrally ; eyes reddish-brown ; proboscis orange ; antennae black
with white ventral spots on each segment; and a tiny orange spot at the tip. Average wing
span 7 cm in males, 7.5 cm in females. Total developmental time varies between 33 and 37
days.

NATURAL HISTORY

The adults of Hamadryas guatemalena in El Salvador frequent wooded areas
bordered by open, low brushy land, and are usually seen perching on tree
trunks with their wings spread open, head pointing down. Several individuals
might thus be seen in neighboring trees and from time to time aerial encounters
occur, with many “click-clicks” emitted while rapidly chasing each other. This
characteristic is limited to the males. The adults do not feed on flowers, but
come often to the ground to suck the juices of fermenting fruits. Mangoes,
guayavas, jocotes (hog plums) and the fruit of a local rubber-tree ( Castilla
gummifera Pittier) seem to be preferred. They also feed from exudations from
various trees. When they feed on the ground, the wings are at times held
perpendicular to the back. When they are feeding on the tree-trunks, the
wings are always spread open. The females ready to oviposit fly close to the
ground, more slowly than usual, until a foodplant is located. They alight
usually under a mature leaf and deposit one egg while the wings are apposed
dorsally. Several eggs might be deposited on a single vine, always on the
underside of a leaf. At times two, and rarely three eggs are deposited one on
top of the other, but never have we seen a female deposit two or more eggs
side by side on the same leaf. When more than one egg has been found side
by side under the same leaf, their hatching is not simultaneous, but separated
by a day or two, indicating successive ovipositions by the same or different
females.

The hatching larvae eat an exit hole from the wall of the shell and might
eat part of a wall. They never consume the whole eggshell. The small larvae
move to the edge of the leaf, bare a vein by eating the tissues around it, and
prolong the bared vein by affixing to it, with silk, small frass pellets, using
this artificial perch as a resting place during first and second instars. Very
often the small larvae affix excrement pellets to their own body, probably
for protective purposes. It is worthy of mention that when two or three eggs
have been deposited one on top of the other the hatching larvae do not damage
the ones on top due to their acquired habit of eating the exit hole on the
side of the eggshell. Damage to the egg on top would be unavoidable if the
larvae should eat the exit hole from the upper part of the eggshell as is usual
in most species of butterflies. The larvae are usually solitary, but when two

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Figs. 12 and 13. Hamadryas guatemalena Bates. 12. Male, dorsal view. Black bar 1 cm.
13. Male, ventral view. Scale in cm.

or three eggs are deposited as described, the ensuing larvae make their restin
perches independently, but on the same leaf, and might stay together durin
the whole larval stage without bellicose interaction. When the third instar is
reached, the resting perch is abandoned and the larvae spend most of the day
motionless on top of a leaf, with the thoracic segments humped and the head
bent so that the horns are parallel to the leaf surface. The larvae of H. guate-
malena are slow moving and rather passive. The spines which cover most of
the body do not have urticant properties. When ready to pupate the larvae

CTQ CfQ

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weave a silken mat on the stem of the vine or under a leaf, clean their digestive
tract and hang from the chosen spot for a day with their thorax and head
incurved ventrally and shed their larval skins. The pupae are firmly anchored
to the silk pad, due to the flat surface of the cremaster. If the supporting
stem is rolled one way or another the pupae will follow the movement rigidly,
“standing” on their cremaster. When disturbed the pupae wiggle laterally and
vigorously for a few seconds and stop moving, usually bent to one side. After
a time, they revert to the vertical position.

The adults emerge rapidly from the pupashell and hang from it while
ejecting a reddish meconium and expanding their wings. When the wings are
rigid enough, the butterflies take flight. From then on the wings are usually
kept spread while at diurnal perching.

This species is subject to heavy parasitism by tachinid flies, which abandon
the host as larvae and pupate on the ground. This happens during the last
larval instar or during pupation of the host.

The larval foodplant in El Salvador is Dalechampia scandens L., an Eu-
phorbiaceae vine which in our own experience is used by other species of
Hamadryas (Muyshondt & Muyshondt, 1974) and other species of Nymphali-
dae, such as Catonephele nyctimus Westwood, at least two species of Dynamine,
and Mestra amymone Menetries. The plant is quite abundant along fences,
ravines and in the borders of wooded land, up to about 1500 m altitude,
which is also the range where H. guatemalena is found. The leaves and bracts
of the plant have urticant properties.

It is to be noted that H. guatemalena , H. februa and H. amphinome share
not only the foodplant but the habitat as well. It is quite common to see
these species, especially guatemalena and februa , fly in the same neighborhood.

DISCUSSION

Descriptions of the early stages of species belonging to this group of
butterflies have been published in the past under the generic name of Ageronia
(Muller, 1886; Seitz, 1916), but to our knowledge this is the first description
illustrated with photographs ever published on the early stages of Hamadryas
guatemalena.

Butterflies belonging to this group have been called by various authors
under different generic names as a whole: Ageronia (Muller, 1886; Holland,
1914), Hamadryas (Klots, 1960; Ehrlich & Ehrlich, 1961) and have been
usually grouped under subfamily Ergolinae (Klots, op. cit.), or tribe Ergolini
(Ehrlich & Ehrlich, op. cit.). The adult shape, coloration and behavior is so
peculiar and similar in all of the species that it is only natural to consider
the various species as forming a well defined group within the Nymphalidae.
Even during the early stages the different species share many characteristics:

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the egg shapes of H. guatemalena and H. februa (Muyshondt & Muyshondt,
1975), and according to Muller (1886) the eggs of other species also, are so
similar as to make it hard to tell apart, if at all possible, one egg from another.
The same thing is true, to a point, with larvae and pupae; they all use the
same group of foodplants ( Dalechampia spp.), and exploit about the same
habitats. Yet there are also marked differences among them, which might
prove true some authors’ contention (Muller, 1886; Burmeister, as cited by
same Muller, op. cit .) that there are marked sub-groups within the genus
Hamadryas Hiibner, which might make it convenient to determine the proper
placement of the species within the group and the use of the names Ageronia
Hiibner, Peridromia Boisduval, Amphichlora Felder in addition to Hamadryas
itself. All these are available generic names according to Hemming (1967).

We will point out the differences we have observed between H. februa , H.
guatemalena and H. amphinome and will use the observations made by Muller
on some of these and on other species to make the point evident. The eggs
of guatemalena, februa, sp. ign. (in Muller), fornax and arete have the same
kind of sculpturing. Not so the eggs of amphinome, which are almost smooth.
The larvae of guatemalena, fornax, epinome and amphinome have dorsal spines
only on segments A- 7 and A-8, whereas these dorsal spines are present on all
abdominal segments in februa, arete and sp. ign. The pupal head prolongations
vary also from species to species: they are about the same in guatemalena
and arete, being laterally divergent and incurved dorsally. In februa they are
partially fused and follow the axis of the body. Then in epinome, sp. ign.,
fornax and amphinome they are divergent laterally, but follow the axis of the
body, as seen laterally.

As for larval behavior, guatemalena, februa, epinome, sp. ign. and arete have
solitary habits and all of them construct the resting perch with frass pellets
on the edge of the leaf. This is not the case with fornax nor with amphinome,
which have acquired gregarious behavior during the larval stage and have
given up the perch-making practice. Amphinome in addition has developed a
very angry and excitable disposition. Pupal behavior is the same in all species
we have observed, and corresponds with Muller’s description except for his
reported light sensibility. They all wiggle violently when disturbed and might
remain bent to one side for some time afterwards. Contrary to this, the adults
we have observed (Muller does not mention adults behavior) of guatemalena,
februa, amphinome, fornax and glauconome Bates, all show the same peculiar
jerky flight, the frantic clicking when males encounter each other, or when
chasing intruders, the feeding on fermenting fruits and tree wounds plus the
characteristic wing-spread attitude while perching on tree-trunks.

According to Muller, Burmeister grouped the species in the following man-
ner: 1) feronia, ferentina and fornax. 2) amphinome, arete, arethusa, related

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to chide. Miiller himself did it as follows: 1). a. — amphinome , b. — epinome ,
sp. ign., fornax ( jerentina) \ 2). — arete , arethusa , proposing to put back in use
the genus Peridromia for the latter. For him chide (the type species of Age-
ronia, according to Hemming) would be an intermediate form between amphi-
nome (which is the type-species for Hamadryas) and epinome (which he
places with fornax). So it looks as though Hamadryas should apply to amphi-
nome and whichever species are found to be congeneric with it; Ageronia to
chide and whichever species are congeneric with it; Amphichlora to feronia
and whichever species are its congeneric and Peridromia to arethusa and its
congenerics. Unfortunately we do not have reference material against which
to compare our species, so we are not in a position to establish which is the
type-species corresponding to guatemalena. We leave that to the taxonomists.

It is worthwhile to point out that there is interbreeding between closely
related species in nature: Limenitis arthemis astyanax Fabricius X L. archippus
Cramer (Klots, 1959; Platt & Greenfield, Jr., 1971), Vanessa atalanta rubria
(Friihstorfer) X Cynthia annabella Field (Dimock, 1973), and as a consequence
hybrids have been found to result under natural conditions. Hybrids have
also been produced in the laboratory from crosses between species naturally
separated by great distances, such as Papilio asterias X P. machaon (Clarke &
Knudsen, 1953), Papilio polyxenes asterias X P. maackii (Clarke & Sheppard,
1964), and several others, what seems to prove close specific relationship
between them, even if living far apart from each other under natural con-
ditions. Yet, even if H. guatemalena , H. februa and H. amphinome dwell in
the same habitat, during all months of the year, and are in addition grossly
similar to each other, we have never found evidence of interbreeding, nor
have we seen interspecific copulations, nor have we known of any report
thereof. That by itself would seem to indicate these species are not so closely
related, as their aspect and other characteristics suggest, as to belong to the
same genus. Unfortunately our efforts to have males and females of the
different species copulate in captivity have failed (actually, even attempts to
obtain copulation with males and females of the same species have proved
unsuccessful), so we can not bring forth proofs in either way.

We emphasize that in H. guatemalena the color of the larva during the
4th and 5th instars becomes very conspicuous by its contrasting colors, which
makes it an easy task to locate the larvae against the green leaves of the
foodplant on which they usually rest quite in the open. This daring behavior
would seem to advertise impalatable conditions, bad flavor or poisonous prop-
erties, to eventual predators. In this respect H. guatemalena seemingly has an
advantage over februa, whose colors are not so gaudy. Probably an increased
impalatability compensates for the loss of the additional mechanical protection
the dorsal spines (missing in guatemalena) provide februa. The pupae of

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179

this species, as in many other protected species, are exceedingly cryptic, imi-
tating to perfection a partly rolled leaf, but rely also on the vigorous wiggling,
which might scare away predators, as protection. The adults, even though
they display an aggressive disposition by rushing at any intruder in their
territory, exploit camouflage to perfection, blending their complicated wing
color pattern to moss and lichen growing on the tree-trunks on which they
rest with the wings spread open.

Euphorbiaceae plants have been historically reputed for their caustic and/or
poisonous fluids. Dalechampia scandens belongs to this family, and the leaves
and bracts have urticant properties. It would seem logical to deduce from
this and from the larval coloration and behavior, that H. guatemalena , which
feed exclusively on that plant, could have developed chemical protection against
predators derived from noxious components of the plant. Furthermore we
find that the species is heavily parasitized by tachinid flies during its larval
stage. We have pointed out in the past (Muyshondt 1973 a, b; Muyshondt &
Muyshondt, 1974) the repeated coincidence of heavy parasitism suffered by
many species generally accepted as protected by poisonous plant derivates
and species suspected as protected. Hamadryas guatemalena is another species
which might be added to the list.

Literature Cited

Clarke, C. A. and J. P. Knudsen. 1953. A hybrid swallowtail. An account of the “cross”
Papilio asterias $ (North American Black Swallowtail) X Papilio machaon $ (The
Swallowtail, of European-Malta-Stock) and a note on the “ machaon complex’'' of the
North American Continent. Entom. Record, 65: 76-80.

and P. M. Sheppard. 1964. The hybrid between Papilio polyxenes asterias Fabr.

female and Papilio maackii Men. male (Lepidoptera, Nymphalidae) [sic]. Ento-
mologist, 131-133.

Dimock, T. E. 1973. Three natural hybrids of Vanessa atalanta rubria X Cynthia anna-
bella (Nymphalidae). Jour. Lep. Soc. 27: 274-278.

Ehrlich, P. R. and A. H. Ehrlich. 1961. How to know the butterflies. Wm. C. Brown
Co. Publishers. Dubuque, Iowa.

Hemming, F. 1967. Generic names of the butterflies and their type-species. Bull. Br.

Mus. Nat. Hist. (Ent.) Suppl. 9.

Holland, W. J. 1914. Butterflies. Doubleday, Page & Co. New York, N. Y.

Klots, Alexander B. 1959. A mixed mating of two species of Limenitis Fabricius (Lep-
idoptera, Nymphalidae). Jour. New York Ent. Soc. 67: 20.

• . 1960. A field guide to the butterflies. Riverside Press. Cambridge.

Muller, W. 1886. Sudamerikanische Nymphaliden Raupen. Versuch eines naturlichen
Systems der Nymphaliden. Zoologische Jahrbuch. 453-461.

. 19736. Notes on the life cycle and natural history of butterflies of El Salvador.

IV A. — Pseudonica flavilla canthara (Nymphalidae-Catonephelinae). Jour. N. Y. Ent.
Soc. 81: 234-242.

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and A. Muyshondt, Jr. 1975. Notes on the life cycle and natural history of

butterflies of El Salvador. I B. — Hamadryas februa (Nymphalidae-Hamadryadinae) .
Jour. N. Y. Ent. Soc. 83 : 157-169.

Platt, A. P. and J. G. Greenfield, Jr. 1971. Interspecific hybridization between Limenitis
arthemis astyanax and L. archippus (Nymphalidae) Jour. Lep. Soc. 25: 278-284.
Seitz, A. 1915. Macrolepidoptera of the World. Vol. 5. Stuttgart.

Vol. LXXXIII, September, 1975

181

Notes on the Life Cycle and Natural History of Butterflies of
El Salvador. Ill B. — -Hamadryas amphinome L.
(Nymplialklae-Hamadryadinae)

Alberto Muyshondt and Alberto Muyshondt, Jr.

101 Ave. Norte 322, San Salvador, El Salvador

Received for Publication November 18, 1974

Abstract: The complete life cycle of Hamadryas amphinome L. is presented in this article

with photographic illustrations, with a record of the foodplant exploited by the larvae in
El Salvador, Dalechampia scandens L., and an account of the larval and adult behavior.
This species apparently is the most evolved of the Hamadryas complex as evidenced by the
notable deviations from the behavior of the other species studied: H. februa and H. guate-
malena, which are solitary during all phases of the metamorphosis, and H. amphinome ,
which is gregarious in the larval stage and shows consistent group behavior of the adults.
Emphasis is made of the behavioral adaptations the different species of the genus have
gone through which gradually change from fully solitary to fully gregarious larval habits.

This is the third article of a series on butterflies of the Hamadryadinae
group of the Nymphalidae, presenting our observations on the developmental
time and behavior of the early stages and the adults of Hamadryas amphinome
L. The field studies were conducted in the area of Los Chorros (a tourist
resort, about 12 km W of San Salvador, capital city of El Salvador), where
the species is fairly abundant throughout the year. During the dry season
1972/73 (November through April), a group of second instar Hamadryas
larvae were found feeding on Dalechampia scandens L., which were different
from those of H. februa and H. guatemalena. It was a surprise to us to notice
that these larvae, unlike the others just mentioned, which have solitary be-
havior, were feeding in a tight group on the underside of the leaf, showing
thus gregarious habits. The resulting adults were H. amphinome. Searches
for eggs were made every weekend from that time with negative results until
26 August 1973, when one peculiar group of “strings” formed by eggs one on
top of the other was found and collected. These were reared and the adults
emerged about one month later. Since then the species has been reared sev-
eral times up to this date. The rearing of the larvae in our insectarium was
in transparent plastic bags, sealed with rubber bands, in which fresh leaves
of the foodplant were supplied every third day until pupation of the larvae.

Acknowledgments: We are greatly obliged to Dr. Alexander B. Klots for his unfailing
guidance in our efforts and for reading, criticizing and otherwise rendering our manuscript
apt for publication. We are thankful also to Dr. A. H. D. Rydon for the valuable reference
material he kindly supplied. We give deserved credit to the youngest member of the team,
Pierre, for his tremendous help in the field work.

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Some colonies were kept together during the whole larval stage; others were
split in half after every moult starting from the second instar, so as to end
up with 4 to 6 individuals per bag. When ready to pupate the larvae were
transferred to a wooden box with windows of mosquito-netting and kept there
until the adults emerged. Records were kept of the developmental time and
size of each instar; photos were taken of all phases of the metamorphosis;
and specimens of eggs, larvae in the different instars and of the pupae have
been preserved in alcohol to be sent to the American Museum of Natural
History, New York.

LITE CYCLE STAGES

Egg. About 1 mm in diameter, round with slightly flattened base, yellowish-white when
recently deposited, turning to gray when ready to hatch. Surface almost smooth except
for faint sculpturing on the upper part of the walls. Eggs in groups of strings of eggs one
on top of the other, totalling 30 to more than 100 eggs per group. Hatch in 4-5 days.

First instar larva. Head shiny black, slightly cordiform, naked. Body whitish before
feeding, turning to olive green, with transverse rows of dark setae. Grows to 4 mm in

2- 3 days.

Second instar larva. Head shiny black with short horns on apices of epicrania. Body
brownish-green with transverse rows of short furcate spines. Grows to 8 mm in 2-3 days.

Third instar larva. Head shiny black with two lateral spines and long slender horns on
apices of epicrania each terminating in a clubbed tip. The shafts of the horns are armed
basally with one secondary spine directed anterad, a little higher a second spine directed
slightly inwards, then higher a third directed outwards, still higher a fourth parallel to the
first and finally, about the middle of the shaft, a fifth spine parallel to the second. The
rest of the shaft is covered by sparse short setae. Body dark greenish-brown with short
furcate spines placed in the following order: on first thoracic segment (T-l) one subdorsal
bifurcate spine, one supraspiracular bifurcate spine and one subspiracular simple spine ; T-2
with one 5-furcate subdorsal spine, one 5-furcate supraspiracular spine sided by a simple
spine, one subspiracular simple spine sided by another very small simple spine; T-3 with
one prominent 6-furcate subdorsal spine, one 4-furcate supraspiracular spine sided by a
simple spine, and one subspiracular simple spine sided by another small one. First ab-
dominal segment (A-l) with one 4-furcate subdorsal spine, one small supraspiracular simple
spine, one bifurcate subspiracular spine, one supraventral simple spine in line with the legs.
From A-2 to A-6, one 4-furcate subdorsal spine, one 4-furcate supraspiracular spine, one

3- furcate subspiracular spine sided by a simple spine, and two supraventral simple spines.
Segments A-7 and A-8 show in addition one 5-furcate dorsal spine. Segment A-9 has only
one 6-furcate supraspiracular spine deflected posterad. A- 10 has an anal plate with a
crown of 4 small simple spines. Grows to 1.2 cm in 2-3 days.

Fourth instar larva. As in third instar, but the head is orange-red, has longer lateral spines
and longer horns, which are incurved slightly caudad. Body spines longer than in third
instar, the subdorsal ones on segments A-2 to A-6 taking an orangish tinge. Grows to
2.6 cm in 4-6 days.

Fifth instar larva. Main change is body color, which is black; from A-2 to A-6 bright
yellow dorsal design of circles and bars forming paired O T O s. Subspiracularly these

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Figs. 1-5. Hamadryas amphinome L. 1 and 2. Typical ovipositions of Hamadryas
amphinome in groups of strings. Note that they are upside down, the strings being actually
pendant from the lower surface of the leaves. 3. Close-up of eggs. 4. Group of first instar
larvae ready to moult. 5. Group of second instar larvae. The rest had dropped to the
ground when disturbed while taking the photo. Notice the reflection of the ring-flash on
the shiny head capsule.

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Figs. 6-9. Hamadryas amphinome L. 6. Group of third instar larvae. Ovals on head
are the ring-flash reflections. 7. Fourth instar larva. 8 and 9. Lateral and dorsal view of
fifth instar larva where subspiracular orange spots and paired O T O marks are clearly
visible.

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185

Figs. 10-12. Hamadryas amphinome L. 10. Pupa, lateral view. 11. Pupa, ventral view.
12. Pupa, dorsal view.

segments show a large bright orange spot which contributes to make the larva very con-
spicuous. The spines on these segments are also reddish-orange. The subdorsal spines on
T-2 and T-3, and to a lesser degree the ones on A-2, A-7 and A-8, as well as the dorsal
spines on the last two, present small secondary spines on the shaft of the scoli. Grows to
4. 5-4. 7 cm in 5-6 days.

Pre-pupa. No change in coloration. It is slightly shorter. Hangs with head and thorax
incurved ventrally for one day.

Pupa. Thickens abruptly from flat cremaster, then gradually to A-2 and A-3, narrowing
then dorsally and laterally to A-l, thickening again to T-2, narrowing finally towards the
head, which is provided with two long, flattened and incurved prolongations. The pro-
longations are diverging laterally and incurved outwards at the tips. Usual color dorsally
light brown with green bordering along wingcases and on thoracic segments ; darker brown,
bordered by very light brown lines along abdominal meson. Ventrally light green, except
cremaster and head prolongations, which are brown. There is a darker morph on which
the green is substituted by light brown, the rest being darker brown than usual. It looks
very much like a rolled decaying leaf. Measures up to 4.5 cm long (including head pro-
longations) and lasts 7-8 days.

Adults. No marked sexual dimorphism in this species, the females having front wings
larger than males. Ground color dorsally black with many dark blue squares, half-
moons, round spots and bars forming a complicated pattern, covering both front and
hindwings, except for a subapical row of elongated light gray spots running from
mid-costal margin to tornus of front wing. The blue color of both wings is highly re-

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Figs. 13-16. Hamadryas amphinome L. 13 and 14. Male, dorsal and ventral view. IS
and 16. Female, dorsal and ventral view.

flectant. Front wing ventrally mostly dull black with orange basal triangle, and a subapical
reproduction of the dorsal gray spots in light yellow. Hindwing mostly bright orange with
a black area along the outer margin, which shows a row of cream colored round spots.
Body dorsally matching the black and blue colors of the wings, ventrally thorax orange
and abdomen black with two longitudinal cream stripes. Antennae and proboscis black.
Wing span varies from 6 to 7 cm. Total development in captivity takes from 27 to 35
days. The colonies kept together during the entire larval stage do not develop as big a
larva as the ones split gradually into smaller groups from the second instar on, consequently
the adults from the large groups are also much smaller than normal, even though the time
elapsed is about the same.

NATURAL HISTORY

The foodplant of Hamadryas amphinome in El Salvador is the same food-
plant used by H. jebrua and H. guatemalena: Dalechampia scandens L., a
vine belonging to the Euphorbiaceae, which we mentioned in detail in previous
articles on other Hamadryas spp. (Muyshondt & Muyshondt 1975 a and b).
It is to be noted that H . amphinome appears to have a more restricted range

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than the species mentioned before, which are found within the whole range
of the foodplant (500 to 1500 m altitude). H. amphinome is rather common
around 1000 m altitude, always in the close neighborhood of wooded land
surrounded by pastures or low-brushy plant communities.

The females of H. amphinome oviposit on the undersides of leaves of me-
dium development of D. scandens, from about two meters to very close to
the ground. The females perform outstanding acrobatics while depositing the
strings of eggs, spending close to 45 minutes to deposit some 50 eggs. The
eggs are laid one by one, one below another, forming perfect pendant strings.
There is no consistent norm of the number of eggs per string. We have found
groups of eggs with strings from 1 to 13 eggs. Muller (1886) records two
groups containing strings from 10 to 15 and from 2 to 13. In our experience
most of the strings consist of 5 to 10 eggs. The range of eggs per oviposition
is from 30 to more than 100, but most of the groups average 40 to 50 eggs.
The strings are rather rigid and keep the same angle in relation to the leaf
surface, even if the leaf is turned upside down, contrary to what Muller (1886)
observed about Ageronia (= Hamadryas) fornax Hiibner, whose eggs are de-
posited in one single flexible string which always hangs perpendicular to the
ground. The eggs of H. amphinome , white or cream-white when recently
deposited, turn to dark gray before hatching.

The hatching larvae eat their way out from the lateral wall of the eggshell
which is an important adaptation acquired by the species to avoid damaging
an adjacent egg and also prevents the rest of the eggs from falling to the
ground, which would occur if the exit hole was eaten from the micropylar
area as is customary in many other species. The small larvae have to crawl
up the string to reach the leaf surface and there the group is formed. They
start eating the lower layer of leaf tissue. Their excrements adhere to the
thin layer of silk formed while the larvae weave a silken foothold as they
move about. The small larvae also affix some pellets to their bodies. The
resultant accumulation of excreta seems to be a protective device adopted by
this species against its enemies. Later, but still during the first instar, the
larvae move in a group to the edge of the leaf, and form a tight line of
individuals perpendicular to the edge. As the larvae grow in the subsequent
moults, the group starts segregating into smaller groups mainly due to space
limitations of the leaf, somewhat similar to what happens with Dione juno
huascama Reakirt (Muyshondt, Young and Muyshondt, 1973), but also due
to the habit of H. amphinome larvae of wiggling convulsively at the least
disturbance, which if continued provokes a massive dropping of the larvae
from the leaf to lower levels of the plant or to the ground. In the act of
crawling back to the plant and reforming the group some larvae end up
forming smaller communities, far from each other. By the end of the larval
stage, the groups are usually reduced to three to seven individuals. When

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ready to pupate the small groups dissolve and the larvae wander about the
plant independently until a suitable location is chosen. This is normally on
the same vine or on a supporting plant, among the thick foliage, where the
larvae weave a silken mat and after cleaning their digestive tract, hang from
their anal prolegs, with head and thorax incurved ventrally, until pupation.
We have never found groups of pupae in the fields. The pupae of H. amphi-
nome , except for the head prolongations and the body color, look very much
like the pupae of H. februa and guatemalena , and behave very much like
them, swinging violently from side to side at the faintest provocation, ending
bent to one side. The vertical position is resumed some time afterwards.
As with the other species mentioned, the pupae are firmly anchored on the
flat cremaster so that they keep vertical to the supporting object even when
this is turned upside-down.

The emerging adults hang from the pupashell while ejecting the reddish
meconium and expanding the wings, which are held folded dorsally. After
the first flight the wings are usually kept spread open, except for night resting.
Even then at the slightest movement of the plant, the wings are immediately
spread, but after some flappings are folded again. The adults are seen, at
times, flying rapidly one following another in groups from five to eight indi-
viduals, as if playing “follow-the-leader.” When so acting, they alight, even-
tually, separately on contiguous fence poles or trees. When one flies again,
the whole group follows and after some fast maneuvers, the group alights
again on the same places or close by. This apparent game has been observed
for considerable periods of time, until the group moves away. When in this
playful mood, no “clicking” has been noticed. At other times, when two males
encounter each other, frantic circumvolations and excited clicking do occur.
Sometimes the fighting males fly up vertically, while circling, more than 100
meters high, coming down vertically also. One of us (A.M., Jr.), timed one
such a fight, and it lasted about 45 minutes, while continuous clicking was
audible. Females ready to oviposit fly more slowly than usual until they
locate a foodplant, and land on the underside of a medium sized leaf. They
stay there for a long period of time until a considerable number of eggs are
deposited as described, moving away afterwards with their customary swift
flight. In both males and females the blue reflection seems to hover over the
flying adults, somewhat as in the blue Morphos. In all other respects the
adults of this species behave like the rest of the Hamadryas group, and it
would be repetitious to further describe their habits. The species is also
subject to heavy parasitism by tachinid flies.

DISCUSSION

W. Muller (1886) describes, briefly, the life cycle of Hamadryas (= Age-
ronia) amphinome and gives the time spent during the larval stage as 19 to

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22 days, not determining the pupal time. He gives for the egg period 3 days,
which in our experience has taken from 4 to 5 days. The times for the dif-
ferent instars of the larva are more in accordance with our findings, which
makes us believe he found the eggs a day or two after they were deposited.
Muller does not mention the foodplant when dealing with H. amphinome , but
when he treats the genus Ageronia Hiibner, he states that all the species in-
cluded therein, feed, in Brazil, on Dalechampia ( triphyla Lam., ficifolia Lam.,
stipulacea Miill. Arg.), which agrees with our findings. Young (1974) records
Dalechampia heteromorpha as the foodplant of H. februa in Costa Rica.
Amazingly Barcant (1970) mentions Aristolochia trilobata (an Aristolochi-
aceae) as the foodplant of H. amphinomel In El Salvador where Aristolo-
chiaceae are well represented by several species, we have never found any
Hamadryas on them. Aristolochiaceae seem to be exploited in El Salvador
exclusively by various species of Battus and Parides (Papilionidae) . Most
probably Barcant’s record is a case of plant misidentification. The present
is apparently the first complete life cycle description of H. amphinome , with
photographic illustrations, ever published.

Hamadryas amphinome , which is the type species of the genus Hamadryas
Hiibner, teste Hemming (1967), is of extreme interest to specialists in evo-
lution studies as it seems to be the most advanced species of the Hamadryas
complex. This is evidenced by the gradual changes in various aspects of the
behavior of the various species, and in the shape of the eggs. H. arete Double-
day deposits one very sculptured egg at a time (Muller, 1886), and the larva
is solitary. H. februa Hiibner and H. guatemalena Bates (Young, 1974; Muy-
shondt & Muyshondt 1975 a and b), usually deposit their sculptured eggs
individually, the larvae then being solitary also, but these two species might
deposit two or even three eggs one on top of the other, the resulting larvae
then leading a loose communal life. All these species share the peculiar perch-
making habit during the first larval instars. H. fornax (Miiller, 1886) de-
posits the sculptured eggs in one string containing up to 10 eggs at a time
(one egg on top of the other), and the ensuing larvae have gregarious habits.
Finally H. amphinome deposits its almost round, little sculptured eggs in
groups of several strings totalling at times more than 100 eggs per group,
and the larvae also are gregarious. The last two species have abandoned the
perch-making habit. In H. amphinome even the adults seem to maintain a
sort of loose gregariousness, as their “follow-the-leader” game seems to in-
dicate.

The shift from solitary to gregarious behavior of the Hamadryas complex,
taken as a whole, has produced the following remarkable deviations from the
probable original solitary behavior: 1) the egg laying technique, which has

affected the shape of the eggs, 2) prerequisite to the massive egg laying, the
larvae must have acquired the habit of eating the exit hole from the side of

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the eggshell instead of the micropylar area, a trait existent already in the
solitary species. 3) The abandonment of the perch-making habit, shown by
the solitary species, in favor of the frass-pellet sticking on the underside of
the leaves. 4) the angry disposition of the gregarious species, contrasting with
the rather passive attitude of the solitary species. 5) The gradual increase in
gaudiness of the larval coloration in the gregarious species, as compared to
the solitary ones, and 6) the ability to produce disagreeable scent, enhanced
by the number of individuals in close association exhibited by the gregarious
species.

It is probable that the acquisition of the gregarious larval behavior is a
relatively recent event, as apparently the species is not yet fully adapted to
some of its consequences (ochlesis), as would be suggested by the readiness
of the groups to part company at the faintest motivation, as if to avoid over-
crowding of individuals. Overcrowding in captivity, has led to a drastic re-
duction in the size of the adults. In other gregarious species of butterflies
we have observed ( Actinote spp., Phyciodes spp., Chlosyne spp., Thessalia
theona Menetries, Microtia elva Bates, Mechanitis isthmia Bates, Colobura
dirce L. Manataria maculata Hopff.) there seems to be a better adaptation
to the crowding resulting from the communal life, and the individuals only
disperse just prior to pupation. In one case ( Dione juno huascama Reakirt)
we have even found communal pupation in the fields (Muyshondt, Young and
Muyshondt, 1973). If in these species there is a certain amount of segregation
of the original group into smaller ones, it is mostly due to space limitation.
It is true that these species, if deprived of sufficient food and space to move
about, do also produce midget adults, but not when they have enough food
and ample space. We emphasize also the contrast between the angry convul-
sions of the larvae of H. ampkinome with the coordinated twitchings of a
mass of Dione juno huascama. All these factors indicate that most of the
gregarious species acquired such behavior much longer ago than Hamadryas
amphinome, and are better adapted to it.

It seems to us that the conspicuous larval coloration of H. amphinome ,
added to the unpleasant odor emitted by the larvae, the adults’ aggressive
disposition and the bright orange coloration of their underwings, suggest im-
palatability to predators (orange color in insects is usually associated with
such condition), even though there is still some crypsis in the dorsal coloration
of the wings. The urticant properties of the foodplant Dalechampia scandens,
which belong to the Euphorbiaceae family reputed to comprise plants with
caustic or otherwise poisonous constituents (Standley, 1923, says about Eu-
phorbiaceae: “The sap usually has purgative and often poisonous properties”),
would seem to sustain our assumption, even though Young (1974) has ob-
served, in relation to Dalechampia heteromorpha Pax and Hoffmann, which

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191

is one of the foodplants of H. februa in Costa Rica: “I chewed several leaves
(both young and old) and found no signs of bitter tastes.” In our experience
some alkaloid (supposedly poisonous) bearing plants are not bitter when chewed,
and on the other hand some which are bitter when chewed do not contain
alkaloids. Furthermore, alkaloids are not the only poisonous materials found
in plants: saponins, cyanogenetic and cardiac glycosides are others (Brower
& Brower, 1964), and are not necessarily associated with bitter tastes.

H. amphinome, as is the case with H. guatemalena (Muyshondt & Muy-
shondt, 1975 b) has lost all the dorsal spines with the exception of the one
on segments A- 7 and A-8, which are present on all abdominal segments (ex-
cept the A-9 and A- 10) of other species of Hamadryas, probably as a result
of the increased protection obtained from the more efficient exploitation of
the noxious components of the foodplant, enhanced in this case by the gre-
garious behavior of the larvae.

Literature Cited

Barcant, M. 1970. Butterflies of Trinidad and Tobago. London. Collins.

Brower, L. P. and J. V. Z. Brower. 1964. Birds, butterflies and plant poisons: a study
in ecological chemistry. Zoologica 49: 137-159.

Hemming, F. 1967. Generic names of the butterflies and their type-species. Bull. Br.
Mus. Nat. Hist. (Ent.) Supl. 9.

Muller, W. 1886. Siidamerikanische Nymphalidenraupen. Versuch eines natiirlichen Sys-
tems der Nymphaliden. Zoologische Jahrbuch 453-461.

Muyshondt, A. and A. Muyshondt, Jr. 1975a. Notes on the life cycle and natural his-
tory of butterflies of El Salvador. I B. — Hamadryas februa (Nymphalidae-Hama-
dryadinae). Jour. N. Y. Ent. Soc. 83: 157-169.

. 1975b. Notes on the life cycle and natural history of butterflies of El Salvador.

II B. — Hamadryas guatemalena Bates (Nymphalidae-Hamadryadinae) Jour. N. Y.
Ent. Soc. 83: 170-180.

, A. M. Young and A. Muyshondt, Jr. 1973. The biology of the butterfly Dione

juno huascama (Nymphalidae: Heliconiinae) in El Salvador. Jour. N. Y. Ent. Soc.
81: 137-151.

Standley, P. 1923. Trees and shrubs of Mexico. Contrib. from the U. S. Nat. Herb.
Vol. 23 Part 3.

Young, A. M. 1974. On the biology of Hamadryas februa (Lepidoptera: Nymphalidae)
in Guanacaste, Costa Rica. Zool. ang Ent. 76: 845-856.

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New York Entomological Society

Notes on the Male Reproductive System in Ants
( Hymenoptera : F ormicidae ) 1

A. C. F. Hung and S. B. Vinson

Department of Entomology, Texas A&M University, College Station, Texas 77843
Received for Publication October 31, 1974

Abstract: The gross morphology of the male reproductive system of Pachycondyla harpax

(Fabr.), Eciton hamatum (Fabr.), Neivamyrmex sp., Pogonomyrmex barbatus (F. Smith),
Crematogaster laeviuscula Mayr, Solenopsis invicta Buren, Atta texana (Buckley), Irido-
myrmex pruinosum (Roger), Conomyrma insana (Buckley), Formica canadensis Santschi,
F. subintegra Emery and Polyergus breviceps Emery was studied. As the ants matured the
spermatozoa descended into the vas deferens and were retained there while the testes
progressively decreased in size. The dilated vasa deferentia where mature spermatozoa are
retained should be called “seminal vesicles” and what was formerly called “seminal vesicle”
should be referred to as “accessory gland.” Two types of accessory glands were found in
ants. In the first type which is found so far only in the Ecitonini, the glands are long, coiled
and both enclosed in a single capsule. In the second type the glands consist of two distinct
bodies and are either ball-shaped, bean-shaped, or elongated.

There have been very few studies on the male reproductive system in ants.
Janet (1902) in his study of the anatomy of the gaster of Myrmica rubra L.
depicted the male reproductive organs. This classic illustration has been re-
produced in such famous myrmecological monographs as “Ants” by Wheeler
(1910), “British Ants” by Donisthorpe (1915), and “Le monde social des
fourmis” by Forel (1921-1923). Forbes (1954) gave a comprehensive review
on this subject. Trakimas (1968) reinvestigated the anatomy and histology of
M. rubra. Unfortunately, only the abstract of her work was published.

According to Janet (1902), the male reproductive system of ants consists of
the testes, the vasa deferentia, the seminal vesicles, the ejaculatory duct and
the external genitalia. An aedeagal bladder was later found in Camponotus
and Formica (Forbes 1954), Eciton (Forbes 1958), Rhytidoponera (Hagopian
1963), N eivamyrmex (Forbes and Do-Van-Quy 1965), Solenopsis (Tice 1967),
and Myrmica (Trakimas 1968). Although a pair of accessory glands was found
in Dorylus labiatus Schuck (Mukerjee 1926), Eciton hamatum (Fabr.) (Forbes
1953), and N eivamyrmex harrisi (Haldeman) (Forbes and Do-Van-Quy 1965),
no mention of accessory glands has been made in other ants (Janet 1902,

Approved as TA 11481 by the Director of the Texas Agricultural Experiment Station.
Supported in part by Sigma Xi Grant-in-Aid of Research award to Hung and by the
Texas Department of Agriculture Interagency Agreement IAC (74-75) -0448. We also
thank Dr. John C. Moser and Dr. Carl W. Rettenmeyer for their generous contributions of
specimens.

New York Entomological Society, LXXXIII: 192-197. September, 1975.

Vol. LXXXIII, September, 1975

193

Fig. 1. Longitudinal section of seminal vesicle and accessory gland of F. subintegra.
(Scale line := 0.1 mm).

Figs. 2-6. Male reproductive system in ants (scale lines = 1 mm). 2. I. pruinosum. A,
newly emerged; B, matured. 3. Neivamyrmex sp., matured. 4. Pachycondyla harpax , newly
emerged. 5. Pogonomyrmex barbatus, matured. 6. A. texana, matured.

Abbreviations: AD, duct of accessory gland; AG, accessory gland; AT, atrophied testis;
BD, bound accessory gland ducts; CP, capsule of accessory gland; GA, genitalia; SP, sperm
plug; SV, seminal vesicle; TS, testis; VD, vas deferens.

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Forbes 1954, Hagopian 1963, Blussky 1967, Tice 1967, Trakimas 1968).
Therefore, according to Forbes (1954) the mature sperm in Camponotus
pennsylvanicus (DeGeer) are stored in the vasa deferentia and prevented from
moving into the seminal vesicles by a granular plug. A similar plug was also
found in Formica subintegra Emery (Fig. 1).

The above citations suggest that army ants differ from other ants in having-
accessory glands and further, that ants other than army ants do not store the
mature sperm in the seminal vesicles like other insects. We suggest the as-
sumption is false. The discrepancy appears to have been created by the use
of incorrect terminology.

According to Snodgrass (1935), the vesicula seminalis is a dilatation of the
vas deferens in which spermatozoa may be retained. Therefore, the seminal
vesicle could be any portion of the vas deferens. For example, in Oncopeltus
fasciatus (Dallas) it is located at the upper portion of the vas deferens imme-
diately following the vas efferens (Bonhag and Wick 1953).

We studied the male reproductive system of the following 12 species: Pachy-
condyla harpax (Fabr.) (Ponerinae) ; Eciton hamatum (Fabr.) (Dorylinae);
N eivamyrmex sp. (Dorylinae) ; Pogonomyrmex barbatus (F. Smith) (Myrmi-
cinae) ; Crematogaster laeviuscuia Mayr (Myrmicinae) ; Solonopsis invicta
Buren (Myrmicinae); Atta texana (Buckley) (Myrmicinae); Iridomyrmex
pruinosum (Roger) (Dolichoderinae) ; Conomyrma insana (Buckley) (Do-
lichoderinae) ; Formica canadensis Santschi (Formicinae) ; F. subintegra Emery
(Formicinae) ; and Polyergus breviceps Emery (Formicinae).

In Pogonomyrmex barbatus , S. invicta (Hung et al. 1974), /. pruinosum,
and F. subintegra in which we had freshly killed male pupae and alates of
different ages, we found that as the ants matured the spermatozoa descended
into the vas deferens and were retained there while the testes progressively
decreased in size (Fig. 2). According to the definition of Snodgrass (1935),
these dilated vasa deferentia (or portions of the vas deferens) should be called
“seminal vesicles.” Consequently, what was previously called “seminal vesicle”
should be referred to as “accessory gland.”

Our studies further revealed that there are 2 types of accessory glands in
ants. In the first type the glands are long, tightly coiled and both enclosed
in a single capsule (Figs. 3 and 7). This type is found so far only in Eciton
and N eivamyrmex. Although Forbes (1958) and Forbes and Do-Van-Quy
(1965) did not mention any capsule enclosing the coiled accessory glands in
their preserved material, our dissection of two fresh specimens of N eivamyrmex
males showed the presence of this capsule (Figs. 3 and 7, CP). In the second
type, the glands consist of two distinct bodies. They are ball-shaped in the
dolichoderines (Fig. 2), but are elongated in Dorylus (Mukerjee 1926) and
bean-shaped in ponerines (Fig 4), myrmicines (Figs. 5-6) and formicines.

Vol. LXXXIII, September, 1975

195

Fig. 7. Diagram of male reproductive system in Neivamyrmex sp. (scale line = 1 mm).

In mature males of some ants the glands are sometimes much smaller than
the seminal vesicles and are easily overlooked (Fig. 6).

Brown (1954) has suggested that Dorylinae might be diphyletic and Gotwald
(1969) goes further to state that the dorylines are tripartite. This preliminary
study on the gross morphology of the male reproductive system in ants cer-
tainly supports the polypheletic nature of the dorylines. As has previously
been pointed out, the coiled, enclosed accessory glands are so far found only

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in Eciton and Neivamyrmex. According to Mukerjee (1926) the accessory
glands of D. labiatus are conspicuous bodies due to their large size and thick
wall. His illustrations further show that the shape of these glands are very
similar to those of the myrmicines and formicines. We have also studied male
alates of Aenictus from Taiwan. Although the entire reproductive system in
our material was beyond recognition due to poor preservation, two distinct
bodies of accessory glands can still be recognized. Therefore, the accessory
gland of the Dorylini appears to have a closer resemblance to that of the
Myrmicinae and Formicinae than to Ecitonini.

There have been both anatomical and behavioral evidences supporting the
phylogenetic affinities between Ponerinae and Dorylinae (Wilson 1958, Her-
mann 1969). As far as the accessory glands are concerned, our study of
Pachycondyla and that of Rhytidoponera by Hagopian (1963) certainly in-
dicate that ponerines are closer related to Dorylini than to Ecitonini.

Literature Cited

Bonhag, F., and J. R. Wick. 1953. The functional anatomy of the male and female
reproductive systems of the milkweed bug, Oncopeltus fasciatus (Dallas) (Heterop-
tera, Lygaeidae). J. Morph. 93: 177-283.

Brown, W. L. 1954. Remarks on the internal phylogeny and subfamily classification of
the family Formicidae. Insectes Sociaux 1: 21-31.

Dlussky, G. M. 1967. Ants of the genus Formica (in Russian). Izdatel’stve “Nauka”,
Moscow. 211 pp.

Donisthorpe, H. St. J. K. 1915. British ants, their life-history and classification. Wil-
liam Brendon and Son, Ltd., Plymouth, England. 379 pp.

Forbes, J. 1954. The anatomy and histology of the male reproductive system of
Camponotus perms ylvanicus DeGeer (Formicidae, Hymenoptera) . J. Morph. 95:
523-556.

. 1958. The male reproductive system of the army ant, Eciton hamatum Fabricius.

Proc. 10th Internat. Congr. Entomol. 1: 593-596.

, and D. Do-Van-Quy. 1965. The anatomy and histology of the male reproductive

system of the legionary ant, Neivamyrmex harrisi (Haldeman) (Hymenoptera,
Formicidae). J. New York Entomol. Soc. 73: 95-111.

Forel, A. 1921-1923. Le monde social des fourmis du globe comparee a celui de l’homme,
5 vols. Libraire Kundig, Geneva. 948 pp.

Gotwald, W. H. 1969. Comparative morphological studies of the ants, with particular
reference to the mouthparts (Hymenoptera: Formicidae). Mem. Cornell Univ. Agric.
Exper. Sta., Ithaca, N. Y. 408: 1-150.

Hagopian, M. 1963. An anatomical and histological study of the male ponerine ant,
Rhytidoponera metallica F. Smith (Formicidae, Hymenoptera). Dissertation, Ford-
ham Univ., Xerox Univ. Microfilms, Ann Arbor, Mich. 124 pp.

Hermann, H. R. 1969. The hymenopterous poison apparatus: Evolutionary trends in
three closely related subfamilies of ants (Hymenoptera: Formicidae). J. Georgia
Entomol. Soc. 4: 123-124.

Hung, A. C. F., S. B. Vinson, and J. W. Summerlin. 1974. Male sterility in the red
imported fire ant, Solenopsis invicta Buren. Ann. Entomol. Soc. Amer. 67: 909-912.

Vol. LXXXIII, September, 1975

197

Janet, C. 1902. Anatomie du gaster de la Myrmica rubra. Georges Carre et C. Naud,
Paris. 68 pp.

Mukerjee, D. 1926. Digestive and reproductive systems of the male ant Dorylus labiatus
Schuck. J. Proc. Asiatic Soc. Bengal, n. s., 22: 87-92.

Snodgrass, R. E. 1935. Principles of insect morphology. McGraw-Hill, New York.
667 pp.

Tice, J. E. 1967. The anatomy and histology of some of the systems of the male of the
imported fire ant, Solenopsis saevissima richteri Forel (Hymenoptera: Formicidae).
Dissertation, Fordham Univ., Xerox Univ. Microfilms, Ann Arbor, Mich. 122 pp.
Trakimas, W. B. 1968. An anatomical and histological study of the male myrmicine ant,
Myrmica rubra L. (Hymenoptera: Formicidae). Dissertation Abstr. 28B: 4360.
Wheeler, W. M. 1910. Ants: Their structure, development and behavior. Columbia

Univ. Press, New York. 663 pp.

Wilson, E. O. 1958. The beginnings of nomadic and group-predatory behavior in the
ponerine ants. Evolution 12: 24-31.

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New York Entomological Society

Species and Numbers of Bloodsucking Flies Feeding on Hogs
and Other Animals in Southern New Jersey1’2

Thomas J. Weiner and Elton J. Hansens3

Department of Entomology and Economic Zoology,

Cook College, Rutgers University, New Brunswick, N. J. 08903

Received for Publication January 20, 1975

Abstract: Tabanidae of 19 species were recorded feeding on hogs of 8 herds in 1973.

Species and feeding location on the animals were recorded. Though small numbers of
tabanids fed on hogs, herds in wooded areas were more subject to attack by Tabanidae
and Stomoxys calcitrans than those in open fields. Stomoxys calcitrans, Haematobia irritans,
and 13 Tabanidae were noted on horses, 2 Tabanidae on goats and 6 on dogs.

In areas adjacent to the New Jersey coastal wetlands, female horse flies
and deer flies are serious pests of domestic animals and man. Recently Tid-
well et al (1972) reported the capability of Tabanidae in transmitting hog
cholera and named 8 different Tabanus species feeding on North Carolina hogs
under field and laboratory conditions. The present study sought to identify
the species and assess the numbers of Tabanidae found on hogs and other
animals under farm conditions in New Jersey.

MATERIALS AND METHODS

Eight New Jersey farms, including 7 located in Cumberland and 1 in Cape
May county, were visited at least twice a week from June 6 to August 10,
1973. At each location, 20 hogs were chosen randomly and counts of feeding
Tabanidae and Stomoxys calcitrans were made. The appearance of the animal
was also noted — clean, dirty or muddy — as well as whether it was in the sun,
shade or shelter. Not only were numbers of flies tabulated, but also on which
part of the hogs’ body a particular fly was feeding. A portion of the flies were
collected and identified in the laboratory to confirm field identification. When-
ever an unfamiliar fly was seen, a special effort was made to capture it. Only
once during the season did such a fly escape. This particular fly was recog-
nized as a Chrysops species.

Black box traps such as that described by Decoster (1968) were set up at
5 farms to monitor the fly population. Beach balls sprayed black were used

1Diptera: Tabanidae, Muscidae.

2 Paper of the Journal Series, New Jersey Agricultural Experiment Station, Rutgers Uni-
versity— The State University of New Jersey, New Brunswick, N.J. This investigation
is part of an undergraduate George H. Cook Scholar Project and was supported in part by
USDA — APHIS, Contract No. 12-16-100-189.

3 Undergraduate Student and Research Professor, respectively.

New York Entomological Society, LXXXIII: 198-202. September, 1975.

Vol. LXXXIII, September, 1975

199

Table 1. Numbers of Tabanidae species collected from hogs and box traps throughout
the summer of 1973 in southern New Jersey.

Species

Number on Hogs

Number in Traps

Tabanus lineola F.

306

1515

T . nigrovittatus Macquart

136

554

T. atratus Fabricius

T. trimaculatus Palisot de Beauvois

Chrysops fuliginosus Wiedemann

C. niger Macquart

C. cincticornis Walker

C. atlanticus Pechuman

C. montanus Osten Sacken

T. nigripes Wiedemann

T. pumilus Macquart

T. melanocerus Wiedemann

Diachlorus ferrugatus (F.)

T. spams milleri Whitney

T. americanus Forster

T. imitans Walker1

T. sulcifrons Macquart

Hybomitra daeckei (Hine)

H. lasiophthalma (Macquart)

C. obsoletus Wiedemann

C. dimmocki Hine

C. flavidus Wiedemann

C. brunneus Hine

C. vittatus Wiedemann

C. celatus Pechuman

C. geminatus Wiedemann

Stomoxys calcitrans (L.)

3119

302

1 The first recorded specimen found north of Maryland — identification made by L. L.

Pechuman, Cornell University.

as targets hung under the traps. Each time these farms were visited, trapped
flies were collected and later identified.

Clean clothing was worn at all times and rubber boots were disinfected
before and after visiting each location.

RESULTS

Hogs. Of the 19 Tabanidae species (Table 1) found to feed on hogs in
southern New Jersey, Tabanus lineola , T. nigrovittatus and T. atratus fed in
the largest numbers. T. lineola made up 48% of the total number of horse
flies counted on the animals and 66% of those Tabanidae collected from the
box traps. T. nigrovittatus contributed 21% and 24% of these totals, respec-
tively. T. atratus fed freely on the hogs but would rarely enter the traps.

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Table 2. Comparison of the number of Tabanidae and Stomoxys calcitrans found on hogs
in woods versus fields, Cumberland County, N.J.

Farms

Tabanidae

Total

Tabanidae/Hog

Stomoxys

Total

Stomoxys/ Hog

A (woods

0.29

1359

4.00

C (woods)

0.04

182

0.61

D (fields)

0.01

0.08

F (woods)

169

0.56

168

0.56

F (fields)

117

0.34

697

2.05

The number of deer flies feeding was generally lower than the horse flies.
Chrysops juliginosus , C. niger and C. cincticornis were those species which fed
most frequently. Chrysops atlanticus, C. montanus, T. nigripes, T. pumilus ,
T. melanocarus and Diachlorus jerrugatus were present in the vicinity of the
hogs in substantial numbers but were not inclined to feed. The stable fly,
Stomoxys calcitrans , was more common on the hogs than any of the tabanids
and made up 83% of the flies found in the box traps.

The stable fly was present throughout the entire season and fed all over the
body of the hog, as did most of the Tabanus spp. However, T. atratus con-
centrated on the back of the animal and T. trimaculatus had a predilection for
the sides and legs. Hybomitra lasiophthalma fed on the legs and Diachlorus
jerrugatus fed only on the lower half of the pig. The feeding activity of most
Chrysops spp. was limited to the hog’s back.

In general, more Tabanidae and S. calcitrans fed on animals kept in wooded

Table 3. Species and numbers of Tabanidae found on hogs in the fields and woods of

Farm F.

Species

Fields (17)1

Woods (15)1

Tabanus lineola

T . atratus

T. nigrovittatus

T. trimaculatus

T. nigripes

T. americanus

Chrysops niger

C. cincticornis

C. atlanticus

Hybomitra daeckei

H. lasiophthalma

Diachlorus jerrugatus

Number of visits.

Vol. LXXXIII, September, 1975

201

Table 4. Numbers of Tabanidae species, Stomoxys calcitrans, and Haematobia irritans
collected from horses during the summer of 1973 in Millville, New Jersey.

Species

Number on Horses

C hr y sops niger

Tabanus lineola

543

T. atratus

1 T. fulvulus Wiedemann

T. nigripes

T. americanus

Hybomitra lasiophthalma

1 T. petiolatus Hine

T. pumilus

1 H. cincta (F.)

T. nigrovittatus

H. daeckei

1 H. trispila (Wiedemann)

1 T. pallidescens Philip

1 T. stygius Say

Stomoxys calcitrans

1376

1 Haematobia irritans (L.)

1 Not seen on hogs.

areas than in non-wooded areas (Table 2). This trend is shown for farms C
and D located in Cedarville and 0.5 mile apart. The wooded farm C had 4
Tabanidae and 61 S. calcitrans for every 100 hogs compared to 1 Tabanidae
and 8 S. calcitrans in the field of farm I).

Farm F was unique in that it had 2 distinct herds separated by 0.7 mile.
The wooded area had 56 tabanids and 56 stable flies feeding on every 100
pigs, whereas the field area had 34 tabanids and 205 stable flies per 100 pigs.
S. calcitrans showed a greater tendency to enter shelters than the tabanids.
This is a possible explanation for the more abundant stable flies found feeding
in the field of farm F. Each time this area was visited counts were taken
from hogs maintained in 2 pens. One pen contained the younger and smaller
animals while the other housed the larger breeding sows which stayed inside a
shelter with one completely open side. The S. calcitrans were largely on the
breeding sows inside the shelter. Throughout the summer only 3 Tabanidae
entered the shelter to obtain a blood meal, namely T. lineola , T. nigrovittatus,
and T. atratus. In addition to finding larger numbers of flies feeding on hogs
housed in wooded areas, more species were also found (Table 3).

Only 5 horse fly species fed on swine in the fields of farm F while 12 species
were found in its wooded area. The smaller number of T. atratus and T. nigro-
vittatus found in the woods is because the hogs in this area were removed on
July 24 (due to theft), whereas counts on the swine in the fields continued

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until August 10 and on that particular day 11 T. atratus and 7 T. nigrovittatus
were seen. The most abundant species in both areas were T. lineola and T.
nigrovittatus.

The 7 Tabanidae species which appeared in South Jersey early in the season
were: Chrysops cincticornis, C. fuliginosus, C. niger, Tab anus nigripes , T. tri-
maculatus , Hybomitra daeckei and H. lasiophthalma. Those ubiquitous species
present throughout the entire summer were Tabanus lineola , T. nigrovittatus
and T. atratus.

Horses. A horse ranch located in Millville, New Jersey was observed 13 times
during June to August 1973. Fifteen horses were chosen at random and feed-
ing flies were counted.

In addition to Stomoxys calcitrans and Haematobia irritans , 15 species of
Tabanidae were recorded on horses (Table 4).

Chrysops niger fed on the head and neck of the horse. Tabanus lineola and
T. julvulus fed on the legs while T. nigripes fed on the upper part of the
animal (head, neck and side). T. atratus was counted on the back and legs
and Hybomitra lasiophthalma in the genital area. S. calcitrans fed all over the
horse but predominantly on the side and legs, while H. irritans fed on the
horse’s belly.

Present throughout the season were Tabanus lineola, T. atratus, T. nigripes,
S. calcitrans and H. irritans.

While riding in wooded areas in the evening the senior author noted feeding
on the horse’s ears by Chrysops macquarti, C. nigribimbo, C. vittatus and C.
celatus.

Other Animals. On hog farm A, where 9 different Tabanidae species were
recorded on hogs, observations were also made of those flies found on 8 goats
and 2 dogs.

The total number of flies attracted to the goats included 2 C. fuliginosus,
1 T. nigrovittatus and 28 S. calcitrans. The dachshund, kept in an open area,
had 1 T. nigrovittatus and 18 S. calcitrans. The German shepherd, kept in a
wooded area, had 24 T. nigrovittatus, 353 S. calcitrans, 2 C. atlanticus, 2 C.
vittatus and 1 each of C. callidus Osten Sacken, C. fuliginosus and C. montanus.

Literature Cited

Decoster, A. A. 1968. How to catch a greenhead fly. Down East — The Magazine of
Maine 13(3): 54.

Tidwell, M. A., W. D. Dean, G. P. Combs, D. W. Anderson, W. O. Cowart, and R. C.
Axtell. 1972. Transmission of hog cholera virus by horse flies (Tabanidae:
Diptera). American Journal of Veterinary Research 33(3): 615-622.

Vol. LXXXIII, September, 1975

203

Speleognathinae Collected From Birds In North America
( Acarina : Ereynetidae ) 1

A. Fain

Institut de Medecine Tropicale Prince Leopold, Antwerpen, Belgium

and

K. E. Hyland

Department of Zoology, University of Rhode Island, Kingston, R. I. 02881

Received for Publication October 21, 1974

Abstract: Fourteen species and subspecies of nasal mites belonging to the subfamily

Speleognathinae and taken from birds are reported, including eight new host records. Two
new subspecies are described: Neoboydaia philomachi thalasseus from Thalasseus maximus
(royal tern) and Sterno hirundo (common tern) from Perry, Florida, and E. Sandwich
Mass., respectively; and Boydaia cyanerpes hylocichla from Hylocichla ustulata (russet-
backed thrush) taken at Big Falls, Newfoundland.

The speleognathine fauna is not exceedingly abundant in North American
birds nor has it been the subject of wide-spread investigation. Except for the
original descriptions there is relatively little literature establishing additional
host records or distributional patterns. Two works, those of Fain and Hyland
(1970) and Pence (1973), have added significant new records, and it is our
intention that the present work augment the existing host and distributional
lists.

The mites which form the basis of this study have been collected from the
nasal passages of a variety of avian hosts and from several widely separated
localities in North America. Two new subspecies are described and several
new host records have been noted. Fourteen species and subspecies have been
recorded.

1This investigation was supported in part by a National Science Foundation Grant (GB-
1295) to the junior author.

Acknowledgments: We wish to thank those persons who assisted with the collection of
specimens, especially W. T. Atyeo, Department of Entomology, University of Georgia,
Athens, Georgia; George West, Laboratory of Zoophysiology, University of Alaska, College,
Alaska; and Arnold Moorhouse, U.S. Department of Agriculture, Animal and Plant Health
Inspection Service, Clifton Animal Import Center, Clifton, N.J.

We wish to acknowledge the assistance of Shashi K. Nagar, University of Delhi, Delhi,
India, in the preparation of the material and for some of the preliminary determinations.

New York Entomological Society, LXXXIII: 203-208. September, 1975.

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New York Entomological Society

Genus Neoboydaia Fain, 1958
1. Neoboydaia philomachi eroliae (Fain & Hyland, 1970)

This subspecies was described from Erolia minutilla and Actitis macularia in Mexico. It
is distinguished from the type female by the following characters:

(1) Setae ic2 and coxal II are toothed (Da), whereas in the types they are very fine
and piliform (Na). See Fain (1970) for setal nomenclature.

(2) In the typical form femora I and II bear three foliate and striate setae (Sd) and
femur III bears one seta Sd in addition to the other setae. In the specimens of
N. p. eroliae collected from the type hosts (E. minutilla and A. macularia) these
setae (Sd) are replaced by cylindrical and dentate setae (Da).

We have collected this form from:

(1) Erolia minutilla (least sandpiper) in Galilee, R. I. (No. H62-08-21-3; Coll. G. West)
11 females, 3 larvae;

(2) Actitis macularia (spotted sandpiper) in Charlestown, R. I. (No. H61-07-24-15 ;
Coll. L. TerBush) 1 female;

(3) Arenaria interpres (ruddy turnstone) in Witless Bay, Newfoundland (No. H62-08-
07-4; Coll. K. Hyland et al.) 11 females and 1 male;

(4) Limnodromus griseus (dowitcher) in Charlestown, R. I. (No. H61-07-24-8; Coll. L.
TerBush) 5 females and in South Kingstown, R. I. (No. H61-08-09-6; Coll. L.
TerBush) 16 females, 2 larvae;

(5) Tringa solitaria (solitary sandpiper) in Rushville, Nebr. (No. A59-08-31-10 ; Coll. W.
Atyeo and N. Braasch) 6 females.

Arenaria interpres , Limnodromus griseus and Tringa solitaria appear to be new host
records.

Those specimens taken from Arenaria and Limnodromus carry two type Sd
and one type Da setae on femora T and II. On femur III one seta of type
Sd is present. The other characters compare favorably with those of the
subspecies eroliae.

Clark (1964) redescribed N. philomachi from specimens which he collected
from the hosts Totanus melanoleucus , Totanus flavipes, and Pisobia melanotos
in Texas. The drawings which he gave with this redescription apparently are
of the subspecies eroliae.

Pence (1973) reported N. philomachi from several charadriiform hosts
without indicating the subspecies. Hosts included the type host, Erolia minu-
tilla, plus Capella gallinago (common snipe), Totanus melanoleucus (greater
yellowlegs) and Limnodromus scolapaceus (long-billed dowitcher). Since the
host genus Philomachus is not represented in North America it seems reason-
able to expect that N . philomachi philomachi is absent from North American
birds and to consider that N . p. eroliae is distinct.

2. Neoboydaia philomachi thalasseus subsp. nov.

This new subspecies can be distinguished from the other two subspecies by the presence
of only three pairs of genital setae in the female (one pair of internals and two pairs of
externals, all of type Na) whereas in both N. p. philomachi and N . p. eroliae there are six

Vol. LXXXIII, September, 1975

205

pairs of setae of which the three externals are of type Na and the three internals are of
type Da. Coxal setae and ic2 are similar to subspecies eroliae. Idiosoma of holotype female
455 microns in length and 142 microns maximum width.

This subspecies was collected from:

(1) Thalasseus maximus (royal tern) in Perry, Florida (No. A60-07-14-4; Coll. W. Atyeo
et al.) holotype and 14 paratype females plus 2 paratype larvae;

(2) Sterna hirundo (common tern) in E. Sandwich, Mass. (No. H61-08-12-3 ; Coll.
K. Hyland et al.) 4 females, 1 male, 1 larva.

Holotype deposited in the U.S. National Museum, Washington, D. C.; paratypes in the
Institut de Medecine Tropicale Prince Leopold, Antwerpen, Belgium and University of
Rhode Island, Museum of Zoology, Kingston, R. I.

3. Neoboydaia colymbiformi Clark, 1964

This species was described from Colymbus nigricollis calif or nicus (eared grebe) in Cali-
fornia. We recorded it earlier from Podilymbus podiceps (pied-billed grebe) in Mexico
(Fain & Hyland, 1970) and Pence (1973) has reported it from the same host in Louisiana.

We have also found it in a new host, Podiceps caspicus (eared grebe) in Rushville, Nebr.
(No. A59-08-31-6; Coll. N. Braasch and W. Atyeo) 9 females, 1 larva.

Genus Astrida Fain, 1955
Subgenus Neastrida Fain, 1963

1. Astrida ( Neastrida ) coccyzae Pence, 1973

We have collected this species from the type host Coccyzus americanus (yellow-billed
cuckoo), as follows:

(1) Hebron, Nebraska (No. A59-07-08-3; Coll. N. Braasch) 3 females; and

(2) North Kingstown, R. I. (No. H62-08-23-4; Coll. A. Hawkes) 8 larvae.

Genus Trispeleognathus Fain, 1958

1. Trispeleognathus womersleyi (Fain, 1955)

This species has been taken from Anas discors (blue-winged teal) collected in both
Rhode Island and Nebraska as follows:

(1) Allenton, R. I. (No. H62-09-03-4; Coll. L. TerBush) 10 females;

(2) Valentine, Nebr. (No. A59-09-02-1; Coll. W. Atyeo) 1 female;

(3) Rushville, Nebr. (No. A59-08-31-1; Coll. N. Braasch and W. Atyeo) 2 females.

To our knowledge it has not been reported from the blue-winged teal previously.

Genus Boydaia Womersley, 1953
Subgenus Boydaia Womersley, 1953

1. Boydaia ( Boydaia ) hirundoae Fain, 1956

This species has been collected from the type host, Hirundo rustica (barn swallow), as
follows:

(1) Richmond, R. I. (No. H62-07-06-1; Coll. A. Moorhouse) 6 females and 1 male;

(2) Waterford, Conn. (No. H62-05-12-1; Coll. D. Blake) 3 females and 1 male.

Pence (1973) has recently reported this same species from the type host in Louisiana.

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2. Boydaia ( Boydaia ) tyrannis Ford, 1959

Specimens of this species have again been taken from the type host, Tyr annus tyr annus
(kingbird), from Michigan as follows: Co. Rd. 400, Kellogg Gull Lake Biol. Sta., Mich.
(No. 59-08-10-5; Coll. K. Hyland et al.) 2 females and 5 larvae.

Although Fain and Aitken (1968, 1970) and Fain and Hyland (1970) have reported this
species from various tyrannids, cotingids and pipnids from Trinidad, Mexico and Brazil,
only the type host has been found infested with this species north of Mexico.

3. Boydaia ( Boydaia ) colini Clark, 1958

We have taken this species from the type host, Colinus virginianus (bob white), collected
in Charlestown, R. I. (No. H62-09-01-1; Coll. A. Moorhouse) 2 females and 4 larvae. This
host is the only host thus far reported harboring B. colini.

4. Boydaia ( Boydaia ) agelaii Fain and Aitken, 1968

This species is distinguished from B. ( B .) quiscali Clark, 1960 by the character of the
claws on tarsi II of the larva. The elongate claw has a different shape and the short claw
is much shorter than in quiscali.

Our collection includes the following hosts:

(1) Spiza americana (dickcissel) from Grand Island, Nebr. (No. A59-06-10-14 ; Coll. N.
Braasch and W. Atyeo) 9 females, 1 male, and 1 larva.

(2) Molothrus ater (brown-headed cowbird) from El Paso, Texas (No. H62-1 1-24-4;
Coll. G. West) 3 females.

(3) Cassidix mexicana (boat-tailed grackle) from Lake Placid, Florida (No. A60-07-25-2 ;
Coll. W. Atyeo and N. Braasch) 1 female.

It should be noted that Pence (1973) placed material collected from Molothrus ater in
B. quiscali rather than in this species. Apparently he did not examine the larvae collected
from M. ater which is the type host for B. quiscali. He also assigned material from
Cassidix mexicanus to B. quiscali rather than to B. agelaii.

The dickcissel ( Spiza americana) constitutes a new host record.

5. Boydaia ( Boydaia ) loxiae Fain, 1963

Eight female specimens collected from Icterus galbula (Baltimore oriole) (No. A60-05-
15-9; Coll. W. Atyeo) taken at Nebraska City, Nebr., have been assigned to this species.
In the absence of larvae it is impossible to distinguish with certainty this species from
others belonging to the “statulata” group ; however, this is apparently the first record of
this species in North America. See Fain, 1971.

6. Boydaia ( Boydaia ) cyanerpes hylocichla subsp. nov.

This subspecies can be distinguished from the type by the following characters in the
female :

(1) Femur I has 6 or 7 setae (compared with 5 in the type):

(2) Pattern of lines on the base of gnathosoma differs from the type particularly in
having two bands in the middle which converge posteriorly (instead of two bands
which diverge posteriorly) ;

(3) Setae on the body and legs are longer: setae dl-d4 are 18 microns long compared
with 13-15 microns in B. h. hylocichla ; posterior intercoxal setae 15 microns com-
pared with 12 microns; setae on tibia I are 21-25 microns compared with 15-18;
and setae on femur I 12-24 microns in contrast to 8-15.

Vol. LXXXIII, September, 1975

207

(4) Palpal solenidion is vestigial, thinner and shorter (1 micron) instead of 3-3.5 microns
as in B. h. hylocichla.

Idiosoma of holotype female is 405 microns long by a maximum of 315 microns wide.

Host: Holotype and 5 paratype females were taken from Hylocichula ustulata (russet-
backed thrush) collected at Big Falls, Newfoundland (No. H62-08-01-9; Coll. K. Hyland
et al.). Holotype in the U.S. National Museum, Washington; paratypes in the collections
of the authors.

Subgenus Coboydaia Fain, 1971

7. Boydaia ( Coboydaia ) nigra nigra Fain, 1955

The type host of this species is Serinus sulphuratus shelleyi (Fringillidae) from Rwanda.
We have recorded the same species from the following hosts in North America:

(1) Carpodachus mexicanus (house finch) from El Paso, Texas (No. H62-1 1-24-10;
Coll. G. West) 5 females, 1 male and 2 larvae.

(2) Spizella passerina (chipping sparrow) El Paso, Texas (No. H62-11-24-14 ; Coll. G.
West) 5 females, 2 males, and 3 larvae.

Both are new host records.

8. Boydaia ( Coboydaia ) nigra icteri Fain and Hyland, 1970

This subspecies was described earlier by us from Icterus spurius (orchard oriole) in
Mexico. We have also identified it from the same host from Lincoln, Nebr. (No. A59-
06-05-1; Coll. W. Atyeo and N. Braasch) one female. We have also recorded it from
Icterus galbula (Baltimore oriole) in St. Joseph Co., Michigan (No. C60-08-23-13 ; Coll.
Unknown) 7 females.

9. Boydaia ( Coboydaia ) sturnellae Clark, 1960

We have recorded this species from the type host, Sturnella magna (meadow lark)
collected at:

(1) Lake Placid, Fla. (No. A60-07-23-10 ; Coll. W. Atyeo et al.) 8 females, 4 males,
and 4 larvae; and

(2) Hope Valley, R. I. (No. H62-07-18-1; Coll. A. Moorhouse) 2 larvae.

B. (C.) sturnellae is close to B. (C.) nigra nigra Fain, 1955. In the female it can be
distinguished from B. nigra principally by the slightly thinner sensillae, which measure 42
microns in our specimens. In the larva, claws I— III resemble those of B. nigra in form
but they are shorter. Claw III is 33 microns long (hook included) compared with 45 to 53
microns for nigra.

B. (C.) sturnellae can be distinguished from B. ( C .) amandavae Fain, 1962, in the larval
form by the shape of claw I which is recurved apically and terminates in a point whereas
in B. amandavae the claws are not recurved apically and they are dilated (see Fain, 1971,
fig. 41). They can be distinguished in the female by the greater elongation of the leg
segment, by the different chaetotaxy and by the sensillae.

B. ( C .) sturnellae appears to be specific for the meadow lark.

Literature Cited

Clark, G. M. 1958. One new and one previously unreported species of nasal mite (Acarina,
Speleognathidae) from North American birds. Proc. Helm. Soc. Wash. 25(2) : 78-86.

208

New York Entomological Society

. 1960. Three new nasal mites (Acarina: Speleognathidae) from the gray squirrel,

the common grackle, and the meadowlark in the United States. Proc. Helm. Soc.
Wash. 27(1) : 103-110.

. 1964. One new and one previously unreported nasal mite (Acarina: Speleog-

nathinae) from North American birds, with observations on speleognathid taxonomy.
J. Parasit. 50(1): 158-162.

Fain, A. 1955. Sur le parasitisme des fosses nasales chez les mammiferes et les oiseaux par
les Speleognathidae. Ann. Soc. Beige Med. Trop., 35(6) : 689-700.

. 1956a. Les acariens de la famille Speleognathidae Worn, au Ruanda-Urundi. Rev.

Zool. Bot. Afr. 53(1-2): 17-50.

. 1956b. Nouvelles observations sur les Acariens de la famille Speleognathidae

parasites des fosses nasales chez les batraciens, les oiseaux et les mammiferes. Ann.
Parasit. Hum. et Comp. 31: 643-662.

. 1956c. Notes sur les Acariens du genre Boydaia Worn, avec description d’une

espece nouvelle. Riv. Parasitol. 17: 27-34.

. 1963. Chaetotaxie et classification des Speleognathinae. Bull. Inst. Roy. Sci. Nat.

Belg. 39(9) : 1-80.

. 1970. Nomenclature des poils idiosomaux et description de trois especes nouvelles

dans la famille Ereynetidae (Trombidiformes) . Acarologia. 1,2(2): 314-325.

. 1971. Cle et liste des especes du genre Boydaia Womersley (Ereynetidae: Trom-
bidiformes). Acarologia. 13(1): 98-112.

, and T. H. G. Aitken. 1968. Les Acariens parasites nasicoles des oiseaux de

Trinidad (Indes Occidentals) . II. Ereynetidae: Speleognathinae. Bull. Ann. Soc.
Roy. Ent. Belg. 104: 80-84.

— , and . 1970. Acariens nasicoles d’oiseaux et de mammiferes du Bresil. IV.

Nouveaux Ereynetidae (Trombidiformes) et Turbinoptidae (Sarcoptiformes) de la
region de Belem (Nord Bresil). Acarologia. 12(2): 1326-1338.

, and K. E. Hyland. 1970. Acariens nasicoles des oiseaux du Mexique. III. Fa-
milies Ereynetidae et Turbinoptidae. Bull. Ann. Soc. Roy. Ent. Belg. 106: 37-46.
Ford, H. G. 1959. Boydaia tyrannis n. sp. (Acarina, Speleognathidae), a new mite from
the nasal cavity of the Eastern Kingbird, Tyr annus tyr annus (Linnaeus). Trans.
Amer. Micros. Soc. 78(4) : 379-385.

Pence, D. B. 1973. The nasal mites of birds from Louisiana. VII. The Ereynetidae
(Speleognathinae). J. Parasit. 59(2): 364-368.

Vol. LXXXIII, September, 1975

209

Elliptochthoniidae, A New Mite Family (Acarina: Oribatei)
From Mineral Soil In California

Roy A. Norton

Department of Forest Zoology, S.U.N.Y. College of Environmental Science
and Forestry, Syracuse, New York 13210

Received for Publication December 27, 1974

Abstract: A new oribatid mite, Elliptochthonius profundus n. gen., n. sp., is described

from mineral soil in a coniferous ecosystem in northern California, and a new family, the
Elliptochthoniidae, is proposed. Relationships with the Parhypochthonoidea and Enarthro-
nota are discussed.

Grandjean (1947) proposed the Enarthronota, without giving it a specific
hierarchic rank, to include all the diverse macropyline oribatid families in
which the notogaster is provided with one to three transverse sutures. In his
revision of major groups in the Oribatei (Grandjean, 1969), the Enarthronota
was divided into seven superfamilies. Another macropyline superfamily, the
Parhypochthonoidea Hammen, was considered as having rank, again unspeci-
fied, equivalent to the Enarthronota. Balogh (1972) more conservatively placed
both these groups, with some deletions from the Enarthronota, in the Arthronota.
The purpose of this paper is to describe an unusual new family, genus and
species of oribatid mite related to these groups, which may prove important in
future studies of higher categories in the Oribatei.

The specimens were part of a quantity of oribatids sent to me for identi-
fication by John M. Wenz, University of California, Berkeley, in conjunction
with a study of the effects of air pollutants on a coniferous ecosystem in
California, sponsored by the Environmental Protection Agency. The site was
a mixed stand of ponderosa and jeffrey pines ( Pinus ponder osa Laws, and P.
Jeffrey i Grev. & Balf.) at Likely Mill, Modoc Co. The new species, collected
in June, 1972, appears restricted to the deeper soil strata. It was never col-
lected in the organic layers or in the upper 10 cm of mineral soil.

The nomenclature and descriptive terminology used below are primarily
those formulated by Grandjean (1935, 1939, 1940, 1947, 1949).

Elliptochthoniidae n. fam.

This family is distinguished from other families of the Macropylina by the following
combination of characters:

1. The notogaster has a single transverse dorsal suture which continues laterally and
ventrally to form a membranous delineation between the genital-aggenital plate and epimere
IV. The result is a completely movable opisthosoma, or pygidium.

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2. The division of the genital and aggenital plates is incomplete, and disappears in the
posterior third. The adanal plates are broadly fused posteriorly.

3. The latero-opisthosomal gland is present.

4. The gnathosoma is stegasime and has undergone structural and chaetotaxic modifica-
tions, including the presence of a single pair of adoral setae, the fusion of the palpal
trochanter and femur, and the reduced setation of the palp (see description).

Type genus: Elliptochthonius n. gen.

The name is derived from the Greek elleipsis , meaning oval, and chthon, meaning earth.
Because of the monotypic nature of the family, I make no attempt here to delineate generic
characters.

Type species: Elliptochthonius profundus n. sp.

Elliptochthonius profundus n. sp.

The specific epithet is the Latin profundus , meaning deep.

Female

Body elongate, oval, dorso-ventrally flattened. Average length of 5 slide-mounted speci-
mens 576 fi (range 565/1-595/0 . Average width at level of seta d3 202/4 (range 200/4-
209/4). Color in alcohol is light yellow.

Prodorsum : Prodorsum roughly triangular in shape from above (Fig. 1) ; rostrum rounded
centrally, but laterally with irregular teeth (Fig. 3). Integument very finely pitted, with
small superimposed tubercles in the postero-medial region. Podocephalic canal ( cpc ) extends
from the point of lateral articulation with infracapitulum to level of acetabulum I.

Normal setation present: rostral setae ( ro ), lamellar setae (le) and exobothridial setae
( exa , exp ) fine, simple, short; interlamellar setae (in) elongate, lanceolate, similar to noto-
gastral setae; sensillus (ss) clavate, distal portion heavily barbed.

Notogaster : Notogaster widest at level of seta d3, tapering posteriorly (Fig. 1). Integu-
mental pitting inconspicuous anterior to setal row e or /, increasing in strength posteriorly ;
strong pitting abruptly stops ventrally at constriction line running parallel to setal row ps
(Figs. 2, 3).

Dorsal suture centrally located between setal rows d and e, continuing latero-ventrally
until joining the ventral membranes. Anterior half (notaspis) with six pairs of minutely
barbed setae (cl, c2 , c3, dl, d2, d3) and one pair of cupules (ia) . An expansion suture
(su) is present laterally. Posterior half (pygidium) with 10 pairs of setae (el, e2 , fl, f2,
hi, h2, h3, psl, ps2, ps3) and four pairs of cupules (im, ip, ih, ips) . Latero-opisthosomal
gland ( gla ) present, its opening dorsal to cupule ip and separated from it by a ridge-like
thickening. Another thickening present dorsal to seta h2. Two expansion sutures present
laterally (sud, suv) on either side of cupule im. Posteriorly, a sharp dorso-ventral con-
striction gives appearance of a thickened rim (Fig. 3).

Ventral Plates : Epimeres I and II separated medially by membrane; epimeres III and IV
completely fused medially (Fig. 2). Laterocoxal seta el present. Setal formula 3-2-3-4 for
epimeres I-IV (not including el).

Genital plates with eight simple setae each, six in paraxial row, two in antiaxial row;
posteriorly fused with aggenital plates, which have one seta (ag). Anal plates each with
three simple setae and one cupule (ian). Adanal plates posteriorly fused; each with four
setae, longer than anal setae, and one cupule (iad ) .

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211

Elliptochthonius profundus n. gen., n. sp.: Fig. 1. Adult female, dorsal aspect. Fig. 2.
Same, ventral aspect.

212 New York Entomological Society

Elliptochthonius profundus n. gen., n. sp.: Fig. 3. Adult female, lateral aspect (slightly
laterally flattened). Fig. 4. Infracapitulum. Fig. 5. Right chelicera, antiaxial aspect.
Fig. 6. Right palp, antiaxial aspect (slightly from below).

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213

Elliptochthonius profundus n. gen., n. sp.: Fig. 7. Right leg I, antiaxial aspect. Fig. 8.
Famulus. Fig. 9. Right leg II, antiaxial aspect. Fig. 10. Left leg III, antiaxial aspect.
Fig. 11. Left leg IV, antiaxial aspect (slightly from below). Fig. 12. Tritonymph, epimera
III-IV and ventral plates. All legs to same scale.

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Gnathosoma: Infracapitulum simple, without secondary articulation (Fig. 4) ; four pairs of
setae on ventral surface (a, ml, m2, h ) ; one pair of adoral setae {or) on lateral lips.
Rutellum (RU) with a large thumb-like projection dorso-laterally. Antiaxial fissure ( a f)
associated with lateral tooth-like structure, Laterocoxal seta e thick, blunt.

Chelicerae chelate-dentate (Fig. S) ; fixed digit bidentate, movable digit tridentate. Seta
cha small, simple; seta chb about four times as long, rapidly tapering distally. Chitinous
barbs present on both antiaxial and paraxial faces ; numbers and placement somewhat
variable.

Palp four-segmented; trochanter and femur fused (Fig. 6). Femur with one seta, genu
with none, tibia with two, tarsus with seven setae (two of them, uk and ul" eupathidic)
and one solenidion (<0 .

Legs: Setal formulas for the legs, from trochanter to tarsus, are as follows (not including
the famulus on tarsus I): leg I (0-6-5-6-18); leg II (1-4-4-4-14); leg III (2-2-4-4-13) ;
leg IV (2-2-3-4-11). Setae distributed as in Figs. 7, 9, 10, 11. Most setae are inconspicu-
ously barbed. Only proral setae (p) on tarsus I appear to be eupathidic, but this is not
certain.

Famulus (e) of tarsus I spatulate, with single long lateral bract (Fig. 8).

Solenidial formulas for genu, tibia and tarsus as follows: leg I (2-1-3); leg II (1-1-1);
leg III (1-1-0); leg IV (1-1-0). Ambulacrum of all legs tridactylous, with a highly reduced
central claw.

Tritonymph

Very similar to adult female with exceptions as follow. Length and width of single
specimen 450/z and 177/z, respectively. Epimeres III and IV longitudinally divided by wide
membranous band (Fig. 12). Genital plate with six setae, only four in paraxial row. Leg
chaetotaxy identical to adult. Ambulacrum of all legs monodactylous.

MATERIAL EXAMINED

Seven specimens, six adult females and one tritonymph, were studied. Depo-
sition will be as follows: holotype female (slide preparation) to the U.S.

National Museum, Washington, D.C.; paratype female (alcoholic) and trito-
nymph (slide preparation) to the Museum of Comparative Zoology, Cambridge,
Massachusetts; paratype female (slide preparation) to the Canadian National
Collection, Ottawa, Ontario; three paratype females retained by author.

REMARKS

1. In five of the females examined there was identical leg setation, and
these were the specimens utilized in the leg descriptions. In the sixth there
was variability, specifically the lack of seta a" on tarsus III, seta d on genu IV
and seta v' on tibia IV. Each loss was restricted to a single leg, the other of
the pair being typical.

2. There does not seem to be a true correspondence between the sutures
which I call expansion sutures ( su , sud, suv) on the notogaster and the supra-
pleural band described by Grandjean (1947) in the Enarthronota. This band
is dorsal to cupule ia in the latter group.

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215

3. Grandjean (1969) discusses at length the types of body articulations
(holoidy, dichoidy, ptychoidy) in sclerotized oribatids. To these I now add
the term trichoidy , defined as the condition of having both the protero-hystero-
somatic articulation and a podo-opisthosomatic articulation, exemplified by the
Elliptochthoniidae .

4. The placement of this family in a major group is difficult. If we assume
that the Parhypochthonoidea and Enarthronota are part of the same monophy-
letic series, the Arthronota, then the placement of the Elliptochthoniidae in
this series seems certain. Based on available information it is more likely that
Grandjean’s (1969) system is correct, that is, the Arthronota is biphyletic.

Grandjean (1969) has listed the principal characters utilized in delineating
his major macropyline groups. To gain insight on relationships let us examine
those characters which differ among the three groups in question from the
standpoint of ancestral versus derived states.

The latero-opisthosomal gland is present (ancestral) in the Parhypochtho-
noidea and Elliptochthoniidae; it is absent (derived) in the Enarthronota. The
cupules iad, ian are present (ancestral) in the former two groups and lacking
(derived) in the Enarthronota. The adult leg ambulacra have a regressive
central claw (derived) in the Elliptochthoniidae and Parky pochthonius ; the
Enarthronota are primarily monodactylous (also derived). The Parhypoch-
thonoidea have three pairs of adoral setae (ancestral) ; the Elliptochthoniidae
have one pair, as do the Brachychthoniidae of the Enarthronota (Grandjean,
1963; Reeves and Marshall, 1971). The latter family also often has a derived
solenidiotaxy identical to the Elliptochthoniidae, whereas that of the Parhypoch-
tonoidea is ancestral in comparison. Sclerotization is a derived state, present
in the Enarthronota and Elliptochthoniidae and lacking in the Parhypochthon-
oidea, but it has obviously occurred in a number of unrelated acarine lineages.

Hennig (1966) states that relationships must be proven on the basis of
shared derived (synapomorphous) characters, not shared ancestral ones. Of
the similarities noted above, synapomorphy can be shown between the Ellip-
tochthoniidae and the Brachychthoniidae for the number of adoral setae and
solenidiotaxy. It is risky, however, to base relationships on similar degrees of
numerical regression, as Grandjean (1935) observed with solenidiotaxies. The
other synapomorphic character to be considered is the tridactylous adult
ambulacrum with a regressive central claw, shared by the Elliptochthoniidae
and Parky pochthonius] the central claw is lacking in the second parhypoch-
thonoid genus, Gehy pochthonius. Although both this and the monodactylous
condition are the result of regression from ancestral tridactyly, they are obvi-
ously of two different lineages.

Inclusion of the Elliptochthoniidae in the Parhypochthonoidea would be
acceptable if the diagnostic criteria of unsclerotized integument and the related

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stegasime condition were omitted. If future workers do not wish to do so, a
separate superfamily for the new family seems unescapable.

5. I am familiar, and often agree, with criticisms of the present “top-heavy”
classification of oribatids caused by the erection of many monotypic higher
taxa. However, such problems are most significant in the Brachypylina. Here
there is extensive development of secondary integumental structures in the
adult stase which confuse relationships, combined with a general lack of
knowledge of immatures, as discussed by Balogh (1972). For the most part,
the Macropylina presents little difficulty in this regard, and monotypic taxa
are more readily accepted. In fact, they are expected in relict groups such as
the one described here.

Literature Cited

Balogh, J. 1972. The oribatid genera of the world. Akademai Kiado, Budapest. 188 pp.
71 pi.

Grand jean, F. 1935. Les poils et les organes sensitifs portes par les pattes et le palpe
chez les oribates. Bull. Soc. zool. France 60: 6-39.

. 1939. Les segments post-larvaires de l’hysterosoma chez les oribates (Acariens).

Bull. Soc. zool. France 64: 273-284.

. 1940. Les poils et les organes sensitifs portes par les pattes et le palpe chez les

oribates. Deuxieme partie. Bull. Soc. zool. France 65: 32-44.

. 1947. Les Enarthronota (Acariens). Premiere serie. Ann. Sci. natur. Zool. (11)

8: 213-248.

. 1949. Formules anales, gastronotiques, genitales et aggenitales du developpement

numerique des poils chez les oribates. Bull. Soc. zool. France 74: 201-225.

. 1963. Sur deux especes de Brachychthoniidae et leur developpement. Acarologia

5(1): 122-151.

. 1969. Considerations sur le classement des oribates. Leur division en 6 groups

majeurs. Acarologia 11(1): 127-157.

Hennig, W. 1966. Phylogenetic systematics. University of Illinois Press, Urbana. 263 pp.

Reeves, R. M. and V. Marshall. 1971. Redescription and chaetotaxy of Brachychthonius
lydiae adults and nymphs (Acarina: Oribatei). Ann. Ent. Soc. Amer. 64(2):

317-325.

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Journal of the

New York Entomological Society

Volume LXXXIII

December 1975

No. 4

EDITORIAL BOARD

Editor Dr. Karl Maramorosch
Waksman Institute of Microbiology
Rutgers University
New Brunswick, New Jersey 08903

Associate Editors Dr. Lois J. Keller, RSM
Dr. Herbert T. Street

Publication Committee

Dr. Kumar Krishna Dr. Ayodha P. Gupta

Dr. James Forbes, Chairman

CONTENTS

A New Species and Review of Sibaria (Hemiptera: Pentatomidae)

L. H. Rolston 218

Predators of the Alfalfa Weevil, Hypera postica in Western Nevada — a Green-
house Study (Coleoptera: Cureulionidae) Manzoor Hussain 226

Calling Songs of Neduba macneilli and N, sierranus (Orthoptera: Tettigoni-
idae: Decticinae) Glenn K. Morris, Ron B. Aiken and Gordon E. Kerr 229

The Effect of Temperature and Humidity on the Amount of Blood Ingested
hy the Stable Fly, Stomoxys calcitrans L. (Diptera: Muscidae)

Corey W. Smith and Elton J. Hansens 235

Abstracts of papers presented at the Eastern Division Annual Meeting,

Entomological Society of America 241

Acknowledgment 241

Book Reviews 234, 288

Index of Scientific Names of Animals and Plants for Volume LXXXIII 289

Index of Authors for Volume LXXXIII i

218

New York Entomological Society

A New Species and Review of Sibaria
( Hemiptera : Pentatomidae )

L. H. Rolston

Department of Entomology, Louisiana State University,

Baton Rouge, Louisiana 70803

Received for Publication November 5, 1974

Abstract: The genus Sibaria is redefined, a diagnosis given for the two species previously

known, and S. englemani new species, which ranges from Mexico to Colombia, is de-
scribed.

Sibaria is distinguished among American genera of Pentatomini by the unique
combination of armed femora and a short rostrum. A pair of preapical spines,
of considerable size on the anterior femora at least, constitute the principal
femoral armament; and the rostrum terminates distally at or just beyond the
mesocoxae rather than at or beyond the metacoxae as is usual in the tribe.

Three species of Sibaria are known: S. armata, inhabiting much of South
America, S. andicola, collected in Bolivia, Ecuador and Peru, and a species
ranging from southern Mexico into Colombia. The latter species has been
confused with S. armata and until now has been unnamed.

A generic description, key to the species, description of the new species and
diagnoses of the other two follow. The three species are so much alike that a
description of more than one would be largely redundant.

Sibaria Stal, 1872

Sibaria Stal, 1872, Sv. Vet. Ak. Handl. 10(4) :23.

Eyes large, together about as wide as interocular distance (Fig. 1) ; width of head little
greater than length; juga subequal in length to tylus, their lateral margins narrowing

Acknowledgements: Specimens pertinent to this study were graciously loaned by Mssr.
W. R. Dolling of the British Museum (Natural History), R. D. Engleman, R. C. Froesch-
ner of the U.S. National Museum, P. Van Doesburg of the Rijksmuseum van Natuurlijke
Historie, and P. Wygodzinsky of the American Museum of Natural History. I am espe-
cially grateful to Dr. G. Petersen of the Akademie der Landwirtschaftswissenschaften for
lend'ng the type series of Sibaria andicola.

Depositories for paratypes are designated as follows: Akademie der Landwirtschaftswissen-
schaften (AL), American Museum of Natural History (AMNH), British Museum (Natural
History) (BMNH), California Academy of Sciences (CAS), R. D. Engleman collection
(RDE), Field Museum of Natural History (FMNH), author’s collection (LHR), Museu
Rio Grandense de Ciencias Naturais (MRCN), Naturhistoriska Riksmuseum, Stockholm
(NR), Rijksmuseum von Natuurlijke Historie (RNH), Texas A & M Univ. (TAMU), U.S.
National Museum (USNM), Universidad Nacional de La Plata (UNLP), Washington State
Univ. (WSU).

New York Entomological Society, LXXXIII: 218-225. December, 1975.

Vol. LXXXIII, December, 1975

219

plates, viewed with anterior and posterior margins of last sternite on same focal plant; first
gonocoxa (gx 1), second gonocoxae (gx 2). Fig. 5. First gonocoxa, viewed with three angles
on same focal plant. Fig. 6. Right paramere. Fig. 7. Genital cup; paramere (p), proctiger
(pr). Fig. 8. Posterior margin of pygophore, ventral view. Fig. 9. Theca and related
structures, lateral view; thecal process (tp). Fig. 10. Same, dorsal view; conjunctiva (c).
Fig. 11. Same, ventral view; penisfilum (pe).

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New York Entomological Society

rapidly before eyes, exposing antenniferous tubercles from above; distal end of first antennal
segment reaching apex of head. Pronotum contiguous with eyes; anterolateral margins
entire, obtusely rounded vertically. Scutellum as long as wide; frena extending along basal
two-thirds. Costal angle of coria acute, surpassing scutellar apex by about one-third length
of scutellum.

Bucculae roundly truncate at base of head, extending to distal end of first rostral seg-
ment; apex of rostrum reaching or just surpassing mesocoxae. Sterna neither sulcate nor
carinate along meson excepting low broad mesosternal carinae produced notably only near
anterior mesosternal margin. Inferior surface of femora armed with stout pair of preapical
spines and pair of tubercles basad of spines, these often reduced progressively on middle and
posterior femora, the latter then armed only with preapical pair of tubercles (Fig. 22) ; all
tibiae sulcate. Abdomen without basal spine or tubercle.

Tubercles of proctiger nearer base than apex (Fig. 21). Thecal processes arising within
theca, compressed, curving ventrad (Figs. 9, 18, and 26) ; penisfilum lying on median
vertical plane, surrounded by median penal lobes.

Spermathecal pump convoluted; spermathecal bulb digitiform (Figs. 3 and 15).

Type species : Mormidea armata Dallas, 1851, by monotypy.

Relationship : The form of the aedeagus and spermatheca, as well as the armament of the
proctiger and femora, suggest a close phylogenetic relationship between this genus and
Ladeaschistus Rolston, 1973.

Key to the Species

1. More than basal half of fifth antennal segment pale S. andicola Breddin

1' Less than basal half of fifth antennal segment pale 2

2. Median emargination in posterior edge of pygophore broad, deep, subquadrate, flanked
on each side by stout tubercle projecting caudad beyond posterolateral angles of pygo-
phore (Figs. 12 and 13) ; first gonocoxae (basal plates) each concavely emarginate pos-
teriorly at lateral limit of second gonocoxae (Figs. 14 and 16) S. armata (Dallas)

2' Median emargination in posterior edge of pygophore little wider than proctiger, flanked
on each side by shallower bisinuate emargination (Figs. 7 and 8) ; posterior margin of
each first gonocoxae sinuous, without emargination at lateral limit of second gonocoxae
(Figs. 4 and 5) S. englemani n. sp.

Siharia englemani, n. sp.

Sibaria armata; Distant, 1880-1890, p. 57 (in part), PI. 5, fig. 17 (1880) and p. 329 (1890)
(nrsidentification) ; Lethierry and Severin, 1893, p. 126 (in part) ; Kirkaldy, 1909, p. 62
(in part).

Overall light brown to fuscous above with black humeri, yellowish white beneath ; punc-
tation of dorsum rather dense, black, on pronotum and scutellum arranged mostly in irreg-
ular rows with a general transverse orientation; usually eight pale spots on dorsum, one
along posterior margin of each cicatrice near mesial limit, three along base of scutellum,
one of these mesial and one beside small black fovea in each basal angle (some or all
occasionally obscure), a spot on apex of scutellum, another on each corium near distal end
of radial vein. Antennae mostly black, ventral and mesial surfaces (except distally) of
first segment, basal ring on segments three and four, basal .2 to .4 of segment five, and
sometimes longitudinal streaks on segments two and three, pale ; length of segments 0.4 to
0.5; 0.8 to 0.9; 0.9 to 1.1; 1.5 to 1.8; 1.5 to 1.7 mm; width of head across eyes 1.8 to 2.0
mm, length 1.6 to 1.7 mm. Humeri acutely produced laterad and somewhat cephalad (Fig.

Vol. LXXXIII, December, 1975 221

Figs. 12-22. Sibaria armata. Fig. 12. Genital cup, parameres and proctiger removed.
Fig. 13. Pygophore, ventral view. Fig. 14. Genital plates; first gonocoxa (gx 1), second
gonocoxae (gx 2). Fig. 15. Distal portion of spermatheca; spermathecal bulb (sb) , sperma-
thecal pump (sp). Fig. 16. First gonocoxa, viewed with three angles on same focal plant.
Fig. 17. Right paramere. Fig. 18. Theca and related structures, lateral view; thecal process
(tp). Fig. 19. Same, dorsal view; conjunctiva (c). Fig. 20. Same, ventral view; median
penal lobe (mpl). Fig. 21. Proctiger. Fig. 22. Posterior face of right front femur and tibia.

222

New York Entomological Society

Figs. 23-28. Sibaria andicola. Fig. 23. Genital cup, parameres and proctiger removed.
Fig. 24. Posterior margin of pygophore, ventral view. Fig. 25. Right paramere. Fig. 26.
Theca and related structures, lateral view; median penal lobe (mpl). Fig. 27. Same, dorsal
view; thecal process (tp). Fig. 28. Same, ventral view; conjunctiva (c).

2) ; pronotal width at humeri 7.3 to 8.3 mm, length at meson 2.3 to 2.7 mm. Scutellar
width 3.2 to 3.6 mm, length subequal; apex narrowly rounded. Boundary of coria and
their membrane slightly sinuous ; membrane fuscous, veins simple or bifurcate, varying
considerably in number. Connexiva narrowly exposed; punctation dense, fine; color pale
at sutures and in subquadrate marginal area in middle of each segment, otherwise dark;
posterior angle of each segment produced as small acute spine.

Venter of head punctate only along bucculae, immaculate but for fuscous mark extending
from eye over superior surface of antenniferous tubercle and continuing briefly cephalad.
Pleura impunctate in irregular areas, most consistently so laterad of procoxae. Evaporative
area on each side matte, sparingly rugose, extending about halfway from ostiole toward
lateral margin of metapleuron. Legs with large fuscous spots and maculae. Abdominal
venter sparsely and weakly punctate about spiracles and large subspiracular callouses,
otherwise virtually impunctate ; black edge of lateral margins interrupted in middle of
each segment.

Posterior edge of pygophore pentasinuate, with a deep median concavity about as wide
as proctiger and on each side two lesser concavities (Figs. 7 and 8) ; margin between
lateral and median concavities expanded, this portion dorsad of intermediate convexity
and with elongate black impression. Anterolateral margins of genital cup produced near
apex of parameres, concealing from above a thin subvertical carina on wall of genital cup.
Head and base of parameres bent rather abruptly from lateral view, shaft entire (Fig. 6).

Vol. LXXXIII, December, 1975

223

Conjunctiva trilobed, a small median lobe above median penal lobes and bifid lateral lobes
(Figs. 9, 10 and 11).

First gonocoxae each evenly sinuous along posterior edge, without emargination above
lateral limits of second gonocoxae (Figs. 4 and 5), Types: Holotype. Male, labeled

Panama, Gatun Dam, 2-IX-1973, D. Engleman, Coll. Deposited in U.S. National Museum,
type no. 72134.

Paratypes : 33 8 $, 23 $ 2. Colombia: (a) Magdalina, 11°10/N,76°08'W, Apr. 1973,

800M, M. Madison, Coll, (b) on piper. (2 AMNH ; 8 BMNH; 28 8, 2 LHR ;

8, 2 MRCN, 8, 2 UNLP). Costa Rica: (a) Dec. 20, 1949, Darwin Norby. (b) Finca
Los Cusingos, San Isidro del General, Quizarra. ( 2 WSU) ; Collection Schild-Burgdorf,
San Carlos (2 USNM) ; (a) Turrialba, (b) Tucurriquel (2 USNM). El Salvador: No.
71458, 10.23.56, Santa Tecla, Col. PAB. ($ USNM). Guatemala: (a) Morales, Jan. 1930,
J. J. White, (b) 103 (c) J. C. Lutz Collection, 1961. (2 USNM). Mexico: Tolosa,

Oaxaca, Aug. 25, 1947, B. Malkin. (8 AMNH). Panama: (a) Bugaba, 800-1500 ft.,

Champion, (b) Ex Godman and Salvin. ( $ AMNH) ; (a) as above (b) P. R. Uhler
Collection (c) Sibaria armata. ( $ USNM) ; Cerro Campina, 800M, Panama Prov., 1
July 72, Coll. D. Engleman ( 8 RDE) ; (a) Portobella, 18.4.12, Pan. (b) A. Busch Coll.
(8 USNM). Panama Canal Zone: Barro Color., V-5-37 (2 USNM); Barro Colorado
So., VII-8-33 ( 8 USNM); Coco Solo Hospital, 9°21/N,79°5l/W, 28-1-73, Engleman. (2
RNH) ; (a) Corozal, 1-21-1929 (b) Collector, C. H. Curran. (8 AMNH); Fort Kobbe,
8°54/N,79°35'W, 22-IX-73, Col: D. Engleman ( 2 AL) ; Ft. Sherman, 30 July 72, Coll:
Engleman, ( 2 RDE) ; Fort Sherman, 9°20/N,79°58'W, 2 June 73, Col: D. Engleman.

(2 BMNH; 3 8 8 RDE); Galeta Is., 9°32/N,79°53/W, 30-VIII-73, Col: D. Engleman.
(2 FMNH ; 8 RDE; 2 TAMU) ; 5 mi. E. Gamboa, 1 Oct. 72 (8 RDE); same data as
holotype (8, 2 CAS; 4 8 8 LHR) ; Gatun Spillway, 9°20/N,79°58/W, 2 June 73, Col:
D. Engleman. (3 8 8 RDE, 8 RNH) ; Madden Reservoir, 29-IX-73, Col: D. Engleman.
(8 RDE); Margarita, 25-28-X-1972, L. H. Rolston. (8 AL; $ FMNH; 8 LHR;
8, 2 NR; 8 TAMU); Pipeline Road, 2 January 72, Col: D. Engleman. (3 2 2 RDE);
Pina Road, 9°15/N,79°57/W, 2-IX-73, Col: D. Engleman. (3 2 2 RDE).

Distribution: From Vera Cruz, Mexico, to Magdalena, Colombia. The southern distribution
may prove more extensive when northwestern South America is better collected.

Comments: Distant mistook this species for its common South American congener, and

all of his records of S. armata in Middle America pertain to S. englemani, as does his
illustration of a specimen from Guatemala.

Adult specimens have been taken feeding on the inflorescence of piper plants in Panama
and Colombia.

This species is named with pleasure for R. Dodge Engleman, M.D., whose interest in
biology extends beyond medicine and continues a tradition that has contributed much to
the systematics of insects.

Sibaria armata (Dallas, 1851)

Mormidea armata Dallas, 1851, p. 125; Walker, 1867, p. 255.

Sibaria armata ; Stal, 1872, p. 23; Distant, 1880, p. 57 (in part); Lethierry & Severin, 1893,
p. 126 (in part) ; Van Duzee, 1901, p. 344 (list) ; Kirkaldy, 1909, p. 62 (in part) ; Becker
& Grazia-Vieira, 1971, p. 20 (list).

From none to basal. 40 of fifth antennal segment pale.

Emargination in posterior edge of pygophore deep, wide, subquadrate, flanked on each

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New York Entomological Society

side by tubercle projecting posteriorly beyond posterolateral angle of pygophore (Figs. 12
and 13). Small tubercle beneath production on anterolateral margins of genital cup par-
tially visible from above. Reticulate face of parameres ovoid, shaft incised shallowly near
head, base greatly expanded and crested on ventral face (Fig. 17). Lateral conjunctival
lobes each with ventral diverticulum near base and toward apex a second diverticulum
directed obliquely mesad, the opposed apical diverticula overlapping when fully inflated;
median conjunctival lobe small, bifid; thecal processes largely concealed except from lat-
eral view (Figs. 18, 19 and 20).

Posterior edge of first gonocoxae concavely emarginate at lateral limits of second gon-
ocoxae (Figs. 14 and 16).

Distribution : Probably present throughout most of South America. Recorded or seen from
Argentina (Missiones), Bolivia, Brazil, Ecuador, French Guiana, Guyana, Paraguay, Peru,
Surinam, Trinidad, and Venezuela.

Comment : The pale basal ring on the fifth antennal segment of a minority of specimens
does not seem characteristic of any region since specimens so marked came from widely
separated places: Guyana, Bolivia, Brazil and Paraguay.

This species has been reared on the inflorescence of piper.

Sibaria andicola Breddin

Sibaria andicola Breddin, 1904, p. 49; Kirkaldy, 1909, p. 62; Gaedike, 1971, p. 79.

Basal .60 to .85 of fifth antennal segment whitish, apex dark.

Posterior edge of pygophore arcuately concave with short posteriorly directed projection
on each side nearer lateral angles than meson (Figs. 23 and 24) ; pygophoral margin at
projection somewhat expanded, not impressed. Carina beneath production on anterolateral
margins of genital cup oblique, directed posteroventrally from production. Shaft of para-
meres incised near head; base crested on ventral face (Fig. 25). Median lobe of conjunc-
tiva quite long, each lateral lobe diverticulate ventrally (Figs. 26, 27 and 28).

Genital plates of female as in S. armata.

Distribution : Known so far from the eastern slopes of the Andes: Napo province in

Ecuador; Amazonas, Cusco and Huanuco departments in Peru; and El Beni department
in Bolivia.

Comment: No distinction between females of S. andicola and S. armata has been found
other than the proportion of pale to dark color on the fifth antennal segment. This pro-
portion varies in both species and too few specimens of S. andicola are available to estab-
lish useful confidence limits on variability. The reliability of this character in separating
all females of the two species is therefore suspect. In describing S. andicola , Breddin men-
tioned the darker dorsum and obscurity of the pale dorsal spots relative to S. armata , but
neither the general color nor clarity of the spots are diagnostic.

In the few specimens examined, the femoral spines are reduced on the middle femora
and represented by small tubercles on the posterior femora.

A single specimen lacking the fifth antennal segments, in the British Museum (Natural
History), is apparently this species, but it differs from the males of the type series in
having the projections on the pygophoral margin more prominently developed.

Literature Cited

Becker, M., and J. Grazia -Vieira. 1971. Contribuicao ao conhecimento da superfamilia
Pentatomoidea na Venezuela (Heteroptera) . Iher. (Zool.) 40: 3-26.

Vol. LXXXIII, December, 1975

225

Breddin, G. 1904. Neue Rhynchotenausbeute aus Sud-Amerika. Soc. Entomol. 19: 49-
50, 58.

Dallas, W. S. 1951. List of the specimens of hemipterous insects in the collection of the
British Museum. London.

Distant, W. L. 1880-1892. Insecta. Rhynchota, Hemiptera-Heteroptera. In Godman, F.

D. and O. Salvin, Biologia Centrali Americana. London, Vol. I.

Gaedike, H. 1971. Katalog der in dem Sammlungen des ehemaligen Deutschen Ento-
mologischen Institutes aufbewahrten Typen V. Beitr. Ent. 21(1/2): 79-159.
Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera) . Vol. I Cimicidae.
Berlin.

Lethierry, L. and G. Severin. 1893. Catalogue general des Hemipteres. Vol. 1. Brussels
& Berlin.

Rolston, L. H. 1973. A new South American genus of Pentatomini (Hemiptera :Penta-
tomidae). J. N. Y. Entomol. Soc. 81(2): 101-110.

Stal, C. 1872. Enumeratio hemipterorum II. Kongliga Svenska Vetenskaps-Akademiens
Handlingar. 10(4): 1-159.

Van Duzee, E. P. 1901. Notes on some Hemiptera from British Guiana. Trans. Am.
Entomol. Soc. 27: 343-352.

Walker, F. 1867. Catalogue of the specimens of Hemiptera-Heteroptera in the collection
of the British Museum. 2: 241-417.

ADDENDUM: Since the author of “A new genus and two new species of Achipteriidae
from New York State (Acari: Cryptostigmata; Oribatei)” in Vol. 82: pp. 177-182 of the
Jour. N.Y. Ent. Soc. failed to indicate a type species for the genus Dentachipteria he wishes
to correct this by designating Dentachipteria ringwoodensis as the type species. He wished
to thank Dr. M. D. Definado of the N.Y. State Museum and Science Service for bringing
this to his attention.

226

New York Entomological Society

Predators of the Alfalfa Weevil, Hyper a postica in
Western Nevada — a Greenhouse Study.

(Coleoptera: Curculionidae)

Manzoor Hussain* 1

Division of Biochemistry and Agricultural Pest Control,

University of Nevada, Reno, Nevada 89507

Received for Publication December 2, 1974

Abstract : Some insect predators of the alfalfa weevil commonly found in the alfalfa

fields of Western Nevada were screened for their predatory efficiency against the alfalfa
weevil larvae and the pea aphids. By offering each predator species a combination of
the alfalfa weevil larvae and pea aphids as diet, their preference for the host insects was
determined.

The three lady beetle species, the big eyed bug and the nabid bug preferred to feed upon
the pea aphids; whereas, the soft winged flower beetle, Collops bipunctatus Say, significantly
preferred to feed upon the alfalfa weevil larvae and hence could be of importance in
the biological control of the alfalfa weevil.

INTRODUCTION

Biological control of the alfalfa weevil, Hypera postica (Gyllenhal), has been
given considerable attention but most of the studies in this field have been
confined to the weevil parasites. Only a limited number of studies have been
done involving insect predators of this pest. In an alfalfa field in Utah,
Webster (1912) observed that the darkling ground beetle, Eleodes sulcipennis
Mann., the soft winged flower beetle, Collops bipunctatus (Say), the imperfect
tiger beetle, Cicindela imperfecta Lee., larvae and adults of Hippodamia sinuata
var. spuria and larvae of Hippodamia convergens and Coccinella nine-notata
fed on the alfalfa weevil larvae. Adults of H. convergens were reported by Essig
and Michelbacher (1933) to feed upon the larvae of the alfalfa weevil. Kaddou
(1960) found that H. quinquesignata (Kirby) preyed upon small alfalfa
weevil larvae in Utah. Clausen (1962) also reported that C. bipunctatus fed on
the alfalfa weevil larvae; and Yada.va and Shaw (1968) studying predatory
behavior of some Coccinellids, found them to prefer pea aphids over the alfalfa
weevil larvae. This report includes a greenhouse study of the predatory ef-
ficiency of some of the entomophagous insects commonly found in the alfalfa
fields of Nevada, against the alfalfa weevil larvae; and if these predators ex-

Acknowledgement : I thank Dr. W. H. Arnett of the Division of Biochemistry and

Agricultural Pest Control, University of Nevada, Reno, Nevada for his valuable advise
and suggestions during this work.

1 Present address: Department of Plant Protection, College of Agriculture, Pahlavi Uni-
versity, Shiraz, IRAN.

New York Entomological Society, LXXXIII: 226-228. December, 1975.

Vol. LXXXIII, December, 1975

227

hibited any preference for the pea aphids, Acyrthosiphon pisum (Harris) or
the alfalfa weevil larvae.

METHODS

Six species of entomophagous insects were used in this study. These were
three lady beetle species; Hippodamia convergens (G-M), Hippodamia sinuata
disjuncta (Timberlake) and Coccinella transver so guttata (Fald.); a soft winged
flower beetle, Collops bipunctatus (Say) ; a big eyed bug, Geocoris pallens pattern
(Stal) and a nabid bug, Nabis americoferus Carayon. The alfalfa weevils and
the predators were collected from the alfalfa fields in Gardnerville, Fallon and
Lovelock, Nevada. The alfalfa weevil larvae were reared on alfalfa plants in
the greenhouse. Adult weevils were allowed to oviposit in the caged alfalfa
stems, eggs were removed and incubated at 81 ± 1 F and 70% relative humidity
in a temperature controlled cabinet. Larvae hatching out of these eggs were
transferred to caged alfalfa plants in the greenhouse and were used for feeding
experiments. The pea aphids were reared on potted alfalfa plants in the green-
house. The feeding experiments were conducted in the greenhouse where the
temperature ranged between 50 F at midnight to 85 F at midday, and the
relative humidity ranged between 30 and 50%. The feeding behavior of the
predators was studied in petri dishes with screened tops.

The predators were starved for a period of 1 2 hours prior to the feeding study.
Each sex of a predator species was individually allowed to feed upon a combina-
tion of 40 pea aphids and 40 alfalfa weevil larvae for a period of 8 hours, but
an observation was made every two hours. Small sized pea aphids were used
in combination with the first instar weevil larvae, medium sized pea aphids with
the second instar larvae and large pea aphids with the third instar weevil
larvae. Ten replicates were obtained for each experiment and the average so
obtained was used to interpret the results. The student “t” test was used to
statistically analyse the results at 0.05% level.

RESULTS

The feeding of the predators on the two host species, in terms of number of
each host eaten, is shown in the table. When a combination of small pea
aphids and the first instar weevil larvae were offered, the three lady beetle
species and C. bipunctatus showed no significant preference for either of the
two hosts and each host was preyed upon at random. The two Hemiptern
predators, however, decidedly preferred to feed upon the pea aphids. Similarly,
H. convergens and C. transver so gut tat a did not show any significant preference
for either host when a combination of second instar weevil larvae and medium
sized pea aphids were offered as diet. H. sinuata disjuncta , G. pattens pallens
and N . americojerus preferred to feed upon pea aphids, while C. bipunctatus
significantly preferred to feed upon the weevil larvae. In case when the third

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New York Entomological Society

Table 1. Average number of host insects eaten by the predator species in a period of 8 hours
when a combination of the host species was offered as diet.

Predator species

H. C. transver- H. sinuata C . bipunc- G. pallens N. ameri-

convergens soguttata disjuncta tatus pallens coferus

Host

combination

Male

Fe-

male

Male

Fe-

male

Male

Fe-

male

Male

Fe-

male

Fe-

Male male

Male

Fe-

male

Small aphids

16.3

21.0

19.5

18.2

9.0

13.3

7.1

5.7

2.9*

6.1*

6.4*

10.3*

1st instar larvae

14.3

17.1

16.1

15.8

7.4

9.7

8.1

7.1

1.3

3.0

3.2

6.3

Medium aphids

4.2

6.3

8.6

8.7

4.9*

5.9*

4.2

4.0

2.0*

2.7*

4.3*

4.4*

2nd instar larvae

3.6

4.4

7.1

7.2

1.7

2.5

6.4*

6.5*

0.9

1.3

2.0

Large aphids

4.2*

5.2*

3.9*

5.3*

: 3.1*

3.2*

1.7

0.8

1.8*

3.1*

2.7*

3rd instar larvae

1.0

0.9

1.8

1.6

0.3

0.9

1.9

3.1*

0.6

0.8

0.0

0.6

* Significant preference at

0.05%

level,

using

student

“t”

test.

instar weevil larvae and large pea aphids were presented together, all the
predator species except C. bipunctatus , preferred to feed upon the pea aphids;
whereas the female C. bipunctatus preferred to feed upon the weevil larvae and
the male did not show preference for either of the two hosts.

Although all the predator species used in this study feed upon the alfalfa
weevil larvae in the presence of the pea aphids, a host of great competitive im-
portance to the alfalfa weevil larvae, the Collops beetles distinctly prefer the
weevil larvae as opposed to the pea aphids and could be of importance in the
biological control of the alfalfa weevil. These studies were conducted in the
greenhouse conditions and hence may not truely represent the behavior of
these predators in the field.

Literature Cited

Clausen, C. P. 1962. Entomophagous Insects. Hafner Publishing Co. N.Y., N.Y. 545 pp.
Essig, E. O., and Michelbacher, A. E. 1933. The alfalfa weevil. California Agr. Exp.
Sta. Bull. 567 : 3-99.

Kaddou, I. K. 1960. The feeding behavior of Hippodamia quinquesignata (Kirby) larvae.

U. of Calif. Pub. in Entomol. No. 5, 181-232.

Webster, F. M. 1912. Preliminary report on the alfalfa weevil. U.S.D.A. Bur. Entomol.
Bull. 112: 1-47.

Yadava, C. P., and Shaw, F. R. 1968. The preference of certain Coccinellids for pea aphids,
leaf hoppers and alfalfa weevil larvae. J. Econ. Entomol., 61: 1104-5.

Vol. LXXXIII, December, 1975

229

Calling Songs of Neduba macneilli and /V. sierranus
( Ortlioptera : Tettigoniidae : Decticinae )

Glenn K. Morris, Ron B. Aiken and Gordon E. Kerr

Department of Zoology, Erindale College, University of Toronto,
Mississauga, Ontario

Received for Publication January 16, 1975

Abstract: The calling songs of decticines remain largely undescribed. Songs of Neduba
macneilli and N . sierranus were recorded and analysed. These species have mirror-image
tegmina and individual specimens exhibit reversed wing overlap. Wing symmetry and the
elaborate pronotum characteristic of this genus are discussed as adaptations which increase
the efficiency of sound radiation.

INTRODUCTION

The calling songs of many shield-backed katydids (Decticinae) remain un-
described. Rentz and Birchim (1968), in a comprehensive revision of nearctic
decticines, indicate the potential value of such calls in the resolution of the
still confused taxonomy of Decticinae. The songs are also of interest in their
own right as elements of communicative behaviour. Rentz and Birchim pro-
vide sonograms of eight decticine species, including Neduba macneilli Rentz
and Birchim. The present paper contains a detailed description of calling song
in N . macneilli and N . sierranus Rehn and Hebard, together with comments
on the tegminal structure of these insects.

MATERIALS AND METHODS

Male specimens of N. macneilli were collected on 24 July 1972, 1 mile
west of Tom’s Place, California, the type locality of this species. They were
located by their stridulation just after dusk (2200 hr) on pinyon pines, 1-2 m
above the ground (temperature 15-16°C).

Males of N. sierranus were taken during the early part of the night (16-18°
C) in Yosemite National Park, California, on 28 July 1972. Singers of this
second species were perched near ground level in an open cedar forest dominated
by sugar pine ( Pinus lambertiana) and incense cedar ( Libocedrus decurrens) .
The collection site was at an elevation of 4000 ft.

Songs were collected in the field with a Uher 4000 Report-L tape recorder
and, subsequently, living males of both species were transported to Toronto.

Acknowledgements: The authors express their appreciation for the help of Dr. Tom

Wood, Dr. Warren Cothran, Dr. D. C. Rentz and Mr. James Fullard. The work was
made possible by a grant (operating 4946) from the National Research Council of Canada.

New York Entomological Society, LXXXIII: 229-234. December, 1975.

230

New York Entomological Society

Neduba macneilli

A\ — - — — song/buzz— — — — — 1 X

X PTG T?

major
. pulse train ,

ipjfHi

111*. hull I

tfSIil!

ilsllff;

minor

pulse train

iilhilflf:

PTG-

200ms

Fig. 1. Oscillographs of N. macneilli calling song, laboratory recorded at 22 °C with
the microphone positioned 4.5 cm dorsal to the insect.

Stridulations were recorded in the laboratory using a Bruel and Kjaer quarter-
inch condenser microphone (4135) and power supply (2801): during the

laboratory recording, the insects were caged as described by Pipher and Morris
(1974). The signal from the microphone was amplified (Keithley 102B) and
then recorded at 76.2 cm/sec on an instrumentation recorder (Philips ANA-LOG
7). Oscillograms were obtained with a Tektronix oscilloscope (564) and
Nihon Kohden oscillograph camera; carrier frequency spectra were determined
with a Tektronix 3L5 spectrum analyser. Specimens of both species were ex-
amined periodically for changes in tegminal overlap.

RESULTS

N. macneilli calling song is a buzz (Fig. 1A) made up of identical pulse train
groups (PTGs), each group comprised of a minor (short-duration) and major
(long-duration) pulse train (Fig. IB). The major pulse train of a specimen
with 72 file teeth contained 52 pulses (based on an average of 10 successive
major trains from a single song). At 16°C, the PTGs are easily resolved by
the human ear, each PTG lasting about 0.5 sec.

One individual, recorded in the field at this temperature, produced buzzes
of 5-12 PTGs at a rate slightly above 1 PTG/sec, with brief pauses of 3-4 sec
between the buzzes. In the laboratory, at temperatures near 23° C, buzzes were
often of much longer duration. The buzz ends abruptly with the completion of
a major pulse train and maintains a uniform level throughout. Slightly in
advance of the beginning of the buzz, there occurs a distinctive pulse pattern
(X of Fig. 1A) which appears to be a minor pulse train together with an initial
few pulses from the subsequent major pulse train.

Vol. LXXXIII, December, 1975

231

Neduba sierranus
A

song

/ \

\l \ i

ticks

buzz

(buzz)

B PTG pulse train

pulse train pulse train

Fig. 2. Oscillographs of N. sierranus calling song, laboratory recorded at 23 °C with
the microphone 4.5 cm dorsal to the insect.

N. sierranus has a bimodal (two-part) song. A specimen recorded in the
field at 16°C repeated its song at a rate of about one song per second. The
human ear resolves the song as a few brief stuttering ticks, leading without pause
into a buzz (Fig. 2A). The buzz is a single pulse train (Fig. 2B). A specimen
with 170 file teeth produced 107 pulses in each buzz pulse train (averaged
over 10 consecutive songs). The tick mode consists of a PTG repeated
(usually) 3 or 4 times as the song is initiated. Each group is comprised
of a low-rate pulse train preceding a sharp-fronted, more intense, high-rate
pulse train (Fig. 2B).

The most intense carrier frequencies of both species lie near 20 kHz (Fig.
3A, B). N. macneilli has a main intensity peak between 15 and 22 kHz with
a lesser peak centred on 35 kHz. The dominant peak of N. sierranus occurs
within the range of 16-23 kHz and is particularly pronounced at 19 kHz. These
frequencies exceed the response capability of most tape recorders and micro-
phones; thus sonograms, such as that given by Rentz and Birchim (1968)
for N. macneilli, may present only a small fraction of the sound energy actually
produced by the animal.

In both species the tegmina are mirror images of each other. Swollen lateral
and mesal veins diverge from the wing base (A and B of Fig. 4) and subtend
a plateau of transparent cells, comprised of relatively stiff, thin cuticle. A
membranous (flexible) skirt (C) hangs ventrally from the lateral vein. The
medial margin of each tegmen functions as a scraper (D). The file (E) lies
toward the midline between the massive veins and is only weakly attached
to them. This stridulatory apparatus encloses a chamber of air on the dorsum

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New York Entomological Society

Neduba macneilli

0 20 40 60

Neduba sierranus

O 20 40 60

Fig. 3. Spectrograms of carrier frequencies in the calling songs of N. macneilli (A) and
N. sierranus (B) ; each record is a series of traces superimposed on the screen of a storage
oscilloscope during a 10 second sample of the insect’s song at a sweep rate of 20 ms/
division ; horizontal scale in kHz ; a 0 kHz marker appears at the extreme left.

of the insect. The floor of this chamber is the terga of the meso- and metathorax
and the first few abdominal segments. Each tegmen contributes the region
bounded by the two large veins as half of the chamber roof. The skirt reaches
and trails out upon the insect’s back, delimiting the sides of the chamber and
closing it off posteriorly. Tegminal structure is essentially the same in both
species; they differ only in the far greater number of teeth occurring on the
file of N. sierranus.

Tegminal overlap was found to be variable in these insects. Of five specimens
of N . sierranus , three exhibited the left over right tegminal orientation typical of
most katydids, but two males had reversed overlap — 'right over left. Of four
specimens of N . macneilli , two exhibited ‘normal’ overlap and two the reverse.
Over a one week period, none of these animals were observed to alter their
original overlap. Manual manipulation of the tegmina of freshly killed speci-
mens resulted in sound production of an identical nature with either orientation.

DISCUSSION

Left on right overlap of dimorphic tegmina has been considered universal
in Tettigoniidae (Ragge 1955). N. macneilli and N. sierranus are exceptions to

Vol. LXXXIII, December, 1975

233

Fig. 4. Ventral view of excised right tegmen of N. macneilli; lateral vein (A), mesal
vein (B), skirt (C), scraper (D), file (E).

this rule, but they are not alone in varying overlap of identical forewings. They
share this distinction with Cyphoderris monstrosa Uhler and C. buckelli Hebard,
survivors of a largely extinct family of primitive katydids (Prophalangopsidae)
(Spooner 1973).

Neduba species have a remarkably enlarged pronotum which projects rear-
ward above the tegmina. When singing the animal adopts a characteristic
posture with the abdomen dorsally concave and lowered and the pronotum
elevated, forming an acoustic horn. The plateaux of the tegmina may be
considered collectively as the driver or diaphragm. Wing symmetry may then
be seen as an adaptation promoting synchronous displacement of the tegmina,
allowing them to function as a single diaphragm. The air chamber enclosed
by the tegmina is very small relative to the wavelength involved (A 20 kHz =
172 mm; chamber dimensions approximately 1X4X5 mm) which means
that the chamber will act as a pure acoustic compliance (Beranek 1954), and
not as a resonating tube, although its presence will affect the resonance fre-
quency of the diaphragm.

The horn is too irregular to be considered an exponential horn so calculation
of its throat inductance is not possible. However, as in the case of the mole
cricket (Bennet-Clark 1970) a properly chosen compliance behind the driver
will improve its efficiency. Since the diaphragm is small relative to the wave-
length (kr = .01) even without the horn the presence of a closed box behind

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New York Entomological Society

(beneath) the diaphragm will greatly increase the efficiency of sound radia-
tion by preventing acoustic short-circuiting between the front and back of
the diaphragm (Michelsen and Nocke 1974).

Literature Cited

Bennet-Clark, H. C. 1970. The mechanism and efficiency of sound production in mole
crickets. J. Exp. Biol., 52: 619-652.

Beranek, L. L. 1954. Acoustics. McGraw-Hill, New York. 481 pp.

Michelsen, A. and H. Nocke. 1974. Biophysical aspects of sound communication in
insects. Adv. Insect Physiol., 10: 247-296.

Pipher, R. E. and G. K. Morris. 1974. Frequency modulation in Conocephalus nigro-
pleurum, the black-sided meadow katydid (Orthoptera: Tettigoniidae) . Can. Ent.,

106: 997-1001.

Ragge, D. R. 1955. The wing-venation of the Orthoptera Saltatoria with notes on
Dictyopteran wing- venation. Brit. Mus. Nat. Hist. London. 159 pp.

Rentz, D. C. and J. D. Birchim. 1968. Revisionary studies in the nearctic Decticinae.
Mem. Pac. Coast Ent. Soc., 3: 1-173.

Spooner, J. D. 1973. Sound production in Cyphoderris monstrosa (Orthoptera: Pro-

phalangopsidae) . Ann. Ent. Soc. Amer., 66: 4-5.

BOOK REVIEW

Geographic Variability in Speyeria. Arthur H. Moeck. 1975 (reprint of 1957 original).
Entomological Reprint Specialists, Los Angeles. 48 pp., 7 maps, 2 photographic plates. $3.50.

The nymphalid genus Speyeria is one of the most distinctively Nearctic of all butterfly
groups. It consists of no great number of species (the count varying greatly depending
on who is doing the classifying) but of a thoroughly bewildering mass of so-called
subspecies, local forms and varieties. Some of these are practically indistinguishable from
some assigned to other species, and can be identified only in the context of the wide-
ranging species to which they are assigned. The basic work in the modern taxonomy of
the group was done by dos Passos and Gray. Arthur Moeck made their study practically
his lifework, collecting widely and accumulating an enormous and highly valuable col-
lection. The article here reprinted, rare in its original form, is very important, setting
forth his chief opinions about the classification and geographic variation of the major
species. It will be essential to all students of the group, and valuable to all interested in
butterfly geography.

Alexander B. Klots

The American Museum of Natural History

Vol. LXXXIII, December, 1975

235

The Effect of Temperature and Humidity
on the Amount of Blood Ingested by the Stable Fly,
Stomoxys calcitrans L. (Diptera: Muscidae)1

Corey W. Smith and Elton J. Hansens2

Department of Entomology and Economic Zoology,

Rutgers University, New Brunswick, N.J. 08903

Received for Publication February 1, 1975

Abstract: The amount of blood ingested by the female stable fly, Stomoxys calcitrans ,
was studied in all combinations of 23, 32, and 38°C and 7, 43, 75, and 97% relative
humidity. No significant differences existed in the amount of blood ingested between
the 12 temperature-humidity combinations. Data show that the percentage of flies
which feed is dependent on temperature-humidity relations. The percentage of flies feeding
is greatest at high temperature and low humidity and lowest at low temperature and high
humidity.

The stable fly, Stomoxys calcitrans L., is an important blood-sucking pest
of man and animals. In resort areas many people recognize that annoyance
varies greatly with temperature and relative humidity. The behavior of flies
and the rate at which food reserves are expended depend to a large extent on
temperature and relative humidity.

While a large body of literature exists showing the effects of the environment
on mortality, growth, and fecundity of insects, little research has been done
to determine the effects of the environment on the amount of food ingested.
The purpose of this research was to determine if high temperature and low
humidity increase feeding and the amount of blood ingested by the stable
fly, S. calcitrans.

The investigations of Voegtline et al. (1965) and Wang and Gill (1970)
on the biting activities of stable flies along Lake Superior demonstrated that
the feeding activity of S. calcitrans is determined largely by the day to day
interaction between temperature and humidity. The constancy of these condi-
tions in the laboratory achieves the same result as the flies changing resting
places under field conditions in order to obtain the best temperature-humidity
combination.

According to Bursell (1964), for insects the interval between meals is longer
at cold than at hot temperatures. Applying that observation to the feeding

1 Paper of the Journal Series, New Jersey Agricultural Experiment Station, Rutgers
University — the State University of New Jersey, New Brunswick, N.J. 08903. This in-
vestigation is part of a George H. Cook Scholar Project by the senior author.

2 Undergraduate student and Research Professor, respectively.

New York Entomological Society, LXXXIII: 235-240. December, 1975.

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New York Entomological Society

habits of S. calcitrans, temperature obviously plays an important role in de-
termining the frequency with which a stable fly feeds.

A portion of the literature has been devoted to defining “bloodmeal” as it
applies to the stable fly. The confusion which exists results in conflicting data.
Thus, Suenaga (1965) found that adults having been previously fed “one or
two times after emergence” ingested on the average 16.43 mg of blood if they
were female and 9.45 mg if they were males while Parr (1962) found that
“hungry adults” from a laboratory culture maintained at 26.6°C and 80%
relative humidity took bloodmeals averaging 25.8 mg, about three times their
mean weight.

A number of laboratory procedures for feeding stable flies have been sug-
gested. Starnes (1949) 3 fed flies horse blood through eyedroppers fitted into
holes in the cages; Granett (1960) used animal membranes in the evaluation of
chemical repellents. Kashin (1965) used an electronic recording device to detect
the various phases of the mosquito bite.

MATERIALS AND METHODS

Tests were conducted using humidity chambers (Fig. 1) consisting of two
clear plastic dishes 13 cm diam X 1.5 cm fitted together to make an enclosed
chamber. In the bottom of the chamber a cardboard disc served as a base
for 1 7 cages glued to its perimeter. The cages, holding one fly each, were made
of 16 X 16 mesh aluminum screen and measured 2 X 1.5 X 1.5 cm. Flies were
inserted through small openings in the tops of the cages which were then
plugged with pieces of cotton. In the center of the disc a small dish was
placed containing 40 ml of a saturated salt solution. Saturated solutions of
NaOH, K0CO3, K tartrate, and K2S04, were used to provide relative humidities
of 7, 43, 75, and 97%, respectfully as recommended by Winston and Bates
(1960). Feeding was accomplished by inserting 3.2 cm lengths of glass tubing
with citrated beef blood through holes in the sides of the humidity chamber
corresponding in position with the cages.

Stable flies were reared under standard conditions of 27 ± 1°C ranging
from 46 to 69% RH. Freshly emerged females were supplied with 5% dextrose
solution for 2 days and then with citrated bovine blood for 10 hours. Then
the flies were anesthetized with C02 and placed in the individual cages in the
humidity chambers. After receiving flies the humidity chambers were equipped
with the saturated salt solutions, sealed with cloth tape, and placed in an
incubator containing a IV2 watt light bulb and set at one of the three tempera-
ture levels, 24, 32, or 38°C. Approximately 10 tests were run at each of the
three temperatures and four relative humidities.

3 Starnes, E. 1949. Ecology and biology of Stomoxys calcitrans in temperate climates.
Ph.D. Thesis. Rutgers Univ. 120 pp.

Vol. LXXXIII, December, 1975

237

Fig. 1. Humidity chamber.

After 17 hours of starvation, the chambers were taken out of the incubator,
one at a time, and the tape removed. Glass tubes containing blood (which
entered by capillary action) were weighed on an analytical balance and the
weight recorded to the nearest 0.1 mg. The tubes were then inserted into
the chambers. Each humidity chamber was returned to the incubator as soon
as it had been supplied with a complete set of feeding tubes. Each humidity
chamber remained in the incubator for a feeding period of 70 min after
which each capillary was weighed again. The difference between the weights
obtained before and after feeding gave a value for each tube.

Controls without flies were included in each test. The average weight of blood
lost to evaporation was determined for each temperature-humidity combination
and subtracted from value for each tube to determine the amount of blood

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New York Entomological Society

13-

12-

^ 7% RH
[H| 43% RH
^75% RH
H 97% R H

TEMPERATURE (°C)

Fig. 2. Effect of temperature-humidity on blood ingested.

ingested by each fly. Each fly tested was crushed on a paper towel to confirm
blood feeding.

RESULTS AND DISCUSSION

In calculating the amount of blood ingested at a given temperature and
humidity, the data from approximately 10 replicates were grouped. Sample
size was not constant because numbers of available female flies varied and high
mortality occurred at some temperature-humidity combinations. Mean values
of blood ingested (Fig. 2) were determined by dividing the total amount of
blood consumed at a given temperature and humidity by the total sample
size for that combination of factors. 95% confidence intervals were calculated
by use of the formula, x ± t.05S, where x is the mean value of blood ingested,
t is the t value determined by the sample size, and s is the standard deviation.
This was done to determine the significance of differences between the means
obtained at the various temperature-humidity combinations and also to de-
termine the significance of consumption differences between temperatures and
between humidities.

Vol. LXXXIII, December, 1975

239

lOO-i

7% RH

[ [43% RH

90-

75% RH

97% RH

~Z_ 80-

Li-

60-

TEMPERATURE (°C)

Fig. 3. Effect of temperature-humidity on percent feeding.

There were no significant differences in amount of blood ingested between
the 12 temperature-humidity interactions except at the 23°C-75% RH combina-
tion. A hypothesis formed prior to the collection of data and based on the
fact that the rate at which food reserves are expended and the drying power
of unsaturated air are temperature dependent was proven to be incorrect, even
though it seemed logical to predict that more blood would be ingested at high
temperature and low humidity than at low temperature and high humidity.

To determine what effect the feeding apparatus had on the amount of
blood ingested as compared to a live host, the senior author fed 15 stable
flies on his arm. Flies were weighed before and after feeding to find the
amount of blood ingested. Results showed that the amount of blood ingested
on a live host (9.6 mg) did not differ significantly from the amount of blood
ingested from an artificial source (10.6 mg).

Though the amount of blood ingested by S. calcitrans was not dependent
on temperature and humidity interactions, the percentage of flies that feed
was dependent (Fig. 3). For temperatures of 23 and 32°C the greatest per-
centage of flies took a bloodmeal at the lowest relative humidity (7%). The
lowest percentage of flies feeding at these same two temperatures occurred
at the highest relative humidity (97%). Though the greatest percent feeding

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New York Entomological Society

at the 38°C temperature occurred at 75% RH, the lowest percent feeding again
occurred at the highest relative humidity. At all humidity levels except for
75%, the largest percentage of flies feeding was to be found at a temperature
of 32 °C. These results compare favorably to those reported by Voegtline et al.
(1965). If we assume that the percentage of flies feeding at a given temperature
is proportional to biting activity, then the percentage of flies that feed must
share an inverse relationship with relative humidity. This is exactly the case
at both 23 and 32°C. Considering the percent feeding data obtained at the 23
and 32 °C levels, percent feeding increased as temperature increased at all
humidities. This, too, agrees with Voegtline’s observations.

Differences in behavior were observed between flies exposed to different
humidities prior to feeding. The least amount of activity occurred in flies
held at the 97% humidity while the greatest activity was observed at the
7% humidity. Flies that were subjected to the 7% humidity and did not feed
during the test were extremely hard to catch when attempts were made with
forceps to remove them from the chambers. However, those flies which did
feed were inactive regardless of the humidity. In fact, engorged flies made no
attempt to escape when forceps were inserted into their cages.

Literature Cited

Bursell, E. 1964. Environmental Aspects: Humidity, p. 324-58. In Rockstein, M. [ed.].

The Physiology of Insecta. Vol. 1. Academic Press, N.Y.

Granett, P. 1960. Use of an animal membrane in the evaluation of chemical repellents
against the stable fly. J. Econ. Ent., 53: 432-5.

Kashin, P. and H. G. Wakeley. 1965. An insect bitometer. Nature (London), 208:
462-4.

Parr, H. C. M. 1962. Studies on Stomoxys calcitrans in Uganda, East Africa. Notes on
life history and behavior. Bull. Entomol. Res., 53: 437-43.

Suenega, O. 1965. A rearing method of stable flies and quantity of blood taken by a fly.
(In Japanese). Endem. Dis. Bull. Nagasaki Univ., 7: 296-301. (Rev. App. Entomol.
B, 56: 709).

Voegtline, A. C., G. W. Ozburn, and G. D. Gill. 1965. The relation of weather to
biting activity of Stomoxys calcitrans along Lake Superior. Mich. Acad. Sci., Arts
Lett. 50: 107-14.

Wang, Tu-hwa E. and G. D. Gill. 1970. Effect of temperature and humidity on mortality
of adult stable flies. J. Econ. Entomol., 63: 1666-8.

Winston, P. W. and D. H. Bates. 1960. Saturated solutions for the control of humidity
in biological research. Ecology, 41 : 232-7.

Vol. LXXXIII, December, 1975

241

ABSTRACTS

FORTY-SEVENTH ANNUAL MEETING
EASTERN BRANCH

ENTOMOLOGICAL SOCIETY OF AMERICA

This year for the first time we are publishing ABSTRACTS of papers, as
well as of symposium presentations, of the Forty-seventh Annual Meeting of
the Eastern Branch, Entomological Society of America, held Oct. 1975, in
Philadelphia, Pa. The New York Entomological Society and the Eastern
Branch, ESA, hope that publication of ABSTRACTS will become a regular
procedure and that future December issues will be devoted, in part, to subjects
presented at the Annual Meeting of the Eastern Branch.

ACKNOWLEDGMENT

The Editors wish to express their appreciation to all those who have helped
in reviewing manuscripts submitted during 1975 for publication in the Journal:
Marion Brooks-Wallace, Elmer P. Catts, Mercedes Delfinado, Robert F. Denno,
C. Clayton Hoff, Alexander B. Klots, Evert E. Lindquist, Arthur H. McIntosh,
Glenn K. Morris, Sally B. Padhi, Radclyffe B. Roberts, John B. Schmitt,
Daniel J. Sullivan, Robert Traub, Asher E. Treat, George W. Wharton, and
Pedro Wygodzinsky.

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New York Entomological Society

Seasonal Variations in Activity of Apanteles melanoscelus
Ratzeburg (Hymenoptera: Braconiclae) Adults as Related to
Seasonal Variations in Age Structure of its Host, Porthetria
dispar (L.) (Lepidoptera: Lymantriidae)

R. M. Weseloh

Department of Entomology, Connecticut Agric. Exp. Station,

New Haven, Conn. 06504

Apanteles melanoscelus Ratzeburg does not develop successfully in large
larvae of the gypsy moth, Porthetria dispar (L.). This study was conducted to
determine if the effectiveness of the second field generation of the parasitoid is
reduced because of this. Field attack frequency of female A. melanoscelus was
monitored weekly by confining laboratory reared 1st and 2nd stadia gypsy
moth larvae to branches of trees and then rearing them in the laboratory to
see if they were parasitized. Also, weekly collections of 100 gypsy moths were
taken in the field, their stadia recorded, and reared to determine percent
parasitism. First and 2nd stadia gypsy moth larvae were abundant in the field
from May 20 up to June 13. Third instars, which A. melanoscelus does attack
but with difficulty, occurred through June 19. Attack frequency of A. mela-
noscelus on confined, laboratory-reared small larvae was low until June 10-13,
when it increased dramatically and then declined in subsequent weeks. This
peak was probably due to second generation emergence of the parasitoid. Thus,
A. melanoscelus adults are most abundant when most gypsy moth larvae are
too large to be suitable hosts (by June 13, 79.3% of the field-collected cater-
pillars were 4th instars and larger). This could be expected to influence con-
siderably the parasitoids, ability to control the pest.

Dimilin Toxicity to Apanteles melanoscelus (Ratzeburg)
(Hymenoptera: Braconiclae) and Effects on Field Populations

J. Granett, R. M. Weseloh and D. M. Dunbar
The Connecticut Agricultural Experiment Station, Box 1106, New Haven 06504

In the laboratory Apanteles melanoscelus (Ratzeburg) larvae were treated
with Dimilin®, l-(4-chlorophenyl)-3-(2,6-difluorobenzoyl)-urea, by feeding
treated artificial diet to the gypsy moth host. The parasitoid EC50 was 0.0059
ppm in comparison with 0.0075 ppm for the unparasitized gypsy moth. At
low treatment rates the parasitoids died during the pupal-adult molt within
the cocoon. At high treatment rates, however, the parasitoids died as larvae
within the gypsy moth larvae. Treatments while the parasitoids were 2nd-3rd
stadial larvae had less effect than treatments earlier in the parasitoids’ develop-

Vol. LXXXIII, December, 1975

243

ment. Dimilin did not apparently affect parasitoids treated as adults. The
toxicity of Dimilin was a direct effect of the chemical and not due to host
morbidity. These results indicate that Dimilin may be used in an integrated
approach to gypsy moth control. To test this, orchard sprays of Dimilin were
made while gypsy moth larvae were at: 1) the lst-2nd larval stadium, 2) 2nd-
3rd stadium and 3) 3rd-4th stadium. No parasitoids emerged from larvae
collected from treated trees 1 week after the first spraying. However, there
was little effect on numbers of emerged parasitoid cocoons when larvae were
collected from trees sprayed thereafter. Dimilin had no marked effect on adult
emergence from cocoons. Thus if timed properly Dimilin should have little
effect on populations of A. melanoscelus and yet still control the gypsy moth.

Effects of the Insect Growth Regulator Altozar on the Parasitoid,
Microctonus aethiops , and Its Host, Hyper a postica

M. E. Ascerno

Pesticide Research Laboratory, Dept. Entomology, Penn State University,
University Park, Pa. 16802

The insect growth regulator Altozar® terminated sexual diapause of adult
alfalfa weevils when topically applied. In addition, it terminated the diapause
of M. aethiops when applied to parasitized adult weevils. Parasitoid survival
and morphological condition were influenced by various Altozar® concentrations.
An interaction was also found between age of non-diapausing parasitoids at the
time of treatment and concentrations of Altozar® employed.

Mass Rearing of Diglyphus isaea (Walker) (Hymenoptera:
Eulophidae) on Liriomyza trifoliearum Spencer
( Diptera : Agromyzidae )

R. M. Hendrickson, Jr.

U.S. Department of Agriculture, ARS, 501 South Chapel St.,

Newark, Delaware 19713

The object of the study was to find a method of rearing large numbers of
the parasite, Diglyphus isaea (Walker), for release against its target host,
alfalfa blotch leafminer, Agromyza jrontella (Rondani). Mass rearing of D.
isaea on alfalfa blotch leafminer proved difficult because of the time and in-
convenience of handling the puparia of the fly which pupates in the soil, the
small numbers of larvae (50-100) available for parasite oviposition per 6"
pot of alfalfa, several troublesome pest contaminants of alfalfa, and the
lengthy time required to grow alfalfa from seed or allow its regrowth after

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cutting. Liriomyza trifoliearum Spencer was found to be a host acceptable to
the parasite. It was reared on Burpee™ Bountiful bush beans, a host plant
on which no contaminant species have as yet been found in culture. The first
pair of true leaves of the bush bean are suitable for oviposition by the fly
6-7 days after planting, at 78 °F. A 5" X 8" pot of 20 plants will produce
1000-1500 larvae. At 78°F the host larvae eclose and reach the final (3rd)
instar in 6 days, at which time they are placed in cages with parasites. It is
important to provide the parasite with 3rd instar host material only. They
sting and kill earlier instars, but no oviposition has been observed. Maturation
of the parasite requires 13-23 days at 73 °F. Thus in about a month it is
possible to go from host plant seed to mature parasite. The use of L. trifoliea-
rum as an alternate host on bush beans (as opposed to using alfalfa blotch
leafminer on alfalfa) saves time and space and results in far larger numbers
of D. isaea .

Control of the Apple Leaf Curling Midge, Dasyneura mali (Kieff)
(Diptera: Cecidomyiidae) in New Hampshire

G. T. Fisher and J. Turmel

Entomology Department, University of New Hampshire, Durham,

New Hampshire 03824

A block of apples in Durham, New Hampshire, heavily infested with the
apple leaf curling midge, was treated with several regularly recommended and
experimental materials. Treatments were made on the following dates: 5/1/75
(Green Tip); 5/8/75 (Tight Cluster) ; 5/15/75 (Pink); 5/26/75 (Petal Fall) ;
6/5/75 (1st Cover); 6/11/75 (2nd Cover), before efficacy data was recorded
(6/23/75). Treatments were randomized, replicated two or three times, and
applied in 350 gal. H20/A at 300 psi. Materials applied were: Zolone 3EC 1
pt/100; Furadan 4F 0.25 lb.ai/100; Parathion 2S 0.5 lb.ai/100; N2596 4EC
1.0 lb.ai/100; Imidan IS 0.5 lb.ai/100; Lannate 0.5 lb.ai/100; Lannate 0.25
lb.ai/100 plus Guthion 50WP 0.25 lb.ai/100; Guthion 50WP 0.5 lb.ai/100;
Mobil 9087 2EC 0.75 lb.ai/100; TH6042EC 0.5 oz.ai/100; TH6042EC 0.132
oz.ai/100; Bayhox 1901 40WP 4oz.ai/100; FMC 3329 3EC 0.1 lb.ai/100;
Bayntn 9306 6EC 4oz.ai/100. Data was obtained by counting the number of
curled leaves per tree. These counts were totaled, replicate averages made and
percent control corrected with Abbotts formula. The commonly used insecti-
cides, Guthion, Zolone and Imidan gave 100% control which probably accounts
for the minor occurrence of this pest in New England commercial orchards.
Parathion 2S and Lannate gave poor control. Treatments with Mobil 9087,
TH6082, Bayhox 1901, FMC 3329, Bayntn 9306 resulted in 100% control,
N2596, 99% control, and TH6082 at 0.132 gave 92% control.

Vol. LXXXIII, December, 1975

245

Is a Black Fly Survey Worthwhile?

Ivan N. McDaniel

University of Maine, Orono, Maine 04473

The town of Jackman, Maine, was surveyed to determine the sources of
black flies that cause severe annoyance from May to July. Prior attempts at
larval control requiring treatment of all swift-water areas had met resistance
from environmentalists since they felt the ecology of the area might be dis-
turbed. It was suggested that a survey might reveal localized areas of high
productivity in certain streams. Therefore, all streams within a 259 m2 area
surrounding the town were monitored during 1970 to determine productivity
and species present. Numerous biting collections supplemented larval and pupal
collections. Results of the survey showed that Simulium venustum Say ac-
counted for more than 98% of the annoyance in the area and an estimated
85% of this species was found to develop in two separate stretches of Wood
Stream. Heald Stream and certain areas of several other streams were pro-
ductive although they are considered rather minor sources of S. venustum.
Some streams did not produce this species and others produced very few.
Results of the study suggest that a black fly survey can be as useful as a
mosquito survey in locating breeding sites for source reduction. The cost of
a survey should be considered minor as compared with the savings realized if
a larval control program should be implemented. Since the amount of insecti-
cide required for control would be markedly reduced, the risks to non-target
organisms also would be minimized.

A Comparison of Malaise Trapping and Aerial Netting for
Research on Houseflies and Deerflies (Diptera: Tabanidae)

Douglass W. Tallamy and Elton J. Hansens
Department of Entomology, Rutgers University, New Brunswick,

New Jersey 08903

Fifty species of Tabanidae were collected near Deer Lake, Booton, N.J.
during the summer of 1974 while comparing the effectiveness of malaise trap-
ping and aerial netting for sampling tabanids. Five sites in the study area
were sampled by malaise traps and by sweeping (netting) about the head with
an insect net. Comparisons were made of 4 community and 4 population
parameters. Differences between the species richness of the malaise-trapped
community, which collected 44 species with a mean of 22.2 species/ site, and
that of the netted community, which collected 44 species with a mean of 27.2

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species/site, were not significant (P0.5,«8). Analysis of trapping data for
species diversity, 2.34 for malaise vs. 1.30 for netting, and the species evenness,
.08 and .40 for malaise and netting respectively, showed both were significantly
different (P.01, «8). The community similarity index, .08, indicated that the
structure of the tabanid community trapped by each method differed sub-
stantially. The relative densities of the species trapped by each method were
significantly different for most Chrysops and the dominant Tab anus and
Hybomitra , although a site X method interaction existed with H. losiophthal-
mus , H. sodalis, T. lineola, and T. pumilus, which statistically masked this
significance. Though the seasonal ranges found by each trapping method did
not closely coincide, the seasonal niche breadths were not significant except
for 4 species: H. sodalis and C. macquarti (P.01,<*8), C. geminatus and T.
lineola (P.05,«8). Neither trapping method should be used alone in studies
concerning the entire tabanid community but can be used effectively together.

Infecting the Gypsy Moth, Porthetria dispar (L. ) (Lepidoptera:
Lymantriidae) with Nuclear Polyliedrosis Virus Vectored by Apanteles
melanoscelus (Ratzeburg) (Hymenoptera: Braconidae)

Bernard Raimo

U.S. Forest Service, NEFES, 151 Sanford Street, Hamden, Connecticut 06514

This study was undertaken to determine the ability of Apanteles melanos-
celus (Ratzeburg) contaminated with nuclear polyhedrosis virus (NPV), to
infect gypsy moth larvae and to determine the feasibility of using this method
of virus dissemination as an alternative to topical foliar application. Three
methods of contaminating the parasitoids with virus were tested. The first
method involved exposing female parasitoids to first, second, and third-stage
gypsy moth larvae that had been feeding on artificial diet containing 1 X 107
polyhedral inclusion bodies (PIB)/ml for a period of 48 hr. In the second
method virus was applied directly to the ovipositor at a concentration of
1 X 109 PIB/ml. A topical application of virus at a concentration of 1 X 10°
PIB/ml by means of an atomizer was the final method of contamination of
parasitoids that was tested. Percent mortality due to virus was found to be
higher among larvae exposed to contaminated female parasitoids than larvae
that were exposed to uncontaminated female parasitoids and the transmission
of virus does not appear to diminish with each successive sting. The most
promising method of contaminating the parasitoids with NPV was found to
be manual application of the virus to the ovipositor, however, none of the
contamination techniques seemed to have any effect on the parasitoids.

Vol. LXXXIII, December, 1975

247

Cassida rubiginosa Muller (Coleoptera: Chrysomelidae), a
Potential Biocontrol Agent of Thistles in Virginia

R. H. Ward and R. L. Pienkowski

Department of Entomology, Virginia Polytechnic Institute and State University,

Blacksburg, Virginia 24061

C. rubiginosa , accidentally introduced into North America and first reported
in 1901, presently has a geographical range encompassing most of eastern
North America north of central Virginia. During 1973 and 1974, life stages
of the beetle were collected from 15 sites in the northern Virginia counties of
Frederick, Clarke, and Warren. The purpose of this study was to isolate this
beetle from a major obligate gregarious exotic larval and pupal endoparasitoid
Tetrastichus rhosaces (Walker) (Hymenoptera: Eulophidae). Immature cassid
life stages were reared to the adult stage and released 100 miles south of its
southernmost range in the southwest counties of Montgomery and Giles. A
total of 7840 adults were released at 9 sites with 3 replicates each for the
three thistle species Carduus nutans (L.) — musk, C. acanthoides L. — plumeless,
and Cirsium arvense (L.) Scop. — Canada thistle. In 1975, cassids were recov-
ered from all release sites with Canada thistle sites having the highest rate of
recovery followed by musk and plumeless thistles. Cassids dispersed less than
30m in any one direction from the release point at most sites. Periodic ob-
servations of cassid life stages at one musk and Canada thistle sites revealed
only one possible case of parasitism, probably by the solitary facultative native
parasitoid Eucelatoriopsis dimmocki (Aldrich) (Diptera: Tachinidae). Preda-
tion by spiders, assassin and stink bugs, chrysopids, and other arthropods was
the major biotic mortality factor.

Growth Form of Host Plant as a Determinant of Feeding
Efficiencies and Growth Rates in Papilionidae and
Saturniidae (Lepidoptera)

J. Mark Scriber and Paul P. Feeny
Department of Entomology, Cornell University, Ithaca, N. Y. 14850

Larval growth performance of butterflies and moths was studied on leaves
of their normal host plants to test whether specialized, stenophagous herbivores
utilize their food resources more efficiently than do generalists. Graphium
marcellus, Battus polydamas, B. philenor , Papilio zelicaon, and P. polyxenes
are specialized primarily on one plant family. P. palamedes, P. troilus, and
P. multicaudatus are intermediate (2 to 5 families), and P. glaucus is very
generalized, feeding on at least 13 families. For both the penultimate and

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final instars the efficiency of conversion of ingested food into larval biomass
of the specialists was significantly higher than that of the intermediate and
generalized species. Growth rates for specialized papilionids were more than
twice as great as those for generalists. Among the swallowtail species the trend
from specialized to generalized feeding paralleled a trend from herb-feeding
to tree-feeding. No significant differences were found in feeding efficiencies
of the final two instars of tree-feeding moths A. pernyi , S. cynthia , C. angu-
lifera, and B. mori (specialized), C. promethea, A. luna, and C. regalis (inter-
mediate), and E. imperialism H. cecropia, A. io, and A. polyphemus (generalized).
Growth rates were also nearly identical for specialists and generalists. Regard-
less of their degree of feeding specialization, tree-leaf feeding larvae of both
butterflies and moths grew at lower rates than herb-feeders. This indicates
that growth form of the host plant has at least as important effect on the
overall ecology and life history of a papilionid or saturnid species as has the
degree of feeding specialization of its larva.

Reproductive Diapause in Notonecta undulata (Say)

( Hemiptera : Notonectidae )

Robert L. Vanderlin and Frederick A. Streams
Biological Sciences Group, University of Connecticut, Storrs, Conn. 06268

Notonecta undulata (Say) completes two generations annually in Connecticut.
The first generation, produced by overwintered adults, reaches maturity in early
July. The second generation of adults develops in late summer and early fall
and enters reproductive diapause. The reproductive activity is regulated by
photoperiod and temperature. Photoperiod seems to be more important than
temperature. Over 75% of the females raised in the laboratory under a 15-hr
photoperiod (LD) and 22 °C oviposited within 21 days after adult emergence,
while less than 25% of the females raised under a 12 -hr photoperiod (SD) and
22 °C oviposited. When half of the latter group were subsequently placed under
LD conditions, 75% of the females commenced oviposition within 21 days.
The remaining half of the SD group was continued under SD conditions and
less than 25% of these commenced ovipositing during the next 30 days. Females
reared under SD and LD conditions were vivisected and terminal oocytes
measured. Females reared under both conditions had terminal oocytes measur-
ing less than 0.4 mm upon emergence as adults. Within 3 weeks LD females
attained an oocyte length of 1.7 mm while females reared under SD conditions
rarely exceeded .9 mm. Oosorption was observed in the few SD females with
terminal oocytes as long as 1.7 mm. Low temperatures tend to inhibit repro-
ductive development. When N. undulata was reared under LD conditions but

Vol. LXXXIII, December, 1975

249

at 15°C, only 50% of the females oviposited within 7 weeks of adult emergence.
Photoperiods under which nymphs are reared appear to have no effect on
subsequent reproductive development of adults.

Environmental Control of Diapause in Three Species of North
American Aecline Mosquitoes (Diptera: Culicidae)

B. F. Eldridge, R. R. Pinger, Jr., J. F. Burger and D. E. Hayes
Department of Entomology, Walter Reed Army Institute of Research,
Washington, D.C. 20012

Environmental factors influencing diapause were studied under field and
laboratory conditions for three species of North American aedine mosquitoes:
Aedes atlanticus (Dyar and Knab), A. canadensis (Theobald) and Psorophora
ferox (Humboldt). To detect naturally occurring diapause, soil samples were
collected in forest depressions in eastern Maryland at five different times of
the year: March, May, September, November and December. Samples were
divided into two equal parts — one part was held under long photoperiod con-
ditions, the other under short photoperiod conditions, both at 25°C. Portions
of each sub -sample were flooded weekly for four weeks. The proportion of
viable eggs which hatched upon flooding varied with the species, the time of
year collected, and the length of time held under experimental conditions. The
results indicate that all three species undergo embryonic diapause which is
terminated only after exposure to chilling temperatures. Photoperiod does not
appear to play a role in diapause termination. Prolonged exposure to moderate
temperatures appears, in addition, to be a necessary antecedent to hatching in
A. atlanticus and A. ferox. Experiments were conducted in the laboratory to
determine the influence of photoperiod on induction of diapause. Results indi-
cated that short photoperiod exposure of the developing embryo of A. cana-
densis induces diapause, whereas similar exposure of P. ferox females results
in the oviposition of diapausing eggs.

An Improved Insect Pest Management Program on Sweet Corn

in New Hampshire

James S. Bowman

Department of Entomology, University of New Hampshire, Durham,

New Hampshire 03824

Three years of field research studies on the detection and control of the
European corn borer Ostrinia nubilalis (Hiibner) and the corn ear worm Heli-
othis zea (Boddie) on sweet corn demonstrated the feasibility of an improved

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insect pest management program for New Hampshire. First brood European
corn borer can be controlled using whorl-stage treatments of either a granular
insecticide applied once or liquid sprays applied twice. Since the arrival time
of the corn earworm’s northern migration can vary from year to year (August
1 to September 1), early detection with light traps for moth flights and in-
spection of silks for newly laid eggs is used to alert growers to the necessity
for spraying. In years when the earworm arrives late, considerable reduction
in the use of insecticides and a resulting savings to the grower is realized.
Small plot research comparing directed granular applications at planting time
(soil applications) and at what stage with spray treatments indicates Dyfonate
10GK, carbofuran 10G, N-2596 lOGk, and Sandose 197 give acceptable eco-
nomic control of the European corn borer. A comparison of granular treatments
applied either directly into the whorl or as a broadcast indicates a slight loss
in effectiveness as a broadcast but acceptable economic control.

Sequential Releases of Rhinocyllus conicus Froelicli (Coleoptera:
Curculionidae) for the Biocontrol of Car duns Thistles

W. W. SURLES AND L. T. KOK

Department of Entomology, Virginia Polytechnic Institute and State University,

Blacksburg, VA 24061

Rhinocyllus conicus , an introduced thistle-head feeding weevil, which has
been established on Carduus thistles in Virginia has shown more effective
control of Carduus nutans (musk thistle) than of Carduus acanthoides (plume-
less thistle). This is apparently due to better synchronization of overwintering
weevil emergence in spring with bud development of musk thistles than with
plumeless thistles. Sequential field releases of overwintered weevils on indi-
vidually-caged thistles were conducted to investigate the advantages of extend-
ing the ovipositional period and improving synchronization of the weevil with
thistle development. Each of the releases produced an initial surge in ovi-
position of progressively decreasing magnitude on both Carduus thistles. The
later releases successfully extended the ovipositional period, but were less
effective due to reduced fecundity. Weevils also suffered increased mortality
during prolonged caging prior to release. Regular monitoring of egg deposition
on the developing thistle blooms revealed that R. conicus preferred the earlier
stages of musk thistle capitula. This was not evident on the plumeless thistles
which had eggs equally distributed on the buds as well as the fully developed
heads. Preference for earlier bloom stages resulted in higher survival of larvae
on musk thistles than on plumeless thistles. Eggs deposited on the later stages
of plumeless thistle blooms failed to complete development, and had high
larval mortality because of the inadequate period for development.

Vol. LXXXIII, December, 1975

251

Survival of Aestivating Adult Rhinocyllus conicus Froelich
(Coleoptera: Curculionidae) at Different Temperatures
and Photophases

L. T. Kok

Department of Entomology, Virginia Polytechnic Institute and State University,

Blacksburg, VA 24061

Successful field establishment of Rhinocyllus conicus, an introduced weevil
for the biological control of Carduus thistles, is best obtained by spring releases
at the initial stages of thistle bud development. To insure an adequate supply
of ovipositing weevils at the time of release, senescent infested thistle heads
collected from an established site in Virginia were held for weevil emergence.
The newly emergent, aestivating weevils were caged with thistle leaves, and/or
thistle heads, or artificial diet. These were exposed to different thermoperiods
ranging from 10°-32°C with either declining or constant photophases of 0-16 h
to determine conditions optimum for their survival until the following spring.
Day-night thermoperiods of 21°-15°C synchronized with short photophases
resulted in relatively lower mortality than treatments of 26°-15°C, and
32°-15°C with similar photophases. Aestivating weevils survived better when
caged with thistle leaves at constant temperatures of 10 or 15°C, and con-
tinuous darkness. Best survival was obtained for weevils placed on artificial
diet subjected to photophase of 0 or 10 h at 10°C. Mortality rate increased
at higher temperatures and longer photophases. In the absence of thistle leaves
or artificial diet, no weevils survived the entire duration of the experiment
(August through April of the following year) ; those weevils subjected to the
higher temperatures of 26°C or 32°C did not survive beyond December.

Increased Gladiolus Spike Growth with Use of Certain
Systemic Insecticides

Roger G. Adams, Jr., John H. Lilly, and Adrian G. Gentile
Department of Entomology, University of Massachusetts,

Amherst, Massachusetts 01002

Previous aphid control experiments suggested that treatments with certain
systemic insecticides might improve gladiolus growth. A replicated field ex-
periment was conducted near Suf field, Connecticut in 1972 to test this hypoth-
esis. The experimental area was divided into 60 5-ft.-long plots, each of which
was planted with 10 corms of the gladiolus cultivar “Peter Pears”. The fol-
lowing 10 treatments were replicated 6 times in a randomized complete block
design: dimethoate 2 EC, oxydemeton-methyl 2EC, carbofuran 4F, carbofuran

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10G, pirimicarb 50WP, acephate 75S, disulfoton 15G, aldicarb 10G, oxamyl
10G, and untreated control. All materials were applied at 1 lb ai/acre. Two
granular soil treatments and 3 foliar spray applications were made with the
respective materials. No significant differences were detected in plant emer-
gence or height 17 and 23 days, respectively, after planting. At peak bloom
granular soil treatments of aldicarb, oxamyl, and disulfoton showed significant
effects on gladiolus growth as indicated by increased plant heights, flowerhead
lengths, and bud numbers. Cucumber mosaic virus disease-infected plants in
both aldicarb and disulfoton-treated plots showed growth benefits. We conclude
it may be possible for gladiolus growers to obtain both increased plant growth
and flower production as well as insect control through utilization of aldicarb,
disulfoton, or oxamyl as soil insecticides.

Evaluation and Control of a Nuisance Fly Problem (Diptera:
Muscidae) at Monmouth Park Jockey Club,

Oceanport, New Jersey

John Milio and Elton J. Hansens

Department of Entomology and Economic Zoology, Rutgers University,

New Brunswick, New Jersey 08903

Throughout the 1973 summer racing schedule, Monmouth Park personnel
experienced an acute fly problem. In 1974, inspections of 10 randomly chosen
horse barns made 3 times per week from July to October, revealed a predomi-
nance of house flies, Musca domestica (L.), and stable flies, Stomoxys calcitrans
(L.), in addition to small numbers of Tabanidae. Potential breeding sites of
the predominant species were manure pits, hay barns, horse corrals, horse stalls,
grass clippings and beached tidal creek vegetation. Larvae or pupae occurred
at all but the latter two sites. Of 92 samples, manure pits and hay barns
yielded the highest proportion of house flies and stable flies per sample.
Layers of moist, decayed hay or alfalfa in front of the hay barns were par-
ticularly productive of stable flies. Manure pits containing STAZ-DRI (horse
bedding derived from sugar cane refuse), horse manure, or a combination of
both, produced large numbers of house flies. Evaluations of ULV applications
with a Cardinal 150 sprayer (Northeastern Associates), using formulations of
1% pyrethrins (Dubois Chemical Co.), 5% pyrethrins (Northeastern Associates),
and 40% resmethrin (S.B. Penick & Co.) diluted 18 fl. oz./gal. with light
mineral oil, showed the latter superior, though variable winds to 5 mph, cool
morning temperatures, and lack of penetration into barns limited control.
Proper disposal of manure and hay barn refuse remains the most practical and
effective means of fly control.

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253

Control of External Parasites on Cattle by
Means of Dust Bags

James E. Roberts, Sr.

Department op Entomology, Virginia Polytechnic Institute and State University,

Blacksburg, Virginia 24061

Dust bags have been used for approximately 15 years for the control of
external parasites on cattle. When properly installed and maintained, use of
this type of self-treatment device will result in near-complete control of horn
flies, Haematobia irritans (L.), and cattle lice and will effectively reduce face
fly Musca autumnalis (DeGeer) populations. Two recent tests with 5% fenthion
in dust bags resulted in 41% and 60% reduction in cattle grub infestation.
Numerous field tests have shown that forced use of dust bags will result in
more effective insect control. A shelter should be provided for any dust bags
that are not water repellent. However, some good water repellent bags are
now available from commercial sources. Protection from rainfall is necessary
to prevent caking of the dust. To obtain the most effective face fly control,
bags must be suspended within 18 to 24 inches from the ground. The efficacy
of insect control is also enhanced when the bag swings freely so that as the
animals pass under them they will bump the bags with their heads and again
with their shoulders, thereby insuring a more thorough dust coverage of the
head and body.

Wing Polymorphism in Salt Marsh Inhabiting Fulgoroitlea

Robert F. Denno

Department of Entomology and Economic Zoology, Rutgers University,

New Brunswick, N.J. 08903

The vegetation of New Jersey tidal salt marshes is composed primarily of
two grasses, Spartina alterni flora Lois. (Smooth Cordgrass) and Spartina patens
(Ait.) Muhl. (Salt Meadow Cordgrass). S. patens occupies a narrow eleva-
tional zone of well drained marsh above mean high water level (MHW). S.
alterniflora, an intertidal species, occurs throughout most of the elevational
range of S. patens , but also extends to levels far below MHW. Near MHW,
where the marsh is flat and poorly drained, S. alterniflora occurs as a dwarfed
form, but along depressed borders of creeks well below MHW, it may grow
to the height of 2m. Structurally (culm height, width, density), S. alterniflora
is more diverse than S. patens and ecologically occupies a more extensive ele-
vational range which receives frequent tidal inundations. Fulgoroids, dimorphic
for wing length, feed upon these salt marsh grasses. The purpose of this work

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was to investigate the ecological significance of wing polymorphism strategies
employed by the fulgorids to cope with the structural variability and stability
of food resources. The delphacid, Prokelisia marginata (Van Duzee), which is
host specific on S. alterniflora, produced highly vagile macropters and brachyp-
ters with less efficient flight capability. In short form S. alterniflora, macrop-
ters and brachypters were sweep-netted in equal numbers. The following
commonalities were evident for 5. patens inhabiting fulgoroids [Delphacodes
detecta (Van Duzee), Tumidagena minuta McDermott, Aphelonema simplex
Uhler] : Brachyptery was extreme, preventing flight. Brachypters significantly
outnumbered macropters. The density of brachypters correlated well with the
seasonal pattern of standing crop biomass of the food resource. Macropter
density was poorly correlated with standing crop biomass; however, seasonal
macropter density occurred just prior to peak standing crop biomass of the
grass. On the high marsh, which is less subject to tidal inundation and is
occupied by a structurally monotonous food resource, fulgoroids have evolved
a brachypterous strategy. On the intertidal marsh, where the food resource is
structurally diverse and occasionally unavailable (tidal inundation), population
mobility is at a premium and fulgoroids have evolved a strategy relatively
skewed toward the production of macropters in order to efficiently exploit
resources.

Isolation and Identification of Entomophthora spp. Fres.

( Phycomycetes : Entomophthorales ) from the Spruce Budworm
Choristoneura fumiferana Clem. (Lepicloptera: Tortricidae)

John D. Vandenberg

Department of Entomology, University of Maine, Orono, Maine 04473

Richard S. Soper

N.E. Plant, Soil and Water Laboratory, USDA-ARS, Orono, Maine 04473

Entomophthora sphaerosperma Fres. and E. egressa MacLeod and Tyrrell
were isolated from field-collected spruce budworm in Aroostook and Washington
counties, Maine. An 18" branch tip was taken from the upper and lower
crowns of 5 trees at each of 13 locations. Living larvae were reared on arti-
ficial diet to determine disease prevalence. Fungi from diseased larvae were
isolated on egg yolk plus Sabouraud maltose agar medium and then identified.
Both species were tested for growth on several media and over a range of pH
and temperature levels. Prevalence of infection was greater in the lower crown
(7.4% vs. 2.6%) with an overall prevalence of 5.0% (90% level of significance).
The conidia of E. sphaerosperma are slender, papillate at the base, rounded and
tapered at the apex; 19.50 ± 2.14 /x X 7.64 ± 1.06 /x (Mean ± Standard

Vol. LXXXIII, December, 1975

255

deviation) from larvae, and 22.43 ± 4.90 /x X 7.24 ± 0.46 /a from specimens
grown on artificial media. Conidiophores are branched and digitate. Rhizoids,
secondary conidia and cystidia are present. Resting spores are hyaline, spherical
and measure 24.90 /a ± 2.18 /a from larvae, and 26.30 /a ± 2.71 /x from media-
reared specimens. The conidia of E. egressa are obovate to pyriform with a
broad papillate base and an evenly rounded apex. They measure 38.56 ± 8.33 /a
X 31.59 ± 7.26 /a, and are multinucleate (8-10). Conidiophores are branched.
Resting spores, secondary conidia, rhizoids and cystidia were not observed.
Mycophil yeast agar (MYA) at 20°C was found to produce the best growth
of E. sphaerosperma. MYA at 20°C was then tested at a pH range of 5.0-7. 5
at intervals of .5. Mycelial growth was predominant above pH 6.5, while
abundant conidia production occurred below pH 6.0. Growth of E. sphaeros-
perma was also tested in shaker cultures on liquid media with Sabouraud liquid
broth (SLB) within the range 5. 5-8.0 pH and 25°C giving the best growth.
Optimum growing conditions for E. egressa as determined in a like manner,
were at 25°C on AK agar throughout a pH range 6.0 to 7.5.

A Sex Pheromone Complex of the Mushroom-Infesting
Sciarid Fly, Lycoriella mali Fitch

J. G. Kostelc, L. B. Hendry and R. J. Snetsinger
Departments of Chemistry and Entomology, Pennsylvania State University,
University Park, Pa. 16802

The sex pheromone complex of the sciarid fly, Lycoriella mali Fitch consists
of a homolog series of saturated, straight chain, aliphatic hydrocarbons. Sciarid
males are attracted to hydrocarbons pentadecane to hexacosane and octacosane
(Ci5-C26,C28) when tested in a biological assay chamber. Heptadecane is the
most active hydrocarbon. In addition, statistical testing of heptadecane and
neighboring hydrocarbons (Ci5-Ci9) support this finding. Concentration studies
of heptadecane show that the best overall response (80%) and the best at-
tractive response (79%) occur at the 1.0 nanogram level. Abdomens of culture
males and females were analyzed for the presence of hydrocarbons pentadecane
to octadecane. Isolation and identification of hydrocarbons was accomplished
by thin layer chromatography, gas chromatography and computerized gas
chromatography-mass spectrometry with mass fragmentography. Hydrocarbons
(Cib-Cib) were found in both male and female abdomens. Female abdomens
had greater concentrations of these hydrocarbons than male abdomens. The
major attractant, heptadecane, was found in the female sciarids at a concen-
tration of 5-6 times larger than found in the male sciarids. Recent results
from our laboratory have shown that some lepidopteran and hymenopteran
pheromones are present in the host plant. Therefore, the cultivated mushroom,

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Agaricus bisporus Lange was analyzed for hydrocarbons. Hydrocarbons pen-
tadecane to heneitriacontane were identified and their relative concentrations
determined. It has been shown that the sex pheromone of the sciarid fly is
not a single component but a complex mixture. Host plants are implicated as
a possible origin of the pheromones.

Results of an Insect Scouting Program in Virginia Soybeans

William A. Allen

Department of Entomology, Virginia Polytechnic Institute and State University,

Blacksburg, Virginia 26061

After 2 years of monitoring insect control decision-making practices in eastern
Virginia, 2 test demonstrations were undertaken to encourage adoption of a
pest management system. Unlike the 2 previous studies, a vigorous educational
program was conducted with farmers. Mean per acre scouting costs were higher
in both counties (Isle of Wight $0.52; Westmoreland $0.59) than in previous
years, reflecting higher labor costs and a difference in scouting efficiency.
Due to a change in objectives, it was not possible to measure the potential
saving or net return on investment as in past years. Premature insecticide
application was reduced from 87.2% in 1972 and 89.7% in 1973 to 0% in 1974.
In addition to the demonstrations mentioned above, the first grower-financed
scouting service in Virginia was adopted in Northumberland and Lancaster
counties in 1974. The program included 11 farmers and 43 fields. A sub-
jective evaluation conducted at the conclusion of the program showed the
following results. Most farmers (72.9%) estimated that they saved $4.98/ acre
using the scouting system. Only 8.3% of the farmers thought yields were
reduced; 85% of the farmers said they would subscribe to a scouting service
on a “pay as you go” basis in 1975 if it were available but only 62.5% said
they would use the system if they had to do the scouting themselves. The
overall response indicated grower approval.

Leaf Feeding Resistance to the European Corn Borer, Ostrinia

nubilalis (Hiibner) (Lepidoptera: Pyralidae), in Tropical Maize

W. M. Tingey, V. E. Gracen, and J. M. Scriber
Departments of Entomology and Plant Breeding, Cornell University,

Ithaca, New York 14853

First generation or leaf-feeding resistance of maize to the European corn
borer has generally been attributed to 2,4-dihydroxy-7-methoxy (2H)-benzox-
azin-3 (4H)-one (DIMBOA), an aglycone that suppresses larval development
and increases larval mortality. Tropical maize genotypes as low in DIMBOA

Vol. LXXXIII, December, 1975

257

as the susceptible inbred WF9, have been shown to be as resistant to leaf
feeding in field trials, as high-DIMBOA inbreds. We studied leaf feeding
activity of 1st instar O. nubilalis on 3 inbred (WF9, B68, B49) and 2 tropical
(6006: San Juan-3 X Antigua-8D; 6008: Puerto Rico-1 X Antigua-2) geno-
types using free-choice and no-choice laboratory caging methods, to determine
the resistance mechanisms. Using no-choice Plexiglas® tube cages or dialysis
tube cages on rolled leaves, feeding rates at 12 hr intervals over a 60 hr post-
caging observation period, were consistently reduced on the high DIMBOA
inbred, B49, and on the tropical genotype, 6006, as compared to a susceptible
inbred, WF9. Feeding was intermediately reduced on 6008 and B68, compared
to WF9. When given a choice between 2 genotypes, larvae consistently pre-
ferred WF9 over the other 4 genotypes. Thus, we concluded that feeding
suppression of O. nubilalis during the 1st stadium is probably a significant
component of resistance in field-resistant tropical and inbred genotypes. More-
over, suppression of larval leaf feeding by the tropical lines, 6006 and 6008,
is mediated by plant factors other than DIMBOA.

Visual Stimuli in the Host Finding Mechanism of the Parasitic Wasp
Itoplectis conquisitor (Say) (Hymenoptera: Ichneumonidae)

David Robacker, K. M. Weaver and L. B. Hendry

Department of Chemistry, Pennsylvania State University, University Park, Pa.

16802

An investigation into the kinds of visual cues utilized by /. conquisitor during
host finding was conducted by measuring attraction, discrimination (antennal
tapping) and acceptance (probing) responses to various host and host-shelter
models. To demonstrate the presence of a visual attractive stimulus, wasps
were tested with sealed glass cylinders each containing a dead host pupa near
one end. The other end, which remained empty, served as a control. The
number of attractions to the pupa containing side was found to be significantly
greater than to the empty end of the same tubes. No discriminatory behavior
was elicited by these glass models. A further elucidation of visual stimuli was
accomplished using construction paper and cellophane models which differed
in shape and degree of transparency. All opaque models were found to be
superior to their transparent counterparts in visual attracting capability. More-
over, all cylindrical models were superior to their flat counterparts in this
property. With regard to discriminatory and acceptance responses, however,
only cylindrical shapes appeared to be of any significant importance. The
possibility that a host finding mechanism other than of a chemical nature is
in operation was confirmed by the following experiment. Newly emerged fe-
males with no ovipositional experience were individually isolated for 7 days

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in a chemically clean environment. After this isolation period, the wasps were
released into glass beakers containing aluminum foil cylinders and their re-
sponses were recorded. Typically, the wasps showed aggressive discriminatory
and acceptance behavior in less than 1 hour of testing.

Dispersal of First-Instar Gypsy Moth Larvae in Relation to
Population Quality

John L. Capinera and Pedro Barbosa

Department of Entomology, University of Massachusetts, Amherst, Mass. 01002

Investigations by D. E. Leonard indicated that the behavioral and physio-
logical ecology of the gypsy moth, Porthetria dispar (L.), varied with the size
of the eggs from which the larvae originated. Also, the ecological patterns
could be shifted by certain environmental stimuli. This principle was termed
population quality. Leonard suggested that dispersal was the most important
parameter affected. Our investigation examined the effect of population
quality on dispersal by first-instar larvae. Eggs deposited by the female gypsy
moth vary in size. The first eggs deposited are largest in diameter and sub-
sequent eggs are smaller. Laboratory dispersal studies indicate that larger eggs
produce larvae that disperse more frequently than larvae from smaller eggs.
However, small larvae unable to locate adequate food will disperse as fre-
quently as large larvae. Dispersal ability of larvae declines rapidly as larvae
feed and become heavier. Except at low wind velocities (less than .9 m/s),
large larvae disperse greater distances than small larvae. Egg size is inversely
proportional to the number of eggs^ per mass. Factors favoring development
of large adults will induce large egg masses and lead to populations of non-
dispersing larvae. Failure of large numbers of larvae to disperse from a site
can lead to localized population outbreaks. In turn, high density populations
produce small egg masses which produce relatively more dispersing larvae,
thereby providing the inoculum for new outlying infestations.

Identification of the Copulatory Sex Pheromone of the Little
House Fly, Fannia canicularis (L.) (Diptera: Muscidae)

E. C. Uebel, R. E. Menzer, P. E. Sonnet, and R. W. Miller
Department of Entomology, University of Maryland, College Park, Maryland 20742
and USDA, ARS, AEQI, Beltsville, Maryland 20705

A sex pheromone that stimulates F. canicularis males to copulate with
females was identified as (Z)-9-pentacosene. Cuticular lipids obtained from
virgin 5-day-old female flies were used for the isolation of the active compound.

Vol. LXXXIII, December, 1975

259

Materials were bioassayed by counting the number of copulatory attempts that
occurred during a 5-minute period when pseudoflies constructed from knots of
black yarn were treated with 100 or 200 /xg of the test material and presented
to unmated males. Isolation, identification, and synthesis procedures were
similar to those reported by Uebel et al. in 1975 (J. Chem. Ecol. 1: 195-202).
All major materials in the female cuticular lipid are hydrocarbons. The major
components of the unsaturated hydrocarbon are (Z)-9-pentacosene and (Z)-9-
heptacosene, which make up 66.5 and 3.4% of the cuticular lipid, respectively.
Approximately 25% of the cuticular lipid is straight chain saturated hydro-
carbon, and 4% is branched chain saturated hydrocarbon. Five-day-old males
have a “non-hydrocarbon” that makes up 27% of the total cuticular lipid.
Approximately 27% of the male cuticular lipid is straight chain paraffin, 20%
is branched saturated hydrocarbon, and 19% is unsaturated hydrocarbon. The
pentacosene present on the male constitutes 7% of the cuticular lipid and is
present as two isomers: (Z)-9-pentacosene (5%) and (Z)-7-pentacosene (95%).
Only fractions containing the unsaturated hydrocarbon from the female stimu-
lated the males to copulate, and tests with the two synthetic monoolefins found
on the female showed that only (Z)-9-pentacosene was active.

Honeylocust Pod Gall Midge, Dasyneura gleditschae Osten Sachen
(Diptera: Cecidomyiidae), Control with Dacamox®

W. R. Harrigan and J. L. Saunders

Department of Entomology, Cornell University, Ithaca, New York 14853

Increased planting of thornless honeylocust as street and shade trees and
nursery production of popular varieties in large blocks has intensified problems
caused by the honeylocust pod gall midge. Three varieties (Shademaster, Sun-
burst, and Skyline) of thornless honeylocust were side dressed with systemic
insecticides on May 1 & 2, 1975 at rates of ozAI per 1000 ft. of row as follows:
aldicarb 7.2 and 14.4, carbofuran- 4.4 and 8.8, disulfoton- 22.5 and 45, acephate-
6.0 and 12, and Dacamox® (3,3- Dimethyl- 1- methylthio- 2 butanone 0-
methylcarbamoyloxime)- 6.0 and 12. The soil, a sandy loam with good field
moisture, was cultivated just prior to treatment. It rained immediately after
treatment. The granules were applied in a 2-in.-wide band and covered with
ca. Vz in. of soil. Counts made June 17, 1975 of the number of galls on 10
shoots (each shoot had 5-10 leaves with ca. 20 leaflets per leaf) on each of 5
trees from the center of each plot indicated that only Dacamox showed promise
of giving economically practical protection. Dacamox, at the high rate, reduced
the number of leaf galls on the different varieties as follows: Shademaster

82%, Sunburst 50%, Skyline 71%. Percent defoliation estimated on July 23,

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1975 for Dacamox vs. check, respectively, was: Shademaster 40% vs. 53%,
Sunburst 17% vs. 52%, Skyline 32% vs. 54%. None of 13 toxicants applied
May 2, 1975 as hydraulic sprays gave economically effective protection.

A Seasonal History of the Variegate*! Leaf roller, Platynota flavedana
Clemens (Lepidoptera: Tortricidae), in Virginia Apple Orchards

J. H. Thomas and C. H. Hill

Virginia Polytechnic Institute and State University, Winchester Fruit Research
Laboratory, Winchester, Virginia 22601

The variegated leafroller overwinters as dormant larvae in leaf litter on the
apple orchard floor. In early March larvae feed on ground cover sprouts and
apple root suckers. Spring moths begin emerging in early May and are present
in orchards until early July. Egg masses are laid on upper sides of apple
leaves from May to July and again in late July to September. A fecundity
study of 21 second generation females indicates an average of 205.7 eggs per
female with 53.4 eggs per mass and 91.8% hatch. The incubation period
averages 12.8 days for first generation eggs and 9.3 days for the second.
Study of 154 individuals shows that most first generation larvae have 5 instars;
however, some have an occasional sixth. The average time for larval develop-
ment is 29.8 days. The pupal period averages 6.5 days. First generation moths
begin emerging the third week of July and are present in orchards until mid-
September. The life cycle averages 42.4 days for the males and 47.1 days
for the females. Fruit is damaged toward summer’s end and early fall by late
instar larvae of the first generation and younger larvae of the following gen-
eration. These second generation leaf rollers reduce their activity but continue
feeding through early November. When autumn leaves fall the larvae range
from second to fourth instar. In 1974 at Winchester, Virginia there occurred
the end of the overwintering generation, one complete generation, and the be-
ginning of a second generation of P. flavedana.

The Milkweed Pod as an Obstacle to the Large Milkweed Bug,
Oncopeltus fasciatus (Heteroptera: Lygaeidae)

Carol Pearson Ralph

Department of Entomology, Cornell University, Ithaca, N. Y. 14850

In the eastern United States Asclepias syriaca is a common host of the
monophagous Oncopeltus. In greenhouse experiments nymphs fed only vegeta-
tive shoots of A. syriaca could not grow or mature, but those provided A. syriaca

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261

seeds readily did. In the field adults and nymphs feed almost exclusively on
pods. However, measurements of nymph mouthparts, feeding punctures, and
the spongy pod walls showed that the mouthparts of the first three nymphal
instars are too short to reach seed in most mature pods. By exploring, these
young nymphs sometimes find sites with thin walls where they can reach the
seed. Usually they live on poorer food tapped from the green tissues and
placenta of the pod. Communal feeding may be especially important during
this stage in increasing food intake. Thus Oncopeltus can exploit this milk-
weed, even though the bug’s critical food, the seed, is inaccessible to the young
nymphs. Although adults and fifth instar nymphs can reach seed through
almost any pod wall, many seeds escape Oncopeltus damage because pods are
abundant and the bugs preferentially feed where the pod wall is thinnest, on
the seeds nearest the suture and the tip.

Growth and Development of Hyposoter exiguae (Viereck)
(Hymenoptera: Ichneumonidae) on Two Instars of Trichoplusia ni
(Hiibner) (Lepidoptera: Noctuidae)

Eugene A. Jowyk and Zane Smilowitz

Pesticide Research Laboratory and Graduate Study Center, The Pennsylvania
State University, University Park, Pa. 16802

Larvae of Trichoplusia ni (Hiibner) were parasitized during phase II of
either their 2nd or 4th instar by the solitary endoparasitoid, Hyposoter exiguae
(Viereck). Beginning at 36 hr post-parasitism and continuing at 8 hr intervals
thereafter, hosts were dissected and measurements of the contained parasitoid
taken. Hyposoter larvae eclosed from eggs between 36 and 44 hr post-para-
sitism in both 2nd and 4th instar hosts. Four larval instars were observed,
the first lasting about 2 days, the 2nd and 3rd lasting 1% days and the 4th
less than 1 day. Parasitoid head capsule widths measured 0.15, 0.34, 0.51 and
0.61 mm for those reared on 2nd instar hosts, and 0.15, 0.35, 0.54 and 0.67 mm
for those reared on 4th instar hosts. Parasitoids reared on 4th instar hosts
emerged to spin their cocoons between 6V2 and IV2 days post-parasitism at
26 ± 1°C, while those reared on 2nd instar hosts emerged a few hr later.
Parasitoid adults reared on both age groups emerged 6-7 days after spinning.
Adult parasitoids reared from 2nd instar hosts weighed 4.81 and 5.17 mg for
males and females, respectively, while those reared from 4th instar hosts
weighed 6.25 and 6.91 mg. Since host age has been shown to affect parasitoid
size and developmental rate, it is important that it be taken into account when
performing experiments on parasitoid fecundity, fertility and development.
Otherwise a wide range of values may occur for these parameters.

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Mite Consuming Capacity of Stethorus punctum (Leconte)

( Coleoptera : Coccinellidae )

L. A. Hull, D. Asquith and P. D. Mowery
The Pennsylvania State University, Fruit Research Laboratory, Biglerville, Pa.

17307

The functional response of the predator Stethorus punctum (Leconte) was
tested at various densities of the European red mite, Panonychus ulmi (Koch).
One S. punctum adult or larva was caged with adult female mites at the fol-
lowing density levels: 4, 8, 12, 16, 20, 50, 80 mites/ cage. The number of
mites consumed was recorded every hour and the mites replenished back to
the original levels. All feeding tests were conducted in a screened insectary
to simulate orchard conditions. The rate of consumption of overwintered adults
(spring feeding rate) rose at first with prey density, but leveled off at higher
densities. The rate of consumption of the 2nd and 3rd generation adults (sum-
mer feeding rate) increased with increasing prey density with no leveling off
attained. S. punctum larvae also exhibited an increased consumption rate as
prey density was increased. At the lower density levels the larval feeding rate
was higher than for male adults. As prey density increased beyond 20 mites/cage
the adults were able to consume more mites. S. punctum adults and larvae
stop feeding at dusk and resume feeding ca. 2 hrs. after sunrise. The ability
of S. punctum to functionally respond to increasing prey density plays an
important role in keeping the European red mite below economic injury levels
in Pennsylvania.

The Influence of a Juvenile Hormone Mimic (JHM) on Trichoplusia
ni (Hiibner) (Lepidoptera: Noetuidae) and Hyposoter exiguae
(Viereck) (Hymenoptera: Iclineumonidae)

Zane Smilowitz, Carol A. Martin k a and Eugene A. Jowyk
Pesticide Research Laboratory and Graduate Study Center, The Pennsylvania State
University, University Park, Pa. 16802

The influence of Altozar®, a juvenile hormone mimic (JHM), on parasitized
and unparasitized cabbage looper, Trichoplusia ni (Hiibner), was studied.
Parasitized and unparasitized T. ni larvae were reared on standard wheat germ
diet until early 4th instar then on diet containing 0.1, 1, 2 and 4 ppm of
Altozar. Larval development was essentially the same on all treatments from
the 4th to the 5th instar. The 1, 2 and 4 ppm treatments of the JHM had a
decided influence on the development of the unparasitized 5th stage larvae.
Individuals on diet containing the higher dosages remained in the larval stage

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263

up to 2 weeks longer than the controls and 0.1 treatment. Approximately 75%
of these began a supernumerary molt into a 6th instar, but none of them
pupated. About 20% of the remainder pupated. No adults were obtained from
2 and 4 ppm treatments and 2.5% from the 1 ppm treatment. T. ni larvae
parasitized by Hyposoter exiguae (Viereck) developed the same on JHM and
control diets. Parasitoids emerged from hosts on both diets at the same time
and immediately began to spin cocoons. When JHM was topically applied at
2 and 20 yg prior to parasitoid emergence, development was extended and adult
mortality ranged from 5 to 95%. JHM generally disrupts insect development
prior to the larval pupal molt. Since the parasitoids’ last larval molt occurs
after the host ceased feeding, apparently insufficient JHM is available to
influence parasitoid development. Topical application of the JHM nearing the
time of parasitoid emergence allows sufficient material to reach the parasitoid
and influence development. Thus the developmental pattern of a beneficial
insect must be known in order to determine the impact of a JHM.

Response of the Alfalfa Weevil Parasitoid, Microctonus colesi (Drea)
(Hymenoptera: Braconidae), to a Recommended Insecticide
Treatment in Pennsylvania

A. A. Hower, Jr., and J. E. Luke

Department of Entomology, The Pennsylvania State University, University Park,

Pennsylvania 16802

Studies were conducted from 1971 to 1974 to determine the impact of rec-
ommended insecticide treatments on Microctonus colesi, a parasitoid of the
alfalfa weevil, Hyper a postica (Gyll) (Coleoptera: Curculionidae) . Two adja-
cent multifarm complexes were chosen for the study. Methyl parathion was
applied on all fields in one region in accordance with the recommended
threshold level of 50 to 75% alfalfa tip injury or 2 wk before normal first
crop harvest date. Alfalfa weevil and M. colesi populations were monitored
weekly in 10 fields in each region during first crop growth. Immediate impact
of a first crop spray of methyl parathion at V2 lb Al/acre 2 wk preharvest
was a substantial reduction in both parasitoid and host populations in each
of the 3 years of application, 1971-1973. M. colesi larvae in the adult weevils
were the most prevalent stages of the parasitoid at the time of spraying. The
impact on M. colesi in the adult weevils was obvious as the sprayed area
contained in excess of 98% fewer adult weevils than the non-sprayed area
one week post spray in each of the 3 spray years. Consequently, each spray
year a significant reduction in the M. colesi larval population in adult weevils
was observed in the sprayed region. M. colesi adults and immatures in weevil

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larvae had just begun to materialize in the parasitoid population when the
spray was applied. However, any postspray decline in adult M. colesi in the
sprayed area resulted indirectly from earlier mortality of adult weevils har-
boring these parasitoids. M. colesi larvae in host larvae were not influenced
by the spray. Their reduced numbers in the sprayed area resulted from few
adult M. colesi available for parasitization and fewer host larvae available to
be parasitized. Better coordination of insecticide sprays and M. colesi devel-
opment is essential if M. colesi is to realize its maximum potential in Penn-
sylvania.

Determination of Seasonal Activity of Four Fruit Pests
Using Pheromone and Other Traps

Richard C. Moore

Department of Entomology, Connecticut Agricultural Experiment Station,

New Haven, Connecticut 06504

Pheromone traps were used for three successive seasons (1972-74) to deter-
mine activity peaks of codling moth Laspeyresia pomonella (L.), red-banded
leafroller Argyrotaenia velutinana (Walker) and oriental fruit moth Grapho-
litha molesta (Busck) while traps combining attractive baits and colors were
used to capture apple maggot Rhagoletis pomonella (Walsh) flies in sprayed
and unsprayed apple trees. Oriental fruit moth, codling moth and apple maggot
were more abundant in unsprayed than in sprayed apple trees. Red-banded
leafroller was as abundant or more abundant in sprayed than unsprayed trees.
Oriental fruit moth was captured over a period of 20 weeks with 3 activity
peaks occurring in May, July and September. Capture of red-banded leafroller
indicated 3 activity peaks for male moths occurring during a 2 2 -week emergence
period. Emergence and peaks of first generation moths occurred one month
earlier in 1974 than in 1972. Codling moth adults were active over a 16-week
period with 2 peaks in early June and August. Apple maggots emerged over a
9-week period with a single peak occurring in mid-July or early August.
Comparison of activity peaks of these moths over a 3 -year period with spray
intervals currently being used in Connecticut orchards indicated that modifi-
cations could be made using alternate middle row or extended interval spraying
to reduce pesticide use while controlling the apple orchard pests. A reduced
spray program in 1973 and 1974 resulted in less than 1.0% fruit damage by
these pests.

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265

Depth Selection in Buenoa (Heteroptera: Notonecticlae)

Steven H. Gittelman

Department of Ecology, Biological Sciences Division, University of Connecticut,

Storrs, Conn. 06268

The coexistence of similar species depends on the division of resources. While
resource sharing between sympatric congeneric species has attracted much
attention, little information exists on how species of Buenoa coexist. This study
documents micro-habitat separation in B. margaritacea and B. conjusa by se-
lection of different swimming depths. Buenoa conjusa swims closer to the
surface than B. margaritacea. Generally, nymphs of these species swim deeper
as they mature. The duration of a dive changes ontogenetically and differs
between species in a manner similar to that of swimming depth. The two
phenomena seem related. It is proposed that swimming depth and dive duration
depend on the amount of hemoglobin carried rather than physical gill efficiency.
Selection of the depth of water for swimming (as opposed to swimming depth
below the surface) differs between life-history stages and species. Nymphs
and species that swim closer to the surface enter shallow water more often.
Swimming depth affects prey selection. In shallow water (12cm) both species
prefer the same prey (Daphnia) . In deep water (28cm) prey selection differs,
with each predator preferring prey items in its preferred depth range.

Rearing the European Corn Borer, Ostrinia nubilalis (Hiibner)
(Lepidoptera: Pyralidae) on a Lima Bean Medium

G. D. Curl, P. P. Burbutis, and C. P. Davis
Department of Entomology and Applied Ecology, University of Delaware,
Newark, Delaware 19713

Our attempts (1971-72) to rear European corn borer, Ostrinia nubilalis
(Hiibner), collected in southern Delaware, on the meridic diet used at the
Corn Borer Investigations Laboratory, Ankeny, Iowa, were unsuccessful. With
modifications, an oligidic diet, previously used to rear cabbage looper, Trichop-
lusia ni (Hiibner), has proven highly satisfactory. The primary ingredient is
dried “baby” lima beans. Brewer’s yeast, agar, methyl p-hydroxybenzoate ,
ascorbic acid, Fumidil B, formaldehyde (37%), and distilled water constitute
the remainder of the diet. The larvae are reared at LD 15:9, 30°C and 20°C,
respectively. The following characterizes the colony under routine rearing
conditions. Egg viability averages over 90%. Eighty-one percent of the larvae
pupate in corrugated cardboard rolls above the media. Adult emergence aver-
ages 91%. Individual females lay an average of 22.1 egg masses (ca. 20 eggs/

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mass) over a period of 12.5 days. A comparison of the first seven generations
of a colony started in 1974 with the last seven generations (i.e. F26-F32) of
a colony started in 1971 showed no significant (p = .05) difference in number
of egg masses produced per female. Larvae, from one colony, were tested for
survival on corn plants after 20 and 30 laboratory generations. Results indicate
no difference in survival as compared to “wild” borers. Diapause induction
and termination studies showed no significant (p = .05) difference between a
colony reared for 31 generations and a colony reared for five generations.

Application of Harmonic Analysis and Polynomial Regression to
Study Flight Activity of Choristoneura fumiferana (Clem.)

(Lepidoptera: Tortricidae) in the Field

G. A. Simmons and C. W. Chen

Department of Entomology and Department of Mathematics, University of Maine,

Orono 04473

Counts of male and female budworm moths captured in aerial Malaise traps
were obtained every hr continuously for 1344 trap hrs over 14 days of study.
Captures averaged 12.6/h and 4.4/ h and ranged from 0 to 183/h and from 0
to 82 /h per trap respectively for males and females. Harmonic analysis showed
the circadian rhythm of male activity could be described by Pt = 12.560 -
11.431 Cos(27rt/24) - 6.156 Sin(27rt/24) + 5.273 Cos(47rt/24) + 6.199
Sin(47rt/24) and the circadian rhythm of female activity by Pt = 4.438 -
5.789 Cos(27rt/24) - 0.558 Sin(27rt/24) + 3.598 Cos(4irt/24) + 0.941
Sin(4?rt/24) where Pt is the mean number of captures in the hr interval and
t = 0 corresponds to 0800 EDST (and t = 0, 1, 2, . . ., 23). Peak activity
occurs at 2152 EDST and 2021 EDST respectively for males and females.
The amplitude of male activity is ca. twice that of the female. Polynomial
stepwise regression of residuals (rt = Pt - Yt) vs weather identified inter-
correlated variables influencing activity. A study of partial derivatives with
respect to weather variables showed complex interactions with humidity, cloudi-
ness, rainfall, wind velocity, and barometric pressure change. Males and fe-
males differed considerably in their response. Circadian rhythms are inter-
preted as gross adjustments to average environmental conditions; and reaction
to weather is interpreted as fine adjustments of the population to specific
environmental conditions.

Vol. LXXXIII, December, 1975

267

Mosquito Control in Unusual Breeding Sites in Southern Italy
( Diptera : Culicidae )

John L. McDonald

Navy Environmental and Preventive Medicine Unit No. 7, Box 41, FPO,

New York 09521

Mosquito problems in the Mediterranean area are characteristically influ-
enced by both climate and culture. Irregular rainy periods have resulted in
the use of a variety of cisterns, irrigation systems (often aqueducts) and
catchment areas for water storage during dry periods. Unusual mosquito
breeding sites are found in many buildings, caves and temples, commonly
referred to as “ancient ruins.” In some instances, some of these “ancient ruins”
have become altered by age or partly sunken over long periods of time due
to nearby geological activity causing the formation of lakes, ponds and slow-
moving streams, all which have become ideal mosquito breeding sites. Because
so many of the mosquito breeding sites are contained within historical land-
marks, mere access to them often requires significant effort. Conventional
remedial means such as draining, filling or spraying with insecticide would
be imprudent, contaminating, or impractical. Use of mosquito fish, Gambusia
afjinis has been useful in resolving some of the mosquito breeding site problems.
In the many elaborate irrigation systems, mosquito breeding sites coexist with
water wastage or poorly managed water. Increasing cost of water for irrigation
has done much to mitigate this problem.

Temefos Residues in the Salt Marsh Snail Melampus bidentatus
Say (Bassommatophora: Ellobiidae)

George Fitzpatrick and Donald J. Sutherland
Department of Entomology and Economic Zoology, Rutgers University,

New Brunswick, N.J. 08903

Uptake of the mosquito larvicide temefos by populations of the salt marsh
snail Melampus bidentatus in the field was measured by gas chromatographic
analysis. Snails exposed in the field to temefos treatments as applied in normal
mosquito control work were placed on ice and taken to the laboratory for
analysis. Temefos was extracted in dichloromethane and cleanup was per-
formed with hexane and acetonitrile. Uptake of measurable levels of temefos
occurred within one day after the first treatment of a 2% granular formulation.
A longer period of time, 3 weeks, elapsed before uptake following treatment
with a temefos emulsion. Residues in samples of snails exposed to the granular

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formulation were generally around 1 ppm, with residues measured as high
as 8.75 ppm in one sample. Residues were considerably lower in snails exposed
to the emulsion. The highest residue was 0.059 ppm in this case. Residues in
snails exposed to the emulsion fell below detectable levels in less than 3 weeks
following cessation of treatments, while measurable amounts were found in
snails exposed to the granular formulation for more than 5 weeks after the
last treatment. Detection of temefos in M. bidentatus for such long periods
suggests the potential for movement of this insecticide through food webs
exposed to the granular formulation.

Bionomics of the Tufted Apple Budmoth, Platynota idaeusalis
(Walker) (Lepidoptera: Tortricidae), in Pennsylvania
Apple Orchards

William M. Bode

Pennsylvania State University Fruit Research Laboratory, Biglerville, Pa. 17307

P. idaeusalis is a leafroller which is a major pest of apple in Pennsylvania
and areas to the south. Larvae consume tissue from leaves and the surface of
fruit. Larvae are difficult to control with insecticides because many are hidden
in protected places which spray does not reach. There are two broods annually
with second brood larvae overwintering. Eggs are laid on the upper surface
of apple leaves in flat green masses which contain an average of 70 to 80 eggs.
Eggs for the first brood are laid during June and the first half of July. First
brood larvae develop during June and July, and some even into August. Eggs
for the second brood are laid during August and September. Second brood
larvae do more damage to apples because they are more numerous and the
apples are larger. Also, growers tend to terminate insecticide applications too
early to protect fruit from all larvae. Larval feeding on apples is finally ended
by the harvesting of the fruit. Larvae of all instars overwinter in leaf litter
under trees; they do not diapause, and may feed on ground vegetation when-
ever the temperature is high enough to permit activity. Natural biological
control agents include some hymenopterous parasites of eggs and larvae, a
tachinid parasite of larvae, some virus diseases, and probably some predators.
A synthetic sex attractant may be used to detect male moths and monitor their
seasonal flight periods.

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269

Parasitization of the spruce budworm, Choristoneura fumiferana
(Clemens) (Lepidoptera: Tortricidae) by Brachymeria intermedia
(Nees) (Hymenoptera: Chalcididae)

David E. Leonard

Department of Entomology, University of Maine, Orono 04473

The introduced parasitoid Brachymeria intermedia attacks the gypsy moth
in Maine, and laboratory studies (Minot and Leonard, p. 269) show that
it will parasitize spruce budworm pupae. No recoveries were made in spruce
budworms after limited field releases of this parasitoid in mature stands of
spruce and balsam fir in 1973 and 1974. To facilitate observations and
sampling, the 1975 release was made in a spruce budworm infested 2 ha
plantation of 3 to 5 m white spruce in Shin Pond, ME. On 23 Jun, when
35% of the budworm had pupated, 14,000 B. intermedia were released along
a 28 m transect. The weather during the period of spruce budworm pupation
and adult emergence was warm, with highs in the 33 °C range, and clear or
partially cloudy days. Activity of the released parasitoids was assessed by
counting the number of adults observed in 10 min searching of trees in the
area of release. Parasitism was determined by collecting and rearing 5453
spruce budworm pupae in the release area, and 2 743 pupae in trees 10 to 20 m
from the release points. Two days after release, B. intermedia were observed
up to 74 m from the nearest release site, but most adults were observed in or
immediately adjacent to the release area, flying about the tree terminals and
occasionally landing. Ten min counts of parasitoids between 24 Jun and 2 Jul
ranged from 1 to 53, with the lowest counts in the morning between 0830
and 1030 hr. In the release area, 18.2% of the spruce budworm pupae yielded
B. intermedia adults, and in the adjacent area, 11.7%. The percentage mor-
tality is considerably higher than the total native pupal parasitoid complex.
The sex ratio of the recovered parasitoids favored males by 3:1, and may be
related to the smaller size and limited food reserves of the host pupae. This
aspect is currently being studied.

Influence of Physical Factors on the Behavior and Development
of Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae)

Mildred C. Minot and David E. Leonard
Department of Entomology, University of Maine, Orono 04473

B. intermedia is a polyphagous endoparasite of Lepidoptera pupae. This
primary, solitary parasitoid has been successfully introduced into gypsy moth
populations in North America. This study tested the response of the parasitoid

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270

to physical factors and developmental rates in gypsy moth pupae at different
temperatures. Adult parasitoids were exposed to a temperature gradient that
ranged from 22° to 31°C. They were exposed to a humidity gradient that
ranged from 11% to 90% relative humidity. Geotactic, photo tactic and olfactory
reactions, and diel periodicity were also investigated. B. intermedia preferred
temperatures between 26.5° and 28.5° and preferred the dry end of the hu-
midity gradient. They were positively phototactic and inactive in the dark.
No geotactic or olfactory responses were detected. They were most active
between 1300 and 1700 hr. There was considerable unexplained mortality
among pupae exposed to the parasitoid. The same number of hosts were killed
at all 3 developmental temperatures, but twice the number of B. intermedia
emerged at 28° as at 23°. At 18° only 3 parasitoids emerged from 120 exposed
pupae. These experiments confirm field observations of several workers that B.
intermedia is attracted to a warm, dry physical environment subjected to high lux.
Adults are most active in the afternoon when the above conditions are more likely
to be encountered. Development is most rapid and successful at the highest
temperature investigated, 28°.

The Use of Autoradiography to Detect RNA in Polyhedral
Inclusion Bodies of Insect Nuclear Polyhedrosis Viruses

Sally B. Padhi and Arthur H. McIntosh

Waksman Institute of Microbiology, Rutgers Univ., New Brunswick, N.J. 08903

Nuclear polyhedrosis viruses (NPVs) which infect Lepidoptera are DNA
viruses of potential use as biological control agents. There have been several
reports that RNA is in the polyhedral protein which surrounds the NPV
particles but definite proof of its presence and/or source is lacking. An at-
tempt was therefore made to apply autoradiography to ascertain whether RNA
is present in PIBs. The Trichoplusia ni (cabbage looper) cell line of Hink
was infected with Autographa calif ornica (alfalfa looper) NPV and labeled
with H3-uridine (RNA specific). Tritium labeled thymidine was used as a
positive control and H3-glutamic acid and H3-glucosamine were used in other
treatments. Procedures were developed for extraction of PIBs from the cells,
and slides were prepared for autoradiography. Because of the small size of
PIBs and their adherence to the cells, difficulties prevented the gathering of
conclusive results. Autoradiograms contained labeled cellular debris which pre-
vented a clear demonstration of whether or not the PIBs are labeled. Indi-
cations are that there is little or no labeling on PIBs produced in the uridine
labeled cells, thus suggesting that PIBs do not contain RNA. However, owing
to cellular debris on the slides it was quite difficult to detect labeled PIBs
in the positive control (thymidine treatment). In conclusion, it is necessary

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271

to improve methods of PIB purification without losing portions of the PIB
which might be labeled. Autoradiography should then be a useful method to
apply to the question of whether or not RNA is present in PIBs.

A Toxic Factor from the Established Cell Line, CP-169 (Hink) :
Carpocapsa pomonella (Lepidoptera: Olethreutidae)

Carol Rechtoris and Arthur McIntosh
Boyce Thompson Institute, Yonkers, N.Y. 10701, and Rutgers University,

New Brunswick, N.J. 08903

Spent media recovered from 5 out of 8 different insect cell lines contain
factors toxic to the established cell line, TN-368 (Hink) ( Trichoplusia ni )
(Lepidoptera). The highest concentration was detected in media from CP-169
cultures. Both the TN-368 and the CP-169 lines have been adapted to TCI 99-
MK (McIntosh et al., 1973). Vertebrate cell lines proved not to elaborate
material toxic to TN-368 cells. The toxic factor, designated CpT, is elaborated
by CP- 169 cells into the growth media. Supernatant media from freshly
washed cultures were non-toxic for TN-368 cells. However, if such treated
cultures were incubated for 24 hours, the toxic factor could be detected in
the media. In addition, extracts from ruptured washed cells proved to be toxic.
TC199-MK incubated at 30°C. for 3 months was non-toxic. CpT is filterable
{.22 fx m Millipore filter), and is inactivated at 80°C. for 30 minutes. It has a
low molecular weight, as shown by centrifugation, and the Cytotoxic Dose Fifty
of a pooled sample is 103-25 units/ ml. CpT is believed to be a non-infectious
agent since it cannot be passaged in TN-368 cells. Furthermore, electron mi-
croscopy of inoculated cultures revealed no virus or other infectious microbes,
and cultures tested negative for mycoplasmas. The early appearance of a toxic
effect 6 hours post inoculation, suggests a toxin.

Changes in Tolerance of Porthetria dispar (L. ) (Lepidoptera:
Lymantriidae) to Insecticides in Relation to Larval Growth
and Mixed-Function Oxidase Activity

Sami Ahmad and Andrew J. Forgash

Department or Entomology and Economic Zoology, Rutgers University,

New Brunswick, N.J. 08903

In vitro investigations have demonstrated mixed-function oxidase (MFO)
activity in several tissues of gypsy moth larvae (Insect Biochem. 3:263, 1973;
Int. J. Biochem. 5:11, 1974). The gut MFO activity, which is 67.8% of the
total activity, rises markedly during larval development. The increase in the

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specific activity of this enzyme system in the fifth instar is ca. fivefold over
the third instar (J. Insect Physiol. 21:85, 1975). The present communication
reports the investigations on the effects of increase in the MFO activity on
the susceptibility of advancing instars of gypsy moth larvae to carbaryl and
diazinon. Topical tests showed that there is a continued increase in tolerance
to carbaryl and diazinon with larval growth, with the result that 5th-instar
larvae can tolerate 25 X as much carbaryl and 50X as much diazinon as 2nd
instars at the LD50 level; on a weight basis the tolerances are 1.9 and 4.5X,
respectively. There is also a substantial increase in tolerance with growth
within instars, but this is largely correlated with size. Piperonyl butoxide and
2,6-dichlorobenzyl-2-propynyl ether, inhibitors of MFO activity, are ineffective
against 2nd instars, but with subsequent instars the activity increases greatly
so that by the 5th instar only one-fifth the usual dose of carbaryl is needed
to kill a larva when applied in combination with the propynyl synergist. The
increase in tolerance to carbaryl and diazinon with larval growth, as well as
increase in synergist efficacy, therefore, correlates with increases in MFO
activity in advanced instars.

Oxygen Consumption of Coleomegilla maculata lengi Timberlake
(Coleoptera: Coccinellidae) Measured in a
Differential Respirometer

Mark E. Whalon and Bruce L. Parker

Department of Entomology, University of Vermont, Burlington, Vermont 0S401

A Gilson differential respirometer was used to measure 02 consumption of
adult female C. maculata. The respirometer had the capability of simultane-
ously measuring 02 uptake in 14 separate vials. Field-collected beetles were
tested singly and in groups of 10. Their respiration was monitored for 6 hr
at 15 min intervals at each of 4 different temperatures; 6°, 12°, 18° and

24°C. At 24°C the respiration of beetles tested singly could accurately be

determined. At reduced temperatures the differential respirometer fluctuated
significantly thus introducing variation in excess of 0.01 /xl Oo/mg/15 min.
If 10 beetles were placed in each vial at 6°C variation was minimized and 02
consumption could be measured. An 02 consumption curve was established. Qn0
values were similar to those reported for other insects. Oxygen consumption
for C. maculata was 0.042 ± 0.013 /d 02/mg/15 min at 6°C, 0.10 ±

0.005 at 12°C, 0.158 ± 0.011 at 18°C, and 0.316 ± 0.046 at 24°C. A

Gilson respirometer can be used to measure 02 consumption (at 24 °C) of
adult female C. maculata tested individually, but accuracy was decreased and
variation increased as temperatures were reduced. If 10 beetles per vial were
used the variation in 02 consumption measurements was minimized. Adult

Vol. LXXXIII, December, 1975

273

female field-collected C. maculata respired at levels from 0.042 ± 0.013 /A
02/mg/15 min at 6°C to 0.316 ± 0.046 /xl 02/mg/15 min at 24°C.

A Phylogeny for Paracymus Thomson (Coleoptera: Hydrophilidae)
Based on Adult Characters

David P. Wooldridge

Pennsylvania State University, Ogontz Campus, Abington, Pa. 19001

An analysis of adult characters reveals that the 69 known world species of
Paracymus fall into 6 distinct groups which, on the basis of distribution data,
appear likely to be monophyletic. The characters examined included the form
of the male genitalia, modifications of the male protarsi, the number of an-
tennal segments, the form of the mesosternal modifications, the extent of
mesofemoral pubescence, dorsal punctation and pigmentation, and the extent
of development of a carina on the first visible abdominal sternite. Two species
groups, restricted to the New World, have the penis very flattened in cross-
section, the antennae 7 segmented and the mesosternal lamina well developed
and reaching the mesosternal crest. In the elegans group, containing 15 species,
the mesofemoral pubescence reach nearly to the knees, while the nanus group,
with 7 species has the mesofemoral pubescence restricted to a basal triangle.
In all other groups, the penis is thickened in cross-section, the antennae have
from 7 to 9 segments, and the mesosternal lamina is less well developed.
The evanescens group consists of 23 Old World species with rounded parameres
and reduced mesofemoral pubescence. Two groups with extended mesofemoral
pubescence are found in both hemispheres, although both are primarily New
World. The aeneus group with 8 species has the parameres flattened in cross-
section. The subcupreus group consists of 15 species with rounded parameres.
The secretus group contains a single New World species with no mesofemoral
pubescence.

Intra-instar Respirometric and Phase Distribution Differences in
Trichoplusia ni (Hiibner) (Lepidoptera: Noctuidae)

Larvae

Douglas G. Baugher and William G. Yendol
Pesticide Research Laboratory and Graduate Study Center and Department of
Entomology, The Pennsylvania State University, University Park,
Pennsylvania 16802

When investigations dealing with lepidopterous larvae are reported, the age,
instar, or weight are used for identification. With Trichoplusia ni, larvae of a

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given age have weights which are normally distributed, but the individuals
may be distributed into more than one instar. Since the physiological state
of a larva is not necessarily represented by age or instar, it is important to
delineate any metabolic differences among larvae, and to determine the daily
distributions of larvae by instar and phase within the stadium. A Gilson
Differential Respirometer was used to measure respirometric rates of 4th
and 5th instar T. ni 8 through 14 days post-eclosion. Larvae were morpho-
logically segregated into 5 phases for each instar. Larval phase distribution
within 4th and 5th instar could not be reliably predicted by age, and random
selections did not yield reproducible phase proportions. Respirometry dem-
onstrated significant differences between instars, and phases within the instars
when rates were determined by /A 02/mg wet weight/ hr. When respiratory
activity was assessed by accumulating the respiratory rates and fitting a re-
gression line, y = bx + c, c = o, the slopes ranged from 0.41 to 0.93, depending
upon the phase and instar. All regression lines had low residual mean squares,
r-squared > 0.95, and were significant at p < 0.01. T. ni larvae did not follow
a general trend of reduced respiration as age or weight increased. Respiratory
rate appeared to be phase dependent. Since the phases within an instar were
neither predictable by age nor were they metabolically homogeneous, experi-
mental precision may be increased by designing tests with similar phase dis-
tributions of larvae in each treatment. This may be necessary to obtain
meaningful comparisons or reproducible results if the physiological states of
the insects can alter treatment effects.

The Oenocytes of Tenebrio molitor Linnaeus
(Coleoptera: Tenebrionidae)

Jack Colvard Jones and Dorothy Hoelzer
Department of Entomology, University of Maryland, College Park, Maryland

20742

The large, conspicuous, naturally yellow-colored, ectodermally derived oeno-
cytes of Tenebrio molitor were studied in freshly dissected larvae, pupae and
adults in unstained saline whole mounts, and were examined with ordinary
light and with phase optics at magnifications of 50 to 1,000 times. The cells
did not pick up any of a series of dyes (neutral red, ammonia carmine, pure
carmine, trypan blue, alcian blue, methylene blue, toluidine blue or India ink)
after these had been injected in varying concentrations into the hemolymph.
On the other hand, the pericardial cells quickly absorbed these dyes. The fat
bodies did not incorporate any of the dyes. The oenocytes showed no con-

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275

spicuous cytological changes in either number, color or configuration relative
to larval ecdysis, pupation, or maturation of eggs. Females with undeveloped,
developing, and fully developed oocytes had oenocytes which were of the same
general appearance as those in females which had recently laid their eggs.
It is concluded from this study that the oenocytes of the mealworm do not
form a part of an athrocytic system. Although the oenocytes are reported to
produce a cuticular material near the time of ecdysis, and are said to be
important in the development of the eggs, and are supposed to secrete a
hemolymph protein, and are claimed to secrete ecdysone in some species, these
cells in the mealworm exhibit no striking changes in their general anatomy
which would lend support to their participation in any of these activities.

Symposium: “Solving Insectary Production Problems”

M. A. Hoy, Moderator

The Genetic Implication of Insect Mass Rearing Programs,
G. Bush, University of Texas, Department of Zoology,
Austin, Texas

Monitoring the Quality of Laboratory-Reared Insects,

M. B. Huettel, USDA, ARS, Gainesville, Florida

Improving the Quality of Laboratory-Reared Insects,

M. A. Hoy, USDA, Forest Service, NEFES,

Hamden, Connecticut

Genetic Changes Occurring in Flight Muscle Enzymes of the
Screwworm Fly During Mass Rearing

G. L. Bush

Department of Zoology, University of Texas, Austin, Texas 78712

A preliminary population genetic study of genetic variation in laboratory and
natural populations of the screwworm fly by gel electrophoresis of enzymatic and
non-enzymatic proteins representing 36 presumptive loci revealed extensive differ-
ences in both allele frequency and the degree of heterozygosity. Further analysis
indicated that the most rapid changes occurred during factory colonization in en-
zymes important to flight muscle metabolism and flight activity such as a-GDH
and PGM. The fact that all factory-adapted strains eventually end up with similar
genetic structure and greatly reduced genetic variability indicates that factory
rearing conditions are exerting strong selection pressures for a “factory type.”

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Because screwworm adults are capable of dispersing long distances and mating
may occur in the air or require some flight activity, monitoring for changes
in enzyme systems essential to normal flight activity could prove to be a
sensitive system for maintaining vigorous factory strains. Lab strains could
also be tailored to fit changing environmental conditions between summer and
winter using specific alleles.

Monitoring the Quality of Laboratory-Reared Insects

M. D. Huettel

Insect Attractants, Behavior and Basic Biology Research Laboratory,
USDA-ARS, Gainesville, Florida 32601

The process of monitoring implies the use of a warning system with an
element of periodicity. To be useful a monitoring system must be applicable
on a routine basis, subject to the constraints of reproducibility, simplicity and
economy. The quality of any particular trait may be defined as the difference
between the trait in a wild insect and a laboratory insect. The overall quality
of the laboratory insect, however, can only be measured in terms of how well
it functions in its intended role when released into the field. It is the quality
of traits which we usually attempt to measure with a monitoring system.

Laboratory insects should possess certain rather specific traits to perform
well in the field. They should have life histories similar to the wild population.
They must be able to disperse from the release site, find and utilize locally
available nutrients, and locate mating sites (host plants and pheromones) or
hosts or prey. They must be able to court and mate successfully in most cases.
During their life span in the field they must also be able to survive local
climatic conditions and avoid predators.

Each of these traits should be amenable to monitoring in the laboratory or
field. Emphasis will be placed on identifying the monitoring system most
applicable to each trait and its state of development as a useful method.
Finally the possibilities for, and difficulties of, extrapolation from trait quality
to field performance will be discussed.

Improving the Quality of Laboratory-Reared Insects

M. A. Hoy

USDA, Forest Service, Northeastern Forest Experiment Station,

Hamden, Connecticut 06514

Genetic improvement of insects has generated only a moderate amount of
discussion in recent years because the problems of maintaining genetic quality
have not been solved. However, certain insects have been improved genetically.

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277

“Domesticated” insects such as honeybees and silkworms have long associa-
tions with man and have been improved in many ways. Some parasitoids have
also undergone selection to improve insectary production and/or field effec-
tiveness.

Parasitoids, or any insects destined to survive and reproduce in a natural
environment, present particularly difficult problems for an improvement pro-
gram. Desirable attributes to be selected must be clearly definable. Adequate
genetic variability must be provided to allow selection to operate. Adequate
selection procedures are a must. Finally, maintenance of the integrity of the
improved strains under field conditions may need to be provided for.

Heterosis has been largely ignored in genetic improvement programs, except
for the spectacular improvements exhibited in silkworm and honeybee im-
provement programs. Some data suggest that heterosis may be useful to improve
the effectiveness of inoculative or inundative releases of parasitoids or predators.
There is yet inadequate experimental evidence to judge the general value of
selection and hybridization for improving insects. Future field testing will
demonstrate the value of such improvement methods.

Symposium : Biosystematics

Gordon Gordh, Moderator

Systematics and Ecology of Chrysopidae (Neuroptera) : Theoretical
and Applied Implications. Catherine A. Tauber and Maurice
J. Tauber, Department of Entomology, Cornell University,
Ithaca, N.Y. 14853

Some Evolutionary Trends in the Chalcidoidea (Hymenoptera) with
Particular Reference to Host Preference. Gordon Gordh,
Systematic Entomology Laboratory, Agr. Res. Serv., USDA.

Territoriality in Male Bees (Hymenoptera: Apoidea).

Edward M. Barrows, Department of Biology,

Georgetown University, Washington, D.C. 20057

Systematics and Ecology of Chrysopidae (Neuroptera) :
Theoretical and Applied Implications

Catherine A. Tauber and Maurice J. Tauber
Department of Entomology, Cornell University, Ithaca, N.Y. 14853

The family Chrysopidae — green lacewings — is a member of one of the most
primitive holometabolous orders (Neuroptera), and clarification of the evolu-

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tionary changes within the Neuroptera is not only of intrinsic value and
interest, but it can contribute to an understanding of the evolution of the
more highly evolved insect orders. In addition to their taxonomic value, many
chrysopid species are useful as subjects in ecological-physiological studies, and
as important predators in integrated control programs in various agricultural
ecosystems.

Our recent investigations with the Chrysopidae encompass 4 categories:
systematics, phenology, behavior, and biological control. The systematics work
is based on a classical, comparative morphological approach and on data de-
rived from experimental studies in phenology and behavior. By combining
the results of morphological and experimental studies, we not only broaden
the basis for the classification and thus advance the systematics of the group,
but we also provide information essential to the use of Chrysopidae (e.g.
chrysopid strains) as biological control agents (1).

Specifically, in the area of systematics, our studies with the Chrysopidae
represent the 3 levels or stages of biological classification:

1. alpha taxonomy — The larvae of most North American species have not been
described. To promote species identification, we have reared and studied larvae
of most North American species and the process of description is well underway
(2,3,4).

2. beta taxonomy — Comparative analyses of the larval morphology (3,4) and
the biological characteristics of adults and larvae (5,6) provide a basis for a
sound classification and for keys to the taxa.

3. gamma taxonomy — The phenological adaptations of geographically diverse
populations are valuable indicators of species-complexes and evolutionary trends
within the genus Chrysopa. Phenological studies, in combination with hybridi-
zation tests, contribute to an understanding of the genetic diversity and the
evolutionary history of geographic populations (7).

Success or failure of biological control projects depends in large part on the
degree to which selected strains of beneficial species are adapted to biotic and
abiotic factors of particular environments. Our recent investigations have led
to the recognition of strains or races within geographically diverse chrysopid
species that are currently used as biological control agents. These strains are
characterized on the basis of morphological, phenological and behavioral criteria
(7,8), and our studies show that some of these strains are better adapted
than others to particular localities and particular agricultural ecosystems (9).

Literature Cited

Tauber, M. J., and Tauber, C. A. 1976. Ann. Rev. Entomol., 21: (in press).

Tauber, C. A. 1969. Univ. Calif. Pubis. Entomol., 58: 1-94.

. 1974. Canad. Entomol., 106: 1133-1153.

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279

. 1975. Ann. Entomol. Soc. Amer., 68: 695-700.

Tauber, M. J., and Tauber, C. A. 1974. Canad. Entomol., 106: 921-925.

. 1974. Canad. Entomol., 106: 969-978.

Tauber, C. A., and Tauber, M. J. 1973. Canad. Entomol., 105: 1153-1167.

Tauber, M. J., and Tauber, C. A. 1973. J. Insect Physiol., 19: 729-736.

. 1975. Canad. Entomol., 107: 589-595.

Some Evolutionary Trends in the Chalcidoidea (Hymenoptera)
with Particular Reference to Host Preference

Gordon Gordh

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

Taxonomically, the parasitic hymenopteran superfamily Chalcidoidea is
among the most poorly known within the Insecta because the number of
systematists working on the group is small, the group is large, and progress
has been slow. Presently, we recognize about 1,220 genera and 9,950 species
of chalcidoids. Hosts for 27% of the genera are unknown.

I believe this superfamily ultimately will be recognized as numerically the
largest and biologically most diverse insect group. Several sources of informa-
tion and lines of reasoning lead me to this conclusion: (1) the chalcidoid host
spectrum extends from ticks and spiders to aculeate Hymenoptera. (2) Rapid
genetic recombination of superior genotypes and subsequent speciation has
been accelerated among chalcidoids because generation time is short (some-
times less than 8 days), they possess several modes of parthenogenesis (ar-
rhenotoky, thelytoky, deuterotoky), and intensive inbreeding via sibmating is
widespread. (3) Chalcidoids demonstrate several host-exploitation strategies:
they develop as obligate egg parasites, larval parasites, pupal parasites, egg-
larval parasites, larval-pupal parasites, and many species are obligate or facul-
tative hyperparasites. Thus, each insect species represents several potential
niches for parasitic chalcidoids. Also, phytophagy has evolved several times in
the Chalcidoidea. (4) Chalcidoids display a finite number of morphological
types, but there are no a priori reasons why morphological criteria must accom-
pany the species status; sibling species are abundant in the Chalcidoidea. (5)
The Neotropical, Ethiopian, Oriental and Australian faunas are almost totally
unknown.

Analysis of generic and suprageneric levels in the taxonomic hierarchy shows
that different taxa of chalcidoids have adopted different progenative strategies,
which are categorized as specialists, generalists and opportunists. Specialists
attack a specific host-taxon, such as a genus or family ( Desantisca spp. on
Latrodectus spp.; Chalcis spp. on Stratiomyiidae) ; generalists prefer a habitat
rather than a taxonomically cohesive group of hosts (Z agr ammo soma spp. on

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leaf mining Lepidoptera and Diptera) ; opportunists also have host spectra that
transcend ordinal and class limits, but they are not restricted to a particular
habitat ( Dibrachys spp., Tetrastichus spp.).

Analysis of chalcidoid biotic potential at the family level, based on parasite-
taxa size, indicates that some exploitation strategies are more successful than
others. Two families (Trichogrammatidae, 64 genera, 369 species; Mymaridae
59 genera, 794 species) are exclusively egg parasites and attack many host
orders. The Encyrtidae (491 genera, about 1,700 species) have focused on the
Homoptera, especially scale insects, and are among the most successful groups
of parasites. The Pteromalidae (233 genera, about 1,400 species), while at-
tacking innumerable taxa of hosts, most commonly parasitize Coleoptera and
Lepidoptera. Extreme specialization does not necessarily reflect much generic
diversity. Thus, the Leucospidae (4 genera, 127 species), parasites of solitary
bees and wasps, and Eucharitidae (11 genera, 193 species), parasites of ants,
have not evolved in great numbers.

Some phylogenetic considerations are made based on host preference, geo-
graphical distribution, taxon size and morphology.

Territoriality in Male Bees (Hymenoptera: Apoidea)

Edward M. Barrows

Department of Biology, Georgetown University, Washington, D.C. 20057

Territoriality in male bees is their continued occupancy and defense of a
topographic area or landmark against conspecific males. Their territories vary
in size and in durations of ownership. Types of landmarks included are nests
of conspecific females, all or certain parts of plants, pebbles, rocks, and patches
of bare earth. Any of these landmarks may be rendezvous places, locations
where a bee is likely to find a mate. Males, conspecific females, or both have
been observed feeding, mating, and nesting in territories. The smallest territory
is held for 3 to 10 days by the sweat bee, Lasioglossum rohweri (Ellis).
It is a microterritory which is only about as large as the bee which defends it.
Male Centris pallida Fox have digging micro territories above virgin females in
soil. Calliopsis andreniformis Smith guards a territory 2 to 5 meters long and
chases away its parasite, Holcopasites. Males of both Calliopsis and L. rohweri
may function in nest defense. Protoxaea gloriosa (Fox), like other species,
defends plants where it is likely to encounter females. The longest territory
(38 m long) is held by Anthidium banningense Cockerell for at least 3
days and males of A. manicatum L. are among the most aggressive of all male
bees. They chase and sometimes harm almost any insect except conspecific
females encountered in territories. In the Galapagos Islands, the carpenter bee,

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281

Xylocopa darwini Cockerell, chases Geospiza finches as well as other insects.
A European mason bee, Hoplitis anthocopoides (Schenck), varies the size of its
territories with regard to its food plant and the numbers of conspecific males
in a given area. These males hold territories for a median of about 16 days.

Factors believed to affect the cost/ benefit ratio of territoriality include
distribution and detectability of female emergence sites, detectability of nest
entrances, distribution and quality of foraging areas, and the number of con-
specific male competitors in an area.

Territoriality in male bees probably has manifold functions. It may function
to increase the efficiency of natural resource utilization and of escaping from
predators because males learn the topography of their territories quite well.
Territoriality may space individuals over the available habitat, reducing com-
petition for food and females. Furthermore, this behavior may reduce the
incidence of disease and parasites and the time spent in agonistic encounters.
Finally, if territories of a particular species cannot be compressed, territoriality
may function in population regulation. Territoriality represents a group of
adaptations that differ from one species of bee to the next.

Bee species with territorial males usually have relatively large males.

Territoriality in male bees is probably a result of convergent evolution.
Because this behavior is found in isolated genera, in 7 of the 9 families
of bees, it appears to be a derived, rather than an ancestral, type of behavior.

Symposium: Ecology of Forest Defoliators

Douglas C. Allen, Moderator

The Importance, Biology and Control of the Birch Casebearer, an
Imported Pest, in Insular Newfoundland. Dr. David G. Bryant,
Newfoundland Forest Research Center, Canadian Forestry
Service, P. O. Box 6028, St. John’s, Newfoundland,
CANADA A1C 5X8

The Role of Defoliators in the Anthropod Community of Red
Maple Crowns. Mr. Jan Volney, Maritimes Forest Research
Center, Canadian Forestry Service, P. O. Box 4000,
Fredericton, New Brunswick, CANADA E3B 5G4

The Douglas-fir Tussock Moth — Influence of Host Foliage.

Mr. Roy Beckwith, USDA, Forest Service, Pacific NW Forest
and Range Experiment Station, 3200 Jefferson Way,
Corvallis, Oregon 97331

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Current Research with Telenomus alsophilae Viereck, an Egg
Parasite of the Fall Cankerworm. Mr. Arnold T. Drooz,
USDA, Forest Service, Forest Sciences Laboratory,

P. O. Box 12254, Research Triangle Park,

North Carolina 27709

The Bimodality of Gypsy Moth Populations. Dr. Robert W.
Campbell, USDA, Forest Service, Pacific NW Forest and
Range Experiment Station, 3200 Jefferson Way,
Corvallis, Oregon 97331

Douglas-fir Tussock Moth, Orgyia pseudotsugata (McD.)

(Lepidoptera: Lasciocampidae) : Influence of Host Foliage

Roy C. Beckwith

Forestry Sciences Laboratory, Corvallis, Oregon 97331

The Douglas-fir tussock moth, Orgyia pseudotsugata (McD.) is one of the
more important defoliators of Douglas-fir, Pseudotsuga menziesii var. glauca
(Beissn.) Franco and true firs, Abies spp., in Western North America. Past
outbreaks have occurred in fir forests from British Columbia south to Arizona.
A resume of tussock moth biology is presented with special emphasis on host
effects.

Tussock moth outbreaks are characterized by a rapid increase, followed by
a sudden and complete collapse; the outbreak cycle usually spans a 3 -year
period in any one location. Endemic populations are extremely difficult to
find by present standard sampling techniques.

The 1972-74 Blue Mountain outbreak in Oregon and Washington prompted
a laboratory study on the effects of host foliage on the tussock moth. Three
common hosts were used in the test; high density field populations were
simulated by forcing one-half the population to feed upon old growth foliage
following the second instar. This “stress factor” had the most significant
effect upon the population, resulting in increased development time, frass
production and mean number of instars, and a decrease in larval size, pupal
weight, and egg production.

It appears that populations can increase equally well on grand fir and
Douglas-fir; the degree of acceptance of old growth foliage may govern sur-
vival rate and ultimate tree damage. Although fed upon, subalpine fir will
not support dense populations.

Only current foliage will be consumed during the release phase of a proposed
hypothetical outbreak model (1), but larvae are forced to feed on old-growth
foliage under high density populations in the outbreak phase which is detri-

Vol. LXXXIII, December, 1975

283

mental to population survival. New foliage depletion in early instars leads
to mass starvation, delayed development, increased exposure to biotic controls
and general population collapse.

Literature Cited

Wickman, B. E., Mason, R. R., and Thompson, C. G. 1973. USDA Forest Serv., Pac.
NW For. & Ran. Exp. Sta., Gen. Tech. Rep. PNW-5. 18 p.

The Role of Defoliators in the Arthropod Community
of Red Maple Crowns

J. VOLNEY

Canadian Forestry Service, P.O. Box 4000, Fredericton, N.B., Canada

The arthropod community on red maple foliage is organized into 5 guilds
each of which remains comparatively simple throughout the growing season.
The simplicity of the community may be a reflection of the comparatively simple
structure of red maple crowns. Lepidopterous defoliators in 2 guilds dominate
this community both in terms of biomass and their impact on the community.
Leaf rollers, principally Cenopis acerivorana (MacK.), provide shelter for a
variety of organisms in the leaf roll and thereby tend to increase the number
of species in the community. I tame pustularia (Gn.), a solitary defoliator, which
occurred in high numbers in one plot, initially reduced the species diversity
of the community. However, the reaction of the host trees to heavy defoliation
resulted in an arthropod community with a higher index of diversity towards
the end of the season. Population levels of those defoliators which dominate
the community also have a considerable influence on the composition of this
community. Trophic bonds between the defoliator guilds are practically non-
existent in the populations studied. Similarly, trophic bonds with the arthropod
communities on other host species in the same stands seem tenuous. The
implications of these results on the process of stand succession in central
New Brunswick is speculated upon.

Current Research with Telenomus alsophilae Viereck, an Egg
Parasite of the Fall Cankerworm, Alsophila pometaria
(Harris) (Lepidoptera: Geometridae)

A. T. Drooz

Forestry Sciences Laboratory, P.O. Box 12254, Research Triangle Park, N.C. 27709

By way of background information, the rationale for our work on Telenomus
alsophilae Viereck takes us back over 20 years, when an outbreak of the elm

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spanworm, Ennomos subsignarius (Hiibner) arose in northern Georgia. This
outbreak lasted 10 years, covering 1.6 million gross acres in the southern
Appalachian Mountains at its peak in 1960. Spanworm populations declined
in 1963, and egg masses could not be located after the hatch of 1964. At
that time an egg parasite determined to be T. alsophilae was found in an
average of 85% of the spanworm eggs; few of these loopers became final
instars, and pupae could not be found. Hardly any information was available
about T. alsophilae at that time. It obviously was time to examine the details
of the biology of so important a parasite. However, the spanworm outbreak
was over and little could be done to initiate work on T. alsophilae. What could
be done would have to be carried out with a more common host, the fall
cankerworm and laboratory-reared spanworm eggs. The inevitable problem
arose when at that time it was impossible to rear the fall cankerworm and
the cankerworm parasites would not attack the spanworm eggs. These prob-
lems, and the observation that cankerworm eggs are attacked in the late fall
through winter in the mountains of North Carolina, while the spanworm eggs
are only attacked in April/ May, prior to hatch, needed to be resolved. There-
fore, a research program was developed whose objectives were:

1. Rear the fall cankerworm for use as host material.

2. Study the relationships between T. alsophilae attacks on the cankerworm

and spanworm in the field.

3. Rear T. alsophilae upon demand.

Work was begun in the winter of 1970/71 to rear the cankerworm and learn
about parasite emergence, sex ratios and longevity. We succeeded in our efforts
to rear the cankerworm, but so far the results are not worth the effort. It
can be reared either on an artificial diet or on host foliage, but prolonged
emergence and the predominance of females, which also occur in natural popu-
lations, defeat the purpose. G. F. Fedde tested the eggs of numerous potential
hosts, and found a number of geometrid and noctuid hosts were satisfactory,
but he found that a relatively unimportant forest geometrid, Abbotana clema-
taria (J. E. Smith) is an excellent laboratory host for T. alsophilae. V. H.
Fedde has reared this geometrid with little difficulty on a diet she developed
for the spanworm. This breakthrough was much needed, and has permitted us
to achieve our goal of producing T. alsophilae and some other species of egg
parasites on demand, as well as investigate details of the biology of T. alsophilae
in the laboratory.

Where are we in our work with the field relationships between T. alsophilae ,
the cankerworm and spanworm? Through our field observations on time of
parasite attack on eggs of these hosts, arena studies in the laboratory, and
reexamination of specimens from the 2 geometrids, USDA specialists indicate

Vol. LXXXIII, December, 1975

285

that the Telenomus attacking the elm spanworm is a new species. Therefore,
we have another potentially useful insect to investigate if we can obtain a
starter culture and can rear it in the laboratory.

Along practical lines, being able to rear T. alsophilae has permitted us to
send quantities of parasitized eggs to A. E. Bustillo in Medellin, Colombia
where he has made successful sleeve-cage tests with T. alsophilae against Oxydia
trychiata (Guenee). This insect is one of several geometrid species killing the
introduced cypress, Cupressus lusitanica Miller.

Within the next few years a number of scientific papers should be published
by our group concerning details of the biology of T. alsophilae. We believe
that they will provide much useful information to the field of biological control
about a valuable species or two which were neglected in the past.

The Importance, Biology and Control of the Birch Casebearer,
Coleophora fuscedinella Zeller (Lepidoptera: Coleophoridae),
an Imported Pest in Insular Newfoundland

D. G. Bryant

Environment Canada, Newfoundland Forest Research Centre, Box 6028,

St. John’s, A1C 5X8

The birch casebearer, Coleophora fuscedinella Zeller, was introduced into
North America from Europe about 1920 (1) and has been found attacking
several broad-leaved tree species such as birch, Betula, and alder, Alnus,
causing severe browning of foliage, branch mortality and occasionally death
of host trees. It was first found in Newfoundland in 1953 and occurs on its
principal host white birch, B. papyrifera Marsh.

The Province is in the Boreal Forest region and white birch comprises about
12% of the standing tree volume. The tree is little used for either lumber or
pulp and has its greatest importance as an aesthetically valued component of
the landscape. Over 30% damage to leaves (defoliation) brings public enquiries
(2). At over 90% defoliation, branch and twig mortality becomes apparent.
There is a high variation in total defoliation among trees in a stand and we
have not been able to identify the cause of the variation. Within trees, defolia-
tion is concentrated in the peripheral 25 cm of crown and is least variable in
the 2 middle crown quarters.

The casebearer has one generation a year. Eggs are laid on the leaves in
July and larvae hatch 3 weeks later. These larvae mine leaves in August and
molt to the second instar and construct a case from the leaf epidermis in
September. The casebearers overwinter at a crevice on the bark and molt to
the third instar in the spring before feeding. The fourth instar larvae construct

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a new case in June, feed for a short period, then pupate on the tree or ground
vegetation (3).

Larvae are distributed throughout the tree but at any instar the majority
are situated in the outer 25 cm of the middle half of the tree crown (4).
The majority of leaves in a tree are in the 25 cm periphery and are the cause
of the casebearer population concentrating in this portion. There is a high
and sometimes significant (P ^ 0.05) variation in larval numbers between
trees. The cause of the variation is unknown and is not related to tree variables
of height, size, form, exposure, or dominance. Within trees, differences in lar-
val numbers are most pronounced (P ^ 0.05) among 25 cm shells and occur
occasionally among crown quarters.

Inter-tree variation was too high to obtain significant correlations of case-
bearer numbers and defoliation within stands. For data from all stands how-
ever, there was a significant correlation (0.48 < r < 0.74; P ^ 0.05) between
defoliation and eggs, overwintering cases and late-instar larvae. The precision
of defoliation prediction is low because of high inter-tree variation. Methods
are being prepared to forecast defoliation classes from casebearer abundance.

Survival of casebearers during a generation appears to be constant at all
population levels. About 40% of larvae survive winter (5) and about one-
fifth of these pupate. This constant mortality suggests that the causes are
intra-specific or relate to the insect-host interaction. High egg mortality of
90%, especially at high population levels, has been recorded (6) but the cause
is unknown. Parasitism of larvae and pupae was extremely low at less than
5%, and parasites, native to European populations, are being introduced (7).
Where defoliation is expected to affect aesthetic values, spraying at first green
in the spring is recommended (8). On large ornamental trees systemic insecti-
cides are suggested.

Literature Cited

1. Reeks, W. A. 1951. Canada Agric., Bi-mon. Prog. Rep., 7(4).

2. Bryant, D. G., and Raske, A. G. 1975. Canad. Entomol., 107: 217-223.

3. Cochran, S. G. 1974. The birch casebearer in Newfoundland. M.Sc. Thesis, McGill Univ.,

126 pp.

4. Raske, A. G., and Bryant, D. G. Canad. Entomol. (In Press)

5. Raske, A. G. 1975. Environ. Can., For. Serv., Bi-mon. Res. Notes, 31: 9-10.

6. . 1974. Environ. Can., For. Serv., Bi-mon. Res. Notes, 30: 1-2.

7. . 1974. Environ. Can., Newfoundland For. Res. Centre, Inform. Rep., N-X-108.

8. Clark, R. C., and Raske, A. G. 1974. Environ. Can., Newfoundland For. Res. Centre,

Nfld. For. Notes No. 7.

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287

The Bimodality of Gypsy Moth, Porthetria dispar (L.)

(Lepidoptera: Lymantriidae) Populations

Robert W. Campbell

Northeastern Forest Experiment Station, Forest Service, U. S. Dep. Agriculture,
6816 Market Street, Upper Darby, Pa. 19082

Populations of the gypsy moth ( Porthetria dispar (L.)) in North America
have 2 numerical phases. A population may remain in the innocuous phase
for many years, and the outbreak phase may continue within a general area
for as much as a decade. Changes from the innocuous phase to the outbreak
one and vice versa, however, usually take place within 2 or 3 years. Our
records, which span a sizeable sample of natural gypsy moth populations in
the northeastern United States between 1911 and 1975, show no tendency
among these populations toward the regular oscillations characteristic of gypsy
moth populations across much of Europe.

A procedure was developed to determine at what point during the generation
the numerical differences appear between a sparse population in an outbreak
area and one in an innocuous area. This study led to the following conclusions:

1. From 50 to 100 times as many eggs were produced by initially sparse
populations in the outbreak area (Glenville, N. Y., 1958-1964) as by equally
sparse populations in the innocuous area (Eastford, Conn., 1965-1968).

2. More than 90% of the difference between the 2 areas in mean number
of eggs produced was caused by differential mortality during instars I to III,
IV to VI, and the pupal stage, and by a lower proportion of females among
the adults in Eastford.

3. The apparent survival during instars I to III was much lower in East-
ford than in Glenville when egg density at the start of the generation was low.
Survival was about equal in the 2 areas, however, when egg density increased.

4. Survival during instars IV to VI was about the same in the 2 areas when
egg density was low, but it decreased rapidly in Eastford when egg density
increased.

5. Survival of pupae was much lower in Eastford across the entire range
of egg densities that was common to the 2 areas. Interestingly, the survival
of pupae was relatively invariant within each area.

6. Differences between the 2 areas in the proportion of females among
adults were minor when egg density was low, but increased as egg density
increased. Since the sex ratio of gypsy moth eggs is known to be constant at
50:50 (2), and since the sex ratio of instar IV larvae is known to have been
relatively invariant in the areas studied at about 65% females (1), it seems
safe to assume that the significant differences between the 2 areas in the pro-
portion of females among adults were determined during instars IV to VI, the

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pupal stage, or both. Thus, not only do substantially more insects in the
outbreak area survive instars IV to VI and the pupal stage, but more of the
survivors are females.

Literature Cited

Campbell, R. W. 1963. Can. Entomol., 95: 465-474.

. 1967. For. Sci., 13: 19-22.

BOOK REVIEW

Man against Tsetse: Struggle for Africa. John J. McKelvey, Jr. Cornell University
Press, Ithaca and London. 1973. 306 pp. $12.50.

Entomologists will be fascinated by the author’s analysis of the development of knowledge
on sleeping sickness and nagana disease of cattle, and how the large continent of Africa was
influenced by contributions of medical entomologists. But this book is not merely an
account of historical and scientific events and discoveries; it analyses in depth the
complicated interrelations that the author, who has spent many years in Africa, has
known from his own experience. The book is written by a scientist, who, as readers will
find, is also an extremely accomplished writer and master of the language. The book
thus provides many enjoyable hours. I am certain that my colleagues who read it
will find perusal of this volume to be both informative and pleasurable. It gives much
useful data to entomologists, epidemiologists, physicians and microbiologists. The author,
a Director for Agriculture of the Rockefeller Foundation, is a knowledgeable medical
entomologist (Ph.D., Cornell U.), a scholar, and an expert on Africa.

This book should be read by anyone who plans a trip to Africa — be it for scientific
reasons, business, or for pleasure. No special training in entomology is required and the
book will be enjoyed by entomologists as well as by laymen who want an authoritative,
up-to-date view of the field. The author has succeeded admirably in bringing together for
the first time the information on the tsetse fly, the trypanosomes, and the development of
African nations. It will therefore serve both as a thorough review of the history and
present status of the problem for college students, and as a guide to the literature for
serious researchers. The Chapter Notes (pp. 239-292) are very valuable for the latter. An
index of 13 pages completes the volume.

There was a definite need for a book on sleeping sickness and at last we have
one, written by an extremely competent authority. It ought to be in every public and
highschool library so as to become for young men and women interested in Africa’s
future and medical entomology what Paul DeKruif’s “Microbe Hunters” became for future
microbiologists — the stimulus to devote one’s life to a deserving cause. I found “Man
against Tsetse” as stimulating as DeKruif’s book, and at the same time more accurate
and up-to-date. Years of careful research have been spent by the author in searching and
checking all the data and facts.

The tsetse fly had an enormous impact on Africa’s development. Great strides have
been made in controlling the insects and in chemotherapy of the disease. Nevertheless,
there remains the dangerous potential for an epidemic, as demonstrated in recent, new out-
breaks of sleeping sickness. This is pointed out succinctly by McKelvey. The book will
remain invaluable for a long time to come to those engaged in biological, medical and
agricultural research.

Karl Maramorosch

Institute of Microbiology, Rutgers University

Vol. LXXXIII, December, 1975

289

INDEX TO SCIENTIFIC NAMES OF ANIMALS AND PLANTS
VOLUME LXXXIII

Generic names begin with capital letters. New genera, species, subspecies, and varieties are
printed in italics. The following are not indexed: Figs 1-8, pp. 4-5, “New or little-known
crane flies from Iran II (Diptera: Tipulidae)” by Charles P. Alexander; Table I, p. 37,
“Parasites reared from larvae of the European corn borer, Ostrinia nubilalis (Hbn.) in
Massachusetts 1971-73 (Lepidoptera: Pyralidae) by F. B. Peairs and J. H. Lilly; “Mites
(Acarina) associated with Popilius disjunctus (Illiger) (Coleoptera: Passalidae) in Eastern
United States” by Mercedes D. Delfinado and Edward W. Baker, pp. 49-59; “Revision of
the genus Endeodes LeConte with a tabular key to the species (Coleoptera: Melyridae)”
by Ian Moore and E. F. Legner, pp. 70-80; Tables 1-7 and Figs. 1-11, “Comparative be-
havior of wasps in the genus Lindenius (Hymenoptera: Sphecidae, Crabroninae) ” by

Richard C. Miller and Frank E. Kurczewski, pp. 82-120; Figs. 1-9 and Distributional Rec-
ords, pp. 124, 126-128, “New or little-known craneflies from Iran III (Diptera: Tipulidae)”
and Figs. 1-8, pp. 132-133, 136-138 (Iran IV), by Charles P. Alexander; pp. 142-156, “An
annotated list of New York Siphonaptera,” by Allen H. Benton and Danny L. Kelly;
pp. 176-179, “Notes on the life cycle and natural history of butterflies in El Salvador.
II B -Hamadryas amphinome L. (Nymphalidae-Hamadryadinae) ” by Alberto Muyshondt
and Alberto Muyshondt, Jr.; Tables 1-4, “Species and numbers of bloodsucking flies feeding
on hogs and other animals in southern New Jersey,” pp. 199-201, by Thomas J. Weiner
and Elton J. Hansens; “Speleognatinae collected from birds in North America (Acarina:
Ereynetidae),” by A. Fain and K. E. Hyland, pp. 203-208.

Abbotana clemataria, 284
Abies, 282

Achrysocharella silvia, 101
Actinote, 166, 190
Adelpha, 167
Aedes atlanticus, 249
canadensis, 249
Aenictus, 196
Agaricus bisporus, 256
Agathis, 101
Ageronia, 165, 187
arete, 166
fornax, 166, 187
Agromyza frontella, 243
Allophylus petiolatus, 16
Alnus, 285

Alsophila pometaria, 283
Anartia fatina, 167

jatrophae luteipicta, 167
Anetia thirsa, 167
Anthidium banningense, 280
manicatum, 280

Apanteles melanoscelus, 242, 246
Aphelonema simplex, 254
Aplomya caesar, 36

Argyrotaenia velutinana, 264
Aristolochia trilobata, 189
Asclepia syriaca, 260
Atta texana, 192
Autographa californica, 270

Battus, 189

laodamas, 166
philenor, 247
polydamas, 166, 247
Betula papyifera, 285
Biblis hyperia, 165
Boophilus decoloratus, 46
Brachymeria intermedia, 269
Buenoa confusa, 265
margaritacea, 265

Caligo memnon, 166
Callicore, 16

meridionalis, 15, 16
sorama, 16

Calliopis andreniformis, 280
Calliphora, 34

290

New York Entomological Society

Calocoris, 86

norwegicus, 86
Camponotus, 192

pennsylvanicus, 194
Cardiospermum, 13
halicacabum, IS
Carduus, 250, 251

Crossocerus, 118

leucostomoides, 118
maculiclypeus, 117
Cuprossus lusitanica, 285
Cyphoderris buckelli, 233
monstrosa, 233

acanthoides, 247, 250
nutans, 247, 250
Carpocapsa pomonella, 271
Cassida rubiginosa, 247
Castilla gummifera, 174
Catagramma pitheas, 10, 16
pygas, 15
titania, 10, 16

Catonephele numilia estite, 17
vyctimus, 165, 176
Cenopis acerivorana, 283
Centris pallida, 280
Chalcis, 279
Cheilotrichia, 123

(Empeda) gnoma n. sp., 121
Chirosomus, 105, 114
Chlosyne, 166, 190
Choristoneura fumiferana, 266, 269
Chrysopa, 278
Chrysops, 198, 246
atlanticus, 200
callidus, 202
celatus, 202
cincticornis, 200
fuliginosus, 200
geminatus, 246
macquarti, 202, 246
montamus, 200
niger, 200
nigribimbo, 202
vittatus, 202

Cicindela imperfecta, 226
Cirsium arvense, 247

Dalechampia ficifolia, 189
heteromorpha, 189
scandens, 164, 170, 181
stipulacea, 189
triphyla, 189
Daphnia, 265
Dasyhelea, 114
Dasyneura gleditschae, 259
mali, 244

Daucus carota, 105
Delphacodes de tecta, 254
Desantisca, 279
Diachlorus ferrugatus, 200
Diaethria astala, 10
candrena, 16
clymena, 16
eluina, 16
salvadorensis, 16
Dibrachys, 280

Dicranomyia demmaculata, 129
Dicranota (Dicranota) capillata, 134
(Dicranota) fuscipennis, 134
(Dicranota) ophidia n. sp., 129
Diglyphus isaea, 243
Dione juno huascama, 166, 187
Dircenna klugii, 167
Dorylus, 194
labiatus, 192
Dynamine, 165, 176

Coccinella nine-notata, 226
transversoguttata, 227
Cochliomyia hominivorax, 34
Coleomegilla maculata, 60, 272
Coleophora fuscedinella, 285
Collops, 228

bipunctatus, 226
Colobura dirce, 190
Conomyrma insana, 192
Crematogaster laevirescula, 192

Eciton, 192
hamatum, 192
Elaeophila, 136
Eleodes sulcipennis, 226
Elliptochthonius profundus n. gen., n. sp.
209

Empeda, 123
Encopognathus, 118
Ennomos subsignarius, 284
Entomophthora egressa, 254
sphaerosperma, 254

Vol. LXXXIII, December, 1975

291

Epiphile adrasta adrasta, 17
Eriborus terebrans, 36

Erioptera (Pseuderioptera) n. subgen., 123
(Pseuderioptera) schmidi n. sp., 121
(Psiloconopa) cancriformis n. sp., 121
(Psiloconopa) idiophalleus, 126
Eucelatoriopis dimmocki, 247
Euphilis rufotaesiatum, 118

Fannia canicularis, 258
Forcipomyia, 114
Formica, 192

canadensis, 192
subintegra, 192

Gambusia af finis, 267
Gehypochthonius, 215
Geocoris pallens pallens, 227
Geospiza, 281
Gonempeda flava, 123
Gonomyia, 2, 121

(Gonomyia) abbreviata, 9
(Gonomyia) basilobata n. sp., 2
(Gonomyia) chalaza, 8
(Gonomyia) ebburzensis n. sp., 2
(Gonomyia) oxybeles n. sp., 2
(Gonomyia) sibyna, 8
(Gonomyia) tenella, 7
(Idiocera) alexanderiana, 3
(Idiocera) curticurva n. sp., 2
(Idiocera) displosa, 6
(Idiocera) jucunda, 6
(Idiocera) laterospina n. sp., 2
(Idiocera) orthophallus n. sp., 2
(Idiocera) phallostena, 6
(Idiocera) punctata, 6
(Idiocera) schrenki, 6
(Idiocera) similior, 9
(Idiocera) spinistylata n. sp., 2
(Idiocera) spinulistyla n. sp., 6
Graphium marcellus, 247
Grapholitha molesta, 264
Gynaecia dirce (h), 166

Haematobia irritans, 198, 253
Hamadryas, 165, 170, 181

amphinome, 166, 176, 181

arete, 189

februa, 157, 170, 181
fornax, 166, 187
guatamalena, 166, 170, 181
Heliothis zea, 166, 249
Hippodamia convergens, 61, 226
quinquesignata, 226
sinuata disjuncta, 227
sinuata spuria, 226
Holcopasites, 280
Hoplitis anthocopoides, 281
Hybomitra, 246
daeckii, 202
lasiophthalma, 200
losiophthalmus, 246
sodalis, 246

Hymenitis oto oto, 167
Hypanartia lethe, 167
Hypera postica, 226, 243, 263
Hypocada virginiana nigricosta, 167
Hyposoter exiguae, 261, 263

Idiocera, 3
Ilisia, 126

Iridomyrmex pruinosum, 192
Itame pustularia, 283
Itoplectis conquisitor, 257

Jenkinshelea magnipennis, 114

Lasioglossum rohweri, 280
Laspeyresia pomonella, 264
Latrodectus, 279
Libocedrus decurrens, 229
Limnobia demmaculata, 129
Limonia (Dicranomyia) chorea, 131, 134
(Dicranomyia) demmaculata, 129
(Dicranomyia) decem-maculata, 129
(Dicranomyia) didyma, 132, 134
(Dicranomyia) flavocincta, 129
(Dicranomyia) lutea, 131
(Dicranomyia) mitis, 131, 134
(Dicranomyia) modesta, 130
(Dicranomyia) nigritorus n. sp., 129
(Dicranomyia) schmidiana n. sp., 129
(Dicranomyia) subdidyma n. sp., 129
(Dicranomyia) vibishana, 129

292

New York Entomological Society

(Dicranomyia) whitei, 129
(Elaeophila) submarmata, 136
Liminophila (Elaeophila) albofascia n. sp.,
129

Lindenius, 82-83, 115-119
montezuma, 82
(Lindenises) albilabris, 82
(Trachelosimus) armaticeps, 82
(Trachelosimus) buccadentis, 82
(Trachelosimus) columbianus errans, 82
(Trachelosimus) panzeri, 82
(Trachelosimus) pygmaeus, 82
Liometopum, 118
Lipsothrix iranica n. sp., 121
nervosa, 122
nigristigma, 122
vobilis, 122

Liriomyza trifolicarum, 243
Lixophaga, 36
Lycoriella mali, 255
Lygus pratensis, 85

Macrocentrus grandii, 36
Manataria maculata, 166, 190
Marpesia, 167

Mechanitis isthmia isthmia, 166, 190
Melampus bidentatus, 267
Meliolotus alba, 105
Mestra amymone, 165, 176
Meterioptera (subgen.), 123
Microctonus aethiops, 243
colesi, 263

Microtia elva, 166, 190

Molophilus (Molophilus) pallidipes n. sp.,

121

(Molophilus) stroblianus, 121
Monomorium minimum, 99
Morinedea armata, 220
Musca autumnalis, 253
domestica, 252
Myrmica, 192
rubra, 192
Myscelis orsis, 15

Nabis americoferus, 227
Narope cyllastros testaceae, 167
Neduba, 233

macneilli,, 229
sierranus, 229

Neivamyrmex, 192
harrisi, 192
nigrescens, 38
Notonecta undulata, 248

Oncopeltus fasciatus, 194, 260
Orgyia pseudotsugata, 282
Ostrinia nubilalis, 36, 249, 256, 265
Oxydia trychiata, 285

Pachycondyla, 196
harpax, 192
Panonychus ulmi, 262
Papilio anchisiades idaeus, 166
glaucus, 247
multicaudatus, 247
palamedes, 247
polyxenes, 247
troilus, 247
zelicaon, 247
Paracymus, 273
Parectecephala eucera, 94
sanguinolenta, 94
Parhypochthonius, 215
Parides, 189
Paullina, 13
pinnata, 15

Paulogramma pyracmon, 16
Peridromia, 165
Perilitus coccinellae, 60
Phaenicia pallescens, 33
sericata, 33
Phormia regina, 34
Phrosinella fulvicornis, 96
Phycoides, 166, 190
Pinus jeffreyi, 209
lambertiana, 229
ponderosa, 209
Platynota flavedama, 260
idaeusalis, 268

Pogonomyrmex barbatus, 192
Pollenia rudis, 34
Polyergus breviceps, 192
Popilius disjunctus, 49
Porthetria dispar, 242, 258, 271, 287
Precis genoveva, 167

Vol. LXXXIII, December, 1975

293

Procystiphora n. sp., 114
Prokelisia marginata, 254
Protophormia tenaenovae, 34
Protoxaea gloriosa, 280
Pseudonica fla villa canthara, 17
Pseudotsuga menziesii var. glauca, 282
Psorophora ferox, 249
Pyrrhogyra hypsenor, 17, 167

Rhagoletis pomonella, 264
Rhinocyllus conicus, 250, 251
Rhytidoponera, 192

Serjania, 13, 15, 16
Sibaria, 217

andicola, 217
armata, 217
englemani, 217
Simulium venustum, 243
Solenopsis, 192
invicta, 192

Spartina alterniflora, 253
patens, 253
Spilochalcis, 17
Stethorus punctum, 262
Stomoxys calcitrans, 198, 235, 252
Sympiesis viridula, 36

Tabanus, 198, 246
atratus, 199
fulvulus, 202
lineola, 199, 246
melanocarus, 200
nigripes, 200
nigrovitatus, 199
pumilus, 200, 246
trimaculatus, 200
Tanytarsus, 114
Tapinoma, 118
Tasiocerodes (subgen.), 123
Teleneura (subgen.), 123
Telenomus alsophilae, 282, 283
Temenis laothoe liberia, 17
Tenebrio molitor, 274
Tetrastichus, 280
rhosaces, 247
Theritas lisus, 166
Thesalia theona, 166, 190
Tithorea harmonia salpadores, 167
Tracheliodes, 118
Tragia volubilis, 165
Trema micrantha, 16
Trichoplusia ni, 261, 262, 265, 270, 271, 273
Tumidagena ruinuta, 254

Xylocopa darwini, 281

Zagrammosoma, 279

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